EFACE
"'"' S ":"i.las of Human Neuroscience combines and provide regio nal and functi o nal co ntexts for
f'C'I)e~~ and beauty of Dr. Frank N etter's illu5tra
;:)a)icting both regio nal and syste mic neuro
'r '.\ ilh updated info rm ation and beautiful new
n:o-ts to reflect our growing understanding of
--e-g.ions and systems of the brain, the spinal
~.'j(j the periphery. This Atlas also includes new
~ns through the spinal cord and the brain
~ \'. e ll as coronal and horizontal sections cor
"::O.,., ....-ith ,'vIRls. We have chosen to let the iIIustra
:J'1J'\ ide the focal point for learning.. rather than
org. detailed written explanations th at cou ld
- ..,:e <: Tull textbook in itself. VVe have provided
..C :lgure legends to point out some of the
func tional aspects of each illustration, particu
as 'rese aspects relate to problems that a cJin i
-.0\ encounter in assessi ng a patient with
"""'0_,<,'"cal symptoms. The figure legends, com
.. 'th th e excell en t illu strati ons, provi de a
=~ =-'" ~n ders tanding of the basic components,
:..JnZ.cloon, and functional aspects o f the region or
Je.- vider consideration .
(J(de - [Q provide an optimal learning experi
::-"01'" jtu dents of neuroscience, we have orga
-5 4t1as in three parts: I. Overview oi the
..>.....: 5\ stem, II. Regional Neuroscience, an d III.
~JC 'euroscience. This organization affords
~ - th e benefit of looking at some of these
-"""neu ral stru ctures and systems in two or
__ cr:erenl contexts, or from two or three
-~i. poi nts of view, so m eti mes as part of an
~ ... so me times with a regional emphasis,
5lJr"""ti mes wi th a view toward understanding
_"'1C"io ning of a speci fic system spanning the
5. Thoughtful repetition from novel perspec
5 a usefu l too l in acq uiring a comfortable
... g 'nowledge of the nervous system that will
di'1ician s w ell in evaluating and treating
..3\\ith neurolo gical problems and th at will
oe neuroscience researchers and educators
-- a broader and more comprehensive under
,,",n:lh ~. oi the ner vous system.
.... ,;,ome information, su ch as th at on upper
'. er m otor neu ron s and th eir control,
d<:.o..ec :actual informatio n must be acquired and
"""-'i"::ood as a preliminary step tow ard under
COt"DrS cli nical aspects of m o tor disorders; w ith
- 4,- un derstanding. the clini cal aspects fall
Onto place. We introdu ce co re information
the information, as appropri ate, to encourage true
understanding rather than ro te memorization. In a
diSCiplin e as complex as the neurosciences, th e
acquisition of a true understanding of the major
regio ns and hierarchies of the nervou s system is
esse ntial for developing a working competence in
basic and clinical neuroscience.
This Atlas illustrates basi c human neuroscience,
no t clinical conditions and disorders, such as multi
pl e sclerosis. Alzheim er's disease, strokes, spinal
cord and brai n injuries, the agnosias, aphasias, and
apraxias, or o ther majo r conditio ns. These topics
are m ore appropriate for a future volume .
Part I. Overview of the Nervous System
We begin with an introductory overview of
neurons and their properties, gross features of th e
nervous syste m, and an understanding of the sup
porting tissues and systems such as the vasculature,
the menin ges, an d the cerebrospinal fluid b ecause
we believe that it is necessary to understand this
body o f information before delving into the details
of re gional and systemic neuroscience. Thus, Part I.
Overview o f the Nervous System first presents the
unique ch aracteristi cs and properti es of the cellular
foundations of the nervous system. including neu
rons and their stru ctural, neurotransmitter, and
electrical properties, and the supporting cells, the
glia. Specific exa mples of primar y sensory neurons,
low er m oto r neuron s, and auton omic pregan
glionic and postganglio nic neurons provide
preparatory examples of th e inputs and outputs of
the nervous system, esse ntial for understandi ng
both regional and system ic neuroscience.
The next po rtion of the Overvi ew introduces the
m ajo r structural features of the nervous system,
including the principal su bdivisi ons and th eir func
tional roles, in a manner that is consistent w ith
introduc to ry labo ratory exam ination o f the human
nervous system. Thus, students are in troduced to
lateral, midsagittal, basal, and some cross-sectio nal
views of the forebrain, with the intention o f obtain
M
ing a "big picture view. This is followed by a similar
in troduction to the brain stem and cereb ellum
and the spinal cord. It is during this introductory
overview that students sho uld become familiar
with the telencephalon, th e di encephalon, th e
brain stem, the spinal cord, and the peripheral
VII
~er\ DUS s y ~t em and acqu ire an understanding of
he major associations and relationships among
r ~e com pon ents .
Subsequent sections of the O verview introduce
~ e men inges, the ventricles and cerebrospinal
'u id, and the vasculatu re, all components of the
"tensiv supportive and protective infrastructure
.i the nen eu s system. Because the vasculature of
ne bra in an d spinal cord is a very important pa rt of
ieurological ass ess ment and understandin& \'\'e
horou ghly in troduce it in the Overview and follow
his \",ith additional reference and explanation of
lOre specific de tails in Part /1. Regio nal
~ euro sc i e n c e and Part III. Systemic N euroscience.
inaih. the Overview presents developmenta l neu
osdence, showing how the mature nervous sys
~r-l devel ops an d how so me of the important
~mction al relatio nships reflect associatio ns that
.egan during neural development.
'art II. Regional Neuroscience
Ian II. Regiona l Neuroscience begins in the periph
'fal nen o us system (PNS ) and moves rostraUy. This
pproach pelm its students to gain a thorough
Inderstan ding of the inputs and outp uts of th e cerr
'al ner\'o us system (eNS), th e brain and spin al
Drd. before attempting to understand more ros tral
E'gions that are involved in processing of inputs to,
nd regulation of outputs of, the eNS.
The in troduction to the PN S establishes a
lorough understan din g of primary sensory inputs
nd motor and au to nomic o utputs of th e CNS.
'onceptua lly, some of the most important
ustrations in the Arras are th ose tha t outlin e
secondary senso ry channels-the reflex, cerebel
t, and lemniscal channels thai further process
imar. sensory input to the CNS, (2) mo tor hierar
~ es, focu sed first on th e lower motor neurons
M "" s that supply sk eletal mu scl e fi bers, regula
>fl 0 1 these LMNs by bra in stem and cortica l
)per motor neuro ns (UMNs), and th e co ordin a
110 and modulation of UMN regulati on by the
~ebellu m and the basa l gangli a, and (3) pregan
onie autonomic connections to peripheral tar
ts through autonom ic ganglia, and the central
O.IllS irom th e brain stem, the hypothalamus, and
;> limbic str uc tures that regulate and coordinate
mautonomi c and neuroendocrine o utflow. A
information to th e brain stem (tri geminal, auditory,
Kough un derstand ing of these hi erarchies-as
vestibul ar, taste, visual), as w ell as somato sen sory
~nted in the in troduction to the PNS-is the
nuclei and chann els, (2) LMNs for the cranial
InOation ior the rest of the Atlas.
nudei, and descending UMN chan nels that control
The section on the PN S next offers a systematic
collection of illustrations of roots, plexuses, and
peripheral somatic nerves, w hich prov ides a
smooth transition between th e central and the
periphe ral aspects of neuroscience. The illustra
tions of the major peripheral nerves are con
ducive to acquiring a better un derstanding of
neuropathies, w hich are so frequently enco untered
in medical practice. 'vVe have tried to bring
together the basics of periphera l nerve anatomy,
fu nction, and cl inical deficits in as simple and suc
cinct a fash ion as possi ble in the figure legen ds for
the periph eral nerves.
The section on the PNS next outlin es the o rgani
Zation and the major d istribution networks of the
auto nomic nefVOU5 system . These illustrations are
important for many reasons, including understand
ing auto nomic disorde rs and the effects o f a ho st of
pharmacological agen ts that infiu ence th e auto
nomic nervou s system and its associated adrener
gic and cholinergi C receptors. In addition, w e are
becoming increaSingly aware that maj or life stres
sors and life style factOfs play an importan t role in
the onset,. progressio n, and o utcome of atheroscle
rosis and cardiovascular disease (including cere
brovascular disease), cancer, diabetes, chronic
pulm onary disease, and even neurodegenerative
diseases and that these factors exert their phys io
logical effect in large part th rough neural o utflow
via the autonomic nervous system and v ia hypo ClL
thalamo-pituitary-neuroendocrine outflow and that or
they have an impact on cytokines, hormones,
infl amm atory mediators, and a va riety of immune
responses. The updates to our und erstanding of
peripheral autonomic mechan isms include th e
depiction of autonomic interactio ns with cells of
-
the immune system and w ith metabo lic cells such
as hepa tocytes and fat cells. ..,jj
The brain stem is usua lly the " terra incogn ita" o f
neuroscience cou rses and can be an exercise in
memorization and m inutiae if not pro perly o rga
nized. This Atlas provides the o rgani zatio n for
rather st raightforward understanding of the brain
stem, despite its com plexities and intricacies. The
brain stem has six major components: (1) sensory
nudei and their pathways- includin g secondary
senso ry nuclei and pathw ays for incoming sensory
 ->,<... _ ,.., """""'e brain stem and the spinal cord,
~-~ - ,.., ~ tic preganglionic neurons, and
-: .a:ulO:l{l'nic pathways and associated
_. "e-~<Jia i e the o utflo",,' o f both the sympa
~:::..,.... ~ paras\ mpalhelic components of the
~ OU 5 sy stem, (4 ) cranial nerve nuclei
..i..()Cl~ ~ \ hieh clirectl y ove rlap with, and
~Jo'''_'d U1. the i irs! three components, (5) th e
=::..:J~.meI its afferent and efferent conn ee
0' the reticular formatio n. Th e crani al
.awJ "netf nuclei are vital compon ents fo r
:':::::=oowi,.,:;: both the brain stem and th e localiza
~ ....,."" ~ in the context of th e neuro logica l his
~na lionJ and they are the appropriate
""'--uneerstanding the first three compo
brain stem, noted above. The Netter
- .a-e lucid, thorough, and readily under
uundations fo r this process.
'Oft to the Netter illustrations, we have
_~ ::-:.ect io ns thro ugh the spinal cord and
!'-er:l. These cross-sections will allow
-va: or" Human Neuroscience to be used
.=..or. and bas ic sc iences guide for stu
3ting th e need for a separate, d eta il ed
-_c:;:.:,....",ic at las. In add ition, an excellent set of
_ _. ..........-izon t.::d and coronal sections, carre
... ~I im ages, is provided. These cross
ct'lfonals, and horizontals emphasize the
Impo rtant struc tures and avoid the use
--":",", , '-ibels and minutiae that get in the way
--.oets ta nding and obscure the " big pic
aDded to this section o n the brain stem
~~rati on s o f Ihe retic ul ar formation, a
~o n 0; the eNS that cuts across all
;:d b invo lved in w idely varied neural
a. cord, brain stem, and fo rebrain COI11
=e:lI;' 0.: the Regio nal Neuroscience section
Ocali zi ng and regional information impor
~ersta nd in g the co nseq uences of lesions
c ( tes. In th e forebrain section, we emp ha
......~'11 U S and the hypotha lamus in the
...." e diencephalon; we include the sub
_""-'" \ JJ th e basal ga nglia and the epithala
..... -'"Ie h\ pothalamus and the limbic system .
_ - -.... > telencephalon has fou r major func
ov-ponenls-cerebral cortex, basal ganglia,
i"Of"din sys tem, and o lfactory system- we
-=c,..~",r . emp hasi ze th e ce reb ral co rtex anatomy
.....-eGio ns in th e Regio nal N eurosc ience sec
- -<It ae~ai l s of basa l ga nglia anatomy, limbic
forebra in struc tures and their connec tions, and the
olfac to ry system are provided Part III, Systemic
Neuroscience. We also have updated the forebrain
section w ith an over view of the chemicaUy specific
systems of the brain, specifica lly the n oradrenergic,
serotonergic, dopaminergic, and c holinergic sys
tem s, which provide extensive inn ervation to \vide
sp read eNS regions and w hic h are so frequently
targeted with neurol ogical and psych iatric drugs.
Part III. Systemic Neuroscien ce
Part III. Systemic Neuroscience provides a more
d etailed understanding of the p eripheral and cen
tral mechanisms of the sensory system s, the motor
system s, the au to nomic-hypo thalam ic-limbic sys
tem s, and some higher func tio ns. The founda tions
for understanding the secondary senso ry and
higher--order processing of sensory p athways begin
w ith Ihe illustrations in the intro du ction to th e PNS
in Part 11. The systemic sensory components
includ e the somatosensory, trigeminal se nsory,
taste, audito ry, vestibular, and visual sys tem s. To
rellect current understandin g, seve ral modifi cation s
to Dr. Netter's artwork have been m ade, as well as
new art crea ted by John Craig, MD and Ji m
Perkins, MS, MFA, particu larly in the secti ons on
the so m atosensory system and the visual sys tem .
A llhough we focus extensively o n the lem n isca l
pa thways for sensory processing and conscious
interpretation of inco ming sensory informa tion,
appropriate diagrams and connectio ns for reflex
and cereb ellar pathw ays are included w hen appro~
priate ((or example, w ith the vestibular sys tem ).
Th e systemic motor components include LMNs,
UMN s, the ce rebellum, and th e basal ga ngli a-of
co urse, with the acknowledgment that th e ce rebe l
lum and th e basal ganglia are involved in coo rdi
natin g and m od ulating ce ntral circuits more
wides pread than just motor system s. For th e
UMN pathways, each sys tem is illustrated, with
emphasis on the functio nal role. The cerebellar
connectio ns included in this section emphaSize the
relationshi ps be tween specific zones of the cere
bellum (vermis, paravermis, la teral hemispheres)
w ith appropriate d eep nuclei (fastigial, globose
and em boliform, d entate) and the UMN systems
with w hich they connect (reticulospinal and
vestibul osp inal, rubrospin al, co rticospinal ).
Similarly, th e basa l ga nglia co nnectio ns show inter
connections w ith the cortex through the thalamu s,
including modu lation of the cells of origin of th e
IX
corticospinal system and other cortical ou tflow.
However, other basal ganglia loops are also
included: even thou gh basal ganglia disorders have
become known for their motor involveme nt, and
are often ca lled "movement disorders," w e now
know th at many other loops exist, linking other
regions of the thalamu s and the co rtex with the
basal ganglia, for cognitive as w eI! as emotiona l
behavioral influences.
The autonomic-hypothalamic-limbic sectio n
inclu des deta iled updates and understanding of this
important set of interconnected circuitry through
the hypothalamus an d limbic system that ultimately
regulates many behaviors, neuroendocrine outflow,
and viscera! activities of the autonomic nervous
system. We begin with the understanding that
many of the connections of th e hypothal amus an d
limbic sys tem focus on auto nomic and neu roen
docrine outflow. In the last decade, the ro le of the
hypothalamu s as a focal point for converging influ
en ces from forebrain sources, brain stem so urces,
x
and circulating mediators has become better eluci
da ted. This is renected in several new illustrations.
The secti on s on limbic forebrain structures (hip
pocampal fo rmatio n, amygdaloid nuclei, septum,
cingulate cortex) have been redone and exp and ed,
reflectin g both the newly discovered complexities
of the circui try and the focus o n converging influ
en ces, particularly through the hypothalamus and
au to nomic o utflow of th e e N S.
It is our hope that the three parts of this Arlas
Overview of the Nervous Sys tem, Regional
Neuroscien ce, and Systemic Neuroscience-will
provide students wi th a comfortab le w orking
kn owledge of the basic com ponents, o rganization,
and functional aspects o f the nervous system and
that this fou ndation w ill serve them well for inter
pretin g neurological exam inations, understanding
most neurologicaJdiseases and di so rders, and
evalu atin g and treating patien ts with neurolo gical
problems.
CONTENTS
P ~ RT I: OVERVIEW OF THE NERVO US SYSTEM
Section A Neurons and Their Properties ........... . . ... .. ... ...... .... ... . .. . ...
4
Section B Brain . ..... ... . < ... . .. .... 21

Section C Brain Stem and Cerebellum ............... .. . . ....... , . , . . . . . . . . . . . .. . . .. 35

Section 0 Spinal Cord ........ . ........ ... .......... .... . ... .............. . . . .... 37

Section E M eninges ... ............ ... , . . . . . . .. . . .. .... . .... . . .. ..... 42

Section F and the Cerebrospi nal Fluid . ..... .. .

Ventricles ... .. ...... ... . .. .... .... 43

Sectio n G Vascul ature ............. ... ........... .. . ........... ...... . ... .. . . ... .. 46

Section H Develo pmenta l Neuro science (DVA) . . . . . . . . . . . . . . . .. . . . . .. . . . . . . . . . . . . . . . . .. 67

P~ RT II : REGIONAL NEUROSCIE NCE

Section A Peripheral Nervous System (PNS) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 86

Section B Spinal Cord .. .. .... . . . . . ...... . ..... ".. . . . . ... " , 141

Section C Brain Stem and Cerebellum ...... ........ .. ... . ........ ... ...... . ........... 150

Section 0 Diencephalon " , . , ' , . . . . . , . . . . . . . . , . . . . . . . . , ' , ,""", . "" I~

Section E Telencephalon ... , .. ..... ..... ....... ...... 179

P~ R T III : SYSTEMI C NEUROSCIENCE

Section A Sensory Syste ms ... , , . , , , , , ..... . ' . . , , ' , , , , . .. ' , , , . ... . . , , , . .. , , , . ..... 214

Sectio n 8 Motor SysLems .. .... . .. ....... . . . ....... . ....... . ..... .... . .. ............. 240

Section C Autono mic1 H ypothalamic, and Limbic Systems ... . , ........... . ,. , " " ' " " , .264

Index ... , , , , , , ... , , , . , , , , , , ...... . .. ' , , , , ' , , , , , .. , . , 299

xv
 = 1\\ OF THE NERVOUS SYSTEM

EL 0 ," 5 A D THEIR PROPERTIES

en.. E
- 'euronal Structure
..,.. -\ pes of Synap ses

:-~.- U, ' eurona l Cell Types

~ Glial Cell Types

l.i: \\\ elination of e NS and PNS Axo ns

= ........ Ua:. Development of Myelination and Axon Enshea thment
""'_ L :-: H igh Magnification Vi e"v of a Ce ntral Myeli n Shea th

OOTRA N5M I5S ION

""':oft '-3: Svn ap ti c Morphology
L9: Chemical NeuTOtransmissio n: Amino Ac id Synap ses

Che mic al Neurotransmissio n: Ca techolamine Synap ses

Chem ical Neurotransmissio n: Seroto nin Synap ses

Chemical Neurotransmissio n: Peptidergic Synapses

Chemical Neurotransmissio n: Cho linergic Synapses

~ ICAl PROPERTIES

"='_ 1.10: ~e uronal Restin g Potential

E:.".., U 1: G raded Potentials in Ne urons

F;o"..,_ 112: -\ct;on Potentials

_ _ 1-13: Propaga tion of the Action Potential

c.~_ 1.1 ~: Con ductio n Velocity
'-."_ 1.15: Presynaptic and Postsynaptic Inhibition
fu..,,_ 1.1 6: Spa lial and Te mporal Summation

B R~IN
.---- 1.17: Interi or View of the Base of th e Adu lt Skull
s".." 1.18 : Foramina in the Base of th e Adult Skull
~= 1.19: Surface Anatomy of the Forebrain: Latera l View

e:.=
&:= 1.20:

i'
1.21 :
1.22:
l atera l View of the Forebrain: Fun ctional Regions
l ate ral View of the Forebrain: Brodmann A reas
A natomy of the Medial (Midsagi ttal) Surface o f the
Brain
""'= 1.23: Medial Surface of the Brain: Lobes and Functional

Areas

re 1.24: Medial Surface of the Brain: Brodmann Are(ls

__ "'e 1.25: Anatomy of the Basal Surface of the Brain: Wi th th e Brain Stem and

Cerebellum Removed
:- :._0-e 1.2 6: Basa l Surface of the Brain: Fun c ti ona l Areas and Brodm ann A reas
:- . '-:'E' 1.2 7: Hor izo ntal Brain Sections Showing th e Basal Ganglia
:- _ eo 1.2 8: Major Limbi c Forebrain Structures
-:- ~ -e 1.29: Corpu s Ca llosum: Horizontal View
':- :. ~--e 1.30: Corpus Callosum: Schematic Vi ew of the Lateral Extent o f Major
Components
- ~ e 1.31: Hippocampal Formation and Forn ix
- ~ e 1.32: Thalamic Anato my
-!;U: 'e 1.33: Thalamic Nuclei

1
 SECTION C. BRAIN STEM AND CEREBELLUM
Figure 1.34: Brain Stem Surface Anatomy: Posterol ateral and Anterior Views
Figure 1.35: Cerebellar Anatomy: Ex ternal and Internal Features

SECTION D. SPINAL CORD

Figure 1.36: Spinal Column: Bony Anatomy

Figure 1.37: Spinal Cord: Gross Anatomy In Situ

Figure 1.3 8: Th e Spinal Cord, Its Men inges, and Spin al Roots

Figure 1. 39: Spinal Cord : Cross-sectional Ana tomy In Situ

Figure 1.40: Spinal Cord W hite and Gray Matter

SECTION E. MENINGES
Fi gure 1.41 : Schematic of the Meninges and Their Relationship to the Brain

SECTION F. VENTRICLES AND THE CEREBROSPINAL SPINAL FLUID

Figure 1.42: Ven tricular Anatomy

Fi gure 1.43: Anatomy of the Fourt h Ve ntricle

Figure 1.44: Circulation of (h e Cerebrospin al Fluid

SECTION G. VASCULATURE
Fi gure 1.45: M enin geal Arteri es: Relationship to Skull and Dura

Figure 1.46: Arterial Supply to the Brain and the Meninges

Figure 1.47: Arte rial Distribution to the Brain: Basal View

Figure 1.48: Arteri al D istri bu ti on to th e Brain: Frontal View and Coronal Section
Figure 1.49: Schema tic of Arteries to the Brain
Figure 1.50: Schematic of the Circle of \'Villis
Fi gure 1.51: Arterial Distributi on to th e Brain: La te ral and Medial Views
Figure 1.52: Color Illustration of Te rritories of the Cerebral Arteri es
Figure 1.53: Angiographic Anatomy of the Internal Carotid Circulation
Figure 1.54: Angiographic Anatomy of the Vertebrobasilar System
Figure 1,55: Vertebrobasilar Arterial System
Figure 1.56: Vascular Supply to Ihe Hypothalamus and the Pitui ta ry Gland
Figure 1.57: Meninges and Superficial Cerebral Veins
Figure 1.58: Veins: Superficial Cerebral, M eningeal, Diploic, and Emissary
Figure 1.59: Venou s Sinuses
Figure 1.60: Deep Venous Drainage of the Brain
Figure 1.61: Deep Venous Drainage of the Brain: Relationship to Ven tricles
Figure 1.62: Venous Drainage of the Brain Stem and the Cerebellum
Figure 1.63: Arterial Blood Supply to the Spinal Cord: Longitudinal V iew
Figure 1.64: Anterior and Posterior Spinal Arteries and Th eir Distribution
Figure 1.65: Arterial Supply to the Spinal Cord: CrossSectional View
Figure 1.66: Venous Drainage of the Spinal Cord

SECTION H. DEVELOPMENTAL NEUROSCIENCE

Figure 1.67: Formation of the Neural Plate, Neu ral Tube, and Neural Crest
Figure 1.68: Neurulation
Figure 1.69: Neural Tube Development and Neural Crest Fo rmati on
Figure 1. 70: Neura l Proliferation and Differentiation: Walls of the Neural Tube

2
.-...we 1.71 : I~ e ura l Tube and Neural Crest Deri vatives
- __:re 1.72: Earl y Brain Development: The 28-Day-Old Embryo
1'e I. 73: Early Brain Development: The 36-Day-Old Embryo
1. 74 : Ea rly Brain Development: The 49-Day-Old and the 3-Mo nth-Old Embryo s
e 1. 73: Forebrain Develo pment: 7 Weeks Throu gh 3 M o nths
'" 1. 76: The 6-Mo nth and the 9-Month Central Nervous System
""=
r:
1. 77: Development of the Ventricles
1. 78: Com parison of SIll-Week and Ad ult Central Nervous System Regio ns
~e 1.79 : Develo pment of M o tor and Prega nglio nic Autonom ic Nuclei in the Brain Stem and
the Spinal Co rd
"""",, 1. 80: Alar and Basal Plate Derivatives in th e Brain Stem
"" 01' 1. 81: Developm ent of the Fourth Ventricl e
""'=:re I. 8 2: Develop ment of Peripheral Axons

3
figure 1. 71: Neural Tube and Neural Crest Derivatives
Figure 1. 72: Early Brain Development: The 28DayOld Embryo
Figu re 1. 73: Early Brain Development: The 36-Day-Old Embryo
Figure 1. 74: Ea rly Brain Development: The 49-Day-O ld and the 3-M onthD ld Embryos
Figure 1. 75: Forebrain Development: 7 Weeks Throu gh 3 M onths
Figure 1. 76: The 6Monlh and the 9-Month Central Nervous System
Figure 1. 77: Development of the Ventricles
Fi gu re 1. 78:Comparison of 51f:z-Week and Ad ul t Cen tral Ner vous System Regions
fi gure 1. 79: Development of Motor and Preganglionic Autonom ic Nuclei in th e Brain Stem and
th e Spin al Cord
Figure 1. 80: Alar and Basal Plate Derivatives in the Brain Stem
Figure 1. 81: Development of the Fo urth Ventricle
Fi gure 1. 82: Development of Peripheral Axons

3
OVERVIEW OF THE NERVOUS SYSTEM
Neuronal Structure
?t'P7;~l\---'~=~::: C:lial
1';
C I@N
FIGURE 1.1: NEURONAL STRUCTURE _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
Neuronal structure reflects the func tional character
istics of each neuron. Incoming information arrives
at a neuron mainly through axonal terminations
on the eel! body and dendrites. These synapses are
iso lated and protected by astrocytic processes. The
dendrites usually provide the grea test surface area
of the neuron. Some protrusions from dendritic
branches (dendritic spines) are sites of speCific
axodendritic synapses. Each neuronal type has a
characteristic dendritic bran ching pattern (dendritic
tree, or arborization s). The neuronal cel! body
varies from a few micrometers to more than
100 )..lm in diameter. The neuro nal cy toplasm con
tains extensive rough endoplasmic reticulum
(rough ER), which reflec ts the massive amount of
protein synthesis necessary to m aintain the neuron
and its processes. The Golgi apparatus is involved
in packaging potential signal molecules for trans
port and release. Large numbers o f mitochondria
are needed to meet the huge energy demands of
neurons, particula rly to maintain ion pumps and
membrane potentials. Each neuron has a si ngle (or
occasionally no) axon . Th e cel! body tapers to the
4
Neurons and Their Properties- Structure
I Jt;:;
~Oendril'C ~pines
, ~ /' (gemmules)
reti cul um I ~ub~tancc)
Axrln
- - Golw body
body (suma)
Al<m
(astrocyte) pro cess
:odendritic syn;)ps.t'
~~p
:" : ,'0';1
axo n at the axon hillock, and the ini tial segm ent of
L
the axon containing the Na channels is the first site
where axon potentials are initiated. The axon
ex tends for a variable (up to 1 m or more) distance
from the cel! body. Axons larger than 1 to 2 ).1m in
diameter are insulated by myelin sheaths provided
by oli godendroglia in th e eNS or by Schwann cells
in th e PNS. A n axon may branch into more th an
500,000 axon terminals and may terminate in a
highly locali zed and circumscribed zone (e. g.,
somatosensory projections for fine discriminative
touch), o r it m ay distribute to many disparate brain
regions (e.g., noradrenergic axonal projection s of
the locus coeruleus). Axons o f macroneurons
(Golgi type! neurons) term inate at a disfa nce from
the cel! body and dendritic tree. Axons o f micro
neurons (Go lgi type !l neuro ns, local ci rcuit
neurons, or ;nterneurons) terminate loca lly, close
to the cell body and dendritic tree. Because each
neuron type has its own speci alization, there is
no "typical" neuro n, although pyramidal cells or
lower motor neurons often are used to portray the
"typica l" neuron.
 ~ an d Th eir Properties Structure OVERVIEW OF THE NERVOUS SYSTEM
B. Dendritic spine synapse
~ <" !

A t

"
u ~bt;~ ~
.(
"
~ I ~
'Ax~C":'~) 8

.--"""'<~~~~"

"";"" " '\

~. f

~ plus E. Combined a)(oaxonic and F. Varicosities ("bouto ns en pOIssant")

-"'~"..,. axode ndritic synapse t

~~
:'>--... ,~ ~.
'
"

, go' ;
.
c'

::>':.~
0,
"

"
/ .
I,
, I. Serial syna pse

- t itie synapse

... L
u Dendro
de ndri tic;

~J~ 'Y"'P"
K. Inner plexiform la ye r of retin a
i~ ' ,,~. r.')l.
~ .
~ ~'a~gli'o n , '" rj '

}~i~~cell " ,.g9'

' t'in%%j')r) \ I
j
,._., " ,'/o'"::<:'P ; ' \
::~,uceU ----,71~ ",;/~ .
M uller cell - - . .,f
!
f'-~
I
. -+......,,..<.....,~
{supprn-tingl j

!
(
M o SSy cell axon
Amauioe cell +_-+____VC""
processes \.

2: TYP ES O F SYNAPSES _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

--e site wh ere an arrivin g action pote n

"5 J. target neuron termin ate as axodendriti c or
::h e\citC'l li o n-secreti o n co upli ng invo lving axosomati c synapses. Specialized syn apses, such
rlis ge rs th e release of one o r more as reciprocal synap ses, or complex arrays of
"'li tters into th e synaptic cl eft (typi cally synaptic interac ti ons provide specifi c regulatory
"'"l-:; neu ro transmitter acts o n receptors on control over th e excitabili ty of th eir target neurons.
neuro nal mem brane, altering th e mem Dendrod endriti c syn ap ses aid in th e coo rdin ated
rial irom it s restin g state. These post firing of gro ups of related neuro ns (sLlch as phreni c
_ "en tia ls are ca ll ed "graded" potenti als. nucl eus neuron s causin g co ntracti on of th e
~ S ca rrying informatio n to w ard a diaphragm),
5
OVERVIEW OF THE NERVOUS SYSTEM Neurons a nd Their Properties- Structure

MulIJpol.,. (pyr~ mjclal) ceU ~ . _ Bipolar ctll ofO.Jnial n. VIII

01 ce.",h,,,l
~Unipola. 'ello(
~SOt'( g.lIlglia of
Associatio n..!' c;r ~",.1 n n, v, VII, I X Of X
commissural and
thollamlC el'ldlrlg~

,I inllied fibers
(J~f' ee ne....e endings
{unmytlina.<."tl fihe.'1

SpeQiI~ ltrll!l'ldin&

of u anl,ll nn u,,'poLa. sen!oOry cel of

HI, IV, V, Vl VII,
dossel spinill tOol ~ndiun
IX, X. XI Of XII ----.l
Multipolar 1:('11
of lowe. b ra in
moIor cenTers ,''')ioafed .,rferent

1\)I..tu~ipotdrYiKe .JJ
/ moro.- (autnn<)m!(;)

- ,/ cd! of ,pinal cord Rl!d : Motnr neuron

Blu e: Sensory neumn
Purpl e: Inlerneuron
"!.'tV", robe< Cr;ay: Glia l and
n... urilen,"l~t
ce ll s ;lnd my~lin

NOle : Ce!cbcll it' cell s no t

shoWrl here
",evl Ii"
";:"""' $.a tp.Ii.p. nl~
RP.n~aw inlempuron
,lin,""
leedbadl
Ii, tie mur.... )(SchwafV1 ce lls ~ Myell n shealh

M}'('r.n $l>eath~
MOtor

a.d'K mlJSCle
<w-oodJlcelk
Srri aK'<I
I
,~Free ne!'>'e e"di oss

~ En(~psu l atE'd end ln ~

~~f!%
','l.!.N
.......... Musck,: spindle

FIGURE 1.3: NEURONA L CEll TYPES _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

l ocal interneurons and projection neuro ns demon
strate characteristic size, dendritic arbo rizatio ns,
and axonal projections. In the eNS, glial cells
(astrocytes, microglia, oligodendroglia) provide
suppo rt, protection, and maintenance of neurons.
Schwa nn cell s and satellite cells provide these func
tions in the PNS. TIle primary sensory neurOns
(blue) provide sensory transduction of in co ming
6
energy or stimuli into electrical signals that are
conveyed into the eNS. The neuronal o utflow
from the e NS is motor (red) to skeletal muscle
fibers via neuromuscular junctions, o r autonomic
preganglionic (red) to autonomic ganglia, w hose
neurons inn ervate cardi ac mu scle, smooth mu scle,
secretory gland s, metaboli c cells, or cell s of the
immu ne system.
 The ir Properties Structure
.-
~. G LIAL CE LL TYPES _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
- - ,...- ~id e structural isolatio n of neurons
_ -'-.a:pses. and they provide ioni c (K' )
~=:a>a' trophic support. and suppo rt for
__-','-'" ~ ;;naling functions to neurons.
c.~ .....
:ro-~ provide myelination o faxons in
roglia are scavenger cells that partici
~ X\ tosis. infla mmatory responses,
- ~ JD: J0 \\ th fac to r secretio n, and some
OVERVIEW OF THE NERVOUS SYSTEM
J. Perki'ns
M S. MFA
C: DN
"....'"
immune reactivity in the e NS. Pe rivascular cells
participate in similar activiti es at sites near the
blo od vessels. Schwann ce ll s provide mye lination,
ensheathment, tro ph ic suppo rt, and actions
for grow th and repair for peripheral neurons.
Activated T lymphocytes can enter and traverse
the eNS for immune surveillance for a period of
approximately 24 hours.
7
OVERVIEW OF THE NERVO US S) STEM '\e urons and Their Pro pert:irs:-St ru ct ure

CENTRAL NERVOUS SYSTEM PERIPH ERA L .' ERVO U S SYSTEM

Sensory neuro n
cell body

Pia m,lter
Oligo
dendro cyte

SchWimn cells associate d w ith

myelin sheaths 01 myelinated axons

Bouto ns of associati o n neuro ns synapsi ng Postgan glionic ne uro n of

f--- - - - t-I with preganglionic aulonomie neuron o f sympathetic or parasympathetic
CwainSlem o r spinal cord s.::a n~li o n

Schwa nn cells assoct<lted w ith

myelin sheaths o f myelinated axons "
.'i;"' " " ,,; /.
<~~~;.-
'....~~-.....S~~i".",._I ; ._~~
aSS()~i~~:Onn~;~rons "<': " ~b7 -,;:,c ';';'4iiP
S~'napsin !\ with somatic
moto r neuron s of brai n
or spinal cord

M5.MfA Axon s terminatin g

~ ,.~ o n mo tor {,Ilel p lal es o f
striated (voluntary) m uscle

FIGURE 1.5: MYELI NATION OF eNS AND PNS AXO NS _ _ _ _ _ _ _ _ _ _ _ _ __

Cent ral myelinatio n of axons is provided by o li go
dendroglia. Each oligodendroglia I cell myelinates a
Sin gle segment of several se parate central axons. In
th e PNS, se nsory, motor, and preganglionic- auto
no mic axons are myelinated by Schwa nn cells. A
Schwa nn cell myelinates only a single segment of
one axon. Unmyelinated sensory and auto nomi c
postganglioni c axon s are ensh eathed by a
8
Schwann cell, which provides a singl e enwrapping
arm of cytoplasm around each of several
such axons. The space betw een adjacent myelin
segments (a node of Ranvier), the site w here
the axon membrane contains sodium chann els,
and allow s the reinitiati o n of action potential s in
th e co urse of propagation, a process called
saltatory conductio n.
 Neurons and Their Properties-Stru cture OVERVIEW OF THE NERVOUS SYSTEM
-

A. Unmyelinaled 'axons>'
of peripher"t neufon,s ::i
(sensory, somatic motor t
or viscp.rai motor,) ""~
beins surroutidec:/ by :"
cytoplasm 01a
neurilemmal (Schwarm) .
cell

~'.
B. Myelinated 011(00.01
peripheral neuron
(sensory, somatic.motor
0 1 vi5c.ernl motor)
of
being surrounded by
Ihetic
a wrapplng'oi cell
memb~eoia
neiJ(ileml)'lal (~wann)
",Jl

c. Mye linated .uon

"1 ;
of eNS neu rons being 1
SUfm~lIldcO by il -1
wrapping of cell 1:
membrane of an :~
oliKodendlOcyte.
Unmyelinated axans
r
)
of e NS neurons ale , Axon
leit unprotected.
'il O ligodendrocyte

FIGURE 1.6: DEVELOPME NT OF MYEliNATION AND AXON ENSHEATHMENT _ _ _ __ _

ng \r\yelination is a cooperative interaction between
the neuro n and its myelinating support cell.
Unmyelinated peri pheral axons are invested with a
5ingle la yer of Schwann cell cytoplasm. When a
peripheral axon of at least 1 ~m in diameter trig
gers myelination, a Schwan n ce ll wraps many
layers o f ti ghtl y packed cell membrane around a
single segment of that axon. The oligodendroglia I
cell ex tends several arms of cytoplasm, whi ch
then wrap multiple layers of tightly packed
membrane around a single segment of each of
several axons.

9
OVERVIEW OF THE NERVOUS SYSTEM Neurons and Their Properties-Structure

Cell body oi an

"
(neurilemmal <:ells
plJ)' similar role
10 periphera l
nervous system )

Cell m embrane of Mitochondrion

myelina ted axon '\ in cytoplasm of
neuronal .. xot~

..;/#d.*';....y"";,.1':'?#.~.

layers o f cell membrane of o ligodendro cyte

Minute ma ~se s of cytoplas m w ra pped JfOUnd ,)XOfl of a myelina ted neuron of
trapped between fu;ed central nervous system (the lipid of lipoprotein
laye rs of cell membrane of coostituting fused ceU membrane is mye lin, w hich
oligodendrocyte _ __ gives mye linated axon a w hi te. glistening appearance)

~~~
119N : ........ : :

FIGURE 1.7: HIGH MAGNIFICATION VIEW OF A CENTRAL MYELIN SHEATH

_ __ _ _ _ _
Fused layers of oli godendroglial cell membrane
w rap aro und a segment of a central axon, prevent
ing ioni c flow across the cell membrane for the
entire myelinated segment. The node between
2 adjacent segments is bare axo n membrane,
which contain s Na'" chann els. These nodes are sites
where acti on potentials are reinitiated in th e con
duction of propagated action potentials. Illustration
after Bunge, Bunge, and Riis.

10
 OVERVIEW OF THE NERVOUS SYSTEM
-
IctUrl: - Their Properties-Neurotransmission

Sc hematic vf ~yn il p l ic rndi ngt

Dend rite'S

Num ero us bo ul o ns (Srnaplic km)b sl

01 presynaptic ne ur () n ~ l~rm in ,'lin g
nn 01 mOtOf nellion arid i'~ riendri te'S

fnlil rgl!d ~rdion vf bo ulo n

Axok!mma ---+--i
G lial p'~"", 1
, '

sYlklpticckh

P,eiynaptic nlembrane
( den~ly ""ii,,,,," j __ ....

Post$y n ~p ( ic membrane ;.
( clC<l~cly st~ inin g l

.. . . f/f"-::,,
. ' F"';;:;'-';:>
. .C":';
. ~~"
---..:--....
.. -
- ' --~
-. - ,.,
;~' .... "'. , . ' .. . . ~~
.

=ICU RE 1.8: SYNAPTIC MORPHOLOGY _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ __

es .. - ,apses are sites where neuro ns co mmuni ca te
-h each o ther and w ith effector or target cells.
- "e upper figure show s a typical neuron that
""E'C eives many synaptic contacts on its cell body
Mld associated dendrites. Incoming axons lose
-reir myelin sheaths, exhibit extensive branching.
.a:1d terminate as synapti c bouto ns (terminals) o n
i"le targe t (in this example, motor) neuron. The
lower fi gure show s an enl arged axosomati c
:errn in al. Chemi cal neuro transmitters are packaged
. '1 synap tic vesicles. W hen an actio n po tential
'nvades the term inal regio n, depolarizatio n triggers
Ca 2 infi ux, and many synap tic ves icle s fuse with
th e presynap tic membrane, releasin g packets of
neurotransmitte r into the synaptic deft. Th e neuro
transmitter binds to recepto rs o n the postsynaptic
membrane, w hich result s in graded excit(ltory or
inhibitory postsynaptic po tentials, or in neuro
modulalory effects on intracellular signalin g
systems, in th e targe t cell. Some nerve te rmi nals
possess presynapti c receptors for their released
neurotransmitter. Acti va ti on of these recepto rs
regul ates neurotransmitter release.
11

OVERVIEW OF THE NERVOUS SYSTEM Neurons and Their Properties-Neurotransmission

Amino acid Catecholamine Serotonin :1 \~

s\'napsl' synapse synapse }JTryplophan ', .
/1
'I
JrrH \",\ )j:..
Gl ut,1 ma !e Pfesyna,Ytic f 5-0H tryptophan\\;)-(
0
00 receptor f ALAADi ~~,
\<\
o l
02. //
;t 5-OH.trypt amlne
,,_ (ser?tonin) \\ \~
gg .~
II M
i
-\\
,,>".- ,
f.;) l
~
__
' .
(~
Metabolism JPf'.. "
I . ,i
~'. . .
d,).,
r
~'

. ~r
l' _,:I) _ 'if~_'--
A'
'-', H';~ffi~~ "_j~"

'lS"!..- ~
~
..I uptake carrier

Peptide
' I' Peptide synthesized Acetylcholine

L~
synapse in cell body synapse

"~l) !
~
r -.,\
j .-
_ 0_ ~.)

FI GU RE 1.9: CHEMICAL NEUROTRANSM ISSION: AMINO ACID SYNAPSES _ _ _ _ _ __

Amino acids are compartmentalized in synaptic
\ esi cles (or release as neurotransmitters. The amino
acid glu tamate (depicted in the diagram) is th e
most abu ndan t excitato ry neurotransmitter in the
( 01 S. Following rel ease from the synaptic vesicle,
some glutamate binds to postsynaptic receptors.
Released glutamate is inac tivated by uptake into
both th e pre- and th e postsynaptic neurons, where
the amino acid is inco rpora ted into the Krebs cycle
or reutilized for a variety of fun c tions.

12
 "T""-er Properties-Neurotransmission
, f 5) 'A PSES _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
_~""'-; C\s are synthesized from th e
..c'" rosine, taken up competitively
-'C." , a carrier system. Tyrosine is syn
r'\3. b\ h rosine hydroxylase (TH ), a
.--etic enzyme. Additional conver
:-IE' 0 '\1occurs in th e cytoplasm via
acid decarboxylase (Al AAD).
~ n.aptic vesicles for subsequent
~,"".....",rgic nerve termin als, dop a
se (DB H) furth er hydroxylates
.--;....~.;""e in the synapti c vesicles. In
" -erminals, norepinephrine is
~ '~ PSES _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
?Sized from the dietary amino
~"en up competitively into the
-.:"I'" ~ stem. Tryptophan is synthesized
!nJ(> an by tryptophan hydroxylase
-g svntheti c enzym e. Conversion
;.:;:o...,.-ophan to 5-hydroxytryptamine
0'. the decarboxylase A LAA D
"'Dplasm. Sero lonin is stored in
..=""-.. =olJow in g release, it can bind to
~ 'APSES _______________________________________________
- ae :ivnthesized from prohormones,
: S' n :,esi zed in the cell body fro m an
-~_ rne larger precursor peptide is
~ ti ona ll y to active neuropeptides,
ed in synaptic vesicles and trans
::radeh by the process of axoplasm;c
\ esicles are stored in the nerve
re eased by app ropriate excitati o n
"'"' '5 induced by an action potential.
=,""",Ie bi nds to receptors on the
-C S) ~APSES _______________________________________________
"-Ch I is synth es ized from dietary
(oA, deri ved from the me tabo
.a the enzyme choline acetyl
';T. '\Ch is stored in synaptic
g ~ele ase{ it binds to cholinergic
OVERVIEW OF THE NERVOUS SYSTEM
methylated to epinephri ne by phenylethanolamine
N-methyl transferase (PNMT). Fo llow ing release,
the CA neurotransmitter binds to appropriate
receptors (OA, alpha and beta adren ergic recep
tors) on the postsynapti C membrane, al te rin g
postsynapti c excitability and/or second messe n
gers. CAs also can activa te presynaptic receptors,
mod ulating the excitabili ty of the presynaptic
terminal. CAs are inactiva ted m ainly by presynaptic
reuptake (hi gh affinity uptake carrier) and by
metaboli sm (MAO deamin atio n and CO M T
methylation) and diffusio n.
receptors on the postsynapti c membran e, alterin g
postsynaptic excitability and/ or second messenger
activation. Seroto nin can act o n presynaptic
receptors (5-HT receptors), modulating exc it
ability of th e presynaptic termin al. Serotonin is
inactivated mainly by presynapti c reuptake (hi gh
affini ty uptake ca rrier) and also by meta bolism
and diffusion.
postsynaptic membrane. In the CN S, there is o ften
an anato mical m ismatch betwee n th e localizalio n
of peptidergi c nerve terminals and the locali
za tion of cells possessing membrane receptors
respo nsive to that neuro peptide, suggesting tha t
the amount of release and the extent of diffusion
are impo rtant factors in neuropeptide neurotrans
mi ssion. Released neuropeptid es (I re inactivated
by peptidases.
recep tors (nicotini c or muscarini c) o n the post
synaptic membrane. influencing the excitability
of Ihe postsynaptic cell. Enzymatic hydrolysis
(cleavage) by acetylcholine esterase (AChE) rapidly
inactivates ACh.
13
OVERVIEW OF THE NERVOUS SYSTEM Neurons and Their Properties-Electrical Properties

F.x(racellular Ilui d + 11111

11 11-1 Axoplasm

N.l-+

if"'''
m+ '-:/
-- ~

~~ ;",,'C

(1 a

(Na t

"" . - !
Res';!Jr>Cc '" t50 mY

,I
I I
Equivalem
~~~
" ICoN
'~ '.-.-.: ::

""",I
I_ ;- EO'';~ .AAA;. di.oll''''''

~ vvv l L~-~'~'~m~vjl--I
\. 'I RM'
-?On .\.'

FIGURE 1.10: NEURONAL RESTING POTENTlAl _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Cations (+) alld anions (- ) are distributed unevenly
ac ross th e neurona l cell me mbrane because the
membrane is differentially permeable to these io ns.
The membrane's permeab ili ty to ions chan ges with
depolari za tion (toward 0) or hyperpo larizati on
(away from 0 ). Ion distri bution depends o n the
forces of charge separation and diffusion. TI)e
lypical neuro nal res ting pote ntial is approxim ately
- 90 mV w ith respect to the extracellular fluid.
The extracellular concentrat io ns of Na+ and CI- of
145 and 105 mEq/L are hi gh compared to th e
14
intracellular conce ntration s of 15 and 8 m Eq/L. The
extrace llular concentration of K+ of 3.5 mEq/ L is
low compared to the intracellular concentration of
130 mEq/L. The resting potential of neurons is
dose to th e equilibrium potential for K+(as if the
membrane were permeable o nly Lo K+)_Na+is
actively pumped out of the cell, in exchange for
inward pump ing of K+, by th e Na+-K+ATPase mem
brane pump. Equivalem circuit diagrams for K+and
for CI-, ca lculated usin g th e Nernst equation, are
illu strated above.
~rties . u rons and Their Properties- Electrical Properties OVERVIEW OF THE NERVOUS SYSTEM

--
Chemical Synaptic Transmission
_ ..." - - ---1 '"....... - - -
._.
~~---- ~~----

"
inS)""";':
"
@ @ '""~
Q ,,.,,...,.
........
~

wr.... """"~ .~..a"" =""I~'Y ".""PI" "'1 in/libiloo S)'~_. ~"""";"rf wbsl3nce
boUlon . .. ( i uS<!" rel",",", ot a ".,,,,,.,,n.. '<"Ieas<'(! br i n Itn~ Inc'~._ ptome
Wbslj"W InlO oynoplk<.lPil Th.. ;"c,~as.H obhW 01 ".,...<).....11'''' m ..ml...ne to '"
p""m~~bihly '" I'O'bvnoplir;: mcmb<~nc I(> IOdcr bu, ~ 10.'1.-. " ~Dutof
,....,. ~ nd I('> ~ ...... N.~ ",,,,... in l0pos~ ~l.
~"C(..II"" " " """.... ou d~1O
....~,#,~~C~ p........l

R~~'" .,," ionic OJn...,' iIow ... in .. R.....c~. ""''' (utrl'lll. .0..... in dire<1ioon
d l'Kllon ..~ Iit.h ~~ 10 dtopoIari>r po". nfll(f>mom to ~ cb'i.I~ pm"_pl i~
lyn3Q\io. ~I II depc>1aru ...., .~ cell. Th.. n',,/'"'' dfIxIbnUlion bo, _'''''''v
fi"", .t"e<hoid i ' Ih(- .on hIIod:. ~n . .. ~ noOo. oJ;i&o.ull..........~ dtvubnz;olion
Hl,,"uM is ~_M~ in post>ynapOC cdL ""O'<IUlfW'O!>ic,h .... ~1II>o1d

.,

,,!

>

<

-"
"
--
FIGU RE 1.11 : GRADED POTENT IALS IN NEURONS
_ __ _ _ _ _ _ _ _ _ _ _ __
" "

)f Excitato ry and inhibitory neurotransmission are
processes by w hich released neuro transmitter, act
in g on postsynaptic membrane receptors, elicits a
loca l or regional perturbatio n in the membrane
potentia l: (1) toward 0 (d epo lari za tio n, excitatory
pos tsy nap lic potential [EPSP J) via an inward flow of
Na+ caused by increased permeabi lity of the mem
brane to positive ions or (2) away from 0 (hyper
polarization. inhibitory postsynaptic potential
(l PSPI) via an inward flow of CI- and a compen
sato ry outward flow of K+ caused by increased
membrane permea bility to CI-. The resu lt ant EPSPs
and IPSPs exert loca l influences that diss ipate over
time and d istance but contribute to th e ove rall
exci tability and ion distribution in the neuron. If
sufficient excitatory innuences bring about depolar
iza tion o f the initial segment of the axon above
threshold, an action potential is fired .
15
OVERVIEW OF THE NERVOUS SYSTEM Neurons and Their Properties-Electrical Properties

+ 20

.
;; 0 Action po tential

1&
\ \ Na + co nductan ce

~
~
E
o
" K- co nduc tance
'"

- 70 I ( .L .......... I" . . . . . . .~ =:s=

ms::....J -l 0~5 "- 1.0 ,
_ _ _ _A A A,_ _ _ ~

,.. " ,. Membrane ..... ,. "

~~:\""" ~f~i'"" 'm

NiI+~ Na+' I Na+ ~ .

=1:
K+
"1:
a - ==::::;: a
K'
K' ~I_
+ -
CI~

+1- - 15
(1 ' .J
-1+
- +
+ 20
+ ~ ~
At firin g level Na+ conduc K+ conductance increases,
tance g reatly iocrea56, causing lepolari zatio n;
givin g rise to sllong iowa rd Na" cond uctance le turn~
Stimu lus current Na + current,. leads 10 exp lo- to no rm al
prod uces depolari zation sive positive feedba ck w ith
depolarizatio n increasing
Na-+ conductance
""--,, +
+

- ~!p

lION
; ..: ..: .
+..J
,
Equivalent circuit diilglarru.

FI GURE 1.12: ACTION POTENTIALS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Action potentials (A Ps) are allor-none, nondecre
mental, electrical potential s that all ow an el ectrical
signal to travel lon g distan ces (a meter or more)
and trigger neurotransmitter release through
electroch emical coupling (excitation-secreti on
co upling). APs are usually initiated at the initi al
segment o f axons w hen temporal and spatial sum
matio n o f EPSPs cause suffi cient excitation (depo
larization) to open N a-+ channels, all owing the
16
membrane to reach threshold. Thresho ld is the
pOint at which N a+ infiux through these Na+ chan
nels cann ot be countered by efflux of K-+. W hen
threshold is reached, an AP is fired . As the axon
rapidly depo/arizes during the risin g phase of the
AP, the axo nal mem bran e increases its K+ cond uc
tance, w hich then allows inAux of K+- to counter the
rapid depolarizatio n and bring the membran e
potential back towa rd its res ting leve l.
erties
- Ne urons a nd The ir Prope rti es- Electri ca l Properties OVERVIEW O F THE N ERVOU S SYSTEM

Inh .Kellul.n Ex llacelluill'

polen]ial -(>0 mY poten tial .. 1 mY

Axop lns m

Intracellular Ex tracellular
potentia l +20 mY po tentiil l - ~ mY

InlrM:ellul31
potffili.:!I -7S mV

Intrac e llular
pOlenlial (mY ) Resti ng
polen!ial
-70 f---~;-t-'===--~--==

.,I
t

----,\1') I
J, ~
Exlrace llular 0 tf. c I~
po ten!ial (mVI_5 1 _ _ _ _=~===~-------
: ~. -.;::
' - 1,OrnsoPc - '

- FIGU RE 1.13: PROPAGATION OF THE ACTION POTENTIAL _ _ _ _ _ _ _ _ _ _ _ __

W hen an AP is initi ated at a specific site o n the
axona l membra ne (usually th e initial segm ent), the
inward flow o f Na-l- alt ers the extracellular ion
environment, causing a local flow of charge from
adjacent regio ns of the axon. This induces a depo
larized state in th e adjace nt node of Ra nvier or
,e patch of axo nal membrane, bringing th at regio n to
thresho ld and resu lting in the reinitiation of th e AP.
The presence of myelination alo ng axo nal seg
ments resu lts in the reinitiatio n of the AP at the
nex t node, thus hastening the velocity of conduc
tio n of the AP. The resultant appearance of the AP
skipping from node to node dow n the axo n is
call ed sa ltatory cond ucti on.

17
OVERVIEW OF THE NERVOUS SYSTEM Neurons and Their Properties-Electri ca l Properties

A. Myelinated fibers

+
L ;./:.J.}
, ..
.. - +/
Site where ac tion potential is reinitiated

Node

B. Unmyelinated fibers

C. Classification of nerve fibers by si;re and conduction velocity /l

12i ~ M yejin.lledfibers/4 '

1 10
1 ~
~~
100 Alpha motor neurons to extlaiusal stria ted
(somatic ) m usc le fibe rs (malOf end plates)
90
IIliN1\'::.'::
Gamma mOtor neurons \
80
to intrafusal l"ibers o f spindles
in striated muscle Group I (Ao: fib ers) Ja fro m primary muscle
70 spindle endings: p roprioceptio n; Ib fro m
Co nduction Goigi tendon o rgans: p roprioception
velocity 60
(meterslsec)
G roup 11 (AP libers) flOm secondary endings of
50 muscle spindles: proprioception; from specialized
U,'U'PIOIS in skin and dee p ti ssues: touch, pressure
40

30 Group III (AS iibeTs) irom iree a nd from some sp ecialiled endings
in mus<:le and joints: pain; irom ski n: sharp pain, hea l, cold, a nd
20 some touch and press ure; a lso man y viscera l afferents

10
IP'" /" Unm yelinated jibers
? - G roup IV (C fibers) 'rom skin <lOO muscle: slow burning pa in; also visceral pain
, 10 15 20
Fiber di<lmeter (microns)

FIGURE 1.14: CONDUCTION VE LOCITY _ _ _ _ __ _ __ __ _ _ _ _ _ _ __

The AP travels down the axo n by dep olarizing presence of a myelin sheath (pa nel C). In myeli
adjacent patches of membrane (panel B), leading nated axons, the AP is propagated from node
to the AP's reinitiati on. The speed of pro pa gation to node, a process called saltatory conduction
increases with larger axonal diameter an d in the (panel A).

18

erties ' eurons and Their Properties-Electrical Properties OVERVIEW OF THE NERVOUS SYSTEM
-
Presynaptic Inhi bi~~~~:;;::"i~::=
tion ;Po
~~st~synaP
~ ~: lic Inhibition
=="-~ ,
(lnhibilo'1' libef\ (ueil,llory iibei' )

MOlor
neuron

Axon
""~
~}ft!%
(lrohobirOf y iih~r)

mV

A. Only E fi rH
oW
-,,, " "lION
.....: : :

9O mV ~ike
,, mV
in E t~r m;n,,1 ~90mV
,,, A', O nly E fi res

- 70
,, EPSPin -6()

r-------
mOlo. neuron
EPSP in -{.o j
mOlar nl"Oron
-70
-7U

B. O nl y I frre~

~~j ~
l l'lng.l,lS ting parlial
d~pol':l.fi l;.ltion
8'. O nl y I fires
in E rerminal
MOIor np.ur()fl
hvper poLlri~ed
-70
NO response in ~
moror neuron - 70 J - 80
C. t firC1 before E
.20 .,
P.arlial ,,
rkpolariwion of
,,
E rermin.J1 reduO!~
spike to ao mV, ,,~80mV C. l ll rHbefo(~E
thus relea$ing less
tl an5mitter ,,
$u!)!ilO1nce OepoLlIil.llion -<0
-70 of molor
Smaller ( PSP in
motor mwroo
-(,()

-70 J ~
neuron reM>
lI~ni" only
f fir M
-70

-B<l
-v----
FIGURE 1.1 5: PRESYNAPTI C AND POSTSYNAPTIC INHIBITION

Inhibitory syna pses modulate neuronal excitability.
Presynaptic inhibitio n (left) and postsynaptic
inhibition (right) are shown in rela tion to a moto r
neuro n. Postsynaptic inhibitio n causes local hyper
po lari za tion at the pos tsynap tic site. Presynaptic
inhib itio n invo lves th e depolari zati on o f an
exci tato ry axon terminal, w hich decreases the
amount of Ca++ influx thaI occu rs w ith depo lari za
tio n of that excitato ry terminal, thus reducing the
resuhant EPSP.
19
OVERVIEW OF THE NERVOUS SYSTEM Neurons and Their Properties-Electrical Properties

:::~~: ~L)t ~'---.J

Excitatory fi bers ,, ~~,

-;.,J (;0 '-::::::::======

==~ A,oo
l: hibiiOry fibers - - Inhib itory hi

A. Rest ing state: mOlor n erve cell sh own w ith syna ptic b o ut nns of B. Partial depolariza tion: im pulse from one excita tory fib er has
exci tato ry an d in hibito ry nerve fibl? rs ending cl ose to il caused partial (below firing thr esho ld) de polar izatio n o r motor
ne uro n

Exci ta tory (ib er5 '" ,,"<:':0,

Exci tatory fib ers '" '-...........~
\ '-

tnh ibito ry 11
Inhi bitory fi
C. Temporal exd t..tory summa ti o n: a series o f impulses in o ne D. Spatial e~ci tatory summation: impuls5 in two excita rory fiber!>
excita tory fiber to gether produce a supra thresho ld depolariZdlion cause tlVO synap tic depolari zatiol1s th at together re ach firing
that triggers an action potential threshold tri ggerin g an actio n po ten tial

Excitatory fibers l :so: -:""'(\ Excita to ry iibe rs

Inh ibitory fi Inhibi tory fibe rs

-
E. Spatial excitatory summation with inhi bition: impulses fro m E. (con tinu ed): motor neuron now receives addition;!1 excitato ry
two exCitatory fibers reach molor neuron but impulses from impulses a nd reaches firi ng threshold d espite a simultaneous
inhibitory fiber prevent depolarization from reaching threshold inhibito r)' impulse; add itio nal inh ib itory impu lse", might still
prevent firing
Axon{s) activaled in each sce nari o
,p ,f$
tT I~~oN
:,.".,.::~

FIGURE 1.16: SPATIAL AND TEMPORAL SUMMATION

_ _ _ _ _ _ _ _ _ _ _ _ __
Neuron s rece ive-multiple excitato ry and inhibitory
inputs. Temporal summation occurs w hen a series
o f sub thresho ld EPSPs in one exc itatory fiber pro
duces an AP in the postsynaptic cell (pan el C) . This
occurs because th e EPSPs are superimposed on
each oth er te mpora lly before the local regio n of
membrane has completely returned to its resting
stale. Spatial summatio n o ccurs w hen subthreshold
20
EPSPs fro m 2 or more synapses trigger an AP
because of synergistic interactio ns (panel D).
Both temporal and spatial sum matio n can be mod
ulated by simultaneo us inhibitory inpu t (panel E).
Inh ibitory and excitato ry neuron s use a wide
variety of neuro transm itters, w hose actio ns depend
o n the ion channels o pened by the ligand-recepto r
inte raction.
perties Brain OVERVIEW OF THE NERVOUS SYSTEM

, Anterior
Sphenoid bone
>-
,
cranial
fossa
r has
Ilor

-/ Sella
: Middle
~ craniill
turcICa I fossil

Groove for int. G1rotid <I .

Jlbers

)
I Posterior

,
!>cranial

-
Suklls for
, I

Occipital bOne
BaSilar

Suki for post

.
v

Internal
iN
,..;: Internal i
Sulcus of sup.

FIGURE 1.17: INTERIOR VIEW OF THE BASE OF THE ADULT SKULL _ _ _ _ _ _ _ _ __

The anterior, middle, and posterior cranial fossae
house the anterior frontal lobe, the tempo ral lobe,
)d and the cerebellum and brain stem, respectively.
I. The fossae are separated from each o ther by bony
structu res and dural membranes. Swelling or mass
~n d lesions can selectively exe rt pressure within the
:or
individual fossae. The perforated cribriform plate
allows the o lfactory nerves to penetrate into the
o lfacto ry bulb, a site w here head tra uma can
result in the tea ring of the penetra ting o lfactory
nerve fibers.

21

OVERVIEW OF THE NERVOUS SYSTEM Brain I

f Olamen ceo:um - - - - - - - - - - Ve in 10 sup. '-<lgi!\al sinus

/ " Am . e lhmoidal (Of.llTlE'n - - - - - - Ant. ethm oidal a.., If. and n.
/ " Forami nil of - - - - - - - - - - - - 0liac(ory f'lefYIi' bun dles
c ribriform plate
I. e lhmoidal forame n - - - - - Po,!. e thm oirl ol a , v. and n.
Optlclill n.
Optio:. ca n.al- - - - - - - - - - {
./ Ophlha lmk a.
Oculomolo l (11 11 n.
ThlOchl(!'a r CIVI n .
......... SU..,. orbilai i'sswe- _ _ _ _ _ _ Ophthalmic n.
{ Abducen s (VII n.
Sup. ophth a lmi c \"
__ fo/;)rnen 'Olundum - - - - - - - - Maxillary n.

Mandib ul,ll n.
. .uessory meningeal a .
............ f Otamen ova.., - - - - - - - - - lesser pelroSdI n .

{
(occasiooal1yl
, . { Middle menin!!eal a. at'ld v.
!0,..----Foramen $pinosom - - - - - - Meningeal branch
-n*-\ of mandibul ar n.
';~;=~~~~'1f" ~'~-
.t~. FOtam e n 01 Vesalius - - - - - - - Sm all e m is.,.. ,y v.
of ."'&;: , (inconsla nli
t";.~;;';;;---~~~
'',_"(.,>,~

.. :~..~".;~'-
' fOX')meI'l lace<um __ __ __ _{ 1m. Cdrolid .1.
" .~ . -:i - 1m. carotid n. plexu s

~Hiatus of canal oi - - - - - - - - - les,er petrosal n.

~ . .,.?'t~~~~ _~, Hia tus of ca nal of - - - - - - - - - G re~ler petrusa l n.
it::"'"~ .... { Facial (V IJ ) n.
. acous tic mealUS - - - - - - Vesl ibulocochlear (VIII) n.
labyrinthine a.
- - Vestibula, aqueduct - - - - - - - - Endolymphatic duc t
M as toid for,lmen ____ __ _ { EmiS>J ry~. "
(incon stant) Bra nch of OCCIpi tal a.
Ini. pelroS<li 5inu~
Glos.s.oph.u yngea i (IX) n.
" Jugular fo ramen _ _ _ _ _ _ _ _ Vagu~ (X) n.
AcCe5SQfY (XI) n.
{ SIgmoid sinus
I'OSl. meningeal .l.

" { Emi ss ary v.

CondylJ( Col nJ I - - - - - - - - Menin geal branch o f
linconstant) asc end ing p/1ar yngeal i1.

'\ Hypoglor.5al <:<In<ll . - - - - - - - - Hypoglo~1 (XII) n.

" Foram en ma!!nvnl - - - - -

{
Medulla oblo ngala:
Meo in ges
Vertebra l aa.
~~p
" 11liN
..
: ::,': \
Sflina l roots of a(Te>$ory nn .

FIGURE 1.18: FORAMINA IN THE BASE OF THE ADULT SKULL _ _ _ _ _ _ _ _ _ __

This illustration of the foramina in the ba se of traction, or masses can damage structures travers

the skull lists the major nerves and blood vesse ls ing in these tightly co nfined spaces.

that course through each opening. Press ure,

22

 OVERVIEW OF THE NERVOUS SYSTEM

---------------------------------------------
Brain Brain

i I margin of cerebrum

Frontal (Fl . irontop;llletal (FPJ

and temporJI (T) opercula
'e<,.p." "",,.'lsu lcus
Supe rior irOOlal gyr u~ In ferior p~rietJI
lobu le
, I
Middle frontal
S~ U \ - _ _ _~;.~~

Inferior frontal
sulcu~ _ __

Inferior
fro ntal
gyru ~

Lateral { Anle< IO<

(syivi,m ) . . , ~III ,IS "'m,,, I
lisw re Posle, ;o< ramus
mar gin 0 1
Temporal temporal cerebru m
gy r u~
Superirn lcmpo ral su lc us OC(lpit~1 ~uk:u ~

Middle temfXIrJI gvrus Preoccipital notch

lnie-rior lemporal ,ukus

Tempo ral
I,..

Short .
los ula limen
{
Lun!: gyru s

ers-
FIGURE 1.19: SURFACE ANATOMY OF THE FOREBRAIN: LATERAL VIEW

The convo luti ons of the cerebral cortex allow a
':i ge expanse o( cortex to be compactly fo lded
, to a small volume, an adaptation particularly
p'ominent in primates. Major dependable land
~ar k s (lateral fissure, central sulcus, pariero
occi pi tal fissure) separate the forebra in into lobes.
n e lateral (sylvian) fissure separates the temporal
" be below from the parietal and fro ntal lobes
above, and the central sul cus separates th e parietal
and the frontal lobes. Several of the named gyri are
associated w ith specific functional activities, such
as the precentral gyrus (mo tor cortex) and the
postcentral gyrus (primary sensory cortex). The
insula, the fifth lobe of the cereb ra! cortex, is deep
to the o uter co rtex and can be see n by o penin g the
lateral fissure.
23
OVERVIEW OF THE NERVOUS SYSTEM
,t<,
FIGURE 1.20: LATE RAL VIEW OF THE FOREBRAIN: FU N CTIONAL REG IONS _ _ _ _ _ __
Some circumscribed regio ns of the cerebral hemi
sphere are associ ated with specific fun ctional
activiti es, includin g the motor cortex and the su p-
plemental and pre motor cortices, the frontal eye
fields, and the primary and associatio n sensory
cortices. Part of th e auditory co rtex is visib le at th e
inferi o r edge of th e lateral fi ss ure (th e transverse
tempo ral gyrus of Heschl). The visual cortex is seen
FIGURE 1.21 : LATERAL VIEW OF THE FOREBRAI N : BROD MAN N AR EAS
_ __ _ _ _ __
Brodmann areas of the cerebral cortex reflect
unique architectural characteri stics of the thickness
and layerin g of the cerebral cortex from histolog
ica l observations origin ally mad e by Korbinian
Brodmann in 1909. This numbering o f cortical
areas is still used as a shorthand for describing func
24
Drain
l
,ttlP
"~
at the occipital po le. Language areas of the left
hemisp here includ e Broca's area (expressive lan
guage) and Wernicke's area (receptive language).
D amage to these cortical regions results in loss of
specific functio nal capabilities. There is some over
lap but not absolute concordance betwee n func
tional areas and th e named gyri (e.g., mo to r cortex
and precentral gyrus).
,tb<
0'-lN
tional areas o r regio ns of the cortex, particularly
those rela ted to sensory functi ons. Some overlap
exists between functi onal areas. For example, the
motor co rtex is area 4/ th e primary sensory cortex
includes areas 3, 1, and 2, and the primary visual
co rtex is area 17.
 Brain Brain OVERVIEW OF TH E NERVOUS SYSTEM
Cingu lale gyrus Pr E'Ceflt ral sulcus
C mgul.l le I Cemral (ro l.l ndic) ~ulcu~
Par.lcentra l lobul~
~co'pc; ca ll osum
Su lcus of corpus ca llosum
Precun e us
Superior sagittal , inu s
Sep{IJm pell uc id um
:K-'''. _~ Cho rOid p lexus of 3rd ventricle
Illlerventrlcular foram en
Parie toccipila l SUlW5
Stria med ullari5 of thalIDllJS
lnterthalamic adhe5ion ClJnelJs
Thalamus _"-'''"
Calcm ine curtex
Anlerior commissu re
IlJpper bank)
Habenul.ll commissure
Calcarine slJlcus
Subcallosal

Lingu;:d gyrus
(pa rolfactoryl are a

C.J. I(.lrine cortex

f'araterm inal g,,", --C'" (lower bankl
Pineal gland
Lamin a Ie/minalis Strai ght sinus

'f
Op tic chiasm

Mammillary

(in tentori um cerebellil

""'"'' o,~I",11 vein (of Galelll
Posterior tepithalamicl commissure
Super io r and inierlor colliculi
'n Cere bellum
Pituita ry gland (anterior and po",,'o,, /
Su periol med ulla ry velum
gel.
Mldbtain
4th ve ntricle and cho roid plex us
s of Cerebral aqued uct In fe rior medu llary velu m
)Ver obIong':lId lof Sylvius)
nc
Irtex Cin gula le

Mammillothalamic .. I
t::5:i::;;~:::~;R_O~~n~ }
':'> Sody
of corpus callosum
Splenit.m
Cuneus
O pt ic (111) ne rve ~<.. ,'--. I
Olfaclory trac t Lingual gyrus

_co~,C"m'""~}
Collate ral "I<,' ''-.;~
offomix
Rhina l ,,"' "'_..:
Medi a l occipitolemporai gy'"' -__-=::=
hippocampus
Ocdpitotemporal >ulru s
Denta te gyros
laler occipitolem(lOfai g\'ft.!s Parah'ppocam paJ gyrus

FfGURE 1.22: ANATOMY OF TH E MED IAL (MIDSAG ITIAl) SURFACE OF THE BRAIN _ _ __
Th e entire neuraxis from the spinomeduUary junc
ti on throu gh the brain stem, diencep halon, and
telen cepha lon is visib le. The corpu s ca ll osum, a
major co mmi ss ural fiber bundle interco nn ecting
the 2 hem ispheres, is a landmark separating the
cerebral cortex above from the thalamus, fornix,
and subcortical forebrai n below. The ventricular
5vstem-including the interventricular foramen, the
third ventricle (diencephalon), the cerebral aque
duct (midbrain), and the fourth ventricle (po ns
and medulla}-provides intern al and external (sub
arachno id [SA] space) fluid protection to the brain.
The thalamus serves as a gateway to the cortex.
The proximity of the hypo thalamus to the median
eminence (tube r cinereum) and the pitui ta ry gland
reflects th e important role of th e hypothalamus in
regulatin g neuroe ndocrine function. The Cshaped
course of the fornix, from th e hippocampa l forma
tion in the temporal lobe to th e septu m and the
hypo thalamus, is shown below. The midsagittal
cut thro ugh the brain stem reveals the midbrain
colliculi, sometimes called the visual (superior)
and auditory (inferio r) tecta.
25
OVERVIEW OF THE NERVOUS SYSTEM Brain

Primary moun .-

limbic,; cingulate Ul<'1ex , _ _ ParKentral lobu lt!

Supplem ental moto r co rte\ ;;~ lali

Corpus callosu m

:::-- Vi~ a l assoc i.ltion (OfttK

P, ima'yvlsuaJ cortex

Calcarine fissure

Pil:\.lit<l.ry

Cefebellum
~~~
IIllN :\ ....... ;:

Meduh obIo~.a

FIGURE 1.23: MEDIAL SURFACE OF THE BRAIN: LOBES AND FUNCTION AL AREAS _ _ _ __
The cingulate cortex is labeled the limbic lobe, motor cortices associated w ith the low er ex tremi
reflecting its association with o ther limbic forebrain ties are located medially and are supplied by the
stru ctures and hypothalami c co ntrol of the auto anterio r cerebral artery. This region is sel ectively
nomic nervous system. Fun ctional areas of the vulnerabl e to vascular or mass lesio ns, resu lting in
cortex, particularly those invo lved w ith vision, are contralateral motor and sensory deficits of the
best seen on a midsagittal view. The sensory and lower extremi ty.

,
~~~ Oll!!l\l
;:,'::: 1
f

R,
FIGURE 1.24: MEDIAL SURFACE OF THE BRAIN: BRODMANN AREAS _ _ _ _ _ __ _ _ a
CI
The Brodmann areas of the cereb ral cortex are co rtices and the continuation of area 4 (motor) and tic
labeled on this midsagittal view. The major regions areas 3, 1, and 2 (primary sensory) o nto the para -'"",
are the primary ( 17) and associative (18. 19) visual central lobule in the midline, l>
26
 Brain Bra in OVERVIEW OF THE NERVO US SYSTEM

Fron tal pole

of (;u rpus Glllosurn

O rbil.:t! ,
-- --- Lamin.:t termin.:t!is

O rbi lal gyri

l .:tler.:tl (sylvian)
/V,"', (1 11) nerve
Pitui t.J.ry gland

Temporal pole

Optic h act

An terio r (rost.:ll)
perforared su hst an(;~

Tuber dnereum

M ammillary body

Posterior

-
:!mi
perforated
substance

Rhinal sulcus Cere b,..1

he pedu ndt'~
,Iy (crU) ct!rebri)
g in l ateral
~e nl culat e bodv

Lat er al
S lIb ~ tan ti ~ n i s r~
occipi to temporal

gyru s
M edial ~ e ni c uliJt t'
bndy
Occipito temporal

sulcus
Reel n!.ldcus
Medi al
Pulvinar
ocdllitotempolal ,,'"' - --"'4[
Supe.ior Icr.lni ..1l t:nllkulu5
CoUateral sulcus - -

PariJhippocampalgyru~ --=~~~td+ Cer~hr ..1aqueduc t (of SVMus)

Splenium oi oorpu~ callosum

Uncus

Cingula te gyr us

FIGURE U5: ANATOMY OF THE BASAL SURFACE OF THE BRAIN: W ITH THE BRAIN STEM
AND CER EBELLUM REMOVED _ _ __ _ _ _ _ _ _ _ _ _ _ _ __

Remova l of the brain stem and cerebeHum with a th e op tic chi asm. The proximity of th e pituitary

-
,nd
cut throu gh the midbrain exposes the underlying
cereb ral cortex, th e base oi th e diencephalon, and
gland to the op tic chi asm is importan t be cause
bitempora l hem ianopia can result from optic
chiasm fiber damage, a possible early indi cation of
th e basal forebrain. Ba sal hypo thalamic landmarks,
1- from caudal to rostral, include the mammillary a pituitary tumor.
bo di es, th e tuber cin ereum, the pituitary gland, and
27

OVERVIEW OF THE NERVOUS SYSTEM Brain

O li,lctor y hulb
Ex ec utiv" an ticipiltory
<Of Ifeat runction

/ OliactOfytr.tct
Tem por al pol

Optic nef\'e (Il ) (tun

,.--- O pliC ,ril el

JI,\ ommlii ary body

.Vluhi.lcnsorv a~s()(]ation artlol 'i

~!~ " lJiliN

',';::: I

Prim<lry "i~"':Jr aortex

FIGURE 1.26: BASAL SURFACE OF THE BRAIN: FUNCTIONAL AREAS AND BRODMANN AREAS _ He
This view provides inform ation about the medial hipp ocampal formatio n, the amygdaloid nudei,
lem porallo be on the left side of the brain, espe and the olfactory system. O n the ri ght side of the
cia lly the cortical regio ns associated w ith the brain, the Bro dmann areas are noted. "'"
.~

L>l'
C~
!)
.,.,
'dJ.....
cor
28

 Brain Brain OVERVIEW OF THE NERVOUS SYSTEM

Genu 01 corpus c.Jllosum

/;ep',"", pelludrlum
Head o f caudale nucll'US

Anterior
_ li mb } 'nlernCl ,
Genu
Posterior limb capsu le

II co rtex

I ( p<llIidus } l l'n lifonl'l

Inlemal ann nucleus
extern.ll segmentS

Extern,,1 CilPsule

Claustrum
~:::::::;:i=- Habenula
Tail of Ciluoolte
nudeu!>
Pineal gland

1.1Ieral vemride
CIUS 0 ; iornix Hippocamp us

Spleniu m o f corpus callosum and fimbrr a

\P,,,,,,.,ioc loccipital)

horn of faleral
ventricle

Cauda te {
nucleus
Head _ _ -,

lev~ ls of {A - - ------ ---- 't .~- - -. ----- - --

~ect10nS B --- ------ ---- - -\+ - -- -------
lentiform nudeus

to p u tamen
{globus )
p.-tllidus med"1<l1 .. ,::;""'~~~~~~~~iM~:.~di:"~;'::~~:: body
::.

Schematic iAIU'm"YI",'di'o'I,~"idh",b"'.i'")g-i-"~"i'.'l ' 'io",hiP ol,h.,I, ","" ',"" 'O~:'eu, body ~ ~,

nud eus. cauOOle nucleus olnd amygdaloid body lviewed from sidl't

.S _ FIGURE 1.27: HORIZONTAL BRAIN SECTIONS SHOWING THE BASAL GANGLIA

_ _ _ _ __
Two levels of horizontal sections through the fo re
Ie bra in reveal the major anatomical features and
relationships of th e basa l gangli a, the intern al
ca psule, and the th alamus (bottom illustration). The
(.-s haped ca udate nucl eus sweeps from th e frontal
lobe into the temporal lobe. In the internal capsule,
the anterior and posterio r limbs and the genu con
tain major connections into and out of the cerebral
cortex. The relatio nships of the basal ganglia, the
internal capsule, and the thalamus are important
for un derstanding imaging studies and the involve
ment of specifi c fllnction al systems in vascu lar
lesio ns o r strokes. Th e extern al capsule, th e clau s
trum, th e extreme capsule, and the insul ar co rtex,
labeled from medial to lateral, are landmarks used
in ima ging stud ies. The fornix is sectioned in the
crus and the column.
29

OVERVIEW OF THE NERVOUS SYSTEM Brain BrdJ

.;>'nte, ;o, n ucleus oi thal amus IntenhJ lamK adhesion

i~'
Intcrv!3 nlrlculal foramen
Slri,l lermina lis
A.nte:IOr commi~",u l e
Stria medu lJ.lris

Cingulate Rvrus

Imgli~eum,- ~

--
..p.~

Corpu~ c<lllo~lIm """"" .'

............. -"

S.. ptum pe Jiuci cilH'll

f'fccomnll$sural iorOl" __ '~

r1
Si2'lllil l rlll
,

S ubcallo~i'llarl'a ~ . '
, ~
RG
-e

wmlO..l ~ Jr =-~
tcm1ITlcli iS
.:;; ?"
b""'
l r ~c l
Q liaclory medial
{ I.Herdl '

Anterior p~ rfor,lted
substan ce -----~/
Optic chiasm /

PO~l commiSSll(al fornix

Mam ll1illMY b ody an d

i11anlmiliothaia mlC IraC I

Medial fort-blain bundle /

Amygdaloid h odv

(nucl ei)

~~~
Parahipp ol:ampal gyru s
Interp edunculM nudeu>
Dest:ending (onne<:tions to reticular
and legmental nud ei of brainslem c :\":.":;:
ll9N
Fasciculul> relr(){lexus I (dOfsal longi tud inal fasciculus)

FIGURE 1.28: MAJOR LIMBIC FOR EBRAIN STRUCTURES

_ _ _ _ _ _ _ _ _ _ _ __
Many of the structures and their pa thways in th e
limbic system form a ring (limbus) aro und th e
diencephalon. They are involved in emotional
behavior and interpreta tio n of external and internal
st imuli, Th e hippocampal formati on and its pat h
ways, th e fornix, curve int o the anteri or pol e of the
diencephalon. The amygdaloid nuclei give rise to
the stria terminaiis, a C-shaped projection (0 the
30
hypothalami c and basal fo reb rain stru c tures. Th e
olfacto ry tract co mmuni cCl tes directly with th e
-'G
cortex a nd othe r forebrain structures, bypassing the
thalamus. Connectio ns from the septal nuclei to
the habenula (stria termin ali s) interconnect the ..3.1
limbic forebrain w ith the bra in stem. The amyg cal;
dalo id nuclei and the hippoca mpal formatio n c"'...
(shaded) are deep to the cortex. ,--,.,
 Brain a rain OVERVIEW OF THE NERVOUS SYSTEM
---------------------------------------------
Anatomy of the Corpus Callosum

:- (:e,,",,1 longitudinal fissure

oi corpus callosum

:- lod,,,'om griseum (on upper

surface o j corpus callosum )

latefal longiludinal striae

_ Bo, of <O',M callosum

Splenium of corpus callo sum

f iGU RE 1.29: CORPUS CALLOSUM : HORIZ ONTAL VIEW _ _ _ __ __ _ _ _ _ __

The co rpu s ca llosum, th e major fiber commissure su rface. Horizontal cuts taken deeper (more
oetween the hemispheres, is a co nspicuous land ventral) will section the genu anteriorly and the
mark in imaging studi es. It is viewed from above splenium posteriorly (see Figure 1. 27),
after dissection of the tissue just dorsal to its upp er

'Jrine
us (tissure) ,
t lris fib ers
Medial
1
stria
Lateral

.j.1)'t:
stria

I[l;N
" ,"c.': :

The Occipilal iorceps (forceps major )

e
ng th e fiGU RE 1.30: CORPUS CALLOSUM: SCHEMATIC VIEW Of THE LATERAL EXTENT Of
MAJOR COMPONENTS _ _ _ _ _ _ _ __ __ _ _ _ _ _ __ __
to
e Many of the commi ssura! fibers of the co rpus midline. These interconn ecti o ns allow communi
g ca llosum, particularly the " forceps" of fibers inler ca tion to take place between the hemispheres,
connec ting the fronta l and the occipital areas, wh ich is essential for coordinating the activity of
curve rostrally and caudally after crossing the these 2 "separate" hemispheres.
31
OVERVIEW OF THE NERVOUS SYSTEM Brain B"
Di ~sect jon of the Hipp oc.amp a l Formation and Fo rnix
~.
,....--- {,,; enu of co rpus C<l li os um
Ilud,
H ead of caudale nl1c1e ... ~
Col"mns of fornix
Stria terminalJs

trody of iomix
1"1:'5 hippocampus

Uncu s

~~:~::,~~fornJX

~ Fimbriol 01 hippoca mpus

~.Hrppocampu~
' .......,;:;. Commi s>ure of fornix
iplen iufll 01 corpu, callosum
lal er<ll ve ntricl e

-"" Pos!elior (occipital) horn

o f 1~leral ve ntricle

Tajl of c~ u d ate nucle us

3D Re cunstructi o n of th e fo rnix Th e Hippo ca mpal Formation
Fimb ri.:l 01 hippocampus
in Co ronal S~ction ,
Columns Com mis 5tJ<"e Choroid
of plexus

CnJlJ of

Hippoc.:.mpJl wlcus
" ,~ -< ." ",

Subioculum

De ntate ventr icl e

ill
boditl~ "' ["ws of hippocampus I

Amysd aloid iii Efltorhinal cort e~

FIGURE 1.31: HIPPOCAMPA L FORMATION AN D FORNIX _ _ _ _ _ _ _ _ _ _ _ __ FIG

The cor tex, w hite matter, and corpus callosum C-shaped bundle is seen in the lower left d iagram. The
have been removed, th e lateral ve ntri cles have The hippocampal formation occupies a large por righ
bee n opened, and the head of the caudate and the tio n of the temporal pole o f the lateral ventricle. has
thalamus have been dissected away close to the The den tate gyrus is adjacent to subco mponents nu el
midline, allowing a downward view of the full of the comu Ammo nis (CA) regions o f the hip and
extent of the hippocampal formation, including the pocampus prop er (the CA 1 and CA3 regions), th e rem
dentate gyru s, and the associated fornix. This view subiculum and the entorhinal cortex. CA l pyrami fro n
reveals the relationship between the hippocampus dal neurons in the CA 1 region are sensitive to in t~
pro pe r and th e dentate gyrus. Th e 2 limbs of the ischemic damage, and their counterparts in pro(
forni x sweep upw ard medially, eventually runnin g th e CA3 regi o n are se nsitive to damage from high eire;
si de by side at their most dorsal positi on ben eath levels of corti costeroids (cortisol).
the co rpus callosum. The full extent of this archin g,
32

";.l, e>1l11c1e
o-.;.aple,us
_"" , _etJ(al v.
-",-~ ,
r, "Heral
.:..It:a.-ine sulcus - _/
~;.....el
_ ~~L ~MICANATOMY ________________________________________
.....:::-::>- 6 \. 'ew ed from above. The entire
- brain, just late ral to the thalamus,
Ed. the head of the ca uda te
IIf"er, secti oned, the corpus callosum
oorsaJ to the th alamus have been
:he th ird ventricl e has been opened
sl.u ia ce. The pin eal gland is present
~.;~. tlS"t caudal to th e third ven tricl e; it
"",...:""i..-1. a horm o ne that he lps regulate
~ . sleep, and immune responses.
OVERVIEW O F THE NERVOUS SYSTEM
Sel>tum p elludrlum
/,Cnl'cm,,, of forn ix
/ ,,,to', , ,, tubercle
,/ " " 0 terminalis
/ II" to"h" I"" " ad hesion
medu ll,u is
lal~ral geniculate body
Medial geniculate body
Brachium of
inferior coliiculu s
Superior co llicu lus
Inferior colliculus
The superior and in ferio r colliculi are shown,
depicting the do rsal surface of the midbrain. O n
the left, the tempo ral horn of the lateral ventricle,
with the hippocampal formation, has been
exposed to show the relationship of these struc
tures to th e thalam us. The terminal vein and the
choroid plexus acco mpany the stria term inali s
alon g th e lateral margin of the thalamu s. The stria
med ull aris runs alo ng the medial surfa ce of the
dorsal tha lam Ll s.
33
OVERVIEW OF THE NERVOUS SYSTEM Brain

3.d

Internal medull':Hy
la mina

, J ld ventricle
'-- Reticular nucleus
o f rhalJmus

Ivi
I,ml i!)')
Thalamic nuclei
" lille!']1 geniculdte body
M idline eM Cenllome(!ian
(medi<ln) nud ei LD l ateral d(){Si)1 '\. Mediill genic ula.!! body
lP Wller,ll posterior
M Medial woop Shem;ttic represe ntation of thalamus
MD Medial dorsal (extern al medullar y Lamina and
VA Ventral anfE'.rior ret icular nuclei removed)

~~p
Schematic section VI Vem fal intermedial
through thalamus \It Venlrallal!!''}/ _ _ Lateral cell mass
(at level of broken VP Venlral posteriOf h 'eolroOOrs.al) _ Medial cell mass
line sho wn in figure VPl Ventral postefOlarerJI
_ Anterior cell mass
CIIiiN
:'.-,.:: : :
alr ighll VPM Venltal poSleromedial

FIGU RE 1.33: THA LAMIC NUCLE I _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

These figures illu strate the subdivision of th e thala
mus into nuclear groups (medial, lateral, and
ante rior), separated by medullary (white matter)
laminae. Many of these nuclei are "specific" thala
mic nuclei that are reciproca lly connected w ith dis
crete regions of the cerebral cortex. Some of the
nucl ei, such as those embedd ed in th e int ernal
medullary lamin a (intra laminar nucl ei such as cen
tro median and para fascicular nuclei), and the ou ter,
laleral shell nucleus (reticular nucleus of the thala
mus) have very diffuse, Nnonspecific" associations
with the cerebral cortex.

34
- Br~ain Stem and Cerebellum OVERVIEW OF THE NERVOUS SYSTEM

Posterol ate ral view

/"Vp '" tract

A "",II genicu late body

(l V) n .

II Vl n.
r' cerebellar l)edun cle
Superior me,,",,II,,yv',,.I" >m'--; peduncle
ti I [II n.
Medi al e mo'o" ,r e -, :-;

Rhomhoid fossa of ,, II

peduncle
4 th veonlricle - - - - ..,

f"",:~::;:::::;J~
Cuneate
Gracile
~
~=~I~GIosS()Phary n 8eal (IX) and vagus (X l nn.
Dorsal roots of HWoglossaltrigo ne
I 5t spina l n. ((1 1 I trigone
AccesslXY (Xl) n.

tract
Anterior view

Optic

1
ill
Tuber
Temp oral lone
I I n.

Cerebral pi
oop:,~,c:::~==::=t~~ ~~1::J4=;::'~"'" UV ) n.
Posterior pE'floralcd
substance In
--=~~~::,~~' {Vt)
:.... :~: o.n.
I (VII) n.
Lateral genlculale
I.
Basilar ,m,,,,_ (Villi

Middle cerebellar ___

ped uncle - - - - -
Ii,

VenTral roots oi
lst spina l n . (C1 I - - - -'k

-
FIGU RE 1. 3 ~ : BRAIN STEM SURFACE ANATOMY: PO STE RO LATERAL AND ANTERIOR VIEWS_

"cen-
Top) Th e entire tel encephalo n, most of th e dien
cep halo n, and th e cerebe llum are rem oved to show
Outer, the dorsa l su rface of the b rain stem. Th e 3 cerebel
lala lar peduncles (superior, middle, inferio r) are sec
Ons tioned, and the cerebellum is removed. The do rsal
roo ts and the cranial nerves provide input to the
sp in al cord and provid e input and output to the
brain stem, respectively. The fourth nerve (troch lear)
is the on ly cranial nerve to exit dorsa lly from the
brain stem . The tubercles and trigones on the floor
of the fo urth ventricle are named for nu clei just
benea th them. (Bottom) The left temporal lobe is
di ssected to show the anterior (ventral ) surface of
the brain stem. Th e cerebral ped uncles, caudal
extensions o f the posterior limb o f the internal cap
sule, ca rry corticospinal and corticobulbar fibers
from the internal capsu le to the spinal cord and the
brain stem, respectively. The decussation of the
pyramids marks the bo undary between the ca udal
medulla an d the cervica l sp inal cord.
35
OVERVIEW OF THE NERVOUS SYSTEM Brain Stem and Cerebellum

Superior surface
~
Q u<,drangula. lobule
Ce ntral lob ule J,1 Pri ma ry tis sure
Hori:w nt.J.lllssu re
Simple x lohule
SUi'>""iOf
vermis ~
Oeclive
f'~tlunJte iissure
Follum Sup<::.iors!'ml
lun ar lo bule

~ t-I o riz()nl al f'~.\ure

- - Inierio. sem ilunar lobull'

Inferior surface ~
Superior (Cemral lobu1r,o --.
verm iS
,be ll

,II,
'1,

Inferior medu lluy ve lum _ _ _

1I'I1".io.
NOO"'. --
Uvula- - - -
vermis
{ Py ram,d
.
-

Tuber- - -
~
~
-- " (post-
, ii,>u re
P051~rior cerebellar no tch
/
tal n, SLln:'
fissure labul<:
Section in plane of superior cerebellar peduncle
- ''''-..~ -. ~~

~CerebralpedullCle

Decuss.ation of
superior cerebell ar peduncles
~ " ft..~-'{,
-.,
w J~~
, -- --Med'al longiru diIlJl l asCirulus
. , r' .,
'iel

1/, ~SupoeriOt cert:~ IIM

a pedunde

rei
:;J~CerebeollJ. r co rtex
De nlate nu cl e u s~
Emboli fornl nucleus....

FIGURE 1.35: CEREBELLAR ANATOMY: EXTERNAL AND INTERNAL FEATURES
_ _ _ _ _ __
(To p) The top 2 diagrams show the superior (dor
sa l) an d inferior (ventral) surfaces of th e cerebel
lum. The ven tral surface of the cerebellum is the
roof of the fourth ventricle. The anterior, middle,
and flocculon odular lobes of the cerebellum are
trad itional anato mical subdivis ions with w ell
descr ibed syndromes from lesions. The vermis, par
avermis, and lateral hemispheres are cerebellar
cortical zones w ith specific projection relationships
w ith deep cerebell ar nucl ei (vermis with fa stigial
nucleus and latera l vestibular nucleus, paraverm is
with globose and emboliform nuclei, lateral hemi
spheres w ith dentate nucleus), connecting w ith
specific upper motor neuronal systems regulating
36
~!p
CIIl!N
".......:'
specific typ es of motor activities. (Bottom) The
major intern al subd ivisions of the cerebellum are
show n. The cerebellar cor tex (3-layeredl, the outer
2.one, is infolded to form numerous folia. The wh ite
matter, carryin g affe rents and efferents assoc iated
with the cerebellar cortex, is deep to th e foli a. The
deep cerebella r nuclei are deep to the w hite mat
ter. These cell groups receive most of the output
from the cerebell ar cortex via Purkinje cell axon
projections and receive coJiaterals from mossy fiber
and cl imbing fiber inputs to the cerebellu m. The
cerebellar peduncles are interior lo the deep nuclei;
these massive fiber bundles interconnect the cere
bellum w ith the brain stem and the thalamus .
 ;'pinal Cord OVERVIEW OF THE NERVOUS SYSTEM

An terior view l eft latera l view Posterior view

, :2J
Axis [( .-="

TI2 ---'~

13 - J:;

Saoum (51-5)

e
ter
li te
,d
he FIGURE 1.36: SPINAL COLUMN: BONY ANATOMY _ _ _ _ _ _ _ _ _ _ _ _ _ __
,t excruciating, radiating pain if dorsal roots are
Anterio r, latera" and posterio r views o f the bony
t involved and loss of mo tor contro l of affected
spinal column show the relationships of the inter
vertebral di scs w ith the vertebral body. The di scs' muscles if ventral roots are invo lved. In the adult,
iber th e spin (l l co rd ex tends caudally onl y as far as
prox imi ty to th e interve rtebral foramina provides
an an ato mical subs trate for understandin g th e pos the L1 vertebral body, leaving the lumbar cistern
:Iei; sible impingement of a hern iated nucl eus pulposus (subarachno id [SA] space) accessibl e fo r w ith
re- drawal of CSF.
on spina l roots. Such impin gement ca n cause
37
OVERVIEW OF THE NERVOUS SYSTEM Spinal Cord

CI
1st cervica l n.
Cervical enlargement
C7
8th cervical n.
TI
hllhoracic n.
Spinal d ura mater
Fila ments of
nerve fool

Conus l1ledullans
Ca uda equina
[5
'-.. Srn lum b~r n.
51
1st sacral n.
Filum terminate

~~~
:;th Scleral n.
CoccVg<l1n.

11!!N Coccy)(
.....
~ ,.,' : :

FIG URE 1.37: SPINAL CORD: GROSS ANATOMY IN SITU _ _ _ _ _ _ _ _ _ _ _ __ FIGU

The posterior portio n or the vertebrae is removed
to show the postefior (dorsal) surface of the spinal
co rd. Ce rvical and lum bosacral enlargem e nts re llect
innervation of the limbs. The spina! cord extends
rostrally through the foramen magnum, continu ous
\\ it h the medulla. The conus medullaris is located
under the L1 verte bral body. The longitudinal
gro\vth of the spinal column exceeds that of the
spinal cord, causing the spinal cord to end consider-
38
ably more rostral in an adult than it does in a new Th e u
born. The associated nerve roots traverse a consid reflec!
erable distance through th e SA space of the lumbar tour c
cistern to reach the appropriate intervertebral extenl
foramina of exit. In the lumbar ci stern, this collec overl)
tion o f nerve rools is the cauda equina (horse's tail). memt
Th e lumbar cistern is a large reservoir from whi ch nerve
( SF can be withdrawn. The filum terminale anchors struct l
the spi nal cord caudally to the coccyx.
_ Spinal C~I Cord OVERVIEW OF THE NERVOUS SYSTEM

Posterior view

Rami communicante5-_ _ _ _i~::;;;

Docsa! rool - -- - - -,

DorSdllOo t (spinal) ganglion

Afachnoid - - - - -

Mesothelial septom in

posterior mediilo ,~',:~~~~:::=-~l

Pia ma Ier love rlying ~na l

Filaments of dors al root - - - - - - --;--:;

Denticulate li gament _ _ _ __ __ +""""'"

Anterior view

Lateral fun iculus

Filaments of dors al rOOl

.3l~"",--- While malter

Dorsal roo t
Dorsal rool
(spin <II) ganglion

I nerve
Ven ltal (001
Filaments of
vent rill root
""--C,---_ An terior median
iissI,lIe
Anterior funiculus

FIGURE 1.38: THE SPINAL CORD, ITS MENINGES, AND SPINAL ROOTS _ _ _ _ _ _ __
lew The upper illu stration shows both intact and The posterior spina l arteries supply the dorsa l
Insid reflecteq menin ges. The pia ad heres to every CO Il spinal cord with bl ood and run ju st medial to th e
mbJ r tour of the spinal co rd surfa ce. The arachnoid dorsal root entry zon e. The lower illu stration shows
extends over th ese contours and ad heres to the the spinal cord with the menin ges stripped away.
~c- overlyin g dura, a very to ugh, fi brous, protective Both the dorsal and th e ve ntral roots co nsist of a
tail). membrane. These meninges extend outvvard to the convergence of rootlets that provide a continuous
ch nerve roots. The denticulate li gaments are fibrous dorsal and ventral array of roo tl ets thro ughout the
hors structures that help anchor th e spinal cord in place. spinal cord.

39
OVERVIEW OF THE NERVOUS SYSTEM Spinal Cord

Sl'ction throogh thoracic vertebra

Dura mater
SrmP<l thetic ganglion
Ve ntral

space
Pia mater
Dor~JI rOOI (adherent to
fsp lnal) sanghon spinal cord)

Spinal nervt.' ' -

Ventral ramus

(intercos tal
nervel_

Section throu gh

lumbar vertebra

Sympathe ti c ga nglion
Ramus r.ommunicans
-" Vf>nlr...1rool

~\ -'" Spinal nerve

--:- r-......
~,~""::.-- Ventral r<lffiU5 (contribu te s
10 lumbar plexus)

~ Filum lelminale

Cauda equiniJ
~ ~p
" 11liN:\":.".'::

FIGURE 1.39: SPINAL CORD: CROSS-SECTIONAL ANATOMY IN SITU _ _ _ _ _ _ _ __ AGt

(Top) The spinal co rd in th e sp inal canal is sur entire spin al cord. Th e sympath etic chai n ganglia -opl
round ed by meninges. Dorsa l and ventral roots (paravertebral), important for fi ght-or-fli ght - ;

course through the intervertebral foramina. The
epidural space, with its associated fat. is sometimes
used for infusion of anesthetics. Arteri es and veins
are associated w ith the spinal nerves and nerve
roots. Some segmental arteries provide anasto
motic chan nels for blood flow from the aorta to
augm ent flow from the anterior and posterior
spinal arterial systems, w hich ca nnot sustain the
40
responses, lie adjacent to the vertebral body (ven i"""dy
trally). The dorsal and ven tral rami of the sp inal
nerves distribute to specific regions. The sp inous '\'
process extends dorsally, w here it can be palpated caud
by physical exa m. (Bottom) The SA space of a lum
bar vertebra, co ntaining the filum termin ale and the desc,
roots of th e cauda equin a, is shown. i-St5
~'"
- Spina' C...I Cord OVERVIEW OF THE NERVOUS SYSTEM

Sections through spinal cord at various levels

, ',. ,
C5 "c'Y""
id -, ,

T2
.-( , I.,
";.;,/. I,
"
.,:J
"

.g \,
"'-'1 .j 4~'K
flllOid
'~

J , . "1<7'
~>'<' '~"Z""'~';W: "
)
!1er
~
/ '" '. I . .

~
~'-~ j~
/
ent 10
:0,(/)
~
- ' - - -/ "
,
.._-'" /'

11

Principal fiber tracts of spinal cord (composite)

E$ Ascending pathwa ys
_ Desce nding pathways
m.~ Fibers passing in both directions

Fasckulu$ ,;" , ililll , _ _ Anterior white commi ssure

r
Lateral (crossed) co rti cospina l
Fasciculus '"'''''' ' - _ _
(pyramidal) trac t
Dorsolateral fasciculus
(of LI",w~I-,

Pos terior
La teriJ l (medu ll ary)
II
reticu lospinat tr ac t

Spinothalam ic Ira ct and Anterior o r medial (pontine)

Spillorelicul ar Iract leliculospinal tract

Anterio r (vemral)

Tectospinal trac t
Spinoolivary tract
_ _ ""'", I ,,, {uncrO$>ed}
Fasciculus p,op'I''' ~~ COllicospinal lracl

19lia
FIGU RE lAO: SPINAL CORD WHITE AND GRAY MAITER _ _ _ _ _ _ _ _ _ _ _ __
Top) Seven spinal cord levels are shown, depicting (lateral horn) wh ere pregangli onic sympathetic
the relative size and th e vari ability in th e amount of neurons reside. Th e white matter is subdivided into
(ven gray matter at each level. Levels associated with dorsal, lateral, and ventral funiculi, each containing
,I the limbs have greater amo unts of gray matter. multiple tracts (fasciculi, bundl es ). The tracts con
JUS W hite matter increases in absolute amo unt from veying pain and temperature information travel
'i !ted caudal to rostral, reflectin g th e level-by-level addi rostrally in the antero lateral funi culus, the spino
lum_ tion of ascendin g tra cts and the term inati on of thalami c/sp in o reticular system. Fin e discriminative
d tile descending tracts. (Bo ttom ) The gray matter con sensation is conveyed throu gh the do rsal funiculus.
sists of dorsal and ventral horns and, in the Tl and Dorsal roo t entry zones and ventral root exit zon es
L2 segments, an intermediola teral cell column are present at each cross-sectional level.
41
OVERVIEW OF THE NERVOUS SYSTEM Me ninges

S<.pt.'rioJ ~ilt'll~1 (pldu",! Sj).)ce ( l'OlCn h~1l

Ar,l rhnn,cI ~(''''u l.llion

inQ('fIling s~ull
(f(Jvw1a)

V~ I l() U '
l a~u l1J
Suba r~ chn o id
'race

Ink'rioI
Middle
>inu5 meninge1

~~
"'lef)' ~n d
ve in

" L1cuna

'> .,,: 1" ";: ; -/;~- :.. ....,"' . ,..

".-~. <"", 'f"

. . . . . . ~.
,. " . __ r. r

,, " "" "u >,, ," r ,'" . Y.""; ~

, , ' ....
:::::::.==- ____~____ _.. ~~w
. , ....
r
.~ ,... ",.," " " ' - - C~I\'a,ia
"'
. ; . ......

~," ,",,-' ,.:

"~;~
!1~ ,,' , ~~~'_~ ~ _
' " . "1~~~
",~ -r
V: .y{>:;_ ",. ..y:"..--- Duramater (ou rer .1nd ," "'" Iav....s)
1'Vf<..
I""i f{
'i7.
S
,,\~Yl\..
uf>durJl lp~Ce IPot~nliJI)
Arachnoid
1, ' , - - - Subarac hno id space
Pi;,. maIer

- -- - Celeb,')1hemiS(lhoe,e
~!?
,-II~

FIGURE 1.41: SCHEMATIC OF THE MENINGES AND THEIR RELATION SHIP TO THE BRAIN _ _
The meninges provide pro tection and support for to provide channels for venous blood, the venous
neural tissue in the eNS. The innermost mem (dural) sinuses. Th e arachn o id granu lations, o ne
brane, the pia mater adheres to eve ry co ntour of way valves, extend fro m the SA space into the
neural tissue, into sulci, fo lia, and o th er infoldin gs. It venous sinu ses, especially th e superior sagittal
adheres tightly to glial endfoot processes of astro sinus, all owin g (SF to drain into the veno us blood
cytes. The arachno id mater, a fine, lacy membrane back to the hea rt. Blo ckage (e.g., acute purulent
ex terna l to the pia, extends across the neural su lci meningiti s) can res ult in increased intracrania l p r~
and foldings. Th e subarachnoid (SA ) space sure. Bridging veins drain into the dural sinuses.
(between the arachnoid and the pia), into w hich These veins are subject to tearing as they enter tre
the CSF flows, provides buoya ncy and protectio n sinus as a result of trauma, especially if there is
to the brain. Arteries and veins run thro ugh the SA some atrophy in the brain, permi tting venous bloo
space to and from the eNS. The rupture o f an arter to accumulate in the subdural space as it dissect:;
ial aneurysm of cerebral arteri es resu lts in an SA the inn er dura from the arachnoid. This subdural
hemorrhage. The dura mater, usually adherent to hematoma can be life-threaten ing from increaseD
th e inn er arachnoid, is a to ugh protective outer intracranial pressure, edema, and the accum ul at:
membrane. It splits into 2 laye rs in so me locations of blood from the hematom a itself.
42
- Meningentricles and the Cerebrospinal Spinal Fluid OVERVIEW OF THE NERVOUS SYSTEM

Anterior (frontal) hom }

l. lateral
Inferior (tempor ~ l ) hom
v entricle

POSleflor locclpllc) l) hm n

Cp.rehral.1queduct (o i Sylvlu s)

4th ventri cle

l.la lelill apeflure

Ifora me n o f l usc hka J

_ - l. lateml rp.r.p. s~
fo rJmen lof Monro)
.1 M edian aperLurE:
d Ilor.lmen of M af\l'ndie)
"
Co' j)US ullOSl lm
R_I,lINal ve"tride
Body of caudate nudeu~

Suprapineal recess
Choroid plc)(u~ of loll e ra I ven tricl E'

I
Stri<l lermin<llis

i R. Ih~ l amostr iate VC II)

,; ;,
Bod\' uf furnix
.; -' . , - -,. "-'
Tela cnOr'OIde'l ()[ :lrd ,elltricie
~::::::::::::: Ch o roid plexu s (J I 3rd ventrl circ
:: Thal amu s

GP", f~'len_'I~d------ ------ -- fl

. O' , U5 P<' I u S
LelltifortYI
I
[in terndllil and ('xlem.~i [el segments) nue cu~
_ _- - Internal capsule

C-----
_ - - - 3rd vcnTricle
HYPolhal 3m us

~r-!2:;;:::~~~==:::::::::::::::: Tall
:::
0 1 c.ludiltP. nucleu~
O ptic trac l
_ - -- Cho roid ple xus of iJleral ventricl e
"'t-- ===== Inlellor (lemporal) hom of latc", 1ven rricl e

lenous
one
---
w--'--::
Fimbria 0 1hij>p0campU5
HillPOc.:J mpus
De ... tate g)'(U5
Suhirulum
he -'-- - - : -- - P.J ra hippo campal (.Ortex
fal ~ Piilmilt e r '(,,;'=:,----:,.,;- - - fl1 10rhlll JI CO"ex
blood,
FfGURE 1.42 : VENTRICULAR ANATOMY _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ __
lent
II pres The lateral ventricl es are (-shaped, reflectin g their then in to th e ce reb ral aquedu ct and th e fourth
es. dssociation w ith the develop in g telencep halon as it ventricl e. Blockage of tlow in the aqu edu ct can pre
'er the sweeps upward, back, and then dow n and forward cipitate int ernal hydrocephalu s, w il'h swelling of
is dS the temporal lo be. The positio n of the lateral the ventricles above. The escape sites for (SF to
blood ventricles in relatio n to the head and body of the flow into the subarachnoid (SA) space cisterns are
ecfs caudate nucleus is an important radio logical land the medial foramen o f M agendie and the la teral
Ira l mark in a va ri ety of co nditions (hydrocephalus, cau foramina o f Luschka, w here blocka ge of (S F tlow
sed date atroph y in Huntington's disease, and shifting can occur. Th e choroid pl exu s, extending into the
lation of the midlin e w ith a tumor). (SF fl ow s through the ventricl es, prod uces the CS F.
foram en of M o nro into the narrow third ventricle,
43

 -
OVERVIEW OF THE NERVOUS SYSTEM Ventricles and the Cerebrospinal Spinal Fluid

PO;;\f . io . "iew ~Hi venlnclC' Habenula. trigon!'

PUI\-jO,l'
l aterJI } Geniculate
Pine ,l l gl.lnd ' ....Iedial bodies
Supe rior colliculus
Media n sulru s
Tro ch lear (IV) nervl!---.. Locus (oeruleus area
Superior medullary velum ~-< ') r

Cere~lJar
_.

I~oondcs {"p
, ,o'd SJJ!.,..\,
C:y'l ~
Sui(us limitans
MIddle . or
Fold,,1c~hcu lus
Inll'rio!
VesTibulilr alea
La l!"....1

~ medull.n e --- ~ "

HyposiosS.l1nerve trigone ----w- ~
Inferior (ovea

Tubf'rcu luon cinE'reum

f- "-<J..:i<~
(spinal trac t oi tri8emin.,' nerve) ' Posterio r mt!di.m ~ulcus

Cunea te tuberd e / f---- F.1Kiculus Cuneatu~

V'l g>ll tfll!one l F.1sck u\us 8 .acill~

Me dian s.lgiuaJ sectio n

Body of fornix Ha benular co mmiSiu(e
~~p
ClliiN
\\':.':: :
ChorOid pll'xUS ()f 1m ventrlc.ie

Interventri cul ar fUfill11en Sple nium oi co rpu ~ ca llosum

. c."' 1.
(OIMOl'lro) ~
~reat cereUr.ll ve in (ofCillel1)
'r hil~ilmu~ ~
.Ct'rehral .lQuedu([ \oi Sy [viu~l
'\Il1. t:omm lssure - "'1'> .'lj

lemin<t It'rmillillis ;':' ;} , I '-" lgula

..

~
Post. co mmis~ure . '" _. ~ Central lubulf'

M~mmi ll arl' uody . . h-t Vermis

OpTiC chiasm
O culomotor lUi ) nerve - - - r

-
Supt'rior r.n lliculus
r Su p. medullar y velum
IIlI. medullary ve lum
Medial Iongi[udi n,)l iil!rOculus./
Choroid plexus of
4 [h vent ricle
4th velltrid e

"Tonsil
,1Ae<lia n dpt'rTur e lo f MaFlendie);' E
Pyramida l decu~~hOfl-TP .' ,;
\ ~
_
" -- --
""m;d }
Uvula
Nudule
Vermis

Cent ral cilflJ l c)/ ~plnil l cord ;. ~ , ;

FIGURE 1.43: ANATOMY OF TH E FOURTH VENTRI CLE _ _ _ _ _ _ _ _ _ _ _ _ __

FIGUI
The rho mbu s-shaped fourth ve ntricl e extends damaged from tumo rs in the region. The lateral
thro ugh the pons and the medulla. The foram in a of margins of the ventricl e are embraced by th e hu ge Cereb
\ \a gendie and luschka must remain pa tent for cerebellar peduncles interconnecting the cerebel late-ra
a s s~
proper flow of CSF into the cisterns. Bilatera lly sym lum with the brain stem and the diencephalon.
metrical pro trusions, depreSSions, and sulci on th e These relationships are important fo r interpreting cou ld
floo r of the ve ntricle defi ne th e und erlyin g anatomy imaging studies in the co mpact brain stem region s, In erea:
01 bra in stem regions. Vital brai n stem centers for w here diagnosis of tumors and vascular lesio ns is fo urth
p rOVIC
cardiovascular, respirato ry, and metabolic fu nc challenging.
tions just below the floor of the ventricle ca n be under:
cisterr
w irhd .
lion o
44
 'Iuid
Ventricles and the Cerebrospinal Spinal Fluid OVERVIEW OF THE NERVOUS SYSTEM

- 5"",,'M""".11 SInus
Dura

I ;

Choroid "Ic,", ,' fl'" "," ",'01.,1

Interpeduncu lM cist ern
N
", : Cereb ral J(jueduct (01 Sylviu;)

La!!:'r,1 ] "pefl llfe (f(> ramcn nf l"",h"'i /' Cistern of g r~ ,lt

ChorOId plt>l<u, oi 4t h "","01,, / ' c:erebralll('in
Dura m",, -
Cerelwlloml:'dullJrv
d, l", rn (cister na m,lgna)
5uhar acnnoid ,pact'
(foramen 01 Molgendit>l

, GURE 1.44: CIRCULATION OF THE CEREBROSPINAL flUID _ _ _ _ _ _ _ _ _ _ __

11
huge r'erebrospinal fl uid now ing thro ugh the ventricles
bel- .uteral to third to ce reb ral aq ueduct to fourth )
~5 es through several point s w here obstructi on
I.
~{J d preci pitate internal hyd rocephalus a nd
ing
~i on s,
ea sed in tracrani al pressure. (SF flow from the
IS IS
'ih ven tricle into the ci sterns of th e SA space
"ides externa l cushioning to prevent injury to
'""'Iderlying eNS ti ssue from minor trauma. Some
-e'r ns (e.g., th e lumbar cistern) prov ide sites for
rxh~ wa l o f CSF (lumbar puncture). The absorp
0 : (S F from the SA space into th e venous
draina ge through the arachnoid granu lations is
dri ven by flow through th ese o ne-way va lves.
Prod ucti on, flow, and absorption of (S F mu st be
in precise balance; disru p tion of this process results
in e xternal hydrocephalus. Flow of the (S F in the
ve ntricl es also can act as a fluid-deli ve ry system
(or dow nstream intluences o f speci fic media tors
(e.g., prostaglandins, interleu kins) and is an internal
paracrin e comm unica tion channel for som e struc
tures close to the ventricles.
45
OVERVIEW OF THE NERVOUS SYSTEM Vasculature

~
. _.' Alilchnoidg, anuJation$
....""~#'s-..r..;.~
r~H i eta ll~,r.J ,In c! fron tal (an t,) . - ____'-,.---
~ t/ ~ ~ Openingoi
br anches of minnie meningeal a.
superior cerebral v.
V('nou~ r..cu na
Middle men inge~I.l.

Ant erior menin&e~J a.

SupPlior
(i/om anterior ethmoidal sag;",,1sinus
D ut ~ mater

MaslOid branch
(Ii occipital a .

A nterior and posterior ml'ningeal

brilllche!> o f vertebral ...
Milstoid branch
il

Mcnin i\fol l branches of asc tmd ing pnaryngeOiI ,1. t,$.-

Temorral branch (cu t) ,lod dor ~1 meningeal branch or
rru:,ningohyp op h)'St'-allrunk

Midcile alld .Jcccssory meningeal3<]. --l,!

Meningenl b ran ch Of ~osteflor ethmoidal il .

AI)le rior n)e nlnge;;tl a.llm m anll'rim Clhmoidal aJ --li!

\I;,
Intem..,1 (,lIOUd a. and Its nlenin ~oh)'pophystc'al trun k (in phanto m)
Mi dd l ~ m~nln 8ea l a.
Accessory memnge,ll a.~
Sup l2'rfici;:tI tem "orai a.

~~ft%
Po steri o r aUricular .1.

hternal carotid a. --!: c lIQN

: .......,' :1

FIGURE 1.45: MENINGEAL ARTERIES: RElATIONSHIP TO SKULL AND DURA _ _ _ _ _ __

Meningeal arteries are found in the outer pOrlion
of the d UI'a an d supply it with blood. They also help
supply adja cent skull and have some anastomoses
vvith cerebra l arteries. The skull has grooves o r sulci
for the men in gedi vessels. This relatio nship reflects
an important fu nctional consequence o f skull frac
tures. Fractures can rip a m en ingeal artery (usually
the middle m eningeal artery) and allow arterial
blood to accumulate above the dura. Su ch an
epidural hemato ma is a space-occupying mass, and
it can produce increa sed intracrani al pressure and
risk of herniation of the brain, particularly across
the tentorium cerebelli . Even very fine fra ctures can ,
have this dangero us co nsequence.

46
 ture Vasculature OVERVIEW OF THE NERVOUS SYSTEM

R. and I. mi dd le cerebral ail.

L mi ddle meningeal a.
R. MId I. anl cerebral iIiI.
:una R. and I. posl
I.:erebral aa. _-<..." AnL COmmumGlling iI.

" ~i n u s R. and I. sup.

cerebell cl r aa.
R. ophthalmic i1.

mdter
Ba.ilar a. - -"4 cC) ll1munica tin g a.

Masloid branch of Cavernous sinus

L. Int. auditory
(labyrinthi ne) , - -"
Post. meningeal branch
of I. ascendi ng
pharyngCill
eoch
,
R. superficial
t emporalJ.

igeal

hra nch c--_ _ _ _ _ Carotid body

lai d.
R. sup. larynlZe" la.

R. asce nding phary ngea l Tlwroid <.iIrli lagc

~ . carotid sinu s R. cornmon c;;lrotid a.
R. int'. thy roid a.
Transverse process of C6 It inL Ihoraeie a.
K deep cervical a. Brachioceptl<llic trunk
R. thyrocervical trunk
R. costocervi c<lllrunk

~
/

' ImN
...-:;:
' ~'

FIGURE 1.46: ARTERIAL SUPPLY TO THE BRAIN ANO THE MENINGES

_ _ _ _ __ __ _
II The internal carotid and vertebral arteries ascend
thro ugh the neck and enter the skull to supply the
s, and brain w ith blood. The tortuous bends and sites of
and branching of th ese arteries (such as the bifurca tion
)55 of th e common carotid artery into the interna l and
~S can the external carotids) produce turbulent blood
Ilow, and atherosclerosis can occur at these loca
tions. The bifurcation of the common caro tid is
pa rti cu larly vu lnerable to plaqu e form ation and
occlUSion, threatening the major anterior part of
the brain w ith ischemia, w hich w o uld result in a
"stroke." Studi es of blood flow through these
arteri es are important diagn osti c tools. M agn etic
reso nance arteri ography (MRA) and Doppl er flow
studies have replaced the older dye studies for
cerebral angiography, for most purposes.
47
OVERVIEW OF THE NERVOUS SYSTEM Vasculature Vas.1

Anterior (ommuoic<lling artery, ~

Anterior cerebral artery,

Recurrent a. (of Heubnerh,.
Circle of
Willis '.,
Medial an d lateral le nticulostriat e aa.

Middl e cereb ral ,

lateral olbitofron tal .... ~
Ascendin g Irontal (unde labra} branch--....J "'I
"1
."

Posterior cerebral artery - - -~

Superior cerebellar
"
Basilar a . and ponline b ra nches
Internal auditory (labyrinth ine)
..,",
""
~

Vertebral iI . - ~~
A':'.s.;,~

Posterior inferior c er~ bellar 01 .-- ~. ~

PoslenOr5P1nala. ~:~, Y\ ;

\ ____ An terio r co mmunicatin g artery

l,.--- Anteriol cerebral ilt1ery

~ Middle cerebral artery

~
'1.. , POS teri.orcomm.u niC<l tin gart ery
~ - Ante nOf chorOId al a.
p-~
. Optic Ifact
,.:~,1

peduncle

,- moo-ot choroidal a.
Poste rior lateral choro id al a .
r--
;f~f'
Choroid plexu s o f late ral ventricle
- - MediClI genicula te b ody
~,.,...,.

~ Lateral ventricle c ,iC~

........"

FI GU RE 1.47: ARTERIAL D I STRIBUTION TO THE BRAIN: BASAL VIEW _ _ _ _ _ _ _ __ Gtcl

Top) The anterior circulation (mid dle and anterior tion of both sides, but it is usu ally not sufficiently
cereb ral arteri es [MeA, ACAJ) and the posterior patent to allow byp ass of an occluded zo ne. ~e co
circula tion (the vertebrobasilar system and its end (Bottom) The circle of Willis and the course of the -'00
.
oranch, the posterior ce rebral artery [PCA]) are ch oro idal arteries are shown. Th e arteries supplying '<ICe
_: e~.aJ
;.hown. The right tempo ral pole is removed to the brain are end arte ries and do not have suffi
.,., COl
s.ho\\ the course of the MeA through the lateral cient anastomotic channels w ith other arteries to
rIss...; re. The circle of Willis (the paired A CAs, M CAs, su stain blood flow. The occlusion of an artery sup t?'air.
- -ne
and PC-\s and the anterio r and th e 2 posterior plying a specific territory of the brain results in func
- i:"L1.
commun icati ng ar teries) appears to allow free flow tional damage that affec ts the perlormance of the
Of blood around the anterior and posterio r ci rcul a- structures deprived of adequate blood now. :.e e:t
-Kri n

~8
:culature \ as culature OVERVIEW OF THE NERVOUS SYSTEM
Frontal view with hemispheres se parated, tilted for a view of the ventral brain stem
Co rpus call05um Paracenlral a.
cleof
Willis Frontal br.lilc ht!~
lateral orbilorrOnl.!1d_
Ascending fro nlal
(candelabral hrandl
Front()poiJ. r ~ .
Ant '~,d':;:~h''',
parietal
Anterior cerebral arteries

P r ecent r~1 (prcrolandic)

~nd cent r~1
Medial orhitoironl<ll "

Ru rr en l a. (oi Heuhnerl

Tempural branches
In t. cmotlO~ .

Middle cerebral
artery and hranche s, Ant. cho ro id<:l l <:I.

deep in laterOlI _ _ -.--j~7J)~

ce re hral (sylvia n) fissur e Posterior cerebral arte ry
AnI. commu nicating

- - '"' audilury tlabyrin rhine) .,.

An I. Inf. cerebellar il. ' -- - Po,t. mi. cerebellar a.

PO, I. spinal a. - -- _-"" - - -- An I. s.p in al ~.

Coronal section through the head of the caudate nucleus

Fro ntal horn of 1",,,,,,, ",.,""",
Co rp.., s striatum
(head o1 lhe ca..,dale aM P""~'")' Callosomargina l aa.
dlld
Medial and latera l '''' ;'"''''''0',.",
~. "",,>fcorpus callosum
Prece nuallprerola ndic l, cerllral (rola nd ie)

Inte rnal c~ p,ul (anlerior li mb)

and parielal b,"""h,, -~
lateral cerehrallsylvi an) 11,,,"
,,/1 ' -.;,.- 5epl\ lm pellucidum

Temperal b ,."d' .,,-~ :>:-~"...- ROSTl um of corp us callosum

'"'. <.;o,~ ,..-\ ~==:=:;;,

Recurrent <I. (oi Heuh ner)

Op li C chias m

-
Ily
FIGU RE 1.48: ARTERIAL DI STR IBUTION TO THE BRAIN: FRONTAL VIEW
A N D COR ONAL SECTION _ _ _ _ _ _ _ _ _ _ __ _ __ _ _ __

The course of the ACA along the midline reflects its
he blood supply to the zone of the sensory and m otor
jying cortices, which are associated with the contra
lateral'lower extremity; an ACA stroke thus affects
to the contralateral lowe r limb. Th e M eA co urses
up laterally and gives branches to th e entire convexity
[une of the hemisphere. End bran ch infarc ts of the MeA
he affect th e con tralateral upper extrem ity and, if on
the left, language functio n. More proximal infarcts
affecting the MeA d istribution to the internal
capsule cause full contralateral hemiplegia w ith
drooping of the contralateral lower face; this results
from damage to cortico spinal and o ther cortico
moto r fibers in the posterior limb and to co rti co
bulbar fibers in th e genu. Th e lenticulostriate
arteri es ("arteries of stroke" ) are thin branch es of
th e M eA that pen etrate into the basal gan glia
and the internal capsule regions in the forebrain.
A stro ke in this area produces the classic contra
lateral hemiplegia.
49
OVERVIEW OF THE NERVOUS SYSTEM Vasculature a

'."

An'erim r.erebJ,1arl ery

Middle cerebr al artery ~~( Antenor comnluniGlling <ltlery

j'c, te ri or c0l11muniC3!ing <lnery O pht ha lmic arte ry

Caroticotympanic bra nch of
~
.. , :;-.<..
Suprao rbi ta l artery
intern,; j caro tid arter~

Pu <;teri or cereb ral artery

:..-:e; n n Supr atrochlear art er y
l acrimal artery
:11
Dor!>al nasal arl ery

Middle meningeill drle' 1f-:1 Middle men in geal "fIery

An gular artery
M 3:.:illar ya rter y
f1. Ii:.. \. " Superficial te mporal arter~'
\ Pm terio r auricular afl ery
Anteri or inierior cerebellar " ' G... ./"\\.
FaCial artery
Posu= ,ior infe rior cerebella r artery / '
Occipital arte ry
lingual artery

Asce nding pharyngeill artery

Superior thyro id
5 ..#1 ... Anterro r ~pj nar artery
Co m mon c.lrOlid ar tery ~i
Spina l segmen ta l med ull ary branch es
Vert ebra l arter y
AKe ndin g cervical arl <, ry ,-
Com mon calO ti d artery
Inferi!)r ,11yro icl art ery ,

Df!ep cer vical arte ry

ThylOcervicaJ trun k

Transve rse ce rvical artery .ti::,:...d

Suprasca pul ar artery
,,"""
.:c~ Supr<, m(' in tercos tal arte ry

~ Co" o" ..,>1~"k

Subda~'ia n artery
Sutx iavian artelY
Internal thoraci c artery
Brac hioce phalic tr unk U 'it
Anastomoses
1 Rlghl-lelt
,,,h
~~p

2 Caroti d- Vertebr<ll
3 I/\Iemal c;arc)[id- EJ,: temal carotid
Aorta { Descending, ~_==:=:
Ascending - 4 Subcl avi an-Ca rotid
. ,.~ i Subdavjan-Vertebr<ll

FIG L RE 1.49: SCHEMATIC OF ARTERIES TO THE BRAIN _ _ _ _ _ _ _ __ __ _ __

Thi s schematic diagram shows the entire layo ut of separation of the anterior (M e A, ACA) and poste
"2 01
me an erial blood supply to the brain, including rior (vertebro-basilar, PCA) circulation are evident

anasto moses. The ci rcle of Willis is present in the in this diagram.

uoper central po rti o n of this schematic. Th e relative

en
011
30
 u

'asculafure \ asculature OVERVIEW OF THE NERVOUS SYSTEM

Vessels dissecloo Qut: infe rior view

AMerior cerebral artery (Az segmenO _--1 1 u".....,-- Medial striate ilTlery (recurrent ':lr1ery 01 He ubne r)

M teromedial centrilil perioraling) arteries

Anterior ce rebral art ery (AI "" m,,"') - --;;;;;ft.p.( Hypo th alamic artery

Ophthalmic artery ----I~f:~' Ii_-<~''''''O'''''''' " ",,,1 ilenticulost ri ale) a,rerit's

Internal carotid ", n..y --::J;;:;;;;~~f<
Middle cerebral arl ery

Posterior WM'""I''''"g"',,,y ---!J-....l~.-:. Superior hypop hyse~1 artery

y rosterio r cere brill il rlt':y Inferior hypophyseal art ery

I segment) AnleriO( choroid,,1 artery

, "h,)"m,,,u),oc,,I(,,,oc,,,,.. mllllll,,y: allery

SUllerio r cerebellilr ilrt ery ' P'"" ,"O,..."i, I ceollal (per'orating) artery
",,~ //'-;"'-~
Basilar Thabmoperlorallng artery

Pontine "',~I,, __..o::;;;~ Posteromedial centl al (pa,,}medi.m) arteries

anches Anterior inferior cerebellar illteo"V ~-- labyrilld'line (I ntemal acoustIC) a!tery

Verlebrill il rtery
~~-.A

Vesse ls in situ : infer ior view

Ante ri or ce rebral artery ---",:-11..11--,--:- Anterior co mmuni ca,in g ..rter y
Cavelno\ls sinus

Su perior hyl~"hy,,,') "" <Y -_~ Iniundibu lum (pituitary stalk) alld
lon~ hypophyseal portal veills
Middle celebral . , .."""
. :.-- Adef)ohypophy-;is
(anterior lobe of pituitary gland)
, (nierior hypophyseal artery
Neurohypop hY'lois Ipmterior lobe of pnuilary gland )
Poitt'llor communicatin8 art~
Efferen t hypophyseal \'eills 0;;;;0;,-.- _ Postelomedial central {perioriltingl arteries
"lal carotid
Posterior cerehral artery Superior ce rebell<lr arte ry

-~- Basilar artery

FIGURE 1.50: SCHEMATIC OF THE CIRCLE OF W ILLIS

_ _ _ _ _ _ _ _ _ _ _ _ __

)oste Th e ci rcl e of W illi s surround s th e optic tracts, the
ident pituitary stalk, and th e basal hypothalamus and is
the most frequent site of cerebral aneurysms. It
includes the 3 sets of paired cerebral arteries plus
the anterior communica ting artery interconnecting
the ACAs and th e posterior co mmunicatin g arteries
int erco nn ectin g th e MCAs and the peAs. The free
flow of arteria l bl ood th rough the communicating
arteri es is usually in sufficient to perfu se the brain
adeq uately in the fa ce of an occlu sio n to a major
cerebral artery. An aneurysm is a ballooning of an
artery resulting from an inherent weakness in the
arterial wall. A rupture can lead to a subarachnoid
(SA) bleed, with loss of perfusio n to vital CNS
territories.
51
OVERVIEW OF THE NERVOUS SYSTEM Vasculature \ ""

Laler.ll view
AnI. p<lrielat bril ll( h

Ce nlrill (ro Jandic l branch ,."

1
Precenlral (prerolandic) branch .' j;. ,1 ?" Termin(ll cortical branc/1...,
Ascending frontJI - ~"'~-.' I _~ oi I. po<;lenor ce rebral ;;l.
{cand ela b ra ) brJ nCh

Tr rrn ir'lil i co rti cal branch es

01 anI. cerebral ilil.

lateral orbitofronlal a.
Post ttom pora l
L. middle bra n che~

cerebral artf'ry~

L anterior An i. tem por~ 1

cerebral arlery ~ branches
A nt. communicating iI .

R. anferior cerebral arterY / '

l. internal carotid il. /

Medial view
Paracent ral iI.
PericalLosal i1.
Precuneal ...
[nl.lrantal { posterior
bril nches Mldrll
Ante,i R. posterior ~rebril l art ery

Post pericOIliosal a.
( J)1owmilrginilJa.

PJrietoocdplta l br~ n ch

Post. tem po ral bra nc h

R. anterior
cerebral artery ~~
All t. temporal h ranch

Medial ...... . ~ _ Ca lC J r101' b ran c h

O'h to iro nta l J . 'ted
~- . (ommuniQ ling a. ~

~~~
ReC\J rre n t ~. (of lil'u b ner )

R. in tern al cJro tid a. /

Post. c ommunic atin g a.
O '.~N

fl GIoRE 1.51: ARTERIAL DISTRIBUTION TO THE BRAIN: LATERAL AND MEDIAL VIEWS _ _ __ FlGL,
lOP The ,'v\ CA sends th e named branches alon g to the midline region o f the frontal and parietal -t:" SJ
the 5wJdace or the hemispheric convexity into the Jo bes. Occl usion disrupts sensory and motor
~ron:a: and parietal lobes and into th e anteri or and
~"'"
functions on the contralateral lower extremi ty. -,
middle regio ns of rhe temporal lobes. Occlusion The PCA distrib utes to the occipital lobe and the
disrupts 5ensor v and motor functions on the con inferior surface of the temporal lobe. Occlusion
trala:efal bodv, especially the upper extremity, or disrupts mainly visual functions from the contra
on the entire contralateral body if the internal lateral visual field.
capsule is aiiected . (Bo ttom) Th e ACA distributes
,-,
culature Vasculature OVERVIEW OF THE NERVOUS SYSTEM

branches
ebral .1.

II Anterior celebr..1,ule ry

II Mid dle ce re bral <!fte ry

II Posterior cerebral ilftery

>ranch

IIN
..".,'::
; \ '.';.': ::

- FIGURE 1.52: COLOR ILLUSTRATION OF TERRITORIES OF THE CEREBRAL ARTERIES _ _ __

Th e specifi c mid lin e and lateral territo ri es of di stri and make particularl y d ear the wa tershed zones
bution of th e ACA, M CA, and PCA illustrate the .
at the junctions of th e majo r cerebral arteries.
exclu sive zo nes supplied by th ese major arteries

53

OVERVIEW OF THE NERVOUS SYSTEM Vasculature Vase!

Femoro cel"('b ral Angiography

-
L>'.... I ... oj " o~ f.......t ~~

~".:..IJlo...l.

C.ilr> "", ,, , ~..,. I '

,;",1
A~:",c< .' It<Ul.od Lo,.....
o', . ~~ . I . k'fII;a,ro"".....,

, .....,,:,01

Op "" ''''''''

"',~ (in terhe1j

Heads d.
f iGU RE 1.5 3: ANGIOGRAPHIC ANATOMY Of THE INTERNAL CAROTID CIRCULATION _ __ Thala1

The left plate is an angiogram lateral view of the opaque contrast agent into the common carotid )a telal I~
in ternal ca ro tid arterial circulation after injection of artery. The majo r branches of this arterial system
a rad io-opaque contrast agent into th e internal are delineated. MR an giogra phy is used commonly
ca rotid artery. The major branches of th e internal to investigate the status of the cerebral arteries, but
carotid artery, particularly th e anterior cerebral and does not provide the same level of detail fo r
,
middle cere bral arteries, are delineated. The right anatomical purposes that standard angiography
j'):ale is an angiogram frontal view o f the internal provides.
carotid arteri al circulation afte r injecti on of a radio
Arteri es of Posterior en nia) Fossa
V~nehral AnV<>&r.I<M:,,~ ria! Ph~
,

fiGURE
The vertl
l ;:""'~O bfMlC~----.J ,CICoN the basil.
......, Ini. CCfeWI<o, . <>1" 1 PQ>I.inr = ct-..:l MI . 1 ~ '.'.-' : : extend if
ing w ed ~
FIGu RE 1.54: ANGIOGRAPHIC ANATOMY OF THE VERTEBROBASILAR SYSTEM _ _ _ _ __
can prod
The upper figures show angiogram s of both lateral circulatio n. The maj or branches of these arteries tralateral
and ironlal views of the vertebra-basilar (posterio r) are labeled. Angiograms with contrast agents pro age abm
circulatio n after injectio n of a radi o-o paque ca n vide more detailed anatomical info rmation, but ipsilatera
lra~t agen t (dye) into the vertebral artery. Th e major are mo re invasive than MR angiography. MR to m s. Thl
2Ilerial branches o j thi s arterial system are delin angiography is used w ith greater frequency, the largel
e2 red. The lower figure is an M R angiogram show although use of contrast agents still occurs for such as tI
ing both the in ternal carotid and vertebro-basil ar specific diagnostic purposes. (PI CA ), tf
54
ulature 3Sculature OVERVIEW OF THE NERVOUS SYSTE M

in,m::.r

Arteries of Posterior Cranial Fossa

Crur~ of fornix Right} (horoid plexuses of
Lateral and medial geniculate bodies of left ,h,l,c-,",_/" Left lateral ventricles

Septum pellucirlum
dnood,
Corpus callosu m , Post. horn of r. lateral ventricle
o~

~.<> oi
An!. cerebral aa.
Longitudinal
\ " Right and
le ft pulvinars
_<or"',"" '_,,(hemispheric) 11 . _ Splenium of corpus (~llosum
-(;o dds of . R, post. pericallosal a.
"'alamogeniculate aa. Parietooccipi ta l and
Medial and Calcarine branches of
ltid OL'erallentlculostriate aa. r, post. cerebral <I,
em Ant. choroidal 0 _ __ L. sup_ colliculus
nonly Ant. cerebral" ___ Sup_ vernlian .1_
Post, medial choroidal a.
'S, but Optic:.: (IIJ n. (to choroid plexus of
and ophthalmic a. 3rd ventricle)
Middle cerebral a.
hy Pos!. latera l choroidal a_
Thalamoperforating aa. L. post. cerebral J. with ant.
Post. communicating a. and pos t. temporal branches
Lint carotid a. Lateral marglllal branch of
sup. cerebella r a.
Sup. cerebellar a.
Inf. vermian a. (in phantom)
Basilar a.
Choroid al point and (huroidal a.
to 4th ventricle
Int. auditory (labyrinthine) il_ TonsHiohemispheric branches
Outline of 4th ventricle (broken lineJ
Post. meningeal hranch of vertebral a,
l. post. spinal a,
L vertebral a_ Ant. spinal J.

FIGU RE 1.55: VERTEBROBASILAR ARTERIAL SYSTEM

_ _ _ _ _ _ _ _ _ _ __ _ __

The vertebral arteries unite at the midline to form
the basilar artery. Medial penetrating branches
extend into medial zones of the brain stem, supply
ing wedge-like territories. Infarcts in these branches
can produce "alternating hemiplegias," with con
tralateral motor deficits (corticospinal system dam
'ro age above the decussation of the pyramids) and
ipsilateral brain stem/cranial nerve signs and symp
toms. The vertebral and basilar arteries give rise to
the larger short and long circumferential branches,
such as the posterior inferior cerebellar artery
(PICA), the anterior inferior cerebellar artery
(AICA), and the superior cerebellar artery (SeA).
Strokes in these arterial territories produce a con
stellation of ipsilateral brain stem sensory, motor,
and autonomic symptoms and contralateral
somatosensory symptoms. For example, a PICA
infarct results in loss of pain and temperature sensa
tion on the contralateral body and the ipsilateral
face. The end branch of the basilar artery is the
peA, distributing to the visual cortex and the
inferior temporal lobe. Occlusion results in contra
lateral hemianopia.
.55
OVERVIEW OF THE NERVO US SYSTEM Vasculature Vas.c-li

,. /.
~

-,,? J Hypolhill"mic vessels ( PefJQ

s...
'"! ;

W
~
l ong hypophyseal Supe rior hypophyseal a rlefy
portal veins \ (from internal caro tid A. or
}
Pos teri()( branch pest. com municating A.)
Shorl hyp0l-'hysea l
portal v e lr'l S .......
Art ery of trilbecula

Trabc<ul a

Effercot "eirt 10

Anterior lobe

Second ary plexus of

hypophysE'ill porlal syste m

L.l!e r.ll brimch / Efferent vein to

,nd c;wernOU$ iilluS

Q; Ca\'ernous sinus
Imenor hypoph yseal ;Jrteryl
Irom the Internal Cilroti d A.I
Inte rn al carotid allery
Posterior commu nica1ing artery
.) ~fJ!:
Superior hypoph \~e3 1 artery ft l'{fI

InieriQr nypoph\'seal arter,' / ~~pClmN

1\..:::

FIGL RE 1. 56: VASCULAR SUPPLY TO THE HYPOTHALA MUS AND THE PITUITARY GLAND _ _
<IGU R
TI-e supe rior hypophyseal arteries (from th e le A or
tt:e posterior communicating artery) supply th e
h\'pothalamus and th e infundibular stalk and anas
to"'10se w ith branches of the inferior hypo physea l
arte'"\ [irom rhe leA). A unique aspect of this arter
ial distribu tio n is the hypophyseal portal system,
\\ ho~e pri mary plexus derives from small arterio les
arrd capillaries that then send branches into th e
~~erio r pitui ta ry gland. Thi s plexus allows neu ro ns
that produce hypothalamic releasing and inhibito ry
56
facto rs to secrete th ese facto rs into the hypo , su~
ph yseal po rtal system, w hich delivers a very hi gh ""ece;\e
concentratio n directly into the secondary plexus in --cludir
the an terior pi tuitary. Thus, anterior pituitary cells -omtfl;
are bathed in releasin g and inhibitory factors in food il
very high co ncentrati on. Thi s private vascular co m .:;.load (I
muni cation channel allows th e hypothalamus to and ou.
exert fi ne control, both directly and throu gh feed
ba ck, over the secreti on of an terior pituitary
hormones.
:ulature ta.sculature OVERVIEW OF THE NERVOUS SYSTEM

Calva ria Arachnoid granula tion s"p,""m sagittal sinus Emissary v.

Skin

- lobe

Falx ce rebri Diploic v.

Epidural space (potential) Pia m aIer

Superior Cf'febral v.

Subdural space Cerebral

Arachnoid
Subarachnoid space

- FIGURE 1.5 7: MENINGES AND SUPERFICIAL CEREBRAL VEINS _ _ _ _ _ _ _ _ _ __

The superi o r sagittal sinus and other dural sinuses
igh receive venous blood from a variety of veins,
.us in including superficial cerebral veins draining blood
ells irom the cor tical surface, meningeal veins draining
n blood from the meninges, diploi c veins draining
:om blood from channel s loca ted between the inner
to and o uter tables of the ca lvaria, and emissary vein s,
ed
which link the venous sinuses and diploic veins
with veins on the surface of the sku ll. These chan
nels do not have valves and permit free communi
cation between these venous systems and the
venous sinuses. This is a Significant facto r in the
possib le spread of infectio ns from foci outside the
cranium to the venous sin uses.

"

OVERVIEW OF THE NERVOUS SYSTEM Vasculalure Vas(

Cerebra l v. penetrating subdura l space 10 ~ ntet sinus

Scalp, skull, meningeal / (bridglng "eim)
;md cerebral blood vessels
gr<lOlllalion

ioll

Fronw l and 1),lfIelal tribUIJ:rit!s

of superrldal temporal v, -........

r rontal ilnd parietal branches

of supertlci,ll temporal a.
ArachnOid gra"ula(;on Middle meningeal
inden tin g skuliliove~i a. and \"
Ve nom lacuna
Deep middle and
~ upe l fi cjal
lemporal
Thalamos\rlilte o1a. and vv.
imd internill
cerebral vv_ ,"d

Ct'rt!bral vv.

""
Tenl!

Diploic and emiSYry veins of skull

Parietal emissary v.
Fron tal d iploic v_ ~
Po~l erior temporal diplok: ....

Occipital emiSSilry v,
Anterior
O ccipita l diploic v.
temporal ~
diplojc ,, ~ ~1 til
M~IOid emiSMry v.

~~p
11!!Nl\': :.'::

FIGURE 1.58: VEINS: SUPERFICIAL CEREBRAL, MENINGEAL, DIPLOIC, AND EMISSARY _ __ FIG UI
Venous blood drains from the skull, th e meninges, able to infectio ns and co ntamin ation from drainage The ia
and the cerebral cortex into the superio r sagittal from superficial venous networks into the central slons (
si nus an d other dural sinuses. This area is vulner venous sinus channels. (onfin
of the
nor sa
thedu
deep r
jugula
the sIr
into th
58

iasculatu n: ulalure OVERVIEW OF THE NERVOUS SYSTEM

Super ior ophthalmic v.

; In"''''''''',I"' (circular) sinus and pi tuit.l ry gla nd
Basilar plexus
Internal caro tid 3.

d space

:e meningeal
11. and,_
) middle dnd
superficial
temporal
aa. and 11\

I Deep ,md
superfiCial n.
middle
Jugular foramen
('rebral V\.
Superior

::::'ft::-- Sigmoid sinu~

Accessor y (XI) n .
and inferior
e.:..:~7-'-_ HypoglosS<ll ( XII ) n.
~trosal sinuses
Trdnsverse sinus

~'Ll:..:.-- O pening of an inierio r cerebrdl v.

Confl uence of ; '''''.; - - - '

Superio r sagina l si nus ---"::::'''::::::':::':''':_--<

Infe ri or sagittal ; """', --:t

Greal ce rebral v. ("f Galenl

Intercallet'nous sinus - - ..j

SuperIo r pelrosa l ~nus
Sllilight sinus
Inierior petros.al "~"~""" - -::'1f!
Sigmoid sinus --"""~

Ocdpital sinus

FIGU RE 1.59: VENOUS SINUSES _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ __

Image The falx cerebri and th e tentori um ce rebelli, protru
tra l sions of fused inner and ou ter dural membranes,
co nfine the ante ri or, mi ddle, and posteri o r fossae
of th e skull. O uter (s uperio r sagittal) and inner (infe
rior sagittal) ve nous chann els found in split layers of
the dura drain blood (rom the superficial and the
deep regions o f th e e NS, respectively, into the
ju gular vein s. The grea t cere bral vein of Galen and
th e straight sinu s merge w ith th e tran sverse sinus
into th e co nfluence of sinuses to drain the dee p,
poste rior regio ns of the eNS. Infection can be
introduced into the cereb ral circulation through
these sin uses. Ve nous sinLIS thro mbosis can ca use
stasis (a backu p of the ve no us pressure), w hich
ca uses in adeq uate perfusion of the regions w here
drain age should occ ur. Th e pro trusions of d ura are
to ugh, rigid mem branes through w hi ch porti ons of
the brain can herniate w hen intracranial pres~u re
in creases.
59
OVERVIEW OF THE NERVOUS SYSTEM Vascu lature 1

.,.,. ~ '~:Jl- longilUdi n<l lli ssurt'

',. , ~, AntE"=OJ (erern-al vv.
E:" 'J\ Rf)5irUm oi ( 01PUS callosum
5~plUm pelluddum
leriOl septal v.
He.ld of c<luddt.:> nucleus
t erminal (caudate) v,

Inlen'entricular foramen (or ,'-'\onro)

Colum ns 0 1 iorn ix
Thalamoo;:tri.1te v.
Superior choroidal \'. and choroid
plexus oi lalera l venllicie
,I
Tela (horoidea oi Jrd venl ricle
Direct I.ltera!,'
Posterior lemlinal (caudate) v.
ternill ceteoral " " .
Sasal 'to (of Rosen l h ",!)
G reat cerebral o:. loi Ga len)
Inf~i nr ~a:; itt,'1 s irlU';
ihl 5i
l entnrium ce rebelli
,~
"'''r

~ ~!
Tran sverse :;inus

~;@N
Cunllu enc:e of ~inu s('s
Superior sagittal sinus
, ':;, ...

Dissection from above Uncal ...,

Ant criOI cerebral v.

Superficial m iddl~ ce.tc'bral v.
(draining to sphenopariet.ll sinus),
Infer ior
cerebra
~

Ba!>dl v. {of ROSt>fllhal,

Med ial genicu lJle bony

Splenium of

corpus cJlio!oum

Great cerebral 1/. (of Galt'fl l

Interior anastomOlic v.

(of Labb e)

Dissection from below

FI GURE 1.60: DEEP VENOUS DRAINAGE OF THE BRAIN _ _ __ _ _ __ _ _ _ __ FI'

(Top) This superior view of the thalamu s and the the brain stem removed illustra tes the drainage of Su
basal ganglia reveals the venous draina ge of forebrain and mesencephalic veno us blood into bIc
deeper forebrain regions into the posterior venous the great cerebral vein of Galen, heading toward g"
sinuses. (Bottom) Thi s basal view of the brain w ith the straight si nu s. dr.
60
:ulature Vasculature OVERVIEW OF THE NERVOUS SYSTEM

Subependym a l Veins ~ V('i n, u n I"'N, I w all 0 ' """' n r.l~

V",n; on me,li,-rI .... ,, 11 J"d flo o ' <.i f v "')l ric l ~
--' ~;r"
=---;z All 'llhlOf , -ei",

roM lcrm,nal fClurl~I"')' (po.t p~" of IhJf.l1llosvial. ~_ ,

In L cc r~ Or" 1 ,,_
,,,f
G<!n u <> 1
corpus ~
C3110Sllm />'
'\
~fe" (",-cb r. ,I"
foIGa "",' :'

D""'I> middle cereb, ~I ,.

In/. lhaL''''OS1r h)lf' ""_
/
B;o,.a IY.lui ,,'_"","
In!. hom o f l" lcd
(,-,r~he<l u m

4th """" ,,<. 1..

Carotid Angiogra ms; Venous Phase

(Su bependyma l an d sup erfi cial vein. opac ifi ed) Sup_ s.~li:jl!~1 sonu,;
\~" . ',e..,",,' IWUd.-.lf') , _

- L;Uer~ 1 projectio n

Fro nta l p roj e<lio n _

Int.IUgut..., ".

[;3<;.,1 v '""~~,""

FIGURE 1.61 : DEEP VENOUS DRAINAGE OF THE BRAIN: RELATIONSHIP TO VENTRiClES

Ige of Subependymal regions of the e NS drain veno us this regio n causes a blockage of drainage and a
into bloo d into th e inferior sagittal sinu s (left) o r the backup of pe rfu sio n, w ith resultant ischemi a oi the
" ard great cerebral ve jn of Galen (right). bo th of wh ich tissue in th e re gion s of drainage.
drain into the straight sinus. O cclusio n o f a vein in
61
OVERVIEW OF THE NERVOUS SYSTEM Vasculature

Veins of Posterior Cranial Fossa P"rt, of c"r~bdillm

L lingula TU tub.>r
CL centTdllobult, P ~yrilmld
l. Slip ilnd inL callicuti ~eft putvinar C culm"" U uvula
o d".cJlv~ N nodule
Right thalamus folilJm T ron,il
Int. cerebral n,
Medial geniculdte Splenium of (orp<JS callosum
Lateral mesencephalic v. Great cerebral v. (of Galen)
Inf. sagittal sinus
}{ ~
\i, . ' _ .,:, -:> ,
Ii
Inf. th~l,lnlostria'e VV.
V~.', ~- /
>
-;7
/'
~
~- .;";';;'1'<"
--"'
.
..
~- ~;: ;' i!i"-,:,::/ ~;- Linconst'lnt)
<-}, / "\:.,
,:.,v;,.'$B .;; .' .' ,." Sup.
vermi~n v.
Ant. cerebral v.
Straight
Optic (tl) 11.
sinus
Falx cerebri
Deep middle
Sup. sJgittal
cerebrcll v.
,inus
Tentorium
Anl. ponto cerehelli (cut)
mesenceph;

Preculminate v.

of sinuses
rOetros~1 v.
L transverse sinus
(dlJilling to sup.
pelrosal sinus) i. vermian v.
\ ~,llx cerebclli (cut)
nti
and occipital sirius
Vestibulocochlear (Villi
~~ " Int. cerebellar hemlsphcricvv.
r-acial (VII) v.
., Precentral v.
Ant. medullary , L.laterJI nrJchial v.

~~~
Vein of laterJI recess of 4th ventricle
Sup., middle and mt. cerebellar peduncle~
4th ventricle
Ant. spinal v.' . spinal v.
IIiiN :W'-.': :

FIGURE 1.62: VENOUS DRAINAGE OF THE BRAIN STEM AND THE CEREBElLUM _ _ _ __
The venous drainage of the cerebellum and the
brain stem is anatomically diverse. The veins of the
posterior fossa drain the cerebellum and the brain
stem. The superior group drains the superior cere-
bellum and the upper brain stem posteriorly into
the great cerebral vein of Galen and the straight
sinus or laterally into the transverse and superior
petrosal sinuses. The anterior, or petrosal, group
62
drains the anterior brain stem, the superior and
inferior surfaces of the cerebellar hemispheres, and
the lateral regions associated with the fourth ventri
cle into the superior petrosal sinus. The posterior,
or tentorial, group drains the inferior portion of the
cerebellar vermis and the medial portion of the
superior and inferior cerebellar hemispheres into
the transverse sinus or the straight sinus.
 lature OVERVIEW OF THE NERVOUS SYSTEM
- Vasculature

Anterior vi ew Posterior view

~A
Pus/eri or cerebral a. Postelio. inferio r
cerebellar a.
Superior cerebellar a.

Basilar <l. Pos terior spin al aa.

..
~ An/c riOI inferior cerebellar a.
filmid
Pos terior inferior ce re bellar
ifu le
Anterior s~ n a l a . Posterior rad icular <la.
' Cerv ical
Vertehral a.
verte Or.:te
Deep cerviC;1J.1.
Anter ior radicular aa.
A~ce ndin Fl c.erviLal a.
Ascending cervical a.

~Iar v. Deep cervical

SI.)nl)
Subclavian a. -'>-.., Subdavian .1 .

~:v, Anterior ra dicular a.

Pos terior intercostal a. "<~7r{~ t

~rebri

19i1lal
.:O>iJ---t-- Posterior in te rcostal a.1 .
Thoracic ,'erte-brae ("'~\~) iiI,r;
n
cut)
~le v. Artery of Ada mkiewicz
(major ameriOlladiculill a.) --"im

Anteri or ra dicular a.
"'--r-- Posteri or rad icular aa.

Anastomotic loops to lumbar aa.

posterior spinal lumbar vertebrae

*.-"-- An astomotic loo p ~ to

anterio r spinal a.
iN
'.'J,: , l,' Ie,,,1Sdcral
(or median saer" l) a,
lateral SGlClal
(or med ian saoal) a. - -H ,j

-
FI GU RE 1.63: ARTERIAL BLOOD SUPPLY TO THE SPINAL CORD: LONG ITU DINAL VIEW
_ __
The major arterial blood supply to th e spinal co rd
derives from th e anteri or spinal artery (ASA) and
ri the paired poster io r spinal arteries (PSAs), bo th
bran ches of th e vertebral artery. The actual blood
e tlow through these arteries, derived from th e poste
rior circulatio n, is inadequate to mai ntain th e spinal
cord beyond the cervica l segments. Radicular
arteries, deriving from the aorta, provide major
anastom oses with th e ASA and the PSAs and
supplement the blood flow to the spinal cord.
Impaired flow thro ugh these critical radicular arter
ies, especially durin g surgical procedures with
abrupt di sru ption of b lo od flow through the aorta,
ca n result in spinal cord infarct

63

OVERVIEW OF THE NERVOUS SYSTEM Vasculature \,

Arteries of cerviCilI
cord exposed
from the rear

Basilar artery
Posterior inferior cerebell ar artery
Verte bral a rtery
Anterior spinal artery
Spin al ramus
Pm/erior spinal ar tery
Posterior radicular artery
Pre-laminar branch

~~p . ,~~

An terior spinal artery ~t -t-'

, 1( ~~

Post-central branch ~
Ii-
!l ij t
' -:"J,'
Anterior centra l arter y - -_ _ _--,

Spinal ramus

N eural bran ch roo' .Jl4jj.~

Anterio r radicular artery ..~:.:;;:;," (-; -}

Posterior ra di cular artery -

Internal .pinal Mteries
Po slerior central artery ~;!". ,!";;!" q:A;7i -'~ .- ,
t.: :..e "" -'. '" ' ._ . ~~+."
Pre-bmina r branch ..:.;;..:.,~' .~. . , _: :",~~:~,:,

Pos terior spinal artery - .,~~

Arteries of spinal cord diagrammatically shown in horizontal section

FIGURE 1.64: ANTERIOR AND POSTERIOR SPINAL ARTERIES AND THEIR DISTRIBUTION _ __
Th e ASA and PSAs travel in the SA space and send
branches into the spinal cord. The ASA sends alter
natin g bran ches into the anterior median fissure to
supply the anterior two-thirds of the spinal cord.
Occlu sion results in ipsilateral flaccid paralysis in
mus cles supplied by the affected segments, ipsi
lateral spastic paralYSis below the affected level
(upper motor neuron axonal damage), and co n
tralateral loss of pain and temperature sensation
64
<I(
below the affected level (damage to the antero
lateral spino thalamic/spinoreticular system). Th e .. ~ I
PSA branches sup ply the dorsal third of the spinal r.;c
cord. Occlusion affects the ipsilateral perception o f
fin e discriminative touch, vibratory se nsati on, and
joint positi on sense below the level of the lesio n
(from damage to fasciculi gracilis and cuneatus, the
dorsal columns).
(asculatu lll ~ ulature OVERVIEW OF THE NERVOUS SYSTEM
-
7.:;:..,,;=;;::;;;:;;;::-::--- Posterior spinal aa.

_ _" """iio, radicular a.

_ _' Po ;(e"io, radicular aa.

r ___ Boa n, h to vertebral

body and du ra mat~r

' - - - S oii ,,11 branch

' - --IDo,,, ramus o f

posterior intercostal a.

L _ -'>,,,.,,,,',,et,oall anastomosis

_A"v,~ .eb..' anastomosis

Section through thoracic spine

Central branches to
~_
~Q~~;~:.:~~::~~,:::: I branchesa, from pial plexus

right side of ,p;n" w,d -----A~r:5: _,Cen'" branches to left side of cord

poste rior spinal a.

Anterior radicular ,,-- - - -
_-Zo" supplied by pen etrating

p' e"'----<l~l[
branches from pial plexus
Pial arterial - - Zo,oo supplied by central branches

r - _ Zone supplied by both central

Posterior radicular a,-- -- - -,/i branches and bra nches (rom
pial plexus

Anterior 'p;' ""a,----~c...-3.\;:;',4.~:::~5

11- P"';'e,;,", radicula r a.

Schema of arterial dist ribu tion

. ;

""--Pia' " ""ii,11 plexus

- FIG UR E 1. 65: ARTER IAL SUPPLY TO THE SPI N AL CORD: CROSS-SECTIONAL VIEW
_ _ _ __
The major contribution to the arterial blood supply strud ures. The penetrating vessels supplying the
of the spinal cord derives fro m the aorta via the spinal cord derive from central branches of the
"I radicular arteries (top). This intercosta l blood sup ASA and from a pial plexus of vessels that sur
I of ply also distributes to adj ace nt bony and muscular rou nds the exterior of the spinal cord.
d

:he

65
OVERVIEW OF THE NERVOUS SYSTEM Vasculature

Veins of Spinal Cord and Vertebrae

~ Anlefior ex!ernal venous plexus
, ..._____Posterior external venous plexu s
inlernal venous plexus
...erlebral v.
'srverlebral \'

An teriOI and posterior

radicular w
Anterio r external
plexus
Anterior internal
,I,

POSlerior internal
plexus

Anterior spinal v. Basivertebral v.

Anterior central v. Ant erior intern al

venous plexus

"j- v - PoSleno r ra dicula r v.

- -Internal spinal vv.

" '17.~
~;c.-~~~,-
~f$~'
.,. ., ....~>.,"1>.~'
;$- ~Posler~;centraJ v.
, .
Po stenOf spInal v. ~~p llliN
" ...
.........:: ~~.~~'b..":~",",N~ ;<~.. Posterior inh~rn,) 1 venous p lexus ".: : ~

FIGURE 1.66: VENOUS DRAINAGE OF THE SPINAL CORD _ __ _ _ _ _ __ _ _ __ f

An ex ternal and an internal plexus of veins extends
along the length of the vertebral column, fo rming a
series of venous "rings" w ith ex tensive anasto
moses around each ver tebra. Blood from the spinal
cord, the vertebrae, and th e li gamen ts drains into
these pl exu ses. Changes in intratho rac ic pressure
66
and (SF pressure ca n be conveyed through these
venous plexuses, affecting the venous vo lume. a
U ltimately, these venous plexuses drain thro ugh
the intervertebral vei ns into vertebral, posterior
intercostal, subcostal, and lumbar and lateral "
Ll
sacral veins.
 OVERVIEW OF THE NERVOUS SYSTEM

Oropharyngeal "~"".......",,
membrane - --,rf"''--. .,
Lens plDcode

Hypophysis
Pa raxial column

Intermediate
"'ppearance of the col umn
"'lj!ural pl ate ~--la'eral plate OP lic area

Neural crest

'\:eulal plate

forming neural Spin'}l

tube

Interm ediale~_ _

mesoderm

Dellelopmental (ates of

'-, traemb ryonic local regio n of ectoderm

of embryonic disc al 18 days

-
ose
AGU RE 1.67: FORM ATION OF THE NE URAL PLATE, NEURA LTUBE, AN D NEURAL CREST _ __
- " e neural plate, neural tub e, and neural crest form destined to become the neural crest with a future
',h e 18-day sta ge of embryonic development. co ntribution to many components of the PNS. At
h - "'e underlying no tochord induces the neural plate, this very early stage of embryonic develo pment,
r d a midline neural groove forms. The eleva ted these neural precursors are vulnerable to toxic or
..a.:eral margins become the neural folds, tiss ue other forms of insult

67
OVERVIEW OF THE NERVOUS SYSTEM Developmental Ne uroscience

Neuril l plate
of forebra in lr;
'0 0-"

Neural groove ~

<
Neural fold s n it,," Fu lure neu ral cre~t, "Neural fold
E
E
q
"!
;,

Primitive streak-- '

Embryo at 20 days
(dorsal vi ew)

Neural ,..-'-' - f'

of fo rebrain----;;-i

Neural groove - - -

Neural folds""-
c
E
~
~
Fu ~d neural folds-

1SI cervical wmile

Caudal neuropore - -
~~;r.:

CIIilI\I
,',":,":::
Embryo a t 21 dayS
(d OlSal view )

FIGURE 1,68: NEURULATION _

_ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ __ fl(
In the 21 to 22-day-old e mbryo, the ne ural plate, system, is in the center o f the neural tube. This no
w ith its midline ne ural groove, thi ckens and begins process of neurulation continues both caudally and
to fold and elevate alo ng eith er side, allowi ng the
2 la teral edges to fuse at the dorsal midlin e to form
rostrally. Disruption can occur because of failure
of fu ll neural tu be fo rmati o n cauda lly (spin a bifid a)
"'"
~o
pia
the comple ted ne ura l tu be. The centra l ca nal, the or rostrally (anencephaly). ;Je'
site of the future develo pmen t of the ventricular len
me
pia
68

~I Neuroscie~ elopmental Neuroscience OVERVIEW OF THE NERVOUS SYSTEM

,

The neufallube wi. form the blain and spinal cord, the two co mponen t.. of th e central nerVOU5
system (eNS)_The n e ur~l cre!11 wi ll give ri..e to all of the neu rons whose cell hnd ics are lo ca ted
outside the eNS in tile pt>riphetal nl'rYOU!i system {PNS) o f n erves, gan glia. an d plexu~e s.

ral fold
Deri vatives of the neural tube ioclude:
Neti rollS oi the eNS
Support in g ce ll s of the eNS
Somatomotor neurons of the PNS
Presyn aptic autO nom iCneurons of PN S

Derivaliv,"s of the neural crest indude;

Sensory ne-utom in the PNS
Po>tSynaplic au tonomic neuron s
Schw ann (neurolemma) cells
Adrenal medull.. cells
Head mesenchyme
Me1anocytes in the ~kin
i'lrachooid and pia mater of meninges
(dura m.:lIer from mesoder m)

Embryo al24 days

(dorsal view)
Dorsal spinal
ga nglion
Ectoderm

--:eural crest
.. ~_ L ------r:......,
__ Sensory ne\Jron o f
Spin al cord
Sympa thetic tronk

-.
gan glion
\ dorsal spinal ganglion
'1eural tube
-spinal c",n>-~ '-.:'.7.
~ ~.>.<
: ~"
':'.,:
___-;r
Visceral mOtOr neuron ,,~.;'r.~~; Cortical p rimordium
01 sympa thetic ga nglion 01 suprarenal gland
4th week Dorsal mesen tery

Germinal epithelium Get

Chromaffin ce ll, or future gonad
suprarena l
medulla ce ll Sero sal li nin g (peritoneum)
01 abdominal celom
(peritoneal cavity)

61hweek

and
FIGURE 1.69: NEU RALTUBE DEVELOPMENT AND NEURAL CREST FORMATION
_ _ _ _ __
Th e dorsal and ventral halves of the neural tube are
separated by the sulcus limitans, an externa l protru
,
sion from the ce ntral canal that demarcates the alar
Ja) plate from the basal plate. This important landmark
persists at some sites in the adult ventricu lar sys
tem , The alar plate is the source of generation of
many neurons w ith sensory function . The basal
plate is the source o f generation of many neu ro ns
with motor or auto nomic function in the spinal
cord and the brain stem. The neural crest cells at
the edge of the neural fo lds unite to becom e a
dorsal crest, th e neural crest above the neural tube.
The neural tube and the neural crest separate from
the originating ectoderm. The neural crest gives
rise to many cell types in the periphery.
69
OVERVIEW OF THE NERVOUS SYSTEM Developmental Neuroscience

~
mater

Ce ntral
ca nal

Ma rginal lonp
zone (gray m (l ll e rj (w hi le maile r)
B. Spinal cord a t J months
Exlern al
Migra ti ng neu roblasts
limiting Pial _ _ <D _ .::v J :::-;.,..., Pia mater
membrane cell @ ./'l>-.......
\ 4th
ventricle

"
...
zone zone ~

, H , m,l\ler

._ 'li -"
Ependym al Ma ntle Marginal "l! ~ =gju .3,
~
layer layer layer
C. Cerebellar hemisphere at 3 months
= , .~ J!
1:l ;
A. Neural tu be a t 5 weeks .Di: !ib ~ ~
;3
Migrating neuroblasb

Latefal
ventr ide

~~fi
j.~
zone zone white
maile r
cor tex laye r

oii O. Cere bral hemisphere at 3 months

T
Marginal zone

FIGURE 1.70: NEURAL PROLIFERATION AND DIFFERENTIATION: CO(

WALLSOFTHENEURALTUBE ___________________________________
~ ::

Early in development (5 weeks), neurob las ts in the
ependyma l layer lining the central ca nal move back
and fo rth from th e epen dymal surface to the pial
su rface, replicating as they go. Neural migratio n fo l
lows distinctive panerns in different regions of the
neural tu be. In the spinal cord, neurons migrate
granul ar layer, from w hich granule cell s then
mi grate inward to synapse w ith other neuro ns pres u"'"
ent in deeper layers of the ce rebellar cortex. In the
cerebral cortex, neuro ns migrate to the outer lone,
where the gray matter remains o n the surface,
external to the white mat1er. These developmental '"
-~

in to the inner mantle zo ne, leaving th e ou ter mar patterns reflect the anatom ical organization of the Jft
gin al zo ne as a site for axonal pathways. In th e
ce rebe llar cortex, some neurons migrate to a
mature stru ctures, their blood supply, and th eir
vuln erab ility to a variety of insults.

--:e
locatio n on the outer pial surface as an external
70
rce Avelopme ntal Neuroscience OVERVIEW OF THE NERVOUS SYSTEM

8lPOi.ll cell of
~g/liof
CI.n~ n. VII'

., O>rom~iiinc d

"
oi .>d,e ... 1mo:tlull~
~
ria mote, cell

late r

eculM
Iver

FIGURE 1.71: NEURAL TUBE AND NEURAL CREST DERIVATIVES _ _ _ _ __ __ _ __

Neural tube ependymal cells give ri se to neura
blasts from which the ne urons of the eNS are
; pres. derived. They also give rise to the glioblasts from
n the which the mature ependymal cells, astrocytes,
zone, and oligodendrogl ia are derived. Microglia, the
"scavenger" cell s o f the eNS, are derived mainly
le ntal fro m mesodermal precu rsors. Cells of glia l origin
,f the are the predominant ce lls that give rise to e NS
tumors. Th e neural crest cells give rise to many
peripheral neural structures, including primary
sensory neurons, postgan gli onic auto nomic
neurons (sympathetic and parasympathetic),
adrenal medullary chromaffin cells, pial and arach
noid cells, Schwann cells (the supporting cells of
the PNS), and other specialized cell types. Neural
crest cells can be damaged selective ly in some
disorders (e.g., familial dysautonomia) and also
can give rise to specific tumo r ce ll types, such as
pheochromocytomas.

71
OVERVIEW OF THE NERVOUS SYSTEM Developmental Neuroscience

Central nervous system at 28 days

Forebrain Midbrain (mesencephalon)

Hindbrain (rhombencephalon)

Optic vesi cle {~___"

Cervical flexure

Spinal cord I
'"
M

/FO"b""
Forebrain
(prosencephalon) ~ ~-- Alar (rooO plate
..... Midbrain (mesencephalon)

~"" .....__Sulcus limitans Optic vesicle

Hindbrain ,- - - - Midbrai n
(rhombencephalon)
Opening of
right Oplic vesicle
_ Alar (roof) plate I--- - Hindbrain
Basal plate / ' ,
~ Sulcus limitans
Spinal cord <::;; Basal plate

In these sections:
~--- Spinal cord
c=:=I A lar plate
.p ~p
Sagittal section Frontal section
~ 8asal plate (venlral to sulcus limitans)
tf ~:H.'rI A.CRAIC...e
c~ r

FIGURE 1.72: EARLY BRAIN DEVElOPMENT: THE 28DAYOLD EMBRYO _ _ _ _ _ _ __ - ..

Some components of the neural tu be expand and the hindbrain (rhombencephalon) caudally.
differentially, resulting in bends or flexures that Th e ventricular system bends and expands to "".i(
separate the neural tube into ca udal to rostral co m accommodate in creasing neural growth. An out ~,

ponen ts. The ce rvical flexure, caudally, and the growth of the caudal part of the prosencephalon .. . '
cephalic flexure, rostrally, result from the differential extends from the future diencephalon to become a>
expansion. Three regions of rapid cellular prolifera the optic vesicle, giving rise to the future retina and :n:
tion develop-the forebrain (prosen cephalo n) ros its ce ntral connections.
trally, th e mesencephal on (midbra in) in th e middle, .
C"1
.... .:::-
72

-l
;cience evelopmental Neuroscience OVERVIEW OF THE NERVOUS SYSTEM

Saginill section
Cranial n. VII (Facia l)
(sensory and
Hindbrain Metencephalon
Ice re bellum, pons)
~ 'h, "w,"',
Thin root of
Cranial n. VI . VIII myelencephalon
(abducens) 'm'''o,, \ ('!',"'dUI, oblong,na)
(se nsory)
(raniill n. 'V~J',~~:~,:~"
(sensory 0'11 n. IX Sulcus 'ti" "it,""
MesencePhalOn \
Cran ia l n. IV mOLOO
(trochle;)r) 'm,,,oO'- Cerebral
Cranial n. X aqueduct
,Midb rain (se nory
a nd motor )
.i. Cranial n.11!
(oculomotor)
(moto r) - - - -
3rd ventricle
Opening of
right telen
cephalic Opening of right optic stalk
vesicle " (,m',,, termina li s
frontal section
(venl ralto sulcus lirnitans)
(myelencephalon) Lamina
ventricle
terminalis
/ ' Telencephalic vesicle
//_ (,,,,,,t' verllricle
Coccygeal n. 1st cervica l n.
(sensory and motor) (sensory
" " mo;tm)-1
15l sacral 11 .
(sensory and mo,",)' -'."
, 1

1st lumbar n. 1st thoracic n.

(senSO ry (Sensory
"d ~o'M" "d moton-' I aqueduct
_ _ _ _ B,__,I plale
Central Nervous System:
Crania l and Spina l Nerves at 36 Days
(cerebellum, pons)
.--.J Sensory neurons and gan glia from neura l crest - - "h ventricle
- - - Myelencephalon
(medull a oblongatiJ)
In sagittal and fron\JI sections:
ventricle
~ Alar (roof) plate
' _ _ _ Spinal cord
t%:.& Basal plale - - - - Central canal

- FIGU RE 1.73: EARLY BRAIN DEVElOPMENT: THE 36-DAY-OLD EMBRYO _ _ _ _ _ _ __

- day 36, the prosencepha lo n begins to expand telencephalon. Th e rhombencephalon further

":JOKIly as the future diencephalon (thalam us and
",alamus) and telencephalon (basal ganglia,
ebrain. olfactory system" and cerebral
_:~<c#rd'#Wpr~
!!."r -rid e for the dien-
iJ"erJJ I 'en/rides /.rom
_ :o;r/"~~....,;<;:?"""""""....6--~
develops into 2 distinct regions, the meten
cephalon (future pons and cerebellum) and the
myelencephalon (future medulla). Distinct spinal
~ford7"a?&7/~dw-d?&P/d'ff87X:;;V
and motor neurons differentiate and begin to con
nec/ with the;/" taifetsin the/JC'/"jolJe-0"
OVERVIEW OF THE NERVOUS SYSTEM Developmental Neuroscience

Central nervous syslem .1t 49 da~'S

Cephulic flexure _ _ __
pons)
___ Roof of 4th ven tricle

113 obI

E Pontine newre

o (cerebral hemi
~ Te Iencepha Ion - <;ni"~1 cord
0 1{actory lobe
1 (paleocortex)
O ptic
I
(pilui

Central nervous system at 3 months

Cerebral hemisphere (neocone)! ) ~

Me~n(epharon

Ou tline of diencephalOfl
(ove rgrcw"" by cerebTa l
hemisphere5) ---~
Cerebellu m (metencephalon)

O lfactolY lobe (pal eocortexl~

MedulliJ oblo nga ta
Optic nerve (crani al nn.1I (myelencephalo n)

Hypo physis (pituitary glandl ./

E

E Pons (metencephalOn)/' ,11 Cervical enlilrgern en t

~ of spinal cord

1 lumboS<l.cral enlargemen t
of spinal cord

. _ ..CRAJC-oo
.IOHN I
,.",.':" J
cr
FI G URE 1.74: EARLY BRAIN DEVElOPMENT: THE 49-DAY-OLD AN D FIG
THE 3-MONTH-OLD EMBRYOS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
'\}eu
By 49 days of age, the diencephalon and the telen covers the diencephalon. The cerebellum forms (OSli
cephalo n differentiate into distinct components: from the rhombic lips of the metencephalon, as cep!
the dorsal thalamus and the ventral hypothalamus, neurOnS travel do rsally to overlie the future pons po le
and the olfactory lobe and the future cerebral and eventu ally most of th e brain stem. The mesen in a I
hem ispheres, respectively. The metencephalon cep hal on ex pand s dorsally, forming the superior ( -sh,
(pon s) and th e myelencephalon (medulla) develop and inferio r co lliculi (quadrigeminal bodies). The leavi
further and fold, sepa rated by the pontine tlexure continuing growth of the sp inal cord as it connects opsi
dorsa lly. Between 49 days and 3 months, massive with periph eral tissues in the developing limbs terml
development of the telence phalon overrides and forms the cervical and lumbosacral enlargements. the s<
74
)science velopmental Neuroscien ce OVERVIEW OF THE NERVOUS SYSTEM
Forebrain at 7 Weeks (tranwersesectiun ) Telencephalon at 7th Weeks (tran w e rsesectiom
oi median teloCl'1e Choroidal ",I" ," ,. """>, Roof of Jrd \'cntl ic1~
''''Ci>phalic vesicle (3rd venlride-t
l lipp0l:ampus
. ~o rJ I hemi sp here;
11,......... / Lateral telocelc !lemisphere;
(iI,chi palliuon)

" (lateral
ven tricle)
neopa llium) _ _ II

- , 11--- betw{"{"n
O Pffling
lal era!
Interven tricular
: pend)'mall',"
, ,, --" iorame n
and medi ,m
M ~n tl e laye r lekJcel0S
(b ~s~ 1 gangli o n)
Marglnn l la yer - - fo ra men I lrd ventricle
(3 .d Y(~nl ricle)
at 2 (nglll "n,,, , ior view)
(coro n ~ 1 ~ecti o n; anteri or view) Right cerebra l hem isphere (n (lp~ lIi lJ m. ClJt edg e)
EpiphYSiS (pineal gla nd ) -:;"ii'''''''~
- ~dJral hem isphe re IO.,'='- Roof 0 1 lrd
ventricle
.',)dlliu m. Cl.H ~Ige)
~ >llp.,"<,",,", (archipa llium)
Choro id p)e,;u s plOl rudinK

=l o roid ,1,,,,, -- III to right lJ, tera l vntfl df'

a lo ng c h(') roi d lissum
Cho roid ,,'" " " -- Corpus ~triatu m
Thalamus _ -4"-; Iba!>al garlglio n)

(orpU$ striatu m ventricle ImClventr!l:lJlar foramen

( b~s~ 1 ga ngli on)
Open ing of (,wily of
l rd ve ntricle fo rame n right n lla<.:tory lobe

lilmin;) lermillah " J If,K"'" lobes (paleo p alliu m)

Right Cerebral Hemisphere at 3 Month s Cerebral Hemispheres at 3 M o nths

(medial as pec\) feo .onal ~ t ion )
Medi a l $uriace o { right ce rebl al

Commissu re o j forllix
(hippocampal
~~""'";'~;:~~~O~P:';i
L (~OO I ; ~"~mi)::~:?~;i~
, I
Lateral \ientri de
,
(ovl!fin~of
rig ht lal eriJl (ho, old plexu$
A nte rior ve n tricl e ;!long Neopallia l/ ,
Cho rQida l ve in
choroid r. ssure cortex"""" .
a nd artery
~' na termin aII S--';~_ ". Hippocampus
Hippocampal Inte rn ,,1capsule-
OJf,lCloo y lobE> (archipallium)
I Stria termin.-dis An terior commi$~lJre
COOI)U$ ( :,"d"~
A
-, /Jf%
I Cl ,I@N
Th alamu s Icut surface)

i of div isi on betwee n

i t

[basal
nucle us

\l ;:;::~~";:J
Interventricu l<lt imame n
i of
ganglia)
........ ::
diencephalon and telencephalon fOof uf )r(1 >'cntride

G U RE 1.75: FOREBRAIN DEVElOPMENT: 7 WEEKS THROUGH 3 MONTHS _ _ _ _ _ __

-

IS
~,Jr o n s of the developing telence phalon move
:-:-ally, dorsall y, and then around the di en
anato mically. Th e caudate nu cl eus also exte nds
around the telencephalon in a ( -shaped pattern,
S 'Y'Phalon in a C shaped course toward the anterior vvith the head of the caudate remaining anterior
ns - e of the temporal lo be. The hippocampus forms and the m uch smaller body and tail follow ing as a
s.e n a dorsal and anterio r positi o n and migrates in a thi nner C-sh aped structure thal ends ventrally in
lr _-5naped course into the anteri or temporal lobe, th e temporal ho rn o f the lateral ve ntricl e. Th e cor
'e e.a . ing the fornix in its w ake. Th e amygdal a devel pus ca llosum and the anteri o r co mmissure co nnect
eels .; in a similar manner, giving rise to the stria the 2 hemispheres. The internal capsule funnels
B'minalis in a C-s hape. Th e lateral ventricles fo llow centrally in the core of the forebrain o n either side
lts. '-e same C-sh aped developm ental process and co ntinu es caudally as th e ce rebra f peduncle.
75
OVERVIEW OF THE NERVOUS SYSTEM Developmental Neuroscience

Brain at 6 montns

'II ItOmm ~

Frontal lobe of left

-...~ ~
CentrallrolandlcJ sulcus
ce rebral hemisphere ............

~
A~ Pa rielal lobe

InsulO1 [island of Reil) in

la le ra llsylvian l sulcus

Occipital lobe

O lfilctory bulb - - -
....-';
Temporal lobe
~ Medulla oblol1 gata
Pons

. _ _ Spinal cord
L-----

Brai n at 9 months (birth)

~.---------- 10.5 mm

Precelllra] (motor) gyrus ____ E, ~ _ Centr,,1(rolandicJ sulcus

P''''en,,_,1 "Ie"~ ;{ .' $ 'ht& Po;<"n" , 1(..nso",1 gyms

Y{
fo Postcentral su lcus
Left o:,rebra l hemisphere ~
'r .

Pari etooccipital sulcus

Lateral (sylvian) sulcus i~ -<>I \" ._~ ...~
Pari ecal lobe
In sula (Island of Reil) \:},;:l "" "'-'-- ~:
~,:;.,

Olfact ory bulb ;. Occipital lobe

y

Tempol'al lobe ~ Cerebellum

Pon, r Medulla oblongata

.JCtf< ...CRAlC....
(l lliiN I Pyra m id :::1
"'....'" Oli,,'
-".- Spinal cord

FIGUR E 1.76: THE 6 MONTH AND THE 9 MO NTH CENTRAL NERVOUS SYSTEM
_ _ _ __ _
At 6 months, the brain ste m has differentiated into co mpo nents of the basal ganglia, th e limbi c struc
the medulla, th e pon s, and th e midbrain, with the tures (e.g., amygdala and hippocampal form ati on
cerebellum overlyin g them dorsally. Even though th e o lfactory system, and the cerebral cortex
the diencephalon is rap idly developi ng, the overly develop rap idly. M ost neurons are present at bin."
ing telencephalo n shows mass ive grow th rostrally, except for some populations of granular cells in
then caudally, dow nward and forward into the ten) cerebellum, the hippocampal dentate gyrus, and
poral lobe. From 6 to 9 months, the cerebral cortex the cerebral cortex, w hich form postnatally in
forms its characteristic convolutions w ith gyri and response to environmental stimuli. The in utero cr
sulci, and the cerebellar cortex forms its distinctive postna tal environments provide major jnf1u e n c~
folds, the folia. W ithin the forebra in, th e major o n neuronal development and function.
76
 troscienu !!'iopm e ntal Neuroscience OVERVIEW OF THE NERVOUS SYSTEM

;oolal sectioll (ve nttal 10 sulcus limi ta os) at 3 6 days Ependymal lining of cavities of brain at 3 monlhs

len lale.,ll venlrid e

3rd ventri cle
Inlervent ricular
- L". ," I v{'ntrid!:! foramen
, ~r pi" . -- - ,," (of Monrol- - ---'iiC

0.1--- Optic 5til lk

_'d ventride
O ptic cu p
Infundibular recess :~;::':~~':~;';;:'i~~~) -~
Cerebr;ll.:aqueduct
:erebr al aq ued uct (01 SylviusJ
Me.>encephdlon

Basa l pJilte Lateral a perture of 4th

\'Nenceph,don
ven tricle (of L",d>'''~/
cN e bo:!IIllm. DO ''')-, in lat eral re('eS5
\l etacoele
..! th ventricle I 0141h
. Mage ndie)

i '.--_ - Spinal cord CeOlral ca nal of spinal cord - - -

I. .1l h vent ricl e
' - -- - Central canal

dymailining of cavities of brain at 9 months <bi rth)

.u lcus

JryJ gyrus
Right lateral ". ""i" J)" - ---c
_------- ~;~~;::i~:
!:.
hOln of left laler.,1
ve ntri cle in fra ntallo be
RegIon of invagina tion oi

choroid p,, 'e,~,"~:,;,'::' ;::,~;'~~~--1

~----- 'C..,,,,, P'''' oi lefllate ral ventricle
iis5ur e of la ........~~

: -- - - - ,,"p~pi ",,'1 rec ess of 3td ventricle

canal
~ lcus 0-- - - I'io" , 'l rece~s
~~~" f,"ii O' hom of Id t la ter al
Foramen in 3rd ven tricle

ventricle in lemporal lob('

fa r inle rth ala mic ad h~ion

_IPo,,,,lo. I,om of left lateral

Thalam ic impression

ventriclE' in Occipi tal lobe

Optic recess of 3rd "e<"" '~"

Infundibu lar recess '--- - - -L" I ,." " ,'1 ap erture (of Luschkal
Region ofirwagination of of 4th ve nlricle
choroid ple)(us alOllg cho.-oid
fis~re of lateral vt!nrricle ------ ~~~~'~:.;.;;:~',~' (of Magendi C' )
Cerebral aqueduct (oi S" "i" " ~-----. (,,,,,,1 c.l nal o f spinal cord
fi GU RE 1.77: DEVElOPMENT O F THE VENTRICLES _ _ _ _ _ _ _ _ _ _ _ _ _ __
)ic stru c n e rapid growth of the brain stem and the fore (medial) an d Luschka (lateral) open the ve ntricu lar
rmation ), """'" ain alters the uniform appearance of the ven tri system into the developing cisterns of the sub
lex d es. The C-shaped lateral ventricles follow the arachnoid (SA) spa ce at th e med ial ancl lateral
t at birth, ;rowth of the telenceph alon, with limited access margins of th e fourth ve ntricle, respectivel\,. (SF
:ells in the lJ"lto the third ventricle through the in terventricular reenters the venous system th rough the arac hnoid
us, and ~ ra m e n of Monro. The narrow cerebral aqued uct granulati o ns, o ne-w ay va lves allowin g drainage
:YIn emain s very sm all in th e upper mesen cephalon from the SA space into the dural (veno us I sinu ses.
utero and and opens into the rhombus-shaped and expand especially the superio r sagittal sinu s.
uences ng fourth ventricle. The foramin a of Magendie

77
OVERVIEW OF THE NERVOUS SYSTEM Developmental Neuroscience

Spinal Cord Mature tl<_~.,....._tIOnJ

5 1/2 we4'kS (tr~n""...,,, """,on,
Central can al
OOl,al alar pia IE.' ~. C'rI tr al canOl Sensory _ Dorsill gray co lumn (horn)
{se ns.o ry and coordi nJ.till ~ J '<'
~ Ep endyrna l1 ilye r ~ Tracts (whi te matter)
--- Mantle la V@I
Lateral gray column Ihom)
'M3rgif'lallil\'er :t{ ,.
Ve ntral bdSill plale Motor ~ Veotral sray <:olurnn (hom)
5uk us Iim; l afl~
(mOlor) - - Tracts (while matter)

Medulla Oblongata Mature (~~n~'@I'ie ",c~on, I .

S 1/ 2 weeks I"an ;,..,.,~ ",,,,,,,n. Vestibular flU e ('I
Roof pl~l e lOC"'Km
::::;l"'-';.y Nucleus tractu s solit~riu s
Dorsal alar plate
,"...,n""" ~~Xt~ ,-- {rode" , oi ,he soli t ~" ,r3cti
, ~g ~.
4th ve ntricle 'OOId;n.~ng
nuc{,,;

nudeu~ of the vagus

( ~ el1sory and 4thventrid e
Ependymal 'g"" mo"<'I'1
l ooruinating) ... " '. _1 Dorsal ', mo ICfi
la)er
l ocalion $ ~/ Hyp oglossal nucleus
Ventral ba~a l platt'
.""lanl)e Iily~r ,,;melo<
<""vol
J4fo >~. ~ Tracts lwhite~a"el)
Ma rwnal i '" . .':>. -......."1
Nucleus amb.suus
(motor) Javer '"""
!5'! ""' tf<:ft
I
~""" In ferior olivary n uc eus
Sulcus iimi'ano; ----- - - Pyramid

.i 1/ 2 weeks f(f,'"~'"..,e ,eCl,<>"' Mesencephalon Mature \"""5\'('(lf !tKtiuo Oi

-
Tectlum- ." .... ___ Superio r colliculu s
[Jors,ll aiM plale Cerebral .1qlleciucl
(sen~or\-' and
Tegmentum
coo rdm<lllng) I~rer

' ,"1anlle Ia... er "~

. ', O
- Nucleus of
VeNial ba$.ll plate Margif1.11 Iarel Edlngerwestphal
{mol o r) Sulcus limilam
B~ _ " Red nucleus
Peduncle

Dience phalon Diencephalon and Telencephalon

~~p
51/2 weeks " ra ,m'''' ..... "'',,n,' Septum peNucidum, Mature 11r' 0"''''5<)50:;(loo )
Choroid plexus (pro;e<: I",S ."I u 1.".,,1,,n";clo .IonH ( hCfoid r,"," r~ '

'~N

Ruof p l ~te
\
,\I 3rd "enlricle

<'/ - {
EpendymallaYeJ
Mantle layer
:..- Claustrum ,~t P"IP-In)
Insula
Dorsal part
o ( ,11Mplate ,) } rMarginal / /~.L~ Lateral sul<:u~
(lhalam us) layer I lobe o(
Th.Jlamus

Hyp o I cortex ISra, """ Ier)

Ventral PM! thal<lmu ;; Iwh'l~ ",.11. "
of .llar pl~te
(h),I)Olh a l a rnu ~ 1
Hypnth,lIJmi( Id body
sulc us
, M,unm,Jlarr rec e!..S

FIGURE 1.78: COMPARISON OF 5'!:!,WEEK AND ADULT CENTRAL NE RVOUS SYSTEM REG IONS _
fI (
At 5 Y1 weeks, th e ventricular system becomes com form s motor and autonomic structures who se
paratively small er as neuronal growth occurs. In axons leave the eNS. The alar plate forms sensory L,-j

adults, th e ce ntral canal of the sp in al co rd is virtu derivatives in the spinal co rd and the brain stem, 50<
ally obliterated. Th e fourth ventricle ope ns up later and stru ctures that migrate ve ntrally (inferior o live, ~ :::

ally; the sulcus limitans demarcates motor nuclei po ntin e nuclei, and red nucleus). The rhombic lips, ,
(medially) and sensory nuclei (laterally). The cere
bra l aqueduc t remains very small. The third ventri
an alar deriva tive of the metencephalon, give rise
to the cerebellum. The diencephalo n and the telen
-'"
,
cle narrows to a sli t. The lateral ventricles expand cephalo n are also alar plate deri vatives. .;ep
massively in to a C-shaped form. The basal plate
78
 Dscien cr ~ elopmental Neuroscience OVERVIEW OF THE NERVOUS SYSTEM

='- II! IGVE) ~~_

.-... III (GS E) ~~

hom)
CN III

" ~ ! ~orn)
CNIV
( NV!
CNVll
CNV
CNVU
horn) CN IX ( NIX
CNX CN X
CN XII
CNXI

"
tIl vaBu ~

eN IV (GSE)

CN v (SVE)

;:>naryngeal CNVII (SVEJ

,arches CN VII (e VE)

C N VI (eSE)

eN IX (CVE)

CN lX (SVE)

"
,
colum n

3racnlomOlor
SV E) column

-
.CSE) colum n
,

"'

,Ve,'"'' horn
Lateral horn

lere
IV

Note: -!- ['nc',,' horn

.
GSE = General somatic efferent
GVE = Ge neral visceral efferent ~A.CR,t,~ fill

SVE = Spe-ciaJ visceral effere nt ;qJll'.-u

Spinal cord
mw"

'5 _ fiGURE 1.79: DEVELOPMENT OF MOTOR AND PREGANGLIONIC AUTONOMIC NUClEI

IN THE BRAIN STEM AND THE SPINAL CORD _ _ _ _ _ _ _ __ __ _
)ry Gray matt er column s develop in the spin al cord for develop in th e rh ombence ph alon as branchio
I, somatic lower motor neurons (ventral horn) and mo to r neurons supplying pharyngeal arch musdes.
ve, preganglio nic autonomic neurons (lateral horn). Both the som atic motor and the branchiomoto r
ps, These columns extend rostrally into the brain stem, neurons are classified as lower motor neu rons and
e maintaini ng th e same general positi onal relati on have axons eXitin g the e NS to syn apse on skele tal
'en ship to each other but organized into a series of muscle fibers.
sepa rate but aligned nuclei. A third grou p of nuclei

,9

OVERVIEW OF THE NERVOUS SYSTEM Developmental Neurosci ence

Superior collicul1.l5 ------"

Te gmen tumy., J gray matter
~ Z - . - - Medial ge niculate bod y
Retl(ulitr format,on ~
. ..'
~ ', -Oculomotornud eus
Sllbslanrl a nigla ~ ;& . . i"
~l;f_.d/:I';" Med ial longil udi nal fasciculus
< "< 'I.IZ
Temporoponlln e fibers ~h' . " M edial, spin al and tri geminal lemni sci

Cms cerebn CortlcosPlnal 3?d , Nucleus of Edinger-Westphal

{
COrTlconud ear l lbers ~ 4 nucleus
Frontopontme li bel) ~" .J ~
~O culomol o r (III) n.
Section through midbrain allevel oisuperior collkuli __________________

medull.ar\'Vel~#~
peodunr~~' ~ '.
Sup. _ 4 th ventricle
Sup. cerebellar Mesencephalic
Ant. spinoce re be ll ar tract
. ,,~ ~1-r
Mediallonglludinal faSCiculus

Reticular fo rm atio n

Mi dd le cerebe ll a r peduncle
- -
Medial, SpiOilI and Irigeminallemnisci

~
. 0-","0.="'=
,,>>::: _~-Trigem i nar (VJ n. and
Cortic ospi nal and cortico nuclear fi be r, ..",. :---.~ gang lio n
~'" . Median ra phe
_ _ _ _ _ _ _ _ _ _ _ _ _ _ Section through pons at level of trigeminal nerves

In!. me dull ary ve lum >m ....,. lei-

Cho ro id plexus of 4 th ventricle .t."'( '. '

La teral cuneate Dorsal vagal nud e us

tn r. cer~bell ar Solitary tract nucl e us
Post. spinoce rebellar trac t Spinal tra ct a nd spinal nu cleus of tri ge minal n.
Ant spi nocerebellar Irac t ~ . . Hypogiossoil t n ucleus
Spi nal lemniscus (~i no thalamic tracts) ;: .'.
Me di al lon gi tu dina l fascic ulus ~ In f. o livary nucl ei
Medial lem ni scus 4!ro. Hypoglossal (XII ,.
Section through medulla obI, .at level of inferior o livary nuclei - - - - - - - - - - - - - -
Cen[ral canal ..........~ Fascic ulu s gracili s
Hy poglossal nucleu s~ Gracile nucleu s
Post ~pi noc erebetla r lril C! ~~~ Fascic ulus (unealus
Spinotha lam ictraclS ~ :~
An t spinoce rebellar trac t in allract ilnd Spi nal nucl e us of trigemin al n.
Medi al le mn iscus - Int arcuate fibers

- - - - - - - - - - Swion through medulla oblongata

Spinal trac t a nd .,pinal nucle us of tri ge minal
Po ~t. spinoce rebe ll a r tract Gracile nucleus
Fascic ul\p.> cunealus

~~p
Dors.:l l ho rn
Spinothala mi c trac ts-= Lateroill corticospi na l tract
Pyramid and ant. cortkos pinal tlbers Pyr<lmidal deCl~tion e llOl\
:~...=
Section through medulla oblongata at level of pyramidal decussati o n

FIGURE 1.80: ALAR AND BASAL PLATE DERIVATIVES IN THE BRAIN STEM _ _ _ _ _ __ _
"The general patt ern of alar and basa l plate
derivatives seen in the spinal cord continues into
the brain stem. The alar plate derivatives are the
se nsory nuclei, the rhombic lip from whi ch the
cerebellum is derived, and nudei w hich migrate
ventrally to fo rm structures such as the inferior
olivary nudei, the pontine nuclei, the red nucle u ~
and o thers. The basal plate derivatives are the
motor an d pregan glio nic autonomic nuclei."

80
Iscience ) evelopmenlal Neuroscience OVERVIEW Of THE NERVOUS SYSTEM

SII2 weeks

Dorsal alar pl.lle

(st;nsor,' and
~-- Mantle layer
coordinating)

SulCU5 limitan~
Venlr.11basal
plale (mOlorl

- ' - - - - - Ma rg in <lliayer

3 1/ 1 months Choroidal artery ,1" 0 ve in

r-- E,,,",, ,,,,, "00' o. 41h ventrid('
Choroid
Epp.ndym,lilloor a/4th ven lricle
Sulcus II
....._ .....~-- ~~:~~: recE.>SS oC
~ 4th ventriclE.>

f-_" "'Ian tle layer

'--- - - Margina l layer

Mature ChOloidal ve~~els -"""'---~

lUclei
i(XII ) n.
Epe ndymal flo or - - ,
Vcstibul,jr nuclei
o j 4t h ve ntricle
~_ _ ".',<'''111 receS5 of
41h venlride

CI"Qc,, ;i c plexu s prutrud lr1 !!

through lilleral apertufe or
4th verlLridc
Descending (spinal ) - --j
l\' udf'uS tractus '>Ol i lani u~
nucleus OlV
Oor\,,1mOlar nucleus or" X

H)'po~kJSS<lI I Xfl) nucleus

Jf% ' ""he " d.' J ,r~

crrG
'.~ r ...
~,."'
() .Imt'\l
~.. : .

FIGURE 1.81 : DEVELOPMENT O F THE FOURTH VENTRICLE

_ _ _ _ _ _ _ _ _ _ __
or The expansion of the fourth ventricle from the
leus, ori ginal ce ntral canal of the rhombencephal on into
its matu re form is a compl ex process. The sulcus
limitans is conspicuous early in development (SY2
wee ks), and th e ori ginal lateral walls expan d out
ward and lay down hori zo ntally (31;2 months) as
the roof plate expands to either side; as a result, the
sulcus limitans becomes a landmark at the dorsal
boundary of the medulla o n the floor of the fou rth
ventricle, separating the motor structures medially
from the sensory stru ctures laterally. Th e latera l
aperture o i the fourth ventricle open s into the SA
space. In their mature form (lower illustration),
these pa irecllateral apertures, the foram ina of
Luschka, are major chann els bet\iveen the in ternal
and the external circulation of the (SF and must
remain open to prevent in terna l hydrocephalus.
81
OVERVIEW OF THE NERVOUS SYSTEM Developmental Neuroscience
II.
Differentiation and Gro\ol.'th of Neurons at 16 Days
~.P

"
Neura l crest - - - - - - - -,

A:
SI)lnalcord {EPeOdYmal la.~::::~
ItholJd c part)
~" ' -~ ~::-
Mantle I,we ,

...\,lI gin.l l laye r i

Spinal (Oro {thoraci<.. pam

Sensory cells I.on, neu ral crest
Commisiural nell. on
ASSOciation neuron
VentlJ! iu olc ulu$
Growing 10 dOfs.J.1surfare oi body

Growing 10 IMeld l and ve ntral

N
'"'""~ ~ surfaces of body

~!~
'...... ob~l~
...... imn , ,*",.>1 ' ..
, """ <7<"f, ! ~. g .........

pn!i""'oc. II
CI19N
, ',\",".' ::
.)'mp."N:t'c,
Growin g to vi)Cera of body
\ """"" I A
Differentiation and Growth of Neurons at 5 to 7 Weeks (righl side of di<lgra m show's neurons acquired since 28th day onlyl

nt'uron
VCrllra l mo tor neurons
',I iunic ulu 5
~ Ii,
Dorsal ramus of ~pm.ll nerve '\
from neur al crestl growirl g to
end on smooth (involuntary)
Ventr al ramus of sp in al
mus.c le and sweat glJn.9,.s

'vVr. lte ramus commurlkans k,

Gray ramus m OIOI
thoracic viscera
Sympatr.el ic trunk

m otor
Colia ter...1~ymPJ I r.elic tfUnk g<tnglion
'I i, I,
(cellae, St lperior, and inierior mesenteriC)
Igarlgliorlic, svmpathetic ~nd mOtor n e urons (derived
viscera from nllural crest) growin g to end on gland s .:md smooth
:Ii (involu ntar y) muscle

FIGU RE 1.82: DEVELOPMENT OF PERIPHER AL AXONS _ __ _ __ _ _ _ _ _ _ __

Peripheral axon deve lopment is a co mplex process ventral roots and terminate o n sympa thetic gan
of neurite extension, trophi c and chemotactic glion cells in the sympa thetic chain or collateral
factors. axonal guidance, and maintenance by ganglia or parasympa thetic intramural ganglia.
innerva ted target tissues. Dorsa l root ganglion cells Postganglionic axons form co nnections w ith target
are bipo lar; a peripheral axonal process associates tissues, including smooth muscle, cardiac muscle,
w ith simple or complex sensory receptor ceils, and secretory glands, some metabolic ce ll s (hepato
a centra l axonal process extends into th e eNS. The cytes), and immune cells in lymph oid organs.
lower motor neurons send axons to th e developing Sensory, motor, and autonomi c symptoms ca n
ske letal muscles through th e ventral roo ts or the occu r in periphe ral neuropathi es based on disrup
moto r cranial nerves, formin g neuromuscular tion of th ese co nnections.
j un ctio ns. Ce ntral preganglionic axons exit in the
82
 -
euroscienr
REGIONAL NEUROSCIENCE
PERIPHERAL NERVOUS SYSTEM (PNS )

'<-I. INTRODUCTION AND BASIC ORGANIZATION

Figure 11.1: Schemati c of the Spin al Cord wi th Sensory, M oto r, and Autonomic
Components of Peri pheral Nerves
Figure 11.2: Relation ship of Spinal Nerve Roots to Vertebrae
Figure 11.3: Sensory Channels: Reflex
Figure 11.4: Sensory Channels: Cerebellar
Figure 11.5: Sensory Channels: Lemniscal
Figure 11.6: M otor Channels: Basic O rganiza ti on of Lower and Upper
Motor Neurons

Figure 11.7: Autonomic Channels

Figure 11.8: Cutan eo us Receptors

Figure 11.9: Pacini an Corpuscl es

Figure 11.10: Interoceptors

Figure 11.11: Skin and Its Nerves

Figure 11.12: Neuro mu scular Neurotransmi ssion

Figure 11.13: Neuroe ffector Juncti ons

- ~.2, SOMATIC NERVOUS SYSTEM

Figure 11.14: Dermatomal Distribution
Figure 11.15: Cervica l Plexu s
neurons Figure 11.16: Phreni c Nerve
Figure 11.17: Th oracic Nerves
loe tic
"ed
Figure IU8: Brachial Plexus
ng lo Figure 11.19: Cutaneous Innerva tion of th e Upper limb
td ryJ
Figure 11.20: Scapular, Axillary, and Radi al Nerves Above
th e Elbow
figure 11.21: Radial Nerve in the Forearm

Figure 11.22: M usculocutaneous Nerve

j
Figure 11.23: M ed ian Nerve
" Figure 11.24: Ulnar Nerve

Figure 11.25: Lumbar Pl exus

, Figure 11.26: Sacral and Coccygeal Plexuses

Figure 11.27: Femora! and La teral Fem oral Cutaneous N erves

Figure 11.28: O bturator Nerve

Figure 11.29: Sciatic and Posterior Femoral Cutan eous Nerves

-
,~
Figure
Figure
A~3~
11.30:
11.31:
Ti bial Nerve

Common Peroneal Nerve

AUTONOMIC NERVOUS SYSTEM (ANS)

Figure JI.32: General Schema of the Autonomic Nervo us System
Figure 11.33: Innervation of O rgans of the Immune System and Metabolic O rgans
get Figure 11.34: Reflex Pathways
e, Figure 11.35: Cholinergic an d Adren ergic Synapses
Figure 11.36: Schematic of Cholinergic and Adrenergic Distributi on to M o tor and Autonomic
Structures
Figure 11.37: Autonomic D istribution to the Head and the Neck: Medial View
Figure 11.38: Autonomic Distributi on to the Head and Neck: Lateral View
Figure 11.39: Schematic of Autonomi c Distributi on to the Head and the Neck

83
 Figu re 11 .40: A uto no mic Dis tributio n to fhe Eye
Figure 11.41 : Innel va tio n of the Limbs
Figure 11.42: Tho racic Sympathetic Chain and Splanchnic Nerves
figure 11.43: Innervation of the Tracheobronchial Tree
figure 11.44: Innervatio n of the Hear t
figure 11.45: Abdom inal Nerves and Ganglia
Fi gure 11.46: Innervatio n of fhe Stomach and the Proximal Duodenum
Fi gure 11.47: Innervation of the Small and Large Intestines
Fi gu re 11.48: Enteric Nervous System: Longitudinal View
Figure 11.49: Enteri c Nervous System: Cross-sectiona l View
Fi gure 11.50: Innervation of the Liver and th e Bi liary Trac t
Figure 11.51: Innerva tion or the Adrenal G land
Fi gure 11.52: Autono mic Pel vic Nerves and Ganglia
Fi gure 11.53: Innerva tion of t he Kidney and the Upp er Ureter
Figure 11.54: Innerva tion of the Urinary Bladder anel th e Lower Ureter
Figure 11.55: Innervation of th e Male Reprod uctive O rgans
Figure 11.56: Innervation of the Femal e Reprodu ctive O rgans

B. SPINAL CORD
Figure 11.57: Cytoa rchitecture of the Spin al Co rd Gray M atte r
Figure 11.53: Spinal Cord Levels: Ce rvica l, Thoracic, Lumb ar, and Sacral
Figure 11.59: Spinal Cord Lower Motor N euron Co ntro l
Figure 11.60: Spinal Somatic Refl ex Pathways
Figure 11.61: Muscl e and Joint Receptors and Mu scle Spin dles
Figure 11.62: The Muscl e Stretch Retl ex and Its Ce ntral Co ntrol via Gamm a M otor N eurons

C. BRAIN STEM AND CEREBE LLUM

Figure 11.63: Brain Stem Cro ss-section al Anatomy (14 Leve ls)
Figure 11.64: Cran ial N erves: Schematic of Distributi o n of Se nso ry, Moto r, and Autono mi c Fib ers
Figure 11.65: Cran ial Nerves and Th eir Nucl ei: View From Above
Figure 11 .66: N erves of th e O rbit and the Ciliary Ganglio n: Dorsa l View and Cross-section
Through the Cave rn o us Sinu s
Figure 11.67: Extraocul ar Nerves (III, IV, and VI) and the Ci li ary Gangli o n: View in Relation
to the Eye
Fi gure 11.68: Trigeminal Nerve (V)
Fig ure 11.69: Facial Nerve (VII I
Fi gure 11. 70: Vestibu locochl ear l\J erve (VIII )
Figure 11. 71: G lossoph aryngea l Nerve (IXI
Figure H.72: Vagus Nerve (X)
Fi gure 11.73: Accesso r ~1 Nerve (X I)
Figure 11.74: H ypoglossa l Nerve (XII)
Figure 11. 75: Afferent Innervat io n of the Mouth and the Pharynx
Figure 11 .76: Reticular Fo rrnatio n: General Pattern of Nuclei in the Bra in Stem
Figure 11 .77: Reticular Fo rmatio n: Nuclei and Areas in the Brain Stem and Diencephalon
Figure 11.78: Major A fferent Connectio ns to the Reticular Forma tion
Figu re 11.79: Major Efferent Connectio ns o f the Reticul ar Formation
Figure 11.80: Sleep- \Nakefu iness Confro l
Figure 11.81: Cerebellar O rganization: Lobes and Regio ns
Figure 11.82: Cerebellar Anatomy: Lo bules
Figure 11.83: Cerebellar Anato my: Deep Nuclei and Cerebellar Peduncles

84
 E'IC EPHALON
=i gure 11 .84: Thalamic Anatomy and Interconnections \lVith the Cerebra l Cortex
=ig ure 11.85: The Hypothalamus and the Pituitary G land
=igure 11.86: Schemati c of Hypothalamic Nuclei

E'IC EPHALON
Figure 11.87: H orizo ntal Sectio ns Through the Forebra in (10 Levels)
Figure 11 .88: Cor0 I1 (1 1Sectio ns Throu gh the Forebrain (10 Levels)
Figure 11.89: Layers o f the Cerebral Cortex
Fig u re 11.90: Cortical N euronal Cell Types
Fig ure 11.91: Vertical Columns: Func tional U nits of the Cerebra l Cortex
Figure 11.92: Efferent Connec tions o f the Cerebral Cortex
Fi gure 11.93: Neuro nal O ri gins of Efferent Connec tio ns of the Cerebral Cor tex
Figure 11.94: Co rtical Association Pathways
Figure 11.95: Major Co rtica l Association Bundl es
Figure 11.96: Noradrenergic Pathways
Figure 11 .97: Serotonergic Pathways
Figure 11 .98: Dopaminergic Path\.vays
Figu re 11.99: Centra l Cho linergic Pathways
Figur. 11.100: Olfacto ry Nerve and Nerves of the Nose

IS

85
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)-Introduction

Sensory ! ,
MOlor ,
PregangliO" ~C
sympa thetic
,
Oor..al
Postganglio nic
sym p.:Uhe tiC
'
,
Dorsal
column

Dorsal root
I

!
I
Vascular smooth
" muscle, sweat
glands, and
arrector pili

Gray ramus communicilns

Sympathe tic chain Rang~on - - ./ ;

/
Spl,,,,hok o",v. 7 /"<", communicans

If 'L
Sympathetic chain

Skeletal
m!Jscle
CoICte,,1

'V;:~::~;C )t1( ./
Preganglionic sympathetic neuror
passing to synapse in anomer
sympaiheli(' chain ganglion

""
J. Pericins
MS, MfA

" '\~r:-!
Sensory neuron oi
abdomina/I'isc'ra
'If.
O
, ~) 5-""/
'., j ~,... . ,
" TO'; '
N euroeffector junctions on smooth
~; muscle, cardiac muscle, secretory
gldnds, metabolic cells, Immun e ce lls

FIGURE 11.1: SCHEMATIC OF THE SPINAL CORD WITH SENSORY, MOTOR, AND AUTONOMI C
COMPONENTS OF PERIPHERAL NERVES _ _ _ _ _ _ _ _ _ _ _ __ _

Periph eral nerves consist ofaxons from primary
sensory neu ro ns, low!;r motor neurons (LMNs),
and prega nglio nic and postganglionic autonomic
neurons. The primary sensory axons have sensory
receptors (transducing elements) at their peripheral
ends contiguo us with their initial segments. The
proximal po rtion o f the axon enters the eNS and
terminates in secondary sensory nuclei associated
with reflex, cerebellar, and lemniscal channels.
LM Ns in the anteri or ho rn of the spinal cord send
86
axons, via the ventral (anterior) roots, to travel ir
peripheral nerves to skeletal muscles, with w hirr,
they form neuro muscular j unctions. A utonomic
preganglionic neuro ns send axons via the ventra
roots to terminate in autonom ic ganglia.
Postganglio nic neurons send axons into splanch.....
o r peripheral nerves and form neuroeffector junc
tions with smooth mu scle, cardiac muscle, secrE
tory glands, metabo li c cell s, and cells of the
immune system.
 ~ ral Nervous System (PNS )-Introduction REGIONAL NEUROSCIENCE

,-,<, ':' {C1abovespiJ),)lC1 nerveexits

vertebra

51

lumbar disc p ro trusion doe, not usuall y affe ct ner ve e )( iling

ab ove disc. liI \(> r<l 1p rot ru sio n.) t el ise levell4-5 affects L5
spi nal ne rve, not l 4 spina l nerve. Protrusion at d isc lev!
L5-5 1 affects S1 spinal nerve, no t L5 spinal ne rve

C()(lUSmedullaris
ft ermin3\ion o f
spinal cord )

.-. terminaJ
- "'" part) - ---..:-'.

S'-+~,;:-

S2 + --:-r'.,-i

"""'"
P""":nal ilium
Pill pan} - -'Ill S4-+~

55- ="
~ Cervical nerves
_ ThOlao c nerves
Me d ial p rotrusio n al disc leve l l 4-5 rare ly affects L4 sp inal
He _
_
lumbar nerves
Sacraland coccygeal nerves
nerve b ut may a ilect l S spina l ne rve and sometimes 5 1-4
spinal ne rv es

m
RE [1.2 : RElAT[ONSH[P OF SP[NAL NERVE ROOTS TO VERTEBRAE _ _ _ _ _ _ __
ich
Dorsal (posterior) and ventral (ant erior) roots of
<=-5pi nal cord segments extend from the spinal
as groups of perip heral axo ns invested initially
- meninges. As the axon s enter the PNS, th ey
Inc iate w ith Schw ann cells for myelination and
, !' .;poort. Th e roots exit through the in tervertebral
-.mi na, compact openings between the verle-
. w here herniated disks (nucleus pulposus) can
impin ge o n th e nerve roo ts, producin g se nsory
and/ o r motor symptoms. Sensory and mo tor axons
enter th e dorsa l and ventral rami of peri pheral
nerves. Auton om ic axons course from th e ventral
roots into the w hite (preganglio nic) rami commun i
cantes/ and po stga ngli onic axon s co urse throu gh
the gray rami co mmuni ca ntes.
87
REGIONAL NEUROSCIENCE Periphera l Nervous System (PNS)-lntrodudi Ol

~n .....)' e h.n ""I. _rfi e.

OXT~AI r>.1 ~v()\..s SY STE,~ \ PfRII' HtJV,1 "' [~VOUS SHTh,

""~""pli<
,

(IIe.Ofl'~ !"-~ ", "'""

M......,.,.... p.k muid~
<'Trim""", ""~~'''Iled

~I
1jlI""'e

I ,,, ,, ~, mol,,,. ""uron ). rClkin'

""'"
""''''A
~!ff'l! Sk.lollal ruu od.

FIGURE 11.3: SENSORY CHANNElS: REFLEX _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Prima ry sensory axons communica te w ith sec muscle spindles directly w ith LMNs involved in
ond ary sensory neurons in renex, cerebellar, and muscle stre tch retlex contraction; this is the only
lemniscal channels carrying transd uced inform ati o n monosynapti c reflex see n in th e human CNS.
from the periph ery into th e e NS. The reflex chan Po lysynapti c reflex channels are directed partic u
nels interconnect primary sensory axons with larly toward fl exo r (withdrawal) respo nses throu g.~
anterior horn cell s (LMNs), thro ugh one or m ore o ne or more interneurons to produce coordinatE":
synapses, to achieve an unconscious reflex m o tor patterns of muscle ac tivity able to remove a por
response to sensory input. These respo nses ca n tion of the body iro m a potentially damaging or
be elicited in an isolated spinal cord devoid of con offending stimulus. This polysynapti c channel car
nections from the brai n. The m onosynaptic retl ex sp read ipsilaterally and contralaterally through
c hannels connect primary senso ry axons from many segments.

Sf<>t<,,~ Ch,nnel, C..,.,b.ILa,

C[ NTRAL -:f.RVOvS SYSIr.~1 I'tRIf'H[R....LNERVOUS S'l'STEM

III (",""~ I,,m

P,imowy ocnry nO""""

',uppOn' mu."''' 'I'<n(He,
Spin!)cO',O'IwIl.,I"ct ~" I ~ '1)I1<it>n or~1n,
Prim".,-""",,,"')'
II" ~.u,.' r><)(>\)"I"'1
........"" cNlIx><t) in

dot.;.:>Itoo'~!!lj!)n

SKOf>d~'l' ...-or'j. " ..........

~
in <pn>;>1 coni
(0) J.. CI.rk.. , nucleus!
'" Iy~jn ,!<'m (...g..
I.*,."I~"MOI" ouc\.,ou, l

FIGURE 11.4: SENSORY CHANNElS: CEREBELLAR _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

-

,. Pet!.in,
c,1!!I!'lI Mu ,cI<
<plnd le

Primary somatosensory axons carrying uncon projectio ns through spinocerebellar pathways.

sciously processed information from m uscles, The dorsal and ventral spinocerebellar trac ts cam
jo in ts, tendons, li gamen ts, and cutaneous sources information from the lower body (16 and belo\.\ ._
enter the CNS via dorsal roots and synapse vvith The rostral spinocerebellar tract and th e cuneo
seco ndary sensory neurons in the spinal co rd or cere bell ar tract carry information from the upper
the ca udal brain stem. These neurons co nvey body (above T6). Polysynaptic indirect spin o
information from the periphery to th e ips il ateral ce rebellar pathways (spino-olivo-ce rebellar and
cerebellum by se nding secondary sensory axonal spino-reticulo-cerebellarl also are present.
88
roduct. 1" eral Nervous System (PNS)-Introduction
Or
E 1/,5: SENSORY CHANNELS: LEMNISCAL _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
sensory axons carryin g information des
'or conscious percepti o n arise from receptors
.....:Qeriicial and deep ti ss ue. These axons enter
='-5 via the dorsal roots and terminate o n sec
...ar\ sensory nuclei in the spinal cord or the
5~e m. Secondary sensory axons from these
..e cross the midlin e (dec ussate), ascend as lem
pa thways, and terminate in the contralateral
us. These thalami c nuclei then project to
':c regions of the primary sensory cortex,
-=rne-grained analysis of incoming co n
perceived info rmation takes place.
o5oensory information is directed into either
- .opa thic or epicritic channels. Epicritic info rma
- "line discriminative sensation, vibratory sensa
oint position sense) is transduced by prim ary
r '""n neu ron s th at send myelinated axo ns to
-50 in the medulla, nuclei gracilis (l ower body,
below), and nuclei cuneatus (upper body,
e -6). Nuclei graci lis and cuneatus give rise to
~i a l l emni sc u s, a crossed secondary sensory
.!\ th at termin ates in the ventral postero
\ Pl) nucl eus of the thalamus. This thalamic
REGIONAL NEUROSCIENCE
1-~
11
,
.,
I. Porli ..
.'"
nucleus has reciprocal connectio ns with cortical
neuron s in the postcentral gyrus (Brod mann areas
3, 1, and 2). Protopathic information (pain, tem per
ature sensa ti on, light movin g touch) is transduced
by primary sensory ganglion cell s that project to
neuro ns in the dorsal ho rn of the spinal cord via
small myelinated and unmyelinated axons. These
spinal co rd neurons give rise to th e spinothalami c
tract (spin al lemniscus), a secondary sensory path
way that terminates in separate neuronal sites in
the VPl nucl eus of the thalamus. This portion of
the VPL nucleus communicates w ith th e primary
sensory cortex and a seco ndary area of the
somatosensory cortex posterior to the lateral post
central gyrus. Some unmyelinated nocicepti ve pro
topath ic axons that termin ate in the dorsal horn of
th e spin a! co rd interconn ect w ith a cascad e of
spinal co rd interneuron s th at project mainly into
the reticular formation of the brain stem (spinoretic
ular pathw ay). This more diffuse pain system is
processed through nonspecific thalamic nuclei,
lead in g to pe rception of excruciating, to ng-lasting
pain.
89
REGIONAL NEUROSCIENCE Periphe ra l Nervous System (PNS)-Introducti o-

CENTRAL NERVOUS SYSTEM PERIPH ERAll\jERVOUS SYSTEM

Conical
upper motor / "'....
neuron -

Corticospinal
Brain stem

tracl
upper motor neuron

(red nudeus,
superior coliiculus,
vestibular nuclei, upper motor
retic:uIM nuclei) neuron tracts

J. Perkins m
MS.MfA,
0 1lON
'--':"."
"
FIGURE 11.6: MOTOR CHANNELS: BASIC ORGAN IZATION O F LO W ER AN D UPPE R MOTOR
NEURONS ____________________________________________ ~I

Lower motor neurons are found in the anterior
horn of th e sp in al co rd or in motor cranial nerve
nuclei in th e brain stem. Their axons exit via the
ventral roots or the cran ial nerves to supply skeletal
muscles. LMN synapses with muscle fibers form
neuromuscular juncti ons and release th e neuro
transmitter acetylcholine (AC h), which acts on
nicotinic receptors o n the skeletal muscle fibers.
A mo tor unit consists of an LMN, its axon, and
the muscle fibers the axon innervates. LMNs are
90
regulated and coordinated by groups of upper
motor neuro ns (UMNs) found in the brain. Brain
stem UMNs regulate basic ton e and posture.
Cortical UMNs (from co rti cospinal and cortiCD
bulbar tracts) regulate consciously directed, or
voli tional, movements. The cerebellum and the
basal ganglia aid in pattern selection and coordI
natio n of movement via connectio ns with these
UMNs; they do not connect w ith LMNs directh.
JG :leral Nervous System {PNS)-Introduction REGIONAL NEUROSCIENCE

CENTRAL NERVOUS SYSTEM PF.RIPI-IfRAL NERVOUS SYSTEM

Limbic forebrain slructures

(e.g~ amygdalo id nudeus,

some conica l 311'0'15)

Hypolhalamus

~ ~tem
-:"I and /
'.-./ -1-".<'--
;>'" .---
/ 1 Hypoph ysea l
portal
sys tem

Anterior
pituitiHY
hormone~

Sensory ncuron in
dors.. 1root ganglion
neuron
In lermediola(eral
.:ell column in laleral Refle)( connections
horn of spinal cord
(T1 -l2)

Ta rge'l t i ~~lI e

Sympathetic C. Jt~;

ins
gan glion cell

~--------~=~~ .:;-"
~ :. -~':,:
I

-- ~ ,

RE 11.7: AUTONOMIC CHANNELS _ _ _ _ _ _ _ _ __ _ _ _ _ __ _ __

" .~
"<"'. p:

_=.=.,glionic neuro ns {o r the sympathe tic nervous
rem (SNS) are found in the lateral horn (inter
{ lateral cell column ) of the thoracolumbar (T1-
a ln _ spina l cord. For the parasympathetic nervous
71 (PsNS), these neurons are found in nuclei o f
.......1nerves (CNs) III, VII, IX, and X and in 52-54
l""mediate gray matter of the spinal cord (a
jOsacral system ). Preganglio nic axons exit the
5 \ ia cranial nerves or ventral roots and termi-
!-e 'n chain ganglia or collateral ganglia (SNS) or in
\. amural ganglia in or near the organ innervated
... 5]. Postganglio nic axons innervate smoo th
muscle, cardiac muscle, secreto ry glands, metabolic
cells, and cells of the immune SYSlem . The SNS is a
fighl-or-lligh t system that respo nds to emergency
demands. The PsNS is a hom eostatic, reparative
system active in the m ore quiescent digestive and
eliminative functions. Prega nglionic responses are
coordinated by autonomic " UMN" equivalents
from the brain stem (autono mic centers), the hypo
thalamus, and limbic forebrain struc tu res. These
central regulatory system s coordinate auto no mic
responses that affect bo th visceral func tions and
neuroendocrine o uttlow from the pituitary gland .
91
 Krause' <;
cY1ffk4

free nerve
ending
.;
,

,~"l{1
, \ ""
Fre':! nerve
ending

Sebaceous gland

Meissner's I
I'.lerve plexus
corpu~de
around
hilir follicle

P,d,;" cmpc,7 M erke l", d'ISC

Free nerve ending Ru fii ni term inals \ Pacinian corpuscle

Ba~ e me nt membr.lne

-- ,
'~.
Axon terminill
M il0chorlddon
(ifl
Lobula ted nucleus
1 $1, ' <:~ O ')
'; ~ ~t;.,. :.~ -' S(.hwann cell
~jf 51S;~ ~l
'~~
~ ;#

Basil
O:! pitheliaJcells ~---

Cytoplasmic
p r Olr u:)ion

M itochondria

Expanded
Axon
axon terminal

~lp
~.,_ .h_
~cv.;/:~:.~~ SchwJnn cells

CIIl!N
"..; .... .' ~
C. Detail of Merkel's disc D . Detail of free nerve ending

FIGURE 11,8: CUTANEOUS RECEPTORS _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ __ _

Glabrous skin and hairy skin contain a variety of
sensory recep tors for detectin g mechani cal,
therma l, or nociceptive (co nscio usly perce ived as
painful) stimuli ap p!ied at the body surface. These
receptors in clude bare nerve endin gs (n ociception,
th ermal sensati on) and enca psulated endings. The
latter includ e pacinian co rpuscles (fast-adapting
mechanoreceptors fo r detectin g vibratio n or bri ef
tou ch), M erk el's disks (slow ly adaptin g mechanore
ceptors for detectin g maintain ed deformation o r
92
sustained touch at the skin); Mei ssner's corpu sclE"5
(fast-adapting mechanoreceptors for detecting
moving touch); Ruffini endings (s lowly adaptin g
mechanoreceptors for detectin g steady pressure
applied to hairy skin), hair follicle receptors (ra pia
adap ting), and end bulbs of Krause (possibly
thermoreceptors). The initial segment of the pri
mary sensory axon is immediately adjacent to the
se nsory receptor.
)duction Peripheral Nervous System (PNS)-Introduction REGIONAL NEUROSCIENCE

, -------::==.::------, --.J Pressure lL-__

To amplilier
/-------~
Generator potential

" ~------1f--l Sl node

"---- - -t---My,, ";i" sheath
Action potential

A. Sharp "on and oif' c hanges in p ressure at stan and end o f pulse applied to lamellated
capsule ca psule are transmilteo to central axon ano p rovoke generator pOlentials w hich in turn
may tri gger action potc ntials; there is no response to a slow change in pre ssu re grad ient
Central core Pres,ure at central core and, <lcc ordingly, generator potentia ls are rapidl y dis~ i pat ed by
yiscoelastic p ropertib o f cap sule. (Action potentiills may he blo cked br pressure ill a
Unmyelinatl"d
axon terminal node o r by dr ugs)

r - - -- ------:.,----- ---- --.J Pressure 1

To amplifier
". . - - - - - - - - - - J- "'._ _
Generato r po tenti,!!

B. In ab sence of capsule. axon

responds to slow as well as 10
rapi d changes in pressure. C ener.'Hor
potential dissipates slo wly. and th ere
i~ no o ff' response

Action potential
Pressure Na+
+ +

Pressure applied to <lxon terminal direcliy' or via capsule c<luses incre ased perme<lbilily o f membrane
to l\ja+, thus set1in g up ionic generator current through 1st f'od e

+ + + + + +
- -
~~p
-.' ,
,
@:@'"
_.,-,",":-- "

1I9N
1""... ::
If resu ll ant depolari zation al 1st (lode is great enough to reach threshold, an action potential appears
which is propagated <lIang nerve fiber

-
:Ies
'lCU RE 11.9: PACINIAN CORP USCLES _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

?acinian corpuscles are mechanoreceptors that
P21sfo nn mech anical fo rce or displacem ent in
oiir"ge-diameter primary sensory axons into actio n
. {entials. The mechanical stimulus is modified by
--;e viscoelastic properties of the contributin g
.-mellae of th e pacinian corpu scle and th e associ
~ accessory cells. An action potential results
wh en a generato r potential of sufficient magn itude
to bring the initi al segment o f the axon to threshold
is elicited. The o nset and cessation of mechan ical
deformation enhance io nic permeability in the
axo n, o ptimizin g the physio logiGll respon se of the
pacin ian corpu scle to vibrato ry stimuli.

93
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)-Introductio r.

A. Carotid body .1nd carotid sinus

.1 Vagus (Xl nerve

ical
G I () ~ $(l philry n ge a l (IX) nerve ~
sympathelic trunk gang lion

CarOlid sinu~ ner...e ~ f- ~el rOfnand ibula r and

Ve ins from carotid bo dy

- l i n~ual vein
External carol id

'- - [ntemil l jugular vein

Caroti d body

A rt ery 10 carot id body ............

'- - Jnle rna l l.:a rol id a rte ry

Superior thyroid Jrtery -............

f-- -Caro tid sinus

Common cMolid 4111ery ___

C. Carotid sinus
B. Carotid body
Synap ti c Nerve
endin g

~!p
Basement
rnembrilnC Type [ iglomusJ cells
I
End othe liJ[ c e lls ::l~
.....: : ;

FIGU RE 11.10: INTEROCEPTORS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Interoceptors, including internal nociceptors,
chemoreceptors, and stretch receptors, info rm the
eNS about the internal state of the body. The
ca rotid body, a specialized chemorecepto r for
detecting carbo n di oxide (in a hypoxic state) or, to
a lesser extent, low blood pH resulting in increased
respira tion, is associated w ith afferents of eN IX
projec ting to caudal nucleu s so litarius in th e
94
medulla. The carotid sinus, a thin-walled region 0:
the carotid artery, contains encapsulated and b c.-~
nerve endin gs th at act as stre tch recepto rs. Th ~
stretch receptors respond to increased art erial
pressure as baroreceptors, eli ci ting a refl ex bract.
cardia and a decrease in blood pressure via
primary afferents of eN IX pro jecting to caudal
nu cleus solitariLl s.
 ------- ------------------- - - - -

Id u - ral Nervous System (PNS)- Introduction REGIONAL NEUROSCIENCE

,~e; ,,",." corpuscle

of sweat gland

r Cuticle
- .:' "n al ' ''0''''' ,

Connective De rma l
:r~s ue layer pap illa
lo r papilla ry
layer)

~ .. ea t " " " " '-

Pac inia n
corpuscl e

Ar(ery

Subcutaneous
artery and vein
Elastic fjbers

Skin liga me nts Ire lin<lcula c Ulis) Cuta neous nerve

MOl o r (auto nomiC! nerve

11.11: SKIN AND ITS NERVES _ _ _ _ _ _ _ _ __ _ _ __ _ _ _ _ __

n b supplied w ith a variety of receptor types
=-- : ..... re 11.8) that transduce slowly and rapidly
~ mechanical sti muli and deform atio n into
.a l impulses in prim ary afferent fibers. The
~r ve endings are associated mainly with
=ptors, peripheral arbo rizatio ns of unmye
Ed axons. Some nocicep tors and thermo
ors are associated w ith small myelinated
axons. These axons contribute somatosensory
info rmatio n collec tively to the spino tha/amicj spin
oreticular lemniscal sys tem for protopathic sensa
ti on. The more com plex encapsulated rece ptors
co ntri bute somatose nso ry informati on to the dorsal
co lumn/ mediallemni scal system fo r epicritic sen sa
tion and are associa ted with larger myelinated
axons.
95
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS) - Introductj o~

SY""ti''''':~::::~~~',,~.,.,
_ r a~~~~~. " r

iI,
Syn~plic ",
Synapli<:
Folds

- - - Mye lin sheath

Active zone
, I Neurilemma
Axoplasm
Schwann cell process Schwa nn cell
Acety lcholine
Mitochondria
I
Basement membrane

Presynaptic membrane
Active zone

Synaptic trough

Sarcolemma
Nucleus o f
Myofibrils musde celJ

SynaPtic deft
Postsynaptic
membrane
Junctional fold ,f.l!'~
Sarcoplasm
Acety lcholine receptor sites
~(~;@\
....-
:

FIGURE 11.12: NEUROMUSCULAR NEUROTRANSMISSION _ _ _ _ _ _ _ _ _ _ __

A xons of lMNs that synapse on skeletal muscle
fibers form expanded termina ls ca lled neur(}
muscular junctio ns (mo tor endplates). The motor
axon loses its myelin sheath and expands into
an extended terminal that resides in a trough in
th e muscle fiber and is covered with a laye r of
Schwann cell cytoplasm. The postsynaptic mem
brane is thrown into secondary folds. When an
action po tential invades th e mo to r terminal,
96
hundreds of synaptic vesicles release their pack f ~
of ACh simultaneo usly into the synaptic cleft. The
ACh binds to nicotinic receptors on the muscle
sa rco lemma, initiating a moto r endplate poten ti""
(EPP), which is normally of sufficient magnitude L:
result in the firing of a muscle ac tion potential, Ie,
ing to co ntraction of the mu scle fiber. A single ~
cl e fiber has only one neuro mu scu lar junction, tL
a motor ax.on may innervate multiple muscle fib
 ra l Nervous System (PNS)-Introduction REGIONAL NEUROSCIENCE

A. Smooth muscle B. Gland (submandibular) C. l ymphoid tissue (splee n)

!d e ce lls (cut) Sy,,,p;,,h, ,;, te rmi nal ending Bloo d vesse l lumen Adv entiti,,1 zon!:'
cell ca p enclosing nerve axons Varicosity n!-' uroe rrec lor

. II Sec,,,,,," t ell ,I Ii lermin ,1ls

cap enclo sing among T lymph oC)!tes in

nerve axons peri,lrteri olar lymp ho id sheath

- ,,"
.-:.~

..,-
,

-
RE 11.13: NEUROEFFECTOR JUNCTIONS
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

-"omic postganglion ic axo ns form neuroeffec
un ctions with cardiac muscle, smooth muscle
secretory glan ds (6), metabolic cell s such as
":Ioa to cytes, and cells o f the immune sy stem (C).
.rse nerve end in gs use mainly norepin ephrin e for
~ >iS and AC h fo r th e PsNS. Th ese endin gs do
:orm classic eNS Of motor endplate synapses
- ;:erminale as neuroeffector junctions, releasing
otra nsm itter into in tersti tial spaces. This per
mils w idespread diffusio n of the neurotransmitter
as a paracrine secretio n, initiating postsynaptic
respo nses on cells with appro pr iate receptors
(includin g many types of immun e ce lls). Som e
close appos iti ons also are found, such as SNS end
in gs o n lymphocytes. Sm ooth mu scle cells th at are
not innervated are cou pled by gap junction s and
can contract together when the innervated smooth
muscle cell contracts.
97
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)-Somatic

~~p CJ(ij(\I
, ',';.',: .
levels of Principal Dermatomes Tt 0 Level oi umbilic us
(5 Clavicles T12 Inguinal or groi n regions
C5, 6, 7 Lateral pa rts of upper limbs ll, 2, 3, 4 Anterior and inner surfaces of lower lim bs
C8; T1 Media l sides of upper limbs L4, 5; 51 Foot
(6 Thumb L4 Medial side of great loe
(6,7, 8 Hand lSi 51, 2 Outer and posterior sides 01 lowe r limbs

(8 Ring and linle fingers S1 l atera l margin 01 fOOl ..nd little toe

T4 l evel of nipples 52, 3, 4 Perineum

FIGURE 11.14: DERMATOMAL DISTR IBUTIO N

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
A dermatom e is th e cutaneous area supplied by a
anesth esia (total loss of se nsation), in th e region
single spinal nerve roo t; the eel! bodies are located
in dorsal root gangli a. The spinal nerve roots are
distributed to structures according to their associa
tio n w ith spinal cord segments. The nerve roots
supplying neighbo ring derma tomes overlap. Thus,
secti on ing or dysfun ction of a single dorsal root
produces hyp oesth es ia (diminished sensation), not
98
predom in an tly supplied by that dermatom e.
Dermatomal anesthesia req uires damage to at I
3 dorsal roots: the central dorsal root and the roo.
above and below it. Knowledge of dermatomes
impo rtant for identifyin g th e location of periphe-"
nerve root lesions and distingu ishing them from
peripheral nerve lesion s.
 -------

- SO rnalll: '"'eral Nervous System (PNS)-Somatic REGIONAL NEUROSCIENCE

Greate r occipital n.
(from dorsa l ramu s of C2)

"'" :ces50ry (XI) n.

l e!.ser ocopital n .

Sternocleidomasto id m.
(upper end)
Geniohyoid m.

Thyro hyoid m.
Nerves to anterior and lateral
rectus capitis mm., and lo n8u 5
~ rool {descendens
capitis and longus colli mm.
~"';~ i) of ansa cervicalis

Nerves 10 longus cap itis, lon8uS

colli and levator scapulae mrn.
n:erior root (descendens
ct-rvicalis) of ansa ce rvicalis Trapezius m.

Nerves to longus colli, scalenus

Transve1secervical n . - - - anterior and scalenus medius

O m o hyoid m. Communicatio n
to brac hial plexus
Sternothyroid m. - - - ---1,

Ansa cervicalis
fa nsa hypoglossil
Phrenic n.
~Iej domas to i d m.
.;.' end)

5",,,,:1,,,,,,:e",,, nn. (medial,

intermed iate and lateral)

Motor fibers
Sensory fibers
Proprioceptive fibe rs

U RE 11.15: CERVICA L PLEXUS _ __ _ _ _ _ _ _ __ _ _ _ _ __ __ __

- 'e cervical plexus lies deep to the sternocle ido
.c:o id muscle. Its branches convey moto r fibers
ilclny cervical muscles and to the diaphragm. Its
_ ry fibers convey exteroceptive inform atio n
'TTl parts of the scalp, neck, and chest, as w ell as
"
ra. :fIri oce ptive informati o n fro m muscl es, tendons,
an d joi nts. Sympatheti c sud o motor and vaso motor
fibers travel w ith this plexus to blood vessels and
glands, The superficial branches perforate the cervi
cal fascia to supply cutaneous structu res; the deep
bran ches su pply mainl y muscles and joints,

99
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)- $omatir

Ventrc1 !
ram i {
CJ

C4

C5
CJ} ven~ral
(4 ram!
C5

An terio r scalt'ne m, ~ An t('rior scalene m.

R. common carotid a. / :
Brachial plexus A Brach ial ple)(us
~

Righ i phrenic nerve ------ left phrenic nerve

R. subclavian a.
L subclavian a .
R. vaguS (Xl n. ~~
L. common carotid il.
IMernal thorJ cic a.---------;
vagus (Xl
Brach iocephalic "
11.

trunk -~-----: Intern al thoracic ,l"

R. periC(1rdia Thor(lcic cardia!.: nn.

cophrenic <I.
L. pericardiaco
Superior phrenic a.
vel),) cava
recurrent
PeriCMdial laryngeal n.

bran ch of

phrenic n.
Root 01
left lung

Root oj

right lung

Phrenicoabdominal
branches ot phrenic nn.
Oiilphragmatic pleur.l
(to inferior surface
(cuO
iasl inal
of diaph ragm)
pleura

Phrenic n n. :~~ /"" ,

(m otor and

sensoryl

Lower intercosl;:li nn.

10
Innervation of diap hragm

~~p c IlfiN
" , 0, ""

FIGURE 11.16: PHRE NIC NERVE _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ __

The left and right phrenic nerves are the 1110 to(
nerves that supply both sides of the diaphragm
from the C3, C4, and C5 ventral roots. The phrenic
nerve also contain s many se nsory nerve fibers,
vvhich supply the fibrous pericardium, the media
slinal pleura, and central areas of the diaphrag
matic pleu ra. Sympathetic postgan glionic nen E
fibers also travel with thi s nerve. Dam age to t h ~
phrenic nerves ab oli shes inspiration, wh ich res.....
in respiratory failure,

100
i)-So "", 'iIIfr.1 1 Ne rvous Syste m (PNS)-Somatic REGIONAL NEUROSCIENCE

Spin;!1 nerve tr unk

spmae m . Co llateral branch o f intercostal n.
Sp in al ganglion SubcostJI mm.
r- Medl,11 and
I r- I" te,,1b ra nches 01
~ ,,",,,,1 filrY! U5
. -;'i1lral

latissim us clUfS; m.
Serratus

,
r

Iransv~e
ligaments Co llateral bra nc h

.J'het ic trun k (p,o>\,,,",, i nlerc.ostal

rejoin ing main ''','leiio' '''''",,' cutilneous
cutaneous br,1nch branch Jnd il nterior
memb rane on externa l
Ra mi commu nicantes cutaneous branc h
inte rcostal m.
~ray and wh ite) _ _-' Communica tin g branch of interco stal n.

Transversus abdominis
Internal intercostal m.

fxteroal
-=01 -
~::: ',"ot;: ':;:'::I~
I m.
membmne

"'ill n.

External
oblique m.

-
E 1.17: THORACIC NERVES _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ __

_ ")airs of thoracic nerves are derived from
.;"aJ and ventral roots of their correspond ing
~. These nerves do not form plexuses; they
e cu taneous branches to the thoracic def
IE'5 anel send other sensory fi bers to deeper
;;:Iruc tures, vessels, the periosteum,
parieta l pleura, the peritoneum, and breast tissue.
Th e th o racic nerves also send motor fibers to mus
cl es o f the thoracic and th e abdominal walls and
carry preganglionic and postganglionic sympa
thetic nerve fibers into and ou t of the sympathetic
cha in.
101
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)-Somati<

5 Roots
3 Trunks !ventral rami)

3 Ve ntral divisions
3 Dorsal divisions

3 Cords To subdavius m.;

(5,6

lenninal
branches
(2lrom
each cor d )

Musculoculaneous n.:
( (4),5,6,7
l ong
tllOracic n.;
Ax illary n.; (5,6,7
(5.6

.l;h..~' '."-~~ - ,.-...7." . .

C5, 6,7,8

'.
Radial n.; (5,

~
6, 7 8; T1 ).4- .," . ~~
~
-.,.~ ~ ~ . 1 st intercosta l :,,_
M edial pectoral n.; C8; T1
M edian n., ((5 ),
6, 7.8:Tl
. ,-'. '
",e, ',-- / "?'"
- .
~ Medial cutaneousn.offorearm;C8;Tl

~~p
Medial c uta neo us n. of arm; T1
Some contrib utions
Ulnar n.; Thoracodorsal n.; (6, 7, 8
inconstant
((7), B; Tl
- U!ill.
... ,~,., ;

Supraclavicular Branches Infraclavicular Brancnes

From plexus rooLS From lateral cord Ulnar 07
To longus colli and SCdlene mm C5, o. 7, 8 latetal pectoral CS,6,7 Medial fOOl of median C
Dorsa l scapula r C, Musculocutaneous C(4). 5, 6,7 From posterio r cord
Branch 10 phrenic C, l a tefill root o f media n 05), 6 , 7 Upper subscapular (5.
Long thora cic C5, 6, 7 From media l cord lower subscapular o
From superior flUnk Me dial pecto ral C8; TI Axillary (d rcumtlex humeral) C;
Suprascapular ( 5,6 Medial c utaneous n. o f arm T1 Thoracodorsal (-:: -
To subclavius m. ( 5, (, Medial c utaneous n. of fo rearm C8; T 1 Radial (S .;, -

FIGUR E 11.18: BRACHIAL PLEXUS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

The brachial plexus is formed by the union of the
ventral roots of C5, C6, C7, ca, and T1, with a
smaller contribution from C4. Sensory and symp a
thetic fibers also distribute with the brachial plexus.
The roots give rise to 3 trunks, 3 ventra l and 3 dor
sal divisions, and 3 cords, as well as numero us ter
minal branches, the peripheral nerves. This plexus
is vulnerable to birth injury (superi or plex us para-
102
lysis), with paralysis of the deltoid, bi ceps, brach.
and brachioradialis muscles, with sparin g of the
hands, and sensory loss over the deltoid area af't,.
the radial aspect of the forearm and the hand.
Pressure of a cervical rib can ca use inferio r ple,"
injury (C8, T1 injury), whi ch results in paralysis (]I
small hand muscles and flexo rs of th e hand, ", i'
ulnar sensory loss and possible H or ner's syndrc."-I'
i)-Soma -'",-",ra l Nervous System (PNS)-Somatic REGIONAL NEUROSClE-. CE

:Onlribuf, '"' Anterior (palmar) view Posterior (dorsal) view

om C4

/
0 0-'"
r am ~
ooIIP'XJavicular nerves
cervical
~
"',
__- -:':'~:~~~~:.:::~I;"
cervical
ne rves
plexus - 0 , 4)
_, - CJ , ' 1- --:'--_

..i.... nerve
. ' ior latera l
::..G..""1eous nerve cuta neous nerve
1(5, 6) _ --ft of arm (C5, 6)

- . nerve
~-iorlateral Rad ial nerve
-o"oous nerve , cutaneous
.rn ICS, 6) nerve of arm ((5, 6. 7, 8)
Intercosto Inferior lateral cutaneous
bfachial
nerve of arm
nerve (T2 )
Posterior cutaneous

----
and medial
nerve of fo rearm
~ c u ta neou s cutaneous (C[5], 6, 7, 8)
_ ~ oi iorea rm nerve of arm
( oo \n!:>l.:. " ;711 (C8, TJ , 2)
:' om T ~ .'lJJ part of
'""\o:!.O.. !oculaneous
Medial Lateral cuta neous
~g u s ( 'J .....e cutaneous
nerve of ne rve of forearm
( aiE'f1e ........
(CS, 6, !7l)
- B forearm
\C8, 11 ) (terminal part of muscu/a
lrco,~" " cutaneous nerve)

Ulnar nerve (C8, T1)

r....---,-A.'---.." Dorsal branch
Radial nerve
Superficial branch
and dorsal digital
branch and dorsa l digital branches (C6, 7, 8)
branches
/f'.I hna, digital
branches
c- .
C'
Median nerve
C5 ~ ~~'- Proper pa lmar
C digital branches
....-,,"
_ :.JJ branches
~ Nale: Division variable between ulnar and radial
C: - 8) innervalion on dorsum 0 1 ha nd and often aligns wi th
middle of 3rd digit instead of 4th digit as shown

:E 11.19: CUTANEOUS INNERVATION OF TH E UPPER LIMB

_ _ _ _ _ __ _ _ __

-" eous innervatio n of the limb derives
-if musculocutaneous, axillary, radial,
dn d uln ar nerves. These nerves are the
- bra nches of th e brachial plexus. Unlike the
re rv e roofS, the cutaneous senso ry distribu-
tio n of these periph eral nerves to th e upper limb
does not overlap. Thus, a peripheral nerve inj ury or
compression results in a zone of anesthesia corre
spondin g to its distribution.
REGIONAL NEUROSCIENCE Periphe ral Nervous System (PNS )-So ma tir:

Dorsa l scapular n('rve Suprascapular nerve Dorsal Scapular Ner \l~ (5 ),

Suprascapular Nerve (5, (6),
A)(iliary Nerve (C5, (6) and
Supra5pil,,,tu s m. Radial Nerve (5, (6, (7, (8; T1)
Above Elbow (viewed from behind )

/Deltoid m.
levator scapulae m.

(supplif'd al<'(J tJy

hranches irom (5 . /Teres minor m.

ilnd C61

..........-- Axillary nerve

UPller l;lIeral
cutaneous n.
Rhomboideus ---- ~ o f arm

!nlnor m.

0.---- Rildial nerve

Rhnmboideus

miljorm. culant!Ous n.

alarm

In(rilspinatu~ m.

Teres major m. c utaneous n.

(If for e Olrm
Lower suh scapuldr n.
late ral
Pos terior cutaneous n. of arm
in termuscular
(brallc h of radial n. in axilla) septum

Brachialis m .
Lo". he,d (Itltera l part)

Tri cep~ bracnii m. la le rcll hl'iUJ _ _ Brachiorad ialis nl .

{
Medial head ~
.tensor ctl fpi
-. ... radialis longus m .
Triceps tendo n 1 ~:

M~ iill eptc.ondvle ,r". Exten sor ca rpi

radia lis b revis m.
O le<:r..rnon ~
~
~~p
Anconeus m .

Ex t"'n~o r digitOrum rn. 1;

UgN
Extensor ca rp i u l n~ris m . ~ '\....111

FIGURE 11.20: SCAPULAR, AXILLARY, AND RADIAL NERVES ABOVE THE ElBOW _ _ _ __
The dorsa l scapular nerve (C5) supplies the levator
scapulae and rhomboid muscles; it aids in elevation
and adduction o f the scapu la toward the spinal
co lumn. A nerve lesion leads to lateral displace
ment of the vertebral bord er of the scap ula and
rhomboid atrophy. The suprasca pu lar nerve ((5,
(6) su pplies the supraspinatus and the infraspina
tus muscles; it aids in lifting and in o utwa rd ro tat ion
o f the arm. A lesion results in weakness in the first
15 of abdu ctio n and in external rotation of the
arm. The axillary nerve (CS, C6 ) su pplies the forearm.
104
delto id and teres minor muscles; it aids in abduc
tio n of the arm to th e horizontal and in outward
rotatio n of the arm. A lesion results in delto id
atrophy and weakness in abduction from 15 to
90. The radial nerve (C S, C6, 0. C8) supplies the
triceps, anconeus, brachioradialis, extensor carpi
radialis, extensor digitorum, and su pinator muscles: ,
and it aids in extension and nexion of the elbow. ZOf
A lesion leads to paral ysis of extension and flexion
of th e elbow and paralys is of supination of the
l,
 phe,al Nervous System (PNSj-Somatic REGIONAL NEUROSCIENCE

-4c------R;,di;,'lne~e
Radial Nerve in Forearm
(5, (6, C7, (6; T1)
(vj~~d from behind and

r ---[)",p terminal bra nc h 51ightly laterally)

i-- Anconeu s m.
Brachior,ldialis m.
Extensor carpi radialis longu s m.
-- Supinator m.
-- Exte nsor carpi radi alis brevis m .
~---- Extensor carpi ulnaris m. Extensor supilliltor
group of muscles
Extensor digitorum m. and
extensor digiti minimi m.
Extensor indieis m.
Extensor pollieis longus m.
' -_ - - Abductor poI rici~ lo ngus m.
Extensor pollicis brevis m".____-->

.l- - Posteri o r inte rosseous n.

(deep bran ch of radial n.)

- - - - Superficial branch
of radial n.

From {upper
auxiliary
lateral
nerv e cutarleous n.
olarm

Lower
lateral

cutaneous n.

of arm - - - - . /

Posterior
From cutar.eotlS n.
radial of arm ____J
P01;tenor
aJianeous n.
o fforearm
s..".-riirioI
Dorsal branch 01

digital nn. radial n. - - - - -_ _ -+

Cutaneous inne f'Villion fro m

r<ldiut and axillary nentl!S

L RE 11.21 : RADIAL NERVE INTHE FOREARM _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

- e forearm, th e radi al nerve (C6, e7, C8 ) sup
s. motor fibers to th e ( 1) exte nsor carpi radialis,
_ extensor digitorum, (3) extenso r digiti V,
- ~\ie nsor carpi uln aris, (5) supinator, (6) abduc
r!JOllicis lon gus, (7) exte nsor po lli cis brevis and
rs:JS, and (8) extensor indi cis propriu s muscles.
R;ppl ies th e po sterio r upper arm, an elongated
~ of th e posterior forearm, and the posterior
hand, thumb, and lateral 21h fingers. A lesio n
results in paralysis of extensio n and flexion of the
elbow, paralysis of sup ination of the forearm, paral
ysis of extension of th e wrist and Angers, paralysis
of abduction of the thumb, and loss of se nsation
over the radial aspect of th e posterio r forearm and
the dorsum of th e hand.

105
REG IONAL NEU ROSCIENCE Peripheral Nervous System (PNS )-So matic

Musculocutaneous Nerve (C5, C6, (7)

(only muscles innervat ed by
musculocutaneous nerve are depic ted)

Musculocuotaneous nerve f8
J;
k
V',~
''tl/{I il/t )
. I. ..
J/

COf3coblachiaiis m . brachial

' - Medial cutaneous nn.

o f fOtearm and arm
Bi<:epsbl-achii m.
(turned back)

Brach ialis m.

Art icula r b ra nch

LaterJ I cu taneous n.
of forearm

Anteri or branch

Posterior branch

innervation ~~p......
01lilN
; ';:

FIGURE 11.22: MUSCULOCUTANEOUS NERVE _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ F

The musculocutaneous nerve (C5, (6) supplies supplies sensory innervatio n to the lateral forearm . T
moto r fibers to the biceps brachii, coracobrachialis, A lesion results in w asting o f the muscles supplied,

and brachial's m uscles; it aids in flexion of the weakness of flexion of the supinated arm, and loss
"
t:
upper and lower arm, supinatio n of the lower arm, of sensation o n the lateral forearm.
a
and elevation and adduction o f the arm. This nerve
t(
n
tr
In

106
 - -- - --- -

atic ~ ripheral Nervous System (PNS}-Somatic REGIONAL NEUROSCIENCE

M edian Nerve (C6, (7. C8i T1 )

(only mU$d es innerva ted
by median nerve are de picted)

Muscu lo( ut<lneou~ n. --- -f-_

Median nerve
Posterior
Pron ator t ere~ rn. (humeri'll head)
Lateral cords of
brachial plexus
Articular branch
Medial cutaneous n.
of arm
Fl exor carpi radialis m.
' M"' -',II cutaneou~ n.
of forearm
Palmaris longus m .
Axillary n.
Prona to r teres m. (ulnar head)

Flexor digitorum 5uperlidaiis m . (Iorned up)

Flexor di)\itorum profun dus ffi.

([<l teral p ortio n su pplied via anterior
interosseous n.: medial portion by ulnar n.)

A flI erior inter osse o us n.

Fle-xor poll io s lon gus m.
Pro nator quadratus m.
Palmar branch

Abductor pollicis hrevi s

Opponens pollieis

Flexor pollicis hrevis

(superficia l he-adi
deep head supplied
by ul nar n.1 Flex/)(
retinaculum

Anastomoric
t st and 2nd lumbrical mm.
branch to
\lln.u n.

[)almilr

Bra nches to dorsum of } digital nn.

middle and distal phala nges

9 GU RE 11.23: MEDIAN NERVE _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

- 1e medi an nerve (C 5-T1) supplies motor fibers
0 the (1) flexo r ca rpi radialiS, (2) pronator teres,
3 pa lmari s lo ngus, (4) ft exor digitorum superfieialis
.l nd profundu s, (5) fl exor po ilieis longus, (6) abdue
'or pollicis brevis, (7) fl exor poilieis brevis, (8) 0ppo
-'"'2n5 poilici s breviS, and (9) lumhrical muscles of
ne index and middl e fingers. It supplies sensory
r nervation to the palm and the adjacent thumb,
th e index and middl e fin gers, and the lateral half of
the fourth finger. A les io n (from carpal tunnel syn
drom e) res ult s in w eakness in flexion of the fingers,
and abd uctio n and opposition of the thumb and
loss of sensatio n, o r painfu l sensation in the radial
distributi o n in the hand. A higher lesion also pro
duces weakness in pronation of the forearm.
107
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)-Somatic

Ulnar N erve {C8; T1 J l"1I1 Ulnar nerve

(only muscles innervated

by ulnar nerve are depicted)
-~ (1l0 blanche; above elbow)

J - Arti(uJ<lT branc h
., (behind medial condyle)

Fle xor digitorum proiundus Ill.

{medial portion on ly; l<lteral
portion ,lIpplied by an terior
intero sseous branch o f median n.)

Flexor capri ulnaris m.

(drawn ilside)

OO(S<'11 bran ch

Pal mar bra nch

/ ' Superficiili branch

Flexor pollj ci~ brevis m.

(deep heJd only; superficia l Deep branch
head and other thenar muscles
~ u ))f.l li e(1 by median n.)

= - - - - : : PalmariS brevis
Ad ductor pollici s m. :.-------Abdu ctor di giti mi nimi Hypothenar
I - Flexor d igi ti min im; brevis mu~ cle <;
,- }
Oppone ns d igiti minim;

Common p.almar digital n.

Palm.1r and dorsal inteross.e i mOl.

:I and 4th lumbrical mOl. (t urn ed down)

~~p
,er palm Jr digitJlnn.
I nerves <He Irom dorsal branch)

"""Branch es to dorsum oi m iddle and distal p h<llanges C IIMI

l'tt.:t

FIGURE 11.24: ULNAR NERVE _ _ _ _ _ _ _ __ _ _ _ _ __ _ _ _ _ _ __

Th e ulnar nerve ((8-T1) supplie s motor fi be rs to
the (1) flexor carpi ul na ris, (2) flexor digito rum pro
fundus. (3) addu elo r po lli eis, (4) abductor digiti V.
(5) opponen s digiti V, (6) flexor digiti brevis V,
medial half of the fou rth finger. A lesio n results in
wasting of hand m uscles, w eakness of wri st fi exion
and ulnar deviatio n of the hand, weakn ess of .
'
abduction and adductio n of the fingers, "claw

(7 ) intero sseus dorsa l and palmar, and (8 ) lumbri cal hand" (hyperextensio n of the fingers al meta ~
muscles to the fo urth and li ttle fin gers. It supplies carpophalangeal joints and flexion at the in ter
d
senso ry innervatio n.to the dorsal and palmar phalangeal jo in ts), and loss of sensa tion in the

medial surfaces o f the hand for the little finger and ulnar distributio n in the hand.
iIb<
108
1m. rp heral Nervous System (PNS)-Somatic REGIONAL NEUROSCIENCE

TIl
x hema
11

12

Ven tral rami o f

spinal nerves
Ll

Gray f(l m i comm u nic a nl es ---,;';f'---"""---iJ'-flf~_~

l4
Muscula r bra nches 10 psoas // ~ ~ An terior division
ilnd i liacus muscl('S ~ "
_ Posterior division
Femora l nerve ...,.., l5

Acces~ o ry ob1urat or

nerve (often absent)

Obturator
nerve ""' - - - lumbosdcra llrunk

While and gray ram i

commun icantes
Diaphragm (cu t)
Subcostal nerve (T1 2!
Subcostal nerve (T12) Iliohypogastric nerve

Sympathetic trunk ilioinguinal nerve

Transversus abdomini s mu scl e
Ilio hyp ogastric nerve

Quadratus lum bor um muscl e

Ilioinguinal nerve

Psoas major muscle

C..eni lotemoral nerve (cut)

Gray ram i communiC;Jn(es

Lateral cutaneous

Genitofemoral nerve
nerve of thigh - -
f emoral nerve - - -, Iliacus muscle
late ra l Cul,l neoU$
Obtu r,lIor n..ve .-- nerve o f th igh

~5 major mU$Cle (cur)

Femoral nerve

>
Lumbosacral tru nks
Genital branch (lnd
Femorill branch of
genitofemoral nerve

Obturator nerve

CA: RE 11.25: LUMBAR PLEXUS _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ __

. u mba r plexu s is formed from th e anterior
f'\ rami of th e L1, L2, L3, and L4 (oats within
";)Osterior substa nce of th e psoas muscle. The
..-xl som e of th e L2) root form s th e iliohypo
-n:::: and ilio in guin al nerves and th e genito
al nerves. These nerves co ntribute
cHion to the transverse and the ob lique
-:li nal muscles. Th e remai nin g roots form the
femoral, obturator, and lateral femoral cuta neous
nerves. Lesion s of the lumbar plex us are unusual
because of the protection of th e pl exus w ithin th e
psoas muscle. Th ey result in w ea kness of flexion,
I,oveakness of adductio n of the thi gh and extension
of the leg, and decreased sensa ti o n on th e anterior
thigh and leg.
109
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)- Somatic

'_~.
---;~- L4 _ l umbosac'" tmo k
. . An terior division Schema
_----::~,
.~
~ Posteri or divis ion " , ' l5 . .
:.........: :?!.. -- /' Gray ram i communicantes

Superior glutea l Pelvic spla nc hnic nerves

(parasympa thetic to
Inferior gluteal nerve inferior h)'pogaslric
fpelvic] plexus)
Nerve to piriformis ___

Tlbi<l lnerve Coccygeal nerve

Scia tic nerve Anococcygeal nerve
Commo n fibular
{
(peroneal> nerve----J !J ;1 ,/ Perineal bra nch of 4th sacral nerve
!I
Nerve to levator ani and (ischia-)
coccygeus muscles
Nerve to qUJdr<l\us fe mo ri s I nerve
(a nd inrerior gemdills)
nerve
N erve 10 o nturator inlt: mus-----.l culaneO\IS nerve of thigh
(and superior gemeJlllsJ

lum bosac ra ll runk / Sympathetic tr unk

Psoas miljar muscle ra mi communicantes

Supe rior gluteal artery a nd ne rve

Pelvic. spla nc hnic

Obluri\l or nerve
nerves (cuI)
Iliacus m uscle _ __ -' (pa rasympa thetic
10 inierio r hypogastric
Inferior gluteal ilncry ~
[pe lvic1 plexus)
Nerve to qu adralus femoris-----------...,

Internal p udenda l 03 r lery _ _ _-I

~~p

Nerve 10 obturator inlernus - -- -I

Pudenda l nerve - -

O btura tor In tt!fO US muscle ---~

" ICoN
.... .
: "." ::
Superior pubi<; ra mu s .) '1

Pi riformi s mu scle ~
(Is<;hio-)c occygeus musc le

Nerve to levalor ani muscle - - - - ~ splanchnic

nerves (cur)
leva lo r an i muscle ------. . (symp.athelic to
infe rior hypogastric
Topography: med ial and slightly an terior view [pelv ic! plexus)
of hcmisec ted pelvis

FIGURE 11.26: SACRAL AND COCCYGEAL PLEXUSES _ _ _ _ _ _ _ _ _ _ _ _ _ __ FI(

The sacral and coccygeal plexuses are fo rmed from nerve (52-54). The pudendal nerve supplies the Th,
the roots of the L4-54 segm ents, located anterior to per in eal and sphin cter muscles, which aid in clos ilio
the piriform is muscle. Th e major bran ches include in g th e sp hincters of th e bladder and th e rectum. cle:
the superior (L4-51) and inferio r (L5-52) gluteal Lesio ns of the sacra) plexus result in weakness of oft
nerves, the posteri or femoral cu taneous nerve the posterior thigh and the muscles of the leg and leg
(51-53), the sciatic nerve (l453) and its tibial and fee l, with decreased sensation in the posterior jOir
common p ero neal divisions, and the pudendal thigh and a perianal/saddle locatio n. ane
ane
110

- ~ r ipheral Nervous System (PNS )-Somatic REGIONAL NEUROSCIENCE

Femoral Nerve (l2, L3, l4) a nd

Lumbar
l ilteral Femoral Cutaneous Nerve
lateral fe moral (L2, L3)
cutaneous nor" ,_ plexus

Femoral nerve

Obtu ra tor n. --- - 11 ~A"mbO"~" "t ru n k

Iliacusm.- - - --'!'4!,;
Psoas majo r m. ---f-'~
(lower part)

Artic ula r !Wig - - --f

Sartorius I
(divided)
~itiSl c utaneous
femoral
Pectineus m. _---C

culaneous
Rectus bt-anches of
" femoris m . (emaral n. - --
" (d ivided)
~ Vastus Sartor ius m
"2 {div idedl
.~ Vastus
~
,
0 medialis m.
0 Vastus
lateral is m.

Inirap.llellar brarxh
Artic ula ris gen us . of saphenous n.

Medial crural
cutaneous branches
of saphenous n.

Cutaneous
innervation
Note: o nly muscl es
Innervated by femoral
nerve are shown

fi GURE 11.27: FEMORAL AND LATERAL FEMORAL CUTANEOUS NE RVES _ _ _ _ _ _ __

-;he femoral nerve (main ly L2-L4) innervates the
to psoas, sartorius, and qua driceps fem o ris mus
d es. It co ntributes to fl exio n and outward rotation
of the hip, flexion and inward rotatio n of the lower
eg. and extensio n of the lower leg around the knee
,oint. It supplies sensory fibers to the anterior thigh
a..,d to the anterio r and medial surfaces of the leg
.L.,d foot. A lesion results in weakness o f extension
of th e leg and flexion of the hip and leg, w ith
qu ad ri ceps atrophy, and in loss of sensation in terri
tori es of se nsory distribu tion. The lateral femoral
cutaneous nerve suppli es sensation to th e skin and
fascia of the anterior and lateral surfaces of the
thigh to the level of the knee; lesions result in loss
of sensation in this distribution.
111

REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)-Somatic

Obturator Nerve (12, l3, L4)

Iliohypoga. tric
"}
II
L3

L4
lumbar
pteX"us

Itioinguinal n.
Lumbosacral trunk
Genitofemoral n.

l atera l femoral
cu ta fleOU~ n.

Femoral " . ~.- '\1

Obturator nerve 1" :. <

Posterior
branch _ _ _.".,
If' :.~f'''-- ObluralOr ex lem us m .
A ITicu lar 1J.
brn nch -
An teriO~
t:
"
_ _ _ _ _ Adduct(ll'" brevis m .
'
./ Adductor longus m .
branch
",... (divided)
Posterior Y' J.C1

branch ~ 1{ " m.

ie II.)

Articular
branch 10
knee joint

H iatus of
ad ductor ..,
(anal t

Cutaneous
innervation

~{ft%
Note: only muscles
innl"fVatoo by obturator
nerve a re shown
CII!!N
"'-'"'-,':'

FIGURE 11 .28: OBTURATOR NERVE _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ __ __

FI
Th e obturator nerve (L2-L4) sup plies th e pectineus, in weakness of adductio n o f th e thi gh, w ith a ten n
add uctor (longus, brevi s, and magnu sj, gracilis, and d ency to abduct the thigh in wa lking. There also L4
exte rnal obturator mu scles. This nerve regul ates is weakness of external rotatio n o f th e thi gh. A n
add uction and rotation o f th e thigh. A sm all cuta smal l zone of anesthetic skin on the medial thigh ne
neous zone on the internal thigh is supplied by sen is present. L
sory fibers. A lesion of the o bturator nerve results tel
tril
ou
up
(L
int
trit
ou
112
",' .
'!r"p heral Nervous System (PNS)-Som ati c REGIONAL NEUROSCIENCE

Sd~ lic Nerve (14, LS; 51, 52, 51) and

1Jsll"I' ior femora l Poslerior Femoral Cutaneous Nerv~
rulaneoos nerve (51,52, 53)

- '~ r ior duneal ..)._ _ -G,,,",, sciatic foramen

:t.,ri neal
"'-.. nc;hes - - - -- Sdalic ne~

TibiOll segment ____ ("m'"," pe roneal segment

oi sciatic nerve of sciat ic nerve

_ong head
,:hidedlof Cul;lnoouS innervation
lemOfj,> m .
ierno,i!> m.
O:;cep~ - 4.

->.dduc;tor long hea d id ivi rlp.d)

"I1<lgnll S m.
of biceps fem ori s m.
21;0 supplied
b\' ohtur.llnr n .)

Semitendinosus
~_ ,emnmon peroneal nerve

~emlm cmbranos us

Tibi al nervI:' _ - -----'0'1 Posteriu r

femoral
cutil neous

7 'l "" ffill sur,,1

cut ilneOU~ 11 .

\Ied ial sural

cutanoous n.
communiC<ltinS
G <lSlrOCne m ius m . branch

Sura l n. - - -

So l e u ~ m. -
- - -C
From Supcriiciill
Tibial n. - - - . .

't<lr i<: n . peroneal n.

Medial calcaneal
bmnchcs --~
SuriJl n. - --jC-
/"",,, 1 calcanea l TibiJ I n.
'Vledia l and lateral branche, ivia med ial
plantar nn . calc Jn eil l
,,~_ l"'ffil dorsal bran chesl
cutolneous n .

FIG URE 11 .29: SCIATIC AND POSTERIOR FEMORAL CUTANEO US NERVES

_ _ _ _ _ _ __
-I"' sciatic nerve is formed from th e roots of the
...":"53 segmen ts. The superior and inferior glutea l
'-~rves branch proximally, just before the sciatic
1Efve's fo rmati on. The superi or gluteal nerve
_-1-5 1) supplies th e gluteus medius and minim us,
~sor fascia lata, an d piriformis muscles. II con
--:ibutes to abduc tio n and inward rotation and some
")Ut\ovard rotation of the thigh and to flexion of the
_oper leg at the hip. The inferior glu teal nerve
_+5 1) suppli es th e gluteus maximus, obturato r
intern us, gemellus, and quadratus mu scles. It con
-:lbutes to ex tensio n of the Ihigh at the hip and to
ou tward rotati o n of the thigh. A lesion results in
difficu lty cl imbing sta irs or risin g from a sittin g posi
tio n. The sciatic nerve proper supplies the biceps
femo riS, semitendinosus, and semimembranosus
mu scles (hamstrin gs); and it regu lates flexion of th e
low er leg. Because it branch es in to the tibial and
common peron ea l nerves, maj or lesions of th e
sciatic nerve result in w eakness of leg flexion, weak
ness of all muscles below the knee, and loss of sen
sation in the posterior th igh, poste rio r and lateral
aspects of the leg, and sole of th e foot. The poste
rior femoral cutaneous nerve (5 1-53) supp li es sen
sory innervation to the posterior thigh, lateral part
of the perineum, and low er portion of the buttock.
113
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)- Somatic P,

Tibial Nerve (l4, 15; 51 , 52, S3)

TIbial nerve Medial

Common
calcanea l
Medial sural
peroneal
nerve
cutaneous n, --l.. ($1,2)
(c ut) . F,~
ieular tibial n.

Arti cular t branch

branc hes . t; ;

Plantaris m.-----rft1 l ateral sural

G~tro
c nemius m .
cutaneous n.
(cut)
\....(51, 2J
k /*,\. .6. . . .
~> I '\ Sural n.
($1, 2)

Nerve to
Saphenous
popl iteus m.
IU , 4)

Crural inter
osseous n. Cutaneous innerva tion of sole

Soleus m . Medial and lateral

Flexor calcaneal branche$
re tinac ulum
(cut)

TibiaJ I _ ,

rQuadratus pl(lntae m.,

Tibialis
posterior m.
nerve :: H~ r Abduclor digiti minim; m.

Medial

Flexor P'ant.IT_ _ _~'

hall uci.
""""
lot : Oeep Sranch
,
longus m. ~i \0:
to:
, 5t, 2 nd, 3rd plantar ,
f lexor inte rossei mm.;
Sural n. digitoriu m
(cut) 3rd,4th
brevis m. 5
lumbrical mm.;
Abductor e
l ateral / Adductor hallud s m.,
hallucis
"
~~p
calcaneal Articular branches
branch I,
Medial II..
llill\l
e
. :: '-Superficia l Branch to:
calcanea l
branch
"
(,

U
Flexor
d
(~
(cu t) d igital nn..
Cu taneous branches ~
d(

FI GURE 11.30: TIBIAL NERVE _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ FI G

The tibia l nerve (L4-S2 ) supplies innervatio n to the hallucis, interosseus, and third and fourth lumbrical The
(1) gastrocnemius and soleus (main plan tar flexors musdes. Sensory branches supply the skin over the the
o( the (oot), 12) tibialis posterior (plantar flexor lateral calf. foot. heel. and small toe (sural nerve), aotE
and invertor), (3 ) flexor digito rum lo ngus (plan tar and the m edial aspect of the heel and the sale of 50 r
flexor and toe flexor), and (4 ) flexor h aUucis lon gus the foot. A lesion can resul t in weakness of plantar extE
(plantar flexor and great toe fl exo r) muscles, and flexion and inversion of th e foot, weakness of toe sian
(5) muscles of the foot, in cludi ng the abductor flexion, and loss of sensation o n th e lateral calf and toru
digiti minimi pedis, flexor digiti minimi, adductor the plantar regio n of the foot. p erc
1 14 .
lie
Peripheral Nervous System (PNS)- Somatic REGIONAL NEUROSCIENCE

Common
Common Peroneal Nerve (L4, LS, 51, 52 )
peroneal n. - --t- -lt:-:
(in phanlom)
Tendon o f biceps

femor is m.
-----i:-+
Art icular branches
Common peroneal
Ant eri or tibial recurrent branch
nerve ~::-=":.1~~i
Head 01 fibula Extensor digilOrum longus m .

Peroneus Deep peroneal nerve

longus m, ' _ . j' t

Tibialis anterior m.

Superficial

peroneal
Culilneous innervation
nerve

Branches of

lateral su ral

cu tan eous n.

digitOfum
Pero neus longus m.
longusm.

Peroneus Extensor
brevis m. - - -4- hallucis
longus m .
Medial dorsal

cu ta neous n .

Inlel mediate lateral sural

dorsal
cutaneous n.
cuta ne<Jus n.

/ l.",,,'1 branch of
Superior
deep peroneal ". Superficial
eKtensor
10 peroneal n.
ret inaculum - - --i / Exleo" " halluds
brevis m. and
Infer ior ExtenSO( digilorum
extensor brevis m.
retinaculum
- - -
(cut)

. / """,di,'1 branch
Laleral dorsal of deep
culaneous n. _ - -1'.\' peroneal n.
(branch of sural n.)
~-P",p.. dorsal

_--~tJ1t
Proper dorso, ,'_ 00 .
di~ital nn. -

FIGURE 11.31: COMMON PERON EAL NERVE _ _ _ __ _ _ __ _ _ _ _ _ _ __

The comm on peroneal nerve (L4-S1) branches into
the deep peroneal nerve, supplying the (1) tibialis
anterior (foot dorsiflexion and inversion), (2) exten
so r hallucis lo ngus (foot dorsiflexion and great toe
extensio n), (3) extensor digitorum longus (ex ten
sion of toes and foot dorsiflexion), (4) extensor digi
torum brevis (extension of toes), and the superficial
peron eal nerve, which supplies the perone us
longus and brevis muscles (plan ta r flexion and foot
evers io n). Sensory branches supply the lateral
aspect of the leg below the knee and the skin on
th e dorsal surface of the foot. A les ion of this nerve
can result in weakness of dorsiflexion of the foot,
weakness of toe extension, and loss of sensation of
th e lateral aspect of the lower leg and the dorsum
of the foot.
11 5
REGIONAL NEUROSCIENCE

Oculomotor (1111n. \
Peripheral Nervous System (PNS)- Auto nomic
-Pe

Facial (VII) n. ~
:' ~ . ~ __ "'~"!._~___ :"'~_- ---~ Intracranial vessels
Gl ossophar~'ngea' {IXl n. 1).
I::
~,<~,
.. ~:-.: -=.,;~:-
CiliJry gall!;lion
, ..
Ey,
Medulla obl on 8 ,~ r a _ _,:':;?' ; (- I _ __ _ .. - ~:;:k
Lacrim al gland
I" .- {M - - ----- - --- ----~-=::'~
Vag us (X) n. _ "' , ~--- --___ p(eryg~~rat~~~
"
( I
( 2
-~
~.:.: ;;,_:!

~ :1!"'-",-
~'I:,
__ __ __~

_ "' . '
i/
... --- :;'1
...
. ~.._......

~ ... "'.._
-- -... ngI'?!J..-.:.--:
. " <.
-:.~ .....
~
Paro tid slanrl

Sublinguill ''lnd
submandibular glands
Peripheral cranial
blood vessels
Larynx
Trach ea
Bronchi a nd lu ngs
Swe.1\

gla nd

H eart

Peripheral
Shown Stomac h
v ess ~ 1
kir only
seg men t live r
Gallbladder
ii,
Pi.tnCfeas
H.,i r Adrenal glands
fo llicl e Brow n f,1t
Ki dnl?Ys

Intestines

l~'m ph oid organs

Dista l co lo n

Bladder

Ii
Coccyx Inf.
I""

Sympa the tic fibers

prega nglio nic
____ postga nglionic
Pa rasympilthetic fibe rs
_ preganglionic
... _ ... _ o osiganglio nic
- Antidrom ic condu ction
~~~
,1liN
:"'; -": 1
j . Pi'!

o~
~

....

FIGURE 11.32: GENERAL SCHEMA OF THE AUTONOMIC NERVOUS SYSTEM _ __ _ _ __ FIG UI

Th e autono mic nervous system (AN S) is a 2-neuron (cranial nerves [CNs] III, VII, IX, and XI and the
chain. The pregan gli on ic neuron arises from the sacral spinal cord (S284 in termedi ate gray). This theA '
brain stem or the spinal cord (eN S) and synapses craniosacral system regulates reparative, homeo musclE
o n postganglio nic neuro ns in the sympathetic static/ and digestive func tions. These au to nomic immun
chain, collateral ganglia (sympathetic), o r an intra system s achieve their actio ns th rou gh innervation d ivisiO!
mural gan gli on (parasympathetic) near the o rga n of smooth muscl e, ca rdiac muscle, secretory patheti
inn ervated. The sympathetic division, derived from (exocrin e) glands, metabo li c cells (h epatocy tes, tiation,
neuro ns in the Tl-L2 (thoraco lumbar) lateral horn, brown faO, and ce ll s o f the immun e system. lymph
prepa res the body fo r fight o r flight-mobiliza tio n Normally, both divisions wo rk together to regulate tivity ar
for emergency respo nses. The parasympathetic visceral activities such as respiration, cardiovascular Im mlJ nl
division is derived from neuro ns in the brain stem func ti on, digestio n, and some endocrine functions. cell-mel
116
!) nom ic phera l Nervous System (PNS )-Autonomic REGIONAL NEUROSCIENCE

Pl-egangliollic sympathet ic
. --- ---- Poslganslioni<: symPiJthelk
Preganglionic vaf<lsympalhetic
- - - - - Postganglionic parasymp.:uhelic
Thymus
.................. --- - - --~
------ ---------" ----- --
............
( 1 Cervi<:al
IA!------ --------------" ---- -----/" --- --- --0 0 Irmph nodes

, 1.,' -- ----- - -_ . . -.. -.. --- ---, . ----~~-.....,-J ~

Upper limb
hont' ma rrow
Sympathetic chain
ganglia Brown fat

_________ _- of
Pulmonary MALT
;.,i,f--~<~~- -------- ------------- - --
--
--
::::fi--~;:".} - --- -- -------- - - ----- - --
~~- ------- ---------- ---------

Liver anrl
ht>paIOCYle~

Spleen
Supenor afld inferior
nle5el"llenc I/,J.ngfla
Mesen te rrc
lymph nodc~

GALT (e. g., Peyer's

patches l wil h
parasympa thetic
intramural gangliJ.
J....---- - --------- --- - --///:><C~~
InS,-Jl nal
- - -- ~~ Iyml)h nodes

'. ~- '
-'. -----<

.-
Lower limb

J. Perkins
"'S, M1'"
-,- bone marrow

,,,... ,,

- FIGU RE 11 _33: ORGANS

INNERVATION OF ORGANS OF THE IMMUNE SYSTEM AND METABOLIC
_____________________________________________

The ANS innervates the vasculature, the smooth
muscle ti ssue, and th e parenchyma of organs of the
immune syste m mainly th rough the sympath etic
division. In the bo ne marrow and the thymu s, sym
pathetic fibers mod ulate cell proliferation, differen
tiatio n, and mobiliza tio n. In the spleen and the
lymph nodes, they modulate innate immune reac
tivity and th e ma gnitude and timing of acquired
r immune responses, parti cularly the choi ce of
ce ll-m ed iated (Th1 cytokines) vers us humoral
(fh2 cytokines) immuni ty. Autono mic fibers
regulate immune respo nses and inflammatory
responses in th e mucosal-associated lymphoid
tissue of the gut and the lung and in th e skin .
Extensive neuropep lidergic innervation, derived
fro m bo th the ANS and primary sensory neurons,
is present in the parenchyma of lymphoid organs.
Postganglionic sympathetic nerves also su pply
hepatocytes and fat cells.
11 7

REGIONAL NEUROSCIENCE
Peripheral Nervous System (PNS)-Auton omic

Sympathetic tru nk
Spinal
gangli.

commuicans
Sympathetic
tru nk ganglion
Gray ramus
communicans

~~~ 0 1liiN
1\'.... : ;

Sympathetic fibers Pia.l'ilSympathelic fibers - Afferent fibers to spinal cOf'd

_ _ preganglionic _ preganglionic - - Afferent fibers 10 brain stem
____ postganglionic __ postganlionic

FI GUR E 11.34: REFLEX PATHWAYS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Autono mic reflex pathways consist of a sensory
(afferent) component, interneuro ns in the ( NS,
and autonomic efferent components that innervate
the peripheral tissue respondin g to the afferent
stim ulus. The afferents can be autonomic (e.g.,
from th e vagus nerve), processed by brai n stem
nuclei such as the nucleus soli tarius; o r they can be
somatic (e.g., nociception), processed by spinal
cord neurons. The preganglionic sy mpathetic or
parasympath etic neurons are activated through
interneu ro ns to prod uce a reflex auto nomi c
respo nse (e.g., co ntractio n of vascular smoot h mus
,
cle to alter blood pressure, increase in heart rate ,
and contractility). The efferent connectivity can be t
relayed via splanchnic o r somatic nerves because ,
of the complexity of auto nomic efferent pathways. t

118
, f
mit "cripheral Nervous System (PNS)-Autonomic REGIONAL NEUROSCIENCE

Salivary glands

Trachea and
Vagus (Xl n. brOnchi

Heart

Striated muscle
~.

,'I
Peripheral arterioles

Visceral arte rioles

Gastroi n te~t inal lrac l

Pelvic splanchnic nn.

Bladde(

C Cholinergic synapse
A Adrenergic synapse

Sympathetic fib ers Parasympathetic fibers - - Somatic fibers

_ _ preganglionic - - preganglionic - - Anlidromic conduction
____ postganglion ic ____ postganglionic

f iG URE 11.35: CHOLINERGIC AND ADRE NERGIC SYNAPSES _ _ _ _ _ __ _ __ __

The AN S is a 2-neuro n chai n. All sympathet ic and no repinephrine (adrenergic responses lAD to

parasympathetic prega nglionic neurons use acetyl activate both alpha and beta recepto rs on target

choline (ACh) as the prin Cipal neurotransmitter in tissues. Alth o ugh ACh and norepinephrine are th e

sy na pses on ganglion ce ll s. These cho linergic (e) prin cipa l neurotransmitters in autonomic neurons,

synapses activate mainly nicotinic (N) receptors o n man y co-localized neuropeptides and other neuro

~h e ganglio n cells. Postga nglionic parasympathelic mediators, including neuropeplide Y, substance P,

neuro ns use ACh at synapses (C) w ith target tissue somatostatin, enkephalins, histamine, and gluta

m activate mainly muscarinic (M) recepto rs. mate, are present.

Postganglionic sympathetic neurons mainly use

119
REG IONAL NEUROSCIENCE Peripheral Nervous System (PNS )-Autonomic

CENTRAL NERVOUS SYSTEM PERIPHERALNERVO US SYSTEM

Luwer

motor

neurons

,t;)
-

Skeletal muscle

Sympathclic
chain .o,~.

<... . " .
ganglion
A '
== ~
. ~aF '~{,
........" ;"'

~ - -",'/J

Collateral
ganglion
Sympllthet ics C N
T1-l2 --l....-tll'i;
,.,.. ,
~
. / '-~ ~- '

T,,' gct tissue

Adrenal '

R ., Epil1ephrine
.. , _.. ~ ~" It NOlepjnephril)t~
.'~
:"'t<~ -,
ChrOm.l.ffin
cell

,,,d . ",,1) ~'

(,,,,,;,, ' ,
p ara)'nlpathcl ic5
, 'S , _
'
;,j
~.;'" g,"~;o"
'm"m",,1 ,c'if:)
~ : ~ ''' .-'l: i
',-. 1 ,""""" _"""'<' ' C
t "/.
. \
"
.. ~,

Targe t tIssue
",:

5yn/l PsK Receptors

.-
~ Adrcnerl4lC (. Of () .l..!vna or betJ il<lrener gic

C Cholinergic N Nicotimc J. Perkins

MS, MfA
M ,"\ t lIS(.lf i r1 i("
..
, ..,."

FIGURE 11.36: SCHEMATIC OF CHOLINERGIC AND ADRENERG IC DISTRIBUTION TO MOTOR

AND AUTONOMIC STRUCTURES _ _ _ _ _ __ __ _ _ __ _ __

All preganglionic neurons of bo th the sympath etic
and parasympathetic nervous system use acetyl
cholin e (AC h) as their neurotransm itter. All gan
glion cells possess mainly nicoti nic receptors for
the fast response to cholinergic release from pre
ganglio nic axons. H owever, additional muscarinic
receptors and dopamine receptors on ganglion
ce ll s help to mediate lo nger term excitabili ty. The
postganglionic sympathetic nerveS use mainly
120
norepin ep hrine (NE) as th eir neurotransmitter, and
target structures in th e periphery possess different
subsets of alpha and beta adrenergic receptors fo r
respo nse to N E. Some postganglio nic nerve fibers l
to sweat glands use ACh as their neurotransmitter.
Postganglio nic parasympathetic nerves use ACh as
their neurotransmitter, and target stru ctures in the
periphery possess mainly musca rinic recepto rs for
response to ACh.
"
"e-'pheral Nervous System (PNS)- Autonomic REGIONAL NEUROSCIENCE

5ensol)' (001
p etf05d I n .
Nasociliary n. root}
Tti genmlnal M n.
{
Motor Deep pelfOSdI n .
ISY;:::~~:I~footn. of ciliary
ganglion

:"""i "i'~ n.
Facial (VII) n. \'Plh, "mi' n.
Veslibulo
coch lear
(V ii I)

nn .

Intern al Otic
carotid n. ganglion

Superior
lal)'ngeal n.
Superior lingual n.
cervical
symp.1lnel ic
trunk
ganglion
Vagu~ (X) n.
Internal Submandibular ganglion
carotid .1.
Cervical
sympa thetic
trunk - - ' I meningeal 3. and plexus

,p~p
Superior 3. aod plexus
cervical
cardiac '(I,m,",," carotid a. and plexus
branch of
Superior ceMc.a1sympathetic card iac n.
4 e lM!
I....:::

FIGURE 11.37: AUTONOM(C DISTRIBUTION TO THE HEAD AND THE NECK: MEDIAL VIEW _
Autonom ic nerve distri buti on to th e head and the
neck incl udes compo nents of both th e parasym
patheti c and th e sympathetic nervous systems.
Parasymp ath eti c co mponents are associated w ith
CN s III (ciliary gan gli on), V II (pterygo palatine. sub
mandibular ga nglia ). and IX (oti c gangli o n). Th e
vagus nerve and its associated ganglia do not
innerva te effector tissue in the head and the neck,
altho ugh they are present in the neck. Sympathetic
components are associated w ith the superi or cer
vical ganglion an d, to a lesser extent, th e mi ddle
cervical gangli on. The geniculate gangli on (eN V II),
the petrosal gangli o n (CN IX). and th e nodose
gan glio n (e N X) process taste inform ation. They
are so metim es th o ught of as autonomic afferents,
but they are no t compo nents of th e auto nomic
efferent nervo us syste m.

121

REGIONAL NEUROSCIENCE Peripheral Nervous .System (PN S)-Autonomic

Int
carotid n.

Closso

pharyng,e_-=-,'_-:

(IX) n.

v~gus
( x. )n . _ _ _~

(cu t)

Sup. cervical

sympathetic

tru nk

ganglion

Pharyngeal plexus
Sup. pharyngeal branch
of vagus n.
Ext. carotid a. and plexus
laryngeal n.
Int. carotid a. a nd carotid
Gray sinus branch of
ram i
glossopharyngeal n.
Carotid body

Carotid 5inus

Syp. cervical cardiac branch

0 1 vagus n.

Sup. cervical sympathetic cardiac n.

Phrenic n.

M idd le cervical sympathetic

(fun k 8<lOglion
Common carotid a. and plexus
Middle cervical 5}'mpathetic cardiac n.
Vertebral ganglion
Vertebral a. and plexus
Recurrenl laryngea l n.
CerviCOlhoracic (stella te) ganglion
Ansa wbclavi3
+- Va2US (Xl n . (c ut)

~~p
Inf. <;ervical sympathetic
cardiac n.

Subclavian a. clliiI\I
....
:\ -:;:

FIGURE 11.38: AUTONOMIC DISTRIBUTION TO THE HEAD AND NECK: LATERAL VIEW _ __ =
The parasympathetic nerve fibers to the head and lingual glands; eN IV, otic gangli o n to pa ro tid
the heck regulate pupiUary constriction and accom gland). Th e sympathetic nerve fibers to the head < ,
moda tion for near vision (CN III, ciliary gangli on to and th e neck derive mainly from th e superi o r cer -~

pupillary constri ctor musde and ciliary muscle), vical ganglio n, w ith synapses to the pupillary dilator -.
tea r production (CN V II, pterygopalatine ganglion muscle, the swea t glands, the vascular smooth m u~ 3.
to lacrimal glands), and salivation (eN VII, sul:r de, and the thymus gland.
mandibular ganglio n to subma ndibular and sul:r
122 'I
 ->1era l Nervous System (PNS)-Autonomic REGIONAL NEUROSCIENCE

CENTRAL NERVOUS SYSTEM I'ERrPHERALNERVOUS SYSTE M

~m pathe ti cs
1 Ciliary ganglion Pupi llary conSlriclor m.

Ciliary m.

....: ellS of
_~- .\ eslpha l
. . .""'."",.~
-I ( ranial n. I!I
"",~;};.-,.,..,-="","", ...;;::::;~~,.."..... '>';'-4'~ ~
""'.

~~ 1
"
\
I.acrimal glilnd~
, ,
Plerygopalilline ganglion Nasal m UC05.)1 gla nd~

:~Derior ( rilni,JI n. V II
... . atory
- ~cleus
4--=-

Subma nd ibula r ganglion Submandibular gland

Sublingu.11 !!Iand
,-.
I'
'~
",. (.-c:-:r~.::-,
I...~: .~ .. ,

Qli<; g,\nglion P,lrolid gl,1nd

'Ierior
-,.- -.
,..w. ato ry /'
~ ucl eu s
'-:"y /,,\
1):., ' I
, '/
\ ..-,-

p,al hl!ti cs Superior ce rvical gans-lion

Tl-n
-:ermedio

I,ateral

~d1 column
,

~weat glands and

J. Pe rkins
.,,~ MW. vilscul,)f smooth muscle
0,"",,,
..,.
.", in hc.1d and nl"ck

CU RE 11.39: SCHEMATIC OF AUTONOMIC DISTRIBUTION TO THE HEAD AND THE NECK

_
' onomie innervatio n to the head and the neck
aerived from parasympathetic neurons in the
n stem, including the Edinger-Westpha l nucleus
" III), the superior saliva tory nucleus (CN VII),
,... the inferio r salivatory nucleus (CN IX), and
from sympathetic neurons in the Tl -T2 inl.ermedio
laleral cell column in the spinal cord. The associ
aled ganglia and targe t (effecto r) tissue are also
illustr<lted.

123
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)- Autonomic

From Hypothalamus
Shofl Orulomolor n. rool o f
c iliarY8il nglion (motor) - - - Edingcr-Wt>Stph<l1
r
J Nucleus
Oculomotor

'e<:'

Dilator
of pupil

Sphincter

of pupil

Ciliary
muscle
Descending Nude!."
ofTrigemil)al N. (\

Nilsociliary n.

g<lnglion
Nasociliary n. rOOI
of ciliary ganglion
caro tid
Ophth<llm ic a.
[)iexus

Intern al ca rotid a.
Thor acic part
Superior ce rvi cal sympatheti c trunk S<'lngJion / of spina l cord

Gray ramus communica ns----....tf

15t thoracic sympath eUc trunk ganglion

Sympathetic Fibers
- - - - - Pre ganglionic
While ramu s communicans
- - - - - - Postgan glionic

Parasympathetic Fibers
- - -- - Pregan glionic
- - - - - - Postganglionic
Afferent Fibe- rs

Pupillary lig ht Reflex Pathway

~~?I""';..
~-
Ot'Scending Pathway < ION
, ~"-.",,

FIGURE 11.40: AUTONOMIC DISTRIBUTION TO THE EYE _ _ _ _ _ _ _ _ _ _ __ _

Pa rasymp(l thetic preganglio nic nerve fibers fro m
the Edinger.Westphal nucleus innervate the ciliary
ganglio n, w hich supplies the ciliary muscle (aiding
in (lccom moda tion to near vision). and the pupil
lary constrictor muscle (constricting the pupil).
Sympa thetic preganglio nic nerve fibers fro m the
T1 T2 intermediolateral cell colum n innerva te the
superior cervical ganglion, w hich supplies the
124
dilator m uscle of the pupil. The pupillary light retle'.
is a major renex in neurological testing. The afferer"
limb is acti vated by light shone in either eye via
eN II, processed thro ugh the pretedum to the
Edinger-Westphal nucleus on both sides (via the
posterior commissure); the efferent limb consists
of automatic outtl ow to the pupillary constrictor
muscles of both sides.
 ~ h e r a l Nervous System (PNS)-Autonomic REGIONAL NEUROSCIENCE

Intern al c.1 rotid n.

Cervical sympathetic
trunk ganglia
," .._------------------
~..,k" cardiac nn.

l owe r lu mbar
and sacral
sympathetic
trunk ganglia .:.::

Sympatheti c railli to lumbar

~" l1Ipa lhet ic fibers sacral plexuses and nerves 10
lower !imb
preganglionic
- -- postgangli onic :\ ...
,': :

-G URE 11.41: INNERVATION OFTHE LIMBS _ _ _ _ __ __ __ __ _ _ _ __

-....Io nomic innervation of the limbs derives (ro m
~ sympatheti c nervous system. Prega ngli onic
Tll pathetic nerve fibers from th e tho ra columbar
-ermediolateral cell column supply sympathetic
"ain ganglia. These ganglia send postgan glionic
_~a dren e r g ic nerve fibers throu gh the gray rami
communi cantes into the peripheral nerves to
supply vascular smooth muscle (vasomotor fibe rs),
sweat glands (sud omotor fibers), and arrector p ili
muscles associated w ith hair follicl es (pilo motor
fibers).

125
REG IONAL NEUROSCIENCE Peripheral Nervous System (PNS)- Autonomic

CerviCOl ho rilcic __ ..,

(~lell ale) ganglion L 1S Cervicotho rOl eiC
{stellate) ganglion
Ansa subc lavia f, - ;.'.. '

R. svmpalhet Cervical
ca rd iac nn.
(sympathetic
Cervical a nd vagal)
c(Hdiac nn.
rS)'mp." hctic
.,nd va&<ll) - -- - - L vagus n. (c ut)

Thoraci c
syml?athctic___~/
cardi.K nn.
R, vaglls n. (Cllt l Thoracic
cardiac nn .
n ,oracic vd.gal (sympathetic
bral1ches to an d vagal)
pulmonary and
<:ardial" pfe.-.::uses
Branches to
5th intercostal n ant. and post.
(a nt. ra mu ~ of pulmonary
Slh thorac ic plexuses
sp inJ. 1n.) - - - - -/
Cardiac plexus
G rOll' and wh ite
rami communicn
L sympathetic trunk
5t h thora cic
symp~th (;ti {; trunk /
g;mg ll on ----~ Thoracic
aorta plexus
R. greater Ihor3cic
splanchni C: n. - - - -- Esophageal
Sympathetic branch pl exus
toesoph<lgea l
plexus ~~----- l. greater thorac ic
spranc hn ic n.
Thoracic duel
Thorac ic L lesser thoracic
aortic plexus splanch nic n,

R. lesser thorac ic Ant.vaga! tru n k

splanchnic o.
R. lowesllhorac ic D ia ph ragm
spla nc hnic n. (pune<! down)

Azygo~ vein
(c ut j - - - -- ,f ,f~
Inf. 4 ~{II""-I
n-,'.-;;:

FIGURE 11 .42: THORACIC SYMPATHETIC CHAI N AND SPLANCHNIC NERVES

_ _ _ _ __
The sympathetic chain is a conection of sympa
thetic ganglia that receive inpu t from the thora
co lumbar preganglionic nerve fibers derived from
the spinal cord_ The ganglia, interconnected by
nerve trunks, are located in a paravertebral array
from the neck to the coccygeal region. Postgan
gli o ni c noradrenergic nerve fibers from the sympa-
126
Ihelic chain supply effecto r tissue in the periphery.
Some preganglioni c nerve fibers do not synapse
as they trave l through the sympathetic chain.
These fibers co ntinue along th e splanchni c nerves
to synapse in co llateral gangli a, wh ich supply
noradrenergic innervation to effec tor ti ss ue in th e
viscera.
 pheral Nervous System (PNS)- Autonomic REGIONAL NEUROSCIENCE

From hypoth alamic !

and higher centers ! , \~~,

i,
~~;t:~f~::;: to smooth mU$(le alld glan&;; alfer ent

, tracheobronchial muco~.a, and alv~oli)

Affer ent nerves from
GI~ ,oph"y n" .. nerve (I X)
1t"C~~ n~e and sinuses (via
~ trigeminal (VI and
glossopharyngeal [IX1
nerves) may also initiate
renex~ in airway~

SuperiOl cervical
Jescending tracts --\ - sympathetic ganglion
in spinal cord

'coo Symp"h.'lk nerves

{adrenergic) Larynx

Caro tid body-\b-lfJJ

-~ ofacic
,a cord

art"" ,..--->. Cough receptors

Sympathetic trunk - --It

.athetic l'ibeN Pulmonary p \ ,'"" "--

- -- - Preganglionic
- - - - Postganglio nic

~""mpathetic l'ibf:rs
- - - - Preganglionic
- - - - - - Postganglionic

~ nt fibers

,energic lerminals (norepinephrine

I'd or epinephrine)

:-'"IOlinergic terminals (acetylcholine)

RE 11.43 : INNERVATION O F TH E TRACHEOBRONCHIAL TREE

_ _ __ _ __ _ __

- 5\ mpa thetic (noradrenergic) and parasympa
M: cholinergic) innervation supply smooth
-'e of th e tracheobron chial tree. Sympath etics
e !rom th e sympatheti c chain, and parasympa
- derive from vagal autonomic input to local
lUral gangli a. Sympathetic influences result in
-oc:hodilati on, and parasympathetic influences
result in bronchoco nstriction. Additional n euro~
peptidergic innerva tio n, some as co-localized o r
in dependent autono mi c fibers an d some as pri
mary afferent fibers, a!so di stributes alo ng the
epithelium and among the alveoli, where it ca n
influence innate immune reactivity and the pro
duct ion of infl amma tory media tors.
12 7
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)- Autonomic

Dorsal vagal Iract nucle us

Su perior cervical sympathetic t

tru nk ganglion

Superior cervical sympathe tic : ~J. '>'.

card iac nerve Supe rior cervical vagal
cardiac branches
Middle cervical sympalhe lic

tr unk ga nglion

Mid dle cervical sympathetic - - - -if(

cardiac nerve

Verte bralgangl ;;,o:O:.::=======~~~J

Ansa subclavia-
Cervicothoracic (5Ie))ale) ganglion
Ventral ramus o f T 1 - - - - -
(intercos ta l n.J

Inferior cervical sympalhelic--

cardiac nerve ~
2nd th oracic sympathet ic
,'
tru nk ganglion
Thoracic vagal cardiac branch - ;:;;.:;

White ram i communicantes<

4th th oraci

trunk ga nglion
Thoracic
sympathetic
cardiac nn.

'.
Cardiac
plexus
Sympathetic fiber;; Parasympathetic fibers
____ Preganglionic

~~~
- - - - Preganglio nic
- - - - - - Postganglionic - - - - - - Postganglionic
Afferent fibers Afferent fibers l rMOo.
o "~
...., "

FIGURE 11.44: INNERVATIO N OF THE HEART _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _

Sympathetic noradrenergic nerve fibers (derived the great vessels and the coronary arteries.

from chain ganglia) and parasympathetic choliner Sympathetic fibers increase the force and rate 01

gic nerve fibers (derived from cardiac intramural cardiac contraction, increase cardiac output, ane

ganglia innervated by the vagus nerve) supply the dilate the coronary arteries. Parasympathetic iibers:

atria, the ve ntri cles, th e sinoatrial node, and the decrease the force and the rate of cardiac contra:

atrioventricular node and bundl e. Sympatheti c tion and decrease cardiac output.

noradren ergic nerve fibers also distribu te alo ng

128
 ~ he ral Nervous System (PNS )-Autonomic REGIONAL NEUROSCIENCE

""pathetic tru nk _ _ __

-W'3.cic duel - - - - - - - ' An terior and

posterior
:-e,lter and - - - - - - , vagal trunks
~""" thoracic - - - - --i!
-"lchnic nn. Cel iac plexus and gangl ia

')"Irenic n. - - ---;; l. grL'atL'r and

lesser thora(;ic
--['rior spanchni(; nn.
'"1llC il. __- - - -
pl exus Sup. rne:.enteric
ganglion
::'eater and - - - --::
~r thorac i c - - -, L. aonimrenal ganglion
':"'1chnic nn.
3drenal
L. IOWl'St thorilcic
o;:'(U 5 ~----~ splanchnic n.

.aorti corena l l. sympathetic tru nk

~~I ion - - -
Intermesen teric
- Iow esl tabdominal
--::~ac ic aort il) plexus
~ .e:nchnic n.
tnf. mesenteric ganglion
C ';enal iI .
.....-.d pl xus tnl. mesenteric J.
.~ ;:\ mpathetic and plexus
-_111..-- - - /
l. colic J. Jnd plexus
~ ',,~ and
~ ie rami
L. common iliac ,,_
.:ommu nica nts
and plexus
C ,,:erna chyli - ---'
Sup. rectill iI
cd [u mb,n and plexus
~.:TI pil theti c
- ...nk ganglion
C-a\ ramus hypogJstric
:~m m u n i can5 plexus
(presJcrJI no:
~ ...m bar
5p:anchnic nn.
H ypogJstric nn.
- ureter to r. Jnd
a.."'Id plexus L inferior
hypogastric
'- testicular a. (pelvic)
.;.-d plexus - --'" plexu~es

.:-i!cral p.:!rl of
'_ ;vmpathetic trunk l. sacral plexu5

FIGU RE 11.45 : ABDOMINAL NERVES AND GANGLIA _ _ _ _ _ _ _ _ _ _ __ _ __

-ne abundance of sympa thetic nerves in the
..bdomen and the pelvis is associated w ith innerva-
n of th e gastrointestinal (C I) and urogenital sys
em s, associated vessels, the peritoneum, and the
- rena l gland. Th e lumbar portion of the sympa
'""'etr c chain and its branches and the splanchnic
"" rv es and th eir collateral ganglia (celiac, superior
and inferior mesenteric, hepatic. aorticorenal.
adrenal, superior hypogastric, others) innerva te
smooth muscle, glands, lympho id ti ~~ u e, and meta
bolic cells in the abdomen and the pelvis. Most
of the collateral ganglia (plexuses) also contain
parasympath etic contri bu tions from the vagus
nerve and associated g(lnglia.
129
REGIONAL NEUROSCIENCE Peripheral Nervous System (PNS)-Auton o lili...

R;.m,
6tll rhoracic left 71h thoracic
sympathetic sy m pathetic trunk
trunk
ga nglio n
--"t
",. ,
ganglio n

7th inter Esoph ageill plexu s

cos l al n. ~
While Aorti c plexu s
"d
left greater th oracic
Spinal splanchnic n.
gan glio n
and ve ntral
roo t of 8 th Posterior vagal trunk
thoracic and celiac bra nch
spin,.r n.
Right Anterior vagal lfunk
/ - -...I ~-l ia c tir<lnch
grea ter ,
andlcsser .
thoracic
splanchni c nn. l eft gastric a.
Celiac
ganglia ", 't""" .s..";\i Splenic a.
and plexus .., ..;~ :
Ce liac
trunk~-----~~.,
Right
lowesl
lhoraci c
spla nchnic n. ~
Common
hepatic a. ;~
Righ t
gastric a. - l
Superio r mesenteric
g,mglion
Aor ticorenal
ganglia
Renal a. l e lt gastro
epiploica.
~ Gastrod uodenal J.

~
Righ i gastro
epi ploic a.
~n
!- Inferior panc reaticoduodenal a.

Sympathetic fibers
_ _ Pregan glionic
Parasympathetic fib ers
_ _ Pregan glionic
- - Afferent fib ers
~~p
lmN
,,\':, : :
_ _ _ _ POSlgangliooic - - - Posiganiion ic

FIGURE 11.46: INNERVATION OF THE STOMACH AN D TH E PROXIMAL DUODENUM - ----1

The stomach and th e proxim al duod en um receive
sympathetic inn ervation in abunda nce from the
celiac and superio r mesenteric ganglia and to a
lesser exten t from the thoracic sympathetic trunk
ganglia. The celiac and superior mesenteric ganglia
receive preganglioni c input from the greater and
lesser thoracic splanchnic nerves. Parasympathetic
iibers distribute to th e stom ach and the proxi'
duodenum fro m the celiac branches of the \ a~
nerve. Sympath eti c fibers decrease pe ri stals i ~ ., .
secretomotor activities . Parasympathetics inc-
peri stalsi s and secreto motor activity (s uch as
gastrin and H e !) and relax assoc iated sphinci

130
~ h e raJ Nervous System (PNS)-Autonomic REGIONAL NEUROSCIENCE

._-
_- -
"-"'<""--
.....- -- -
..... 003_'
....- ...... ----

RE 11.47: INNERVAT ION OF THE SMALL AND LARGE INTESTINES _ _ _ _ _ _ _ __

..aonomic inn ervation of the small and large ganglia and plexuses via eN x and pelvic splanch

-.:.ti nes is suppli ed by extrin sic sympatheti c and ni c nerves. Sympatheti cs generally decrease peri.

..IJ2S\ mpath eti c fibers. Sympathetic innelVation stalsis and secretomotor function s (i.e., decreased

.es irom the TS-L2 intermediolateral cell fluid secretion). Parasympathetics in crease peri

-n n of the spinal cord, and it distributes to co l stalsis, relax involuntary sphincters, and increase

e d! ganglia (superior and inferio r mesenteri c,
..eel. Parasympathetic innervation derives fro m
e .a. gus nerve and from 5254 interm ed iate gray
-e spinal co rd, and it di stributes to intramural
secretomotor activities. The extrinsic innervation
of the intestines is integrated with the intrinsic
(enteri c) innervation.
131
REGIONAL NEUROSCIENCE Periph e ral Nervous System (PNS)-Autonomi c

Myenteric; p lexus lyi ng

Subserous plexus on longitud inal muscular
layer. Fi ne secondary
bu ndl~ crOSSing meshe$
l ongiludino l (duodenum of gu inea
inlJ<l muscular pig. Champy-Cou jard,
plexus o sm ic stai n, )( 20)

M yenteric
(Auerbach's)
plexus

Group of multipolar
Circu lar neurons, type II, in
intmmuscular ganglion of myenteric;
plexus plexus (ileum of cat.
Bielschowsky. silver
stain, x 200)

{Meissner's}
plexus
Periglandulnr
plexus

Submucous r>1~xus
(ascending colon 01
guinea pig. Stained by
gold impregnation, x 20

Lumen

\ f--- Mucosa and mucosill gla nds

- - Muscul"ris mUCOSiJe
:---Clands
............. Submucosa
,........... Circular muscle layer

I muscle ~lp
" 11iiN
I peritoneum
:~ .; :. ; .

FIGURE 11.48: ENTERIC NERVOUS SYSTEM: LONGITUDIN AL VIEW _ _ _ _ _ _ _ __ _

Th e ent eri c nervou s syste m prov ides intrin sic inn er than 20 distinct neurotransmitters have bee n

vation to th e small and large intestin es. The myen ide ntified in enteric ne urons (e.g., AC h, substance

teric (A uerbach's) and submu cosal (Meissner's) P, serotonin, vasoactive intes tinal peptide [VIP],

plexu ses are its main el ements. The myenteri c so matosta tin, nitri c ox ide INO]). ACh and sub

plexu s mainly controls motil ity; th e submu cosa l stance P are excitatory to smooth muscle, whereas.

plexu s mainly con trols fluid secretio n and absorp VIP and NO are inhibitory, Optim(l l functioni ng 0;

ti o n. Neurons of th is system intercon ned with the GI tract requires coordinated interactions

o ne another and w ith neuronal processes of the between end ocrin e, paracri ne, and neurocrine

ANS, although man y neuronal co mpo nents of this mediators.

network are free of auto nomic influ ence. More

132
_ pheraJ Nervous System (PNS)- Autonomic REGIO NAL NEUROSCIENCE

Vagus ne rves Preganglionic sympathetic

.. - - -- - Poslga nglionic sympath e tic

)
.. - - - - - -
- - -
Preganglionic parasympil thellc
Postg.m gl ionic p.:l rilsympathetic
Inlrinsic e meric ne urons !Y.t "-_ Spl.:mchnic
t Symllathetic - - - "'"(!fenIS
nc r\'e)
\ chain \
!, Celiac or

J mt!'Se nteric
g,1nglia

Mesen teric
nerves
[oI!owing ~'--l+--i l
blood
vessels

Mesentery

Petvic nerves

plexus

Cir cu lar - - ----:

muscle

plexus

mucoSde

Muc05a - - - - -

GURE 11 .49: ENTERIC NERVOUS SYSTEM: CRO SS-SECTIONA L VIEW _ _ _ __ _ __ _

- .he myenteric and submucosa l plexu ses, some
rons are innervated by sympathetic nerve fibers
m the sympath eti c chain and collateral ganglia
'"'W:I by vaga l or pe lvic splanchnic parasympa th etic
~ 'e fibers; o ther neuro ns are independent of
ionomic regulation. Auto nomic postganglionic
and intrinsic neufopeptidergic nerve fi bers supply
macrophages, T lymp hocytes, plasma cells, and
other ce ll s of th e immune system with innervation.
Thi s provides a regulatory network that mod ulates
GI tract host defenses and immune reacti vity of
gut-associated lymphoid tissue (G ALT).

133
REGIONAL NEUROSCIENCE
Periph eral Nervo us Syste m (PNS)- Auto nomk

Sympathetic fibert
- - - - - Preganglionic
~r"".~
" '' ..
- - - - - Postga nglionic

Pa rasympathetic fibers
f I' I Dorsal roOi
ganglion
Hj,J
.=%\ '11'
- - - - - Preganglio nic
- - - - - Postganglionic
Affe re nt fibers
left grealer
Ihorolci<

n. ~
spklnchnic:; n.
Righi gre<ller thoracic spl,mchnic
Poste rior v.agal lrunk -\----\ nV t-- A nterior vagal ''\(
l x.
.)
,
RighI phrenic lru""

~,J
,r'
\
I

Common areas of
rererred pain in
biliary di5eases

tt\ t
L
t <';ji. ,w,~-;",,;

,
r - Anterior vagal tfonk
Celiac ganglia
H~ ....llc tri;>d
j

' - '- - Splenk artery

~
. Aorta

artery aocl plexus

~~~
,.......
c~
.
FIGURE 11,50: INNERVATION OF THE LIVER AND THE BILIARY TRACT
_ _ _ _ _ _ _ __

Sympathetic nerve fibers to the liver derive from
T7-10 spinal cord and distribute main ly via th e
ce liac ganglion and its associated plex us. Parasym
pathetic nerve fibers to the liver derive from the
abdom inal vagu s nerve. Postgangli onic n o r adre n er~
gic sympathetic nerve fibers end directly adjacent
to hepatocytes and initiate glycogenolysis, increase
blood glucose for fight-or-flight respo nses, and
induce gluconeogenesis. Autonomic innervation
helps to regulate vascu lar, secretory, and phago
cytic processes in the liver. The gallbladder, espe
cially the sphincter ampull ae and the sphin cter ex
the choledochal duct is also supplied by au!o
nomic nerve fibers. The sympatheti cs ca use
contractio n of the sphincters and dilation of the
gallbladder; the parasympathetics cause open i~
the sph incters and contractio n of the ga ll blad ~

134
 -oheral Nervous System (PNS)-Autonomic REG IONAL NEUROSCIENCE

cortex
OH
,.,,,. !
CH.CHr t-.;H
A CH J
OHJ...y' Conversion
OH

;
sti mulated
by cortisol ,
-;

pressure
elevation
OU/pUi

Glycogenolysis;
hyperglycemia

Urinary
excretion
of metabolites
FfA
(free
fatty
acid s)
- ?}~
lipolysis

-:C U RE 11.51 : INNERVATION OF THE ADRENA L GLAND _ _ _ _ _ _ _ _ _ _ _ __

-" path etic preganglionic nerve fibers from neu
III1Sin th e rl O-L1 intermediolateral ce ll column
~s throu gh the sympathetic chain and travel in
~ nc hni c nerves to directly innervate adrenal
-edullary chromaffin cells, which fun ct ion as sym
~.h e ti c ganglion cells. An adrenal portal system
D)I1 \ eys blood fro m the adrenal cortex direc tly to
~ adrenal medulla. Highly concentrated co rtisol
-;:.e~ved from action of the hypothalamo-pituitary
ildrenal (HPA) axis bathes the chro m affi n ce lis,
inducing the enzyme phenylethanolamin e
N-methyl tran sferase (PNMT), which is responsible
for the synthesis of epinephrine. Approxim ate ly
70% to 80% of th e adrena l medull ary o utput of
catecho lamines is epinephrine; the remainin g
output is norepinephrine. Both epinephrine and
norepinephrine can be taken up into sympathetic
postganglion ic noradrenergic nerve termi nals by
the high-affinity uptake carrier and subseq uently
released.
135
REGIONAL NEUROSCIENCE Peri phe ral Nervous Syste m (PNS)- Aut on or-"

~ 2nd lumbar sympatheTic tfunk gan&lion

Gray and
wh ile ram i [nterrnesentenc (abdominal aortic) p lexus
communlc,:mles ~
~ Infenor mesen teric gan glion
Right __
Lumbar splar1chn ic nn.
svrnp all1elic (w l1k
a'nd il~ . IJ
ird lu mb'!l gilnghon ~
':r lnferio. mesen teric.i'l.
G ray ra mi and plexus

/ Superior hypogastric
plex us Ip resac rill n.)

Right and lei! /Superior rectal a.

hypgaslric 1) 1'1 . i.Jnd plexu!>

1st sacral Nerves 'rom inferior

hypogastric plel(u~!t
s\'mpathetic
trunk f.!<l n~lf
to sigmoid ilnd
descenc1ing colon
Gray fil m!

ureteral plexus
Sacril l part o f /Seminal vesicle
sv mpil thetic
Ductus deferens

Solcral

~ ~p
c lIl!N
,,","'/; :
Pelvlt,;
spl.mchn ic n n.
(S.1C1i1 1 p a rasympa
............Vesical ple)(u~
TheTic OUlnOW) ~ ~
~ _
f - InferiOr rectal plexus

Pudendal ...
~Prostat ic pleKus
Cavernuu) plexus
Right iniC'rior
hypo g.ls tri t r - DorSJI n. of ptnis
(pelvic)

FIGURE 11.52: AUTONOMI C PELVIC NERVES AND GANG LI A

_ _ _ _ __ _ _ _ __ _

Th e sympath eti c nerve fibers supply the pelvis
th ro ugh the sympCl thetic tru nk gangli a and the
superior hypogas tri c pl exus. The fibers travel
along visceral and vascular nerves to the colo n,
the ureters, and the great vessels such as rhe
inferior mesen teric and common iliac vessels.
Parasympatheti c nerve fibers ari se fro m the 525";
int ermediate gray of the spinp.1co rd and trave l
via th e pe lvic splanchnic nerves to distri bute witf"-.
the branches of the inferior hypogastric plexuses.
The intramural parasympa thetic ganglia are in, or
adjacent to, the wall of the o rgan innervated .

136
 eral Nervous System (PNS)-Autonomi(" REGIONAL NEUROSCIENCE

p;lthelic fibers Solitary tract "'""""' ---::i~;'=:=--+-

' Dorsal vagal nucleu~
_ __ Preganglionic
. ... ...... Postganglion ic
.JIIII. ..F - - MeduJla oblongata

vasympalhetic fi b e~
- - - Prega nglion ic ) -- -- - - - - 1/'8" ' (X) n .
- ......... .. Postganglionic
Gray ramus , - -- - -- Spi na l ganglion
"'"(' r(' nt fibe rs
1-- - - - - D,,,,,""", m>ers

- - - S'P;",'I cord (TI O to l 1)

Ventra l ram us o f T J 1 - -- - --c."'-
(intercos tal".) ~~~

- pollhelic; Irunk .." . ';" - --< i-- - l",",'ho,,,;, splanchnic n.

~t5~'---- CO",,'I thoracic splanchnic n.

H lumbar spJ.m cnnic". ---,t"-;

~:--- A,,"oo,,,,, ganglion

1<>- -- R,", ganglion

U:~~j:--- Intermese nteric plex u~

,-f-"~~su perior hypogastric pleKus

(pre SaCr.l1 n.)

--::-~-- Hypogastric n.

Sacral plexus ---<:t~

Pe lvic splanchnic nn. - - - + -<E

~ - - - Infe rio r hypogastric (pelvic) ple)(U5

C URE 11 .53: INN ERVATION OF THE KIDNEY AND THE UPPER URETER _ __ _ _ _ __
"T"'l pathetic innervation of the kidney and the
4!JPer ureters arises from the T1 0ll intermedio
fal ("ell column preganglio nic neuro ns in the
nal co rd and travels th ro ugh the lower th oracic
axf upper lumba r splanchnic nerves to synapse in
~ celiac or aorti corenal gan glia, Postgangli onic
IDers travel in fascicles that accompany the upper
..r.:>teric, renal, pelvic, ca lyceal, and segmental
.ra nches o f the renal vessels. Parasympathetics are
. ~ rri b uted to renal ganglia by th e vagus nerve and
pelvic splanchni c nerves via a lo nger co urse
through o ther plexuses. The sympathetic nerve
fibers stimulate renin secretio n (and the renin
angiotensin-ald ostero ne system), decrease the
glomerul ar filtrati on rate (GFR), stimulate proximal
tubu le and co ll ecti ng du ct N aCI reabsorption, and
stimulate co ntractio n o f the ureters. Parasym
pathetic nerve fibers cause relaxatio n of smoo th
m uscle in th e pelvis, the calyces, and the upper
ureters.
137
REGIONAL NEUROSCIENCE Peri pheral Nervous System (PNS)-Autono mir

ndi Descending fihers Sympathetic fibers

fibE?rs _ _ _ _ Pregan glioilic
- - ............ Postga ngli onl(

Spinal
:Ii. Parasympathetic fibers
C rayramus Renal gangJi()('l
- - - - Preganglion ic
... ..... _ - - Postga nglior
ganglion
Somatic motor fibers
Renal artery
and plexus
Affe rent fibers

lumbar parI
spin., 1n.
of spinal cord
splandlnic nn. ~ J l '~ I Jl Imermesenleric plexus

' ~I

/ ' Superior hypogastric plexu s (pre 5acr~

'I
Ascending 9f-Des<:ending fibers
...,..H ypogastric nil.
fibers

Sacral
splanchnic nn.

,. l'rO';.:..
-" p.e'

no.

~1p
I'ude ndal n. mus ~
Sphincter urethrae in

.
.....
1. .... kI..
,~
" ,~~, ;;
deep perineal space
between layers of
urogenital diaphragm

FIGURE 11.54: INNERVATION OF THE URINARY BLADDER AND THE LOWER URETER _ _ __
Sympathetic innervati on of the bladder and the bladder via pelvi c splanchnic nerves. 5ympa th~

lower ureters derives m ainly from the L1-L2 pre nerves relax th e detrusor muscl e and contract

ganglioniC neurons in th e spi nal co rd and travels trigone and th e internal sphincter. Parasympa... ....

through sacral splanchni c nerves to the hypogastri c nerves contract the detrusor muscle and re la\. ~

plexus. Parasympathetic innervation derives (rom trigone and the internal sphincter, thu s stimu la

the 5254 intermediate gray o f the spinal cord and emptyi ng of the bladder.

distributes to intramural ganglia in the wall of the

138

 "~-:"'''r al Nervous System (PNS)-Autonomic REGIONAL NEUROSCIENCE

~:1! etic fibers

- - - - Preganglionic
-- - - - Postganglionic
splanchnic n.
=- - - CeliilC !\i1nglion
"1r.li5".m pathetic fibers (from T5 T9)
Superior mesenteric
Preganglionic
Lesser thoracic - -- - 7 ganglion
-- - - - Postganglionic splanchnic n.
:':C_---Aorlicorenal
~I iibers Lowest thoracic -----:000: ganglion
splanchnic n.
~-- Renal ganglion

~"---Interrnesenteric
plexus
;;;;tii~'---Inferior mesenteric
ganglion

Right testicular a. and p,,'"'---_j~

,---Ir- - - - Superior hypogastric

plexus (presacral n.l

splanchnic 1111.

Sacral Sacral
plexus

Pudendal 0.------\
~r- Pcd'od," o.
- -- - oors,ll n. oj penis
Inferior hypogastric (pelvic I I i-- - - Epididymis
Vesical - - - Ductus deferens

'-----Testis
Penile cavernous

GU RE 11.55: INNERVATION OF THE MALE REPRODUCTIVE ORGANS

_ _ _ _ _ _ _ __

- oathetic innervation of the male reproductive
,"":2J1 S derives from T1 O-L2 intermediolateral cell
~m n neurons and reaches the superior hypogas
.. - ole xus via thoracic and upper lumbar splanch
ne rves. Parasympathetic innervation derives
-;:r-1 the S2-4 intermediate gray of the spinal cord
travels to the inferior hypogastric plexus via
- .ic splanchnic nerves. Sympathetic nerves
~e contraction of the vas deferens and the pro
'aJc capsule, and contract the sphincter to the
bladder, which prevents retrograde ejaculation.
Sympathetics also contribute to vascular responses
in the penile corpora cavernosa related to erection;
beta-receptor blockade can result in erectile dys
function. Parasympathetic nerves regulate the vas
cular dilation that initiates and maintains penile
erection. Sympathetics and parasympathetics
must work together to optimize sexual and repro
ductive function.
139
REGIONAL NEUROSCIENCE Peripheral Nervous System (PN S)-Autonomic

Sympathetic fibers
Pre ganglionic

5th thoracic

--""-"",-,,
T 4 -~~ ;.W'

,ympathetlc I ;) ~
- - - - - - Postganglionic trun k gan~l ion - ...:.::...=---: . ,~_ _ Ll:fr svmpa\hc
'::5"_ trunk '
Parasympathetic fibers
- - - - - Preg<lnglioni {- Gray ramus - - - -
- - - - - - Postganglionic communiC<lns
Afferent fibers \-Vhi te ramus----""""
communicans
... ::::::::. Celiac gan~ ;i""

" , -_ _ Aor licorenal

Greater 1horacic ./.... ;;;:<""'- ganglio n
splanchnic 11.

I.e~se( th~l aciC .;:;q-~ mesenteric

splanchnIC n, T12 -:---_~ ganglion

loweSI lhof3CC ---...--:?.

-.=-:o.",
.:i L -... Ren;,1g'lngl,
splan<.:hnic 11 .

Inferio r
if:7 mesenteri C
ganglio ll

-',.- - n l "" lJ ~ (presacr,l .

Ov arian il. and plexus

.,. l&: Hypug..1 wk

Ovary
~ U terovaginal pie

Pelvic 5planchni<: nn. ltV" _ '" 'icsr ,

~{t'
Sacral plexus

1.1'<ri<." P\ldendal n. - - - '"ts.

~iii -9t ~ J Pudel1dal !\.
~il!!N
,,-'c."

FIGURE 11.56: INNERVATION OF THE FEMALE REPRODUCTIVE ORGANS _ _ _ _ _ _ __

Autonom ic nerves supplying the female reproduc
tive organs have a similar o rigin to those supplying
their male counterparts. Sympathetic nerves
sti mulate contraction o f the uterus, but the extent
of this action depends also o n hormonal receptor
responsiveness and neurotransmi tter receptor
expression. Sympathetics supply the vaginal
140
arteries, the vestibul ar glands, and the erectile
tissue. Parasympath etics supply the muscular and
mucous coats of the vagina and the urethra, stim..
late erecti le tissue of the vestibular bulb and the>
corpora cavernosa of the clitoris, and supply the
vestibular glands.
 .....I Co,d REGIONAL NEUROSCIENCE

Nuclear cell columns Laminae of Rexed

,-~-----------~------------~~~-------~,------~~

N udt:us posterior margin<JHs

(mllrgin<ll zone)
Subs tantia ge1atinosa (lamina II)
-'
\\,
N urieus propriu ~ of pCl slerior horn

'\.!uciell s do rsalis;
Clarke's colu mn
LT1-13 ) I
,:\',
L.;lI(:!T,ll basai lludeus

~;::>lI1 a l reticlIl;Jr zan",

iof"mcdio lilleral cell

o"iITln; sympathetic
:-:.lnglio nic neurons
--r-------=
~~iecliom('di<l cell
mn: parasympathetic
_-1-------
-;;:anglionic neu ro ns
:_~I

Moto r FlexOfS
neurons

of li mb, Extensor s

[ce rvi cal

-nd lumbar D i ~I'1 1

pari of limb _-'-:t,,~,_
.lrgeme nts
01cord) Proxim.11p.lrl of li mb

Moto r neurons of trunk ~nd m:c.k

(Cl-3 and T2 12)

RGU RE 11 .5 7: CYTOARCH ITECTURE OF THE SPI NAL CORD GRAY MATTER _ _ _ _ _ ___

~e spinal cord gray matter is found in the interior
j the spinal co rd in a "butterfly" patte rn. It is sub
\'ided into 3 horn s: ( 1) dorsal horn-site of major
~ n sory processing, (2) intermediate gray w ith a la t
eral horn- site where preganglionic sympathetic
th ora columbar) and parasympathetic (sacral) cell
bodies resid e and interneuron al processin g occ urs,
.and (3) ventral horn-site where lower motor neu
rons (l MNs) reside and where convergi ng refiex
.and descending control of lMNs occ ur. Neuronal
cell groups appea r homo geneous in som e regions
01 gray matter, with some discrete nuclei (e.g.,
Clarke's nucl eus, su bstanti a gelatinosa). laminae
01Rexed, an alternate system of cytoarchitectural
classificatio n establi shed in the 19505, subdivides
the sp in al cord gray int o 10 lami nae. Th is system is
used extensively for the dorsal horn and the inter
mediate gray, laminae I to VII, particularly in con
junction w ith anatomical derails of nociceptive
processing, and for so me reflex and cere bellar pro
cess in g. Althou gh these laminae have distinctive
characteri stics at each segrnentCl lleve l, they show
some similarities across segments. The absolute
amount of spi nal cord gray is more ex tensive in
the cervical and lumbosacral en largements of the
sp in al cord, wh ich co rrespond to zones associ ated
\Nith limb innervation, than in th e upper cervica l,
thoracic, and sacral regions.
141
REGIONAL NEUROSCIENCE Spinal Cor<

Second cervical Fasciculus sracilis

Fa~ciCI.J lus cune;JIUS ~
Dorso lal !!ral ra$ciCulu S ~ ' " .. ...- "
(lis~ uer's Zone) \..

DOlsal spinocere bella r traCI--crr"

/:'
/ .... ,. Subslanlia gel<llinosa

/
- - - -- - - , - Nu cleus p ro pri us
Rostral spinocerebe llar tra ct - ~ /J.
lateral corti c[)~pi n a l traer ! . _ I.
Ruhrosplnal II<lel it '.." '"
Ven tral spinoc(>rebellar lIaCI -:\
-;--'-- - --f-- Spinal accessory nucleus

Anterolah:ral system (Spinothalamic .

trac l ami ~p;noreljcu'a r Iracl) - -- - - -",,""

:,.,1,'---
AnteriOf horn
Lateral (n le<iull,lfyl reticulospin.a l lract .,., ...._
u teral v~l l bu lospinal lracl ;:,
Medidl (po ntine) rt.'tiCulospi n.,l lfacl ---~
Arll E'rior corlicospinal trac t Anterior while commissu rE'
.
Seventh cervical

Medial100giludinal i<lsOculus
(with medial ves tibulospinal tract,
interstiliospinal tract, and lecto!>J)inal tract)
Descending monoamine <1.\
(norad renl!fgic, serotonger
'.
L

Oesceoding fibers from hyPolhala .......

r asnculus gradlisi - - -- - -
and br..in sl em 10 ~pinal Co
Fasc.iculus cuneatvs \

Dorsolateral fasc icul us \ \.~----\:.- -r---"",

(Li5S,HI~r's Zonel .,.-.'.. ,. '\ I -. . . ,

.,/ , ----~---MiI 'I~in al zone
DorSill spinocerebella r \rilCI / /
"'" -.~-------Substan\ia gelC!ti.......

..
.'.
1 tv - -'---- -- - --,;c,- Nucleus proprous
ROSlra l spinocerebellar trilCl--
lCl telal cortK.ospin.1l u.lC1
.
! ,' .
", ..,- ' " __

v
Rubrosptn"lt rilCI ~ "- V1 - - - -- - - - - ,, Intermediate F";

~\ ~~:i "I
V" x
Ve ntrill spinocere bellar trac t ~
'x IX _ l ower malar net.:
Anterolale ral system \ "'. VHI

) ~ :$ f# i in anterior horn
{SpinothalamiC t.ael and "

'r;"'~"w"""~" ' ~~'

~_ -~
. .,
.~ '- -,-' "~ #\ i

l ateral { medu ll.u ~'l reliculospinal lraCI

"' ..,,' v.,,;bu""p;",' ".c,
Medial (pOntine ) feticuiospinai lraci
~
. - - ..- .-

/
I

An" ';O""';'e comm'''"''
Medial longitud inal fasciculus
,. Pe rkiM
, MS..-o-
o,~
Amerior corticospinal trJ ct

FIGURE 11.58: SPINAL CORD LEVELS: CERVICAL, THORACIC, LUMBAR, AND SACRAL _ _ __
The o rganization of the gray matter info laminae of matter increases in absolute amount from caudal
Rexed is retained throughoul the spinal cord. The rostral. The dorsa l columns contain only fascicL
dorsal an d ventral horn s are larger and w ider at the gracili s be low T6; fasciculu s cu nea tus is added Ia:.
levels of the cervical and lumbosacra l enlarge erall y above T6. The sp inothalamic/ spino reticu!a
ments. The lateral horn is present from L1 to T2 . anterolateral system increases from caudal to ro;
Some nuclei, such as the intermediolateral cell fral. The descending upper motor neuron {U ,\\,
colum n w ith preganglio nic sympathetic neurons pathways diminish from rostral to caudal. Th e la....
(Tl -L2 lateral horn), Clarke's nucleus IC8-L2), and eral corticospinal tract loses more than half oi i:::
th e parasympathetic prega nglioni c nucleus (S2-4), axons as they synapse in the cervical segments -
are found only in ci rcumscribed regions. The w hite tract Ihen diminishes in size as it extends cauc~
142
 REGIONAL NEUROSCIENCE

Fasciculus 8r..olt5 - -- - - ,
FasCiculus (unealu$ \
Dorsolateral fasCiculus \ \.. ," -'
(Llssaue,'s Zone) ~\:
-, ,
""""""/),11,, '' ' ~
// - - -- -- -'.-- - M arginal zone

~ COrl icos pinal HilCI -f''-----i'' - - -- -- -- - -- Subslanri<l gelatinosa

f h. A.
- -- -- -- -- -- Nu cleu~ propriu s
Rubrospinal traCl-~\i-,o,--c" c -i t ) ____-------- -+1- Nud~u, dorS<l liso( Clarke
,\
\ - - - - - --f'- - latel al horn
-p, noct-,ebella( I,act ----"";;:"
"'" I
In te rmediolateral cell colum n
"'nlerolatt!ra lsystem '- >. .." !If
'.e - - - - ---:::;,:!---- l ower mOIOT neurons
_: ' Olhal,lmic trael and in an terior hom

r
~
'--
- .enulia' y) rehcuiospin,-,Itraci
Laleral v~hbulospina l lracl
\\~' a l (pontine) reliculosplnaJ Iract
' -~~~~ ;
Anle rior white commissure
Medial longitudinal iasciculus
Anterior corticospinal lract
,
' ..
....
..
D escending monoamine axon s
(nnradrenergk, sero ton~ergic)
Descending fibers irom hypot halam us
and hrain stem 10 spi nal cord

thoracic Fasciculus gradli.. - - - - - ,/\ .~. ___ .__

Dorw laleral folSciculus - - - , / \

(U,,,,,,,', Zo", ) //A '
-~I spinocerebellar Iract ------,~

~2====~~~===
I
" .,---- ~ ij' Substan
Milrginaltiazone
ge la tino 5"
I h. II. "til N udeu~ IJroprius
. ll eral cor ticospinal lr<lCI _ _, / L_-:'-:-.

Rubrospinal lract l"

--;--,-----:---
, '
. IV
V

"
VII
~_--------- Nucleus ool'Sdlis oi Clarke
t '- ] late ral ho rn
X" "- /f Intermed io lateral cell column
~tr.J1 spinocerebellar Irao - --"'" 'j~.

\ I
'"O
w (j_ _ ---i"'--_"o wer motor Jl~urOIlS
"metOlaleral sy51em _ ___C'''~
ISpi no thal,lmic. tract ,lnd
"" in ,Interior hom

spinore tieular (fac tt -:...: _ :.' Jl "

'77 ,;YI'\
L.i.' ('ral (medullary) ft!liculosplnallract
Lateral vestibulospinal tract
M edial (ponllnel retlcu l()sprn al tract ? / / riiY
AnterIor while comnussurt:!
Medra110nglludlnal i:)sticulu s
AnteuorcortlCospmaltract
J. Perkins
M5,MfA
" lION
.,,""

143
REGIONAL NEUROSCIENCE Spinal Cor..:.

First lumbar Fasciculus gracilis ~/_/ '~ __. " -y- ______
-~ ~,

,/' ~
~
~"
Dorsolateral fasciculus \ 1/ ';j '. .
(Lissauer's Zone) X
~ \
.,;:'~ ' . ,
': ! -:'%4" .;.
Marginal zone

Dorsal spinocerebellar tract I /!:__,:fi:,\,,~,., . :~ .' '_. .~, \

'.
Substantia gelatinosa
Nucleus proprius

Lateral corticospinal tract / / / h~ k . '{g;;~/ ;;~~ tJ?lt~~~_____'-'c Nucleus dorsalis of Clarke

. r . \..- -;--; -{ A~ .-):". __ __ .ffJjJ(" : :~
Rubrospinal tract :, '-. ___ '~' - . ,~-"t: :;>I:~_'~'>'~

\\~\:'..' \ l~-\\ ' "~"'\

' ' \$, .k

~j
Lateral horn

Ventral spinocerebellar IracI--- 'o,

, n~ '.. <p'\ } Intermediolateral cell COlli'

Anterolateral system
\\"
\ .....' \ \.
'X
i '~"
\@<@
. "-
JL
,,,
Lower motor neurons

(SpinothalamiC tract and

splnoretlcular tract)
\,;'i:; ',-....
~~:: - ..J~: ~
~ ....
I;~
_
~1 1 .;~q{'

'
in anterior horn

~<~ l , _ J"/.I,jfo"""'<

Lateral (medullary) retlculospmal tract

Lateral vestibulospinal tract
Medial (pontine) rctlculospmal tract
/ : ; ",:J' \. ~.
Anterior white commissure
Medlallongltudmal tasoculus
Anterior corticospinal tract Descending monoamine 2VP'
,--

~
..
,i .. .

{noradrenergic. seroton ge' f

Descending fiber; from hypoth a l ~- _
and brain stem to spin21Cl

Third lumbar Fasciculus graciliS -----~

Dorsolateral fasciculus \" /-" -\"-'-~r---" - '- ~'"'.

(Lissauer's Zone) .'
" " :"
.:' . ., ~
.~ Marginal zone
/ "<' .,
/," .~ .Il.. "';;" .;. "':",, ' \ Substantia gelatinosa

Dorsal spinocerebellar tract , ~~~- ~ - ..... ::~luW:'$

. ' Nucleus propriUS

Lateral corticospinal tract t
"~ ~
" Nucleus dorsalis of Cl ar>.e-
Rubrospinal trac t

Ventral spinocerebellar trac t -----L

"".\ " , ,..... , ~

~ ..
'

Anterolateral system .; / Lower motor neurons

(Spinothalamic tract and in anterior horn
spinoreticular tract)

Lateral (medullary) reticulospinal tract ;r.

Lateral vestibulospinal tract~ Anleriorwhite commissure
Medial (pontine) reticulospinal tract------./ J. Pe rlci
Medial longitudinal fasciculus MS, ' ''' '
C'l l@l'\l
corticospinal tract "'," "

144
...,1Cord REGIONAL NEUROSCIENCE

~' , filsOrulus8rac;' i~ ~

Dorsolateral fasciculu~
/ "\
/"

__
--... --..."

(Ussauer's Zone) \
,.,,-"-,~----- Marginal zo ne
/ /_'

DoHill spinocere be ll ar tra ct ----~. ~

-'''---''', - - - Substan ti~ g elil tinOSJ
i 11-:-
l.ller,,1 c()rti<."OSpinal tract ----+-~. --'----l,- - -- Nucleu~ proprius
_
Rubrospinal 'ract ___71l!.12\.-"7--. -/
f ~. - -- - ---'\;-- - - Intermediale gray

V,,,,,I " '"o<.,.b.lI" "''''-----(\

Ante ro liJ teral sy~lem ~' \ \
j
~I__--f_---low{' r motor ne uro ns
(Spino thalamic trar.! and ~)... ..:~ in an terio r horn
spinorelicula r I racl) ~"-. ...:

~." ,,,,,,,"",,,, ,,,,,,,10>00001 ",0<' '. ~

latelal vestibulospinal tract / /
,_._--- ....
MW.JI (pontinel reliculospinal tract Amerio( while commtssure
Medial l(lfl~itul linal (asciculus :4
A
Anterior oonicospinclt Ir,let :.-~- .......:
Descendins monoamine ,1xon~
(noradrenerglc, serotongergi<:l
Descending tlber$ flom hypoth'll"mu~
and I.Iraul stem to spi n~ 1 COld

F",.dculus grac ilisl - - - -_

,~-
Dorsolateral fasciculus - - -
(Liss~ uer's Zone) )< .I ,, ~ . --c,,--- -- ,lvl<lfginal Z<"Ine
~
/
Dorsalspinocerel>ellar 'r <1C'----,
f'/~
",/ - I~ -c;-'"',--- -- Substan ti a gelalinosa

\V. . - - - - - ' : -- -- Nucleus p ropri us

lateral corticosl'ina l 1'.lCI / ::' ......
V
Rurnospinal lracl ... " x -"----'~-- Sacral parasympclthetic nucleus
VI

---+
--- -
VII

Vf'ntral spinocerebellar tract

VIII
IX
Anterolateral system IX ;
~~-----,#----lower motor neurons
(Spinolhal.1mic " ad and \,:., ,
in ,1ntefiOr horn
spinme ticular Iracl) ~ _, _ ~

". llm,doII,cy) ",'k"IO>P'"" ',"d ~'

.
lateral vestibulo spin.ll lrad / /
An terior while (.;ommissure
Medial (pontine) rel iculospin~ 1 II.KI ,. Perk ins
Mediili iongi ludinal faSCIculus MS,Mf"
Anlen O. corl,cosI'inallr.lCt '''--,''

145
REGIONAL NEUROSCIENCE Spinal Con:

Dorsal horn
interneuron

mOlor neuron

'<om

Dorsal horn ,
interneuron .

a motor axOll

Schematic representation of motor neurons

" ~r~-T

,,
i i
'

\'/

~~
In cervical In lumbar
enlargement enlargement
o f spinal cord of spinal cord
CON
: ......:::

FIGURE IL59: SPINAL CORD LOWER MOTO R NEURON CONTROl _ _ _ _ _ _ _ _ __

Lower motor neurons are localized in the ce rvi ca l,
thoracic, lumbar, and sacral segments in th e ventral
(anterior) horn of th e spinal cord. LMNs also have a
med ial to latera l and dorsal to ventral organization.
LMNs supplying trunk musculature are found med i
ally and vent rally; LMNs innervating more distal
musculature are found dorsally and laterally. Thi s
o rganization also is apparent in the topography of
UMN control of LMNs. UMNs regulating fi ne
hand and finger movements are sent from the
146
co rticospinal system to terminate o n dorsal and
lateral LMNs. UMNs regulating basic truncal ton e
and posture are sent from the reticul osp in al and
vestibu losp inal systems to terminate on ventral
and med ial LMNs. Reflex pathways regu late LM i\.
activity through monosynaptic (muscle stretch
reflex la afferents) or polysynaptic (flexor or cuta
neous reflex aHerents) pathways. Superi mposed 0 :"
this o rganizatio n is the descending UMN control
and coordinatio n of LMNs.
 Co rd REGIONAL NEUROSCIENCE

8. Stretch reflex
'- '-....l ere n! inhibilion (reciprocal inhibition) C. Realrren! inhibition O. Tendon organ reflex
- - - -- -- 1'I-'-''----'-'--'--- . -- --'-- ' - -- - - 1' rl--'--"'-===
...... ~O r spindle From exte05()( spindle From eJC l ensor tendon
..i. iibers) receptor (Ia fibers) organ (Ib ~be rs )

.... ,-,, --r-j'
\
I i
-. . ... . .

.
,. / J
'. \;' ' '.
",
f ~ I~ . ' ;
. I

\
Rl?OsOaw --'--F
cell s '!
~ ~
\

.
\
\
,, ___ J
'
,
~ Inhibitory
synapse

Excita tory
synapse
To flexors __i1~t To flexo rs

Collateral s
oors To extensors
To synergistic muscles
To extensors

E. Flexor withdrawal reflex F. Renshaw cell bias

-----~

~i;ve fi bers

\
" \ Exdte s

".Jr__ "',.;
I , '""T- - phas ic
~, f, r{ ' / j O"o~
synapse
',,'h:;;

.
l.ewrs Toilexors '.... ~ ~ To fl exors
J. Perkins
/tis. MfA Inhibits to nic ~ _ __ _ _
To extensors :v~ '" exten sors To extensors

URE 11.60: SPINAL SOMATIC REFLEX PATHWAYS

_ _ _ _ _ _ _ _ _ _ _ _ _ __

to ":Ie muscle stretch reflex, la afferents excite the
lonymous LMN pool directly and inhibit the
;:a gonist LMN pool reciprocally via la inhibitory
e-neurons. The Golgi tendon organ (GTO) reflex
........bits the homonymou s LMN pool disynaptically
.rrl excites the antagonist LMN pool reciprocally.
=aor reflex responses excite a larger pool of
_ .\\!s, with reciprocal inhibition of the antagonist
_., ~s, to bring about a protective w ithdrawal
~p on se from a noxious stimulus. These refl exes
can extend throu ghout th e spinal cord. When an
LMN fires an action potential, it excites a Renshaw
cell, w hich inhibi ts the LMN, thereby ensuring a
cl ean slate for the next set of inputs to it. Rensh aw
cells receive input from axon collaterals of both
fiexor and extensor LMNs to exert an inhibitory
bias that is mainly directed toward inhibition of
extensor LMNs and reciprocal excitation of flexor
LMNs. Thus, the Renshaw cells favor flexor move
ments and help to inhibi t extensor movem ents.
14 7
REGIONAL NEUROSCIENCE Spinal

Muscle ilnd joint receptors

Alpha motor nel.J ron~ 10 e)(traTusai
striated mu~cle end plates - - - -- - ----
G a mma molor nellro ns [0 inl! <Iiu5al
stri~t8d musci!:! end pla te s - - - -- - ------
I~ (Au ) iiber~ from <lnnulospiral
endi n R~ (pJOprioceptloni - -- - -- - - "

11 (AIlI fibers fwm flo w er $pray

end ings (proprioception): - - -- - - ' " /

fro m paciniiorm co rpu_Kl es (p ress ure) - . / /

and pacinian c orpu scle~ (pressure) /

III (Ao) fillers from free nerve endinl\s

a nd from ,orn e specialized end ings'
(pain .md some press ure) /

IV lunmve hnatedl fiht'r~ from iree

nerve e~dings (pain) /

Ib (Aa) !iber~ from G oigi tend oll WI An fib ers hom Golfl j-tvp e (: nd ing,
organs (pro prioception) - -
1/1 All fibe rs from pacini/orm
corpu'>(;les c:md Ruffin i terminals

- - AS and C tibers irom fre!? nerve ~nd ings

Alphil illo tor r'u=uron to extrafu sal

muscle fibe r end piOlte5 ---

Gilmma motor neuron to inlraiu ;al

Intra(u sal muscle fib ers
muscle fiber end pla tes /"

II (A ~) fiber irom fl o wer spra y e ndi ngs

la IAn) tIber from annulospiral endings " - yl plilleend lngs

- - - - s h l ? ath
- . ,pace
, - - - - l vmph
Nudear bag fil ter
~fp
c lCuN
"'--;,' ,:
' Nuclear <.h ain ti ber
Detai l of
muscle sp indle Effere nt fibers
Affe re nt fibe rs

FIGURE 11.61; MUSCLE AND JOINT RECEPTORS AND MUSCLE SPINDLES _ _ _ _ _ __

Joints are innervated by a host of afferent recep
tors: bare nerve endings, Golgi-type endings,
pacin iform endin gs, Ruffini-like endings, and other
encapsulated endin gs. GTOs inn ervate tend ons
and respo nd to stretch w ith increased discharge,
causing disynaptic inhibitio n of the LMNs that con
tract the homonymous m uscles. The mu scle sp in
dles, com plex sensory receptors w ithin th e muscle,
are arran ged in parallel w ith the extrafusal (skeletal)
muscle fi bers. These receptors contain small
intrafusa l muscle fibers that are stretched when
the muscle is stretched. The la afferent fro m the
muscle sp indle excites the homonymou s LM N
pool mon osynapti cally and respond s to both the
length and the velocity (change in length w ith
respect to time) of the extrafusal muscle fi ber.
These muscle reflexes ass ist in maintainin g hom p,:",
stasis durin g contraction and help to regul ate m L~
cle tone during movement.

148
 Cord NEGIONAL NEUROSCIENCE

B.-Jsetine fiung ...

-
Extra/usal m uscle tiber

neuron s ..........

Gam ma mo tor neurons

A. Pas~'~ . lretch. Both ;mra fu, ai 3r1d f'xlloru S<ll mu 5d1:! fibe rs , Ire tc ned,
spindle-.; 3C[' \'';lIed. Rp,lcx\'ia 1:1 libers and ol\Pha mOtOf noWron~ c~uses
Golgi te ndo n s.eco nrlu r)' COrltrxtion (b~sis of Stfe l<;h ,eilE-xes, slI<:;h iii knee jerlo.
o rgan - - - -*It: Stretch is too w eak to ac tivate Golg; len dOl'1 organs

Al pha activation
from brain

Exlra/u sal muscle fiber

' nlTa ru~al muscle liber
Inhibitor), intern euron
neurons -+

Gilmm<l m OlOr neurons

B. Ad ~ contraction. Cen1ral ~(jtJ.tion or alpha nIotor "eurOn~ 0111"
caUWi con l T~ cljon 01 "\{,,,iu!.iI1muS(le fiber s " iln tO n~ l'quenl rel ,lx~"on
Ge lg' tendo n of inlrafuSllltihcr,; spind les nOI actj,' atE'd, Tension j< 10\\; (I t'~ 1 not M!IUSI
to inut:'dsed r(>~isIJn Cf;". Tenoon organ aCli\B Ie<:! , (dUJonS fel~xation Th,s
olg.ln -----<OO':j;Ji.~ is theoretical only. Act",. (oO't ra<:1;on imol\E'~ n! coar ln alion

Jbfibe r;;- _ ~ , Alpha ;Iod 8<l mma

aClivJlio ll from nrain
13 iiber5
(maintains bJseline)

Ext raru 5<l1 muscle iiber

InlrafusaJ muscle fj IJer

Gamma motor
neurons
-
C. Ad hot Contra ct ion with alphag~mma coactivation.l nlrJ iu s.a1as
...-ell as Io'xrraiusill tiber5 corur.J.c t spindle~ ,u:;ti..ated, reinforcing
contraction Jl;muluJ ,~a 1;'1 iibers in accord with resIJlanf.t.
Te ll(i()n organ ~ l iva tE'd. causi ng rr. IOIxation if load is Io n !!reJ I

fiG URE 11 .62: THE MUSClE STRETCH

MOTORNEURONS REFLEX AND ITS CENTRAL CONTROL VIA GAMMA
_________________________________________

..Juring passive stretch, a muscle stretch reflex
of'\ci tes ho monymou s LMN s, whi ch results in mus-
e contraction to restore homeostasis. If active
co ntracti on occurs without y.LMN activati on, th e
"'1u scle spindle "unloads" and the tension in the
'f'Irrafusa\ fibers is redu ced, resultin g in diminish ed
ring of bo th la and group II afferen l s. Ho\vever,
",n en LMNs co ntract becau se of brain stem UMN
activity o r voluntary corticosp inal activity, a.-LMNs
:ond y-LMNs are adiva ted together. This process,
a-"fCoactivation, ensu res that the tensio n on the
muscle sp indle (throu gh th e intrafusal innervati o n
by y.fibers) adj usts immediately, that is, as the
extrafu sal mu scle contracti o n (thro ugh a-fiber
innerva tio n) occurs. In norma l physio logical cir
cumstances, a-LMN s and "tLMNs are coactivated,
although they can be modulated separately by cen
tral nellronal circuits. If y.LMNs are differenlially
activated in pathol ogical circum sta nces, increased
muscle tone and spasticity ensue.
149
REGIONAL N EU ROSCIE NCE Brain Stem and Cerebellu

Fasciculus cuneatus

~
FaSCI<.UIUSgracil i S
Spinal nucle us eNv

'-..

level of section
Spinal tracl eNv

DOf'NI l
spinocerebellar
tract - -- _ __
y l
I
I
,.... -~ "\ I"
,I

"

~
:,>;,~:J.9:
~~~ '-

'.
. tract

, -
X l a tera l ( on icospinal

Central canal

Decussation oi
'f:;'if
<;.~

Ven tral '%fW pyramids

spinocerebellar ' !?
tract

Spinothalamic! XI
"...

-- IJ
j'::1J,~
spinoreticu1ar
Inll:;l /

NudeusCN XI ..,...
__ ..cow:...,
I
PyramId "w'"
Section 1: Med ulla-Spinal Co rd Transition- Decussa ti on of the Pyra mids

Dorsal Fasciculus cuneatus

('~:t)

~' ,T'J'
spinocere bellar
tract

f, '.f
'
.~y-
~ Nucleus gracilis
,
, ':
.,
'-.
Spinal trac t 1/ >\, Nudeus cuneatus

AY

i\
C NV ~
Spinal nucleus

'iF -',\
~"",--.c;'r\- Tra c tu 5 soli tanus
CN V

-
Level of se ction
---f-- D o rsal m Olor
Nucleus ambiguus ---.I.
- 't-__
, nucleus of X
Ven tral
spinocerebellar -J- Nucleus eN Xli

'l
lrael ./
SpinOl halamiC/ V"- Medial lo ngiludinal
fasd culus
'tract
",""'eli",'"
- ---',
/~'l'
Do rsal accesso ry olive

Medial accessory o live

pyramids" .I_"COW:~
._ I
Py ramid
Nucleus solilarius 'Wi m

Section 2: Medulla- Level of the Dorsal Column Nuclei

FIGURE 11.63: BRAIN STEM CROSS-SECTIONAL ANATOMY _ _ _ _ _ __ _ _ __ _

150

 Sie m and Cerebellum REGIONAL NEUROSCIENCE

Nucleus runeatus
External (lateral)
cu nea te nucle us Nucleus gracilis
Nucleus
Inferior S(IIit.Jrlius
cerebella r
peduncle ~vit
do rsal
spinoce rebellar ',.
r . . / - (:eo,,,1canal

nucleus X
trac! ~ ~."l'1 Nucleus eN XII
Tractus '
~
,

Level of section SOlitar~jUs: <;:' :.)..._- Imern al

arcuate
Spinal tract . ~'.
(NV ~,
fibres

CNV - -J
"
Spi nal nucleus ,:.,
;~~b~%~~J2~[
Medial lo ngi tu dinal
fasciculus
Spino thala micl
Tectospinal tract
spinore ticular
Medial lemniscus
Inferior olivary nucle us

ivledial accessory olivary nucle us

...., 3: Medu lla-level of the Obex

Exle rn')l c uneale Nucle us cu neiltus

nucleus r - N,,,I,,,, so!itari us

_ T""" " solitarius

Infe rior
cere bellar ~. 000 , 1mo tor nucleus o f X
ped uncle
~.;'~!"J!~~;;~;: Choroid plexus

0-- - fe,""" ve ntricle

Leyel of sectio n
CNX
Spinotha!.amic/
spinoreticula r
,~- , ,,,!;,l longitudinai
tract
fasc iculus
Nucleus ambiguus
Tectospinal lracl
Inferior olivaI)' nucle us
~ M,di,'lemniscus
Dorsal ilccessory oliva ry nucleus

CNX II
L ",,,fj, 1accessory olivary nucleus

!Clion 4: Medulla-Level of th e Inferior O li ve

1 ': 1
REGIONAL NEUROSCIENCE Brain Ste m and Cerebell lr"

Tra c tu s so lil.)rillS Nucleus solitariu!>

Inlerior r- Medial vestibular nucle us
ve sTibular
nucleus
formation
Inferior

cerebellar

peduncle

Dorsal
mOlo r
nucleus Dorsal m O':
level of sedion
nucleus 0'

longitudina
Spinal tract eN v fasciculus

Spinal nucleus eN \I TeCiospinal tra.,.

Spino thal am ic/spinarelieular Iracl ' - Medial lemniscus

Infe ri or olivary nucleus
.,jOHN"'CI'f"'IC.~
e l(DN I
,w,'",

Section 5: Medulla- level of eN x and the Vestibu lar Nuclei

N ucleus prepositus
Dor~3JcochleM Med ial
ve stib ular iormalion Medi'll longitu dinal
nucleus
nucleus fasciculus
Inferior

traci
Ven lral
cochleM
nucleus Raphe
nuclei
level of !iection (obs (l-~;
(NVIII pallidm

Spina l tract eN v
Spln,l l nucleus eN v
Inferior vestibu lar nucleus
Spinothalam ic/spinoret icular tract
~ Inferior olivary nuclew.
Med ial lemniscus

Central te gmenl alu3cl Corticospinal ~HN ACR4.'1 c.

Middle> cerebellar pedunde L Pon tine nuclei tract e~

Section 6: Medullo-Pontine Junction-Level of the Coch lear Nuclei

152

 . Slem and Cerebellum REGIONA L NEUROSCIE NCE

Superior vestibular nucleus l <lteral vestibula r nucleus

N ucle us eN VI
Superio r cerebell.tr peduncle ~ \
Supedor o livary nucleus
Dent.}le nude us ______ _ \

Middle cerebellar Medial

peduncle _ _ _ longitudinal
fasci cu lus
In(erior cerebellar ___
peduncle- - Fibers
( NVII

Level of section
Spinal tract eNv
Spinal nucleus eN v
---=========' tract

Nucleu s eN VII -~"'""------

CNVII" 7<
CN VI I - -
Spinolhalamic/spinoretlcular Iract
"--- R,>ph,. n ucleus tmagnu!.)
Central tegmental tract

Tr.lpezoid body
~"'- I~e,j;a' lem n i >CU5
Corticospinallract
"' ( NVI
Po ntine nuclei

IOn 7: Pons-level of the Facial Nucleu s

Dentate nudeus \ Superior cerebellar peduncle

Superior vestibular
nucleus
Inferior
cerebellar
pedunde _ _ _
,~
' r Globose 3r1 d emboliform nuclei
i CNVII

Nucleus
M iddle cerebellar ~ CNVI
pedunde (MCP ) _
Med ial

Level of section vestibula r

lateral
nudeus
~
' \fl~:; ;'?<?2i~~~~
.
longitudinal
fasciculus

Fibe rs
Spinal tract CN V (NVI
Medial vesti bula r nude us
Tecto spinal
Spinal nucle us C N V tract
Sp i no th ala mic/spi noreticular tract --C" ot,, 'l tegmenl al
tract
Trape zoid body
Media l le mniscus

Cort icospinal lrac t

Pontine nude i

Section 8: Pons-Level of th e Genu of the Facial Nerve

153

REGIONAL NEUROSCIENCE Brain Stem and Cerebell

l ateral Medial l ocus

(' r.
~k....
......
~,
t,"
para brachial
nucle us
parabrachial
nucleus
coeruleus Fourth
ventricle
Jr Super ior
cerebellar

- \ ,,,
peduncle ~~ . /' 'Yl '
f .JrT,;r--P--
Medial
longituo
fascicul\..,
M iddle
cerebellar

level of section
peduncle >-' tr;) c t

Mesencephalic Raphe
nucleusCN V nu deu~
(ponl lS
CNV~'~

Main !chief) en~ly nucleus eN v J

.\I\olor nucleus eN ' - Centrat legmer
Ira e!
5pinOlha tamiC/spinoreticular Iract
' - Medial lemniscus
Crossing fibers of middle cerebellar peduncle - 1
............IC_
Pontine nuclei
. W'::::
""" I

Section 9: Pons-l evel of Trigeminal Motor and Ma in Sensory Nuclei

Supe rior cerebd lar peduncle

locus coeruleus

, -:=
~ PeriaQueductal gray maltfc:

,., !,
-Z! 1
; Aqueduct
\ ./
l evel of section 1, ...,
- - - - Dorsal raphe nud'h .
Spino thalamid
spinoreticular tract
i:'
-,.. '<
Media l longiludina j
Medial lem ... i~cus -..../" fasciculus
Corticospinal
tract - - -
Central supelior

.-

Pontine raphe nucleus

nudei

~"CRAc..... c:
Cenlrallegmental tlael ,.......:,'

Section 10: Pons-Midbrain Junction- level of eN IV and locus Coeruleus

154
Jain Stem an d Cerebellum REGIONAL NEUROSCIENCE

Brac hium 01 in ferior !n feri()( coUiculu ~

Re ticular formation

Periaqu educlal gray mailer

-'_
;::;========;'2,=~N~'Ucieus eN IV
d Oor,,,1raphe nudell'
-----A
f)}~,
- .'vted iiliiongitu dinal fasck ul us

- -<-- Superio r cerebellar peduncle

~ de( u $S<l tl o nl
Cenlral
tegmen tal tract _ _/

Interpedu ncular nuclei

Medial lernniscus

Sobstanti;a nigra

Kti on 11: Midbrain-level of the Inferior Colli culus

Brachi um ot inferior Superior culilculll S

colliculus

Mesencephalic nucleus eNv

/~.--_ _- -\"- PeriQqueduClal gray mailer

I ;''....- . - - - - - --Aqueduct

l evel of section
:iC'-----~"'=::-- Nucleus of [ dinger-Westph<ll
Spinothalamid
spinoreticular tract Nudeus CN [II
Medial lemniscus
Medial lo ngitud inal
Cen(ral fasciculus
tegm ental
trac t
Ventral tegmental
Substantia decussation
nigra .,IOHN A"C:RA1i-""
e N II[ area II!lN
"....... :"

Red nucleus

:!Ection 12: Midbrain-Level of the Superior Colliculus

155
REGIO NA L NEUROSCIENCE Brain Stem and Cerebel

Sr i11 0lhala mic J Brac hiulll oi Inferior colliculus

sp ino reti cular
tract - -
Periaqueduct,,1gra y m;:ltter
Medial
lemniscus Aqueduct
Nucleus of
Edj l)ger-W~lphil

l evel of sec tio n

Cef~bra l peduncle
Optic Iract
SubSla,)lia nigra

Cerebe lloru b rot ha lam k IraCI

Central tegmenwl v act

Red nucleus
Medial longitudin al fascicu l u~

Sectio n 13: Midbrain- Level of the Med ial Ge ni culate Bod y

.-
,.jOHN A,CR4JC.,..w;'
,',w,""

Superior Peria<.jueduclal gr ay matter

Pul"inar colliculus
Posterior commissure

$p rnotha:
sprnorel'C::"
tract

Level of secti on

Laleral geniculate body (nucleu s) ~

lerr',""

Cerebri'll peduncle
Darksch
O pile. tract ---'
Cerebelloru broth alamic tract Medial Ion:;;
fascicu lu s
Re-d nucleus
~-, '?::%\ - -:;;~ Centra l tegmt:!nw,1
Subsl.1ntia nigra
~ - tract ..IOHNACAAc.
l(lIer.ll hypothalamic area
l . h}'PO thalamic area
Postenor
.,.....,
'We:;:

Section 14: MidbrainOiencephalon Jun ctio n-level of the Posterio r Commi ss ure

156
m Brain Stem a nd Ce rebellum
REGIONAL NEUROSCIENCE
- Molnr fibers JJ[ Oculomotor
- Se nso ry fibers (vII eye mU$cles excep t IV Trochle ar V Trigeminal
tho se b elow. Also ci1iarr Sensory to fa ce, sinuses, teeth,
iris, 5phiflc1erJ gen era/ scns"tro n to ,in /errOr 2;, of
o f al <:<I \,l [\,. lOn gu e .

Motor to muscl es
of mas'ical ion

Ner v u s intermediu.q
A,u/Q n(JfOlC - suu,,,.v<tlli,,y
sublingual, )"o imal glilnds
Sensory - t.lSle 10
anterior 2/3 oi to ngue,
soft palate
expressi on,
stapedius

VIII Vestibulocochlear

.,
IX G lossoph aryngeal
Sen )Qr~'- t.l ste to pOStNior
1/) 01 IOngUl?, general s e n~ ati O Il
to lo nsil, pharynx, midd le eilf
Motor - styl upharynge us,

pha ryngea l rnusc:ulillu re

x Vagus
Motor - to pharynx, Itlrynx
Autonomic /0 hf'an,
lu ngs, bro nchi, G I tra c t
Sensory - hear!, lungs
bra nch;, trachea, lafrnX,
p har ynx, G I tract,
external ear

Sternocle idomastoid.
t rapezi u~ (upper 2/ 3)

URE 11.64: CRANIAL NERVES: SCHEMATIC OF D ISTRIBUTION OF SENSORY, MOTOR, AND

AUTONOMIC FIBERS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

iiIf<al nerves (eNs) I and II, both sensory, are
_ :> trac ts derived fro m the neural tube. eNs III-XII
~rge from the brain stem and supply sensory
"-5V, VII-X), motor (e Ns III-VII, IX-XI I), and auto
lie (eNs III, VII, IX, X) nerve fibers to structu res
~ e head and the neck. All th e crani al nerves that
~' erge from the brain stem distri bute ipsilaterally
th eir target structures. Th e cranial nerve nudei,
except CN nucleus IV (trochlear) and some motor
co mpo nents of eN nucleus III (ocu lomotor), are
located ipsilateral to the pOin t of emergen ce of the
cranial nerve. Th e spin al accesso ry portion of CN
XI emerges from motor neuro ns in the ro stral spinal
cord, but it ascends throu gh th e foramen magnum
and th en exi ts w ith eNs IX and X; thus, it is con
sidered a cranial nerve.
157
REG IONAL NEUROSCIENCE Brain Stem and Cerebellu-'

OcUIOnlOtOt (III) n.

~ed nucleus

Superior ICldo ia l1(oilicul / / ' Ed inger-Westphd l nucleus

/
Termina tion si te<; fo r ." OculomOIOI nucleus
fibers in optic trac t, -\
C -I __ Trochlear nucleus

Trochlear( IV1 n.
./ nucleus of
,"'-10101
trigeminal n.
Tfigemlnal lV) n. inalM n.
and ga ngli()n ngli,
Principal (pontine)

sensory nu<:lelh

o( trigemi,lal n.~
AbdUCMS nucleus

Geniculate ganglion
of fada l n.

Facia! nucleus

_ _ __ _ _ _ _ Superi
or OInd inferior
Veslibulo
cochlea , (VIII) saliyatory nuclei

Ven tral (Ochl ear

nucle us /"
:lil",;o;'''''O\lr -- Nucleus ambiguus
Dorsal cochlear
nucleus ----""" ............ Glossopharyngeal (I X) n.
G los'>Oph'-lIyng~a l (I X)
~ Vagus (Xl n.
Veo;tibular nuclei ........... Accessory (XI) n.
V;)gus (X) n.
Dorsal (motor) vagal nucleus
Spina l tract and spina l Hypoglossal nucle us
nucleus of trigeminal n.
;,

~1p
_ Efferent fibers _ Motor
_ Affe rent fibers
_ Effe ren t fibers _ AutOllOmic c~

FIGURE 11.65: CRANIAL NERVES AND THEIR NUCLEI: VIEW FROM ABOVE
_ _ _ _ _ __
Lower motor neuro ns (lMNs) of the brain stem are
localized in a medial column (CN s III, IV, VI, XII I
and a lateral colun1n (eNs v and VII, nu cle us
ambiguu s ICNs IX, Xl, CN XI). Pregangli oni c
parasy mpathetic nuclei are fou nd med ially in the
Edinger-Westphal nucle us (e N III) and the dorsal
vagal nucleus (e N Xl and laterally in the superior
(eN VII) and inferior (eN IX) sa livatory nuclei.
Secondary sensory nuclei include the main sensory
and descending nuclei of eN V, the vestibular
nudei and the cochlea r nuclei (eN VIII ), and the
nucleus sol itarius (eN s VII. IX, and X). The superior
co!licul us and the lateral genicul ate body rece ive
seco nd ary sensory axo nal projectio ns fro m the
op tic tract; the inferior co lliculu s receives input
fro m th e cochlear nu clei and o th er accessory
auditory nuclei. The nuclei graciliS and cuneatus.
located in the medulla, receive input from dorsa
root ganglion cells, w hich convey epicritic
somatosensory modalities (fine discriminative
touch, vibrato ry sensation, joint position sense).

158

bell Stem and Cerebellum REGIONAL NEUROSCIENCE

;~~~I~~~"i"'~h;~;'i~"";:r "'O"'i'"

Supratroc hlear n. I ~'l . i In.

M ed ial re<l u. m. superi ori, m.
Superior obl iq ue m .
Nasoc:ilIi.)ry n. - - --:::::-_
Cnbifo rm pl,Ue oj ethmo id bone
CommOll ..nnu!ar tendon

O pticlili n. - -- ---
Opti c .;hi a,m - - - _ _
O phthalmic n
Pituit ary , talk (infundi bulum)
~:;;;;:==-; Maxillory n.
::;: M~nin !l~al brMICh oi ma~ill Jry n.
O cul omoto r (illi n.

Ma nd ibu lar n.
Tm ..:t.I..ar (IVI n. -==::::::::::~~
_ _- Men inge al brand ,
Abducens (V I) n . -
(ner~us SplnOSIJ ,) of m andibul ~r n.
Tentorial (meningeal)
S;:::::::::::-G.eluer
les5e' poel!osal n
n.
branch of ophthalmic Il.
f'~:'::~__--=::-;,~:, ~::::;: sanglio n
- - - -- \

;:. TO! n lOriu m ({'Iebelli

Supratrochlear n. (cul l _ _ _ _ - '

p~l pebl ae $upenOriS m. leu !)
Me dial and lateral
Ii I
branlh.., oj supraorbi tal n, (cut)
Lacrim al r"I. leut)

Infratro chleM n. --;:-:=====;~ Short cill ~ ly r"ln .

An leno r ethmoi dal A. Bllln<..h or" oeulOI'llOIOl n . to mfen"r ()1 ,lique m.

l ong ciliary nn. ","'=--C;I;. ~ S~ngJjoo

Oplic (Ill n. : ; : -- Mo'."p"" ym'P . ".~i,,) ",.,,'mm oculomotor o .
Sym~lhelic ' 0 0 1 ('(>011 iOl emal (,]roud ple~ u~
Nasociliary n. Sl'"f\j.QI)' ' 001 i,om nasociliary n.
Op hlhalmi'; n. -------"'=-'l Branch es 10 n-..edi.:d .:m d inferi", 1("<luS mm.

(e~~=====3!
"
I n (10 I~teldl re ctu s m .l
Ocu lomotor
Trochlear (IV)(IIn.I) n iii f ocu lomotor n.
Abducen, (VIr n.

lolemal c<I.Olid <I.

o.aphr<rgma scl ae
OculomolOf IIU) n. ~=:;;~
Trochlear (I V ) n. -
Pitultarv glan d

Internal urotid 3.

Abducen s (VI) n.
Ophthal mic n.
C.",~nou s

M..,ib'1 fI .

"l- RE 11.66: NERVES OF THE ORBIT AND THE CILIARY GANGLION: DORSAL VIEW AND
CROSS-SECTION THROUGH THE CAVERNOUS SINUS _ _ _ _ _ _ _ __
II ca rries visual information from the ipsilat eral
:.d . Axons from th e temporal hemiretin as
-'""kljn ipsilateral, wh ereas axons from the nasal
retinas cross the midline in the optic chiasm.
:1.\ons then enter the optic tract. eN s III (fro m
~ m otor nuclei), IV, and VI innervate the extrin
m uscles of the eye. Sensory portion s of th e oph
v
.mic division of e N supply gen eral sensation
.... e cornea and th e eyeball and provide th e affer
limb of the corn eal refl ex. Motor fibers of eN
nnervate the o rbicularis oculi muscle, closing
eye; these fibers constitute the efferent limb o f
~ corneal reflex. Parasympathetic preganglionic
.:lETS irom the Edin ger-Westphal nucl eus distribute
to the ci liary ganglion, which suppli es the pupillary
constri ctor mu scle and th e ci liary muscle (accom
modati on for near vision). Prega nglionic parasy m
pathetic axons from the superior salivatory nucleus
distribute to the pterygopalatine ganglion, w hich
supplies the lacrimal gland (tea r production).
Sympath etic postganglioni c nerve fibers from the
superior cervical ganglion supply the pupillary dil a
tor mu scl e and the superi o r tarsal muscle (dam age
results in m ild ptosis). e Ns III, IV, VI, and th e oph
thalmic and maxillary divisio ns of e N V traverse the
caverno us sinus and are vulnerable to dama ge
fro m cavernous si nus thrombosis.

159
REGIONAL NEUROSCIENCE Brain Stem and CerebeU......

Short ,iliary nn.

Lo ng (Wary n. Edin8e r-W~~tphal nll cleu. }
(autono mic) Oculom otor (111 :1 n.
Oculomotor nucle us

SenSOly ' 001 01 ciliary SlPng!ion

Superior oblique RI.

SuperiQr reclU. m . divi5ioo of oculomOlDf n. Abducen s nucleus

Levator SupetlOf and

p~ lpebrae inferim c.ollicuh
MJperia. i. m.

Su perio r
tarsal m.
(involuntary)

Sphincter

pup illae m.

D ilalor

pupillae m.
'\ Abducens ( VI) nerq
\TrOChl~ar{lV) nef\-e
\ Oculomoto r (JII) nerve
Inierlor oblique m. In ternal ca rotid a. and p lex~;

Infraorbilal n.
Inferiof reclllS rn. \ \ MaxilL-uy n .
Zygoma tic n.
MOlo< \pa' a5~mpathetic) . 001 oi ohl)' ganglion
la teral reetu. m. a.nd abducens n.
(turned bad )

Common annular ten oon

Levalor palpebrae
superioris m. ---~- Supel'lo obliq u e m .
Trochlear
{ (IV) nerve
Superio r l ec tus m. - - - -
Oculomoto r J

~ ~' 112'.
{ Abducen~ .Jt...~
(III) nfrve Medial rec:tu s m, - '
later,ll rectu5 m.
Illferior rec lus m. ~ (VI) nerve
. I
10ie li00 obItQu!;l m . d
,','.
MOlorfib en
Sen.ory fibt>rs

Sympathetic fibers

Parasympathetic fibe,~

FIGURE 11.67: EXTRAOCULAR NERVES (III, IV, AND VI) AND THE CILIARY GANGLION: vim
IN RELATION TO THE EYE _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _

CN VI innervates the lateral rectus muscle; damage results in paralysis of ipsilateral medial gaze}, aJ)I"
results in ipsilatera l paralysis of lateral gaze. CN IV the levator palpebrae superioris muscle (dama:?
innervates the superior oblique muscle; dam age results in profound ptOSiS). The ciliary ganglion
results in inability to loo k in and down (most con gives rise to postga ngli onic pa rasympathetic .l>.i.
spicu ous w hen climbing stairs, ste pping off a curb, that supply the pupillary co nstrictor muscle a
reading in bed). eN III (ocu lo mo tor nuclei) inner the ciliary muscle, Damage results in a fixed anG
va tes the medial rectus, superior rectus, inferio r dilated pupil that will not co nstrict for the pup
rectu s, and inferior ob lique muscles (damage light reflex and will not accommodate to near \ ~

160
 Stem and Cerebellum REGIONAL NEUROSCIENCE

Tri!emin~1 (VI ne rve and

- MOlor fibers t (sefTInuoJrl gJ ngllon
Tenlorial (mening.,....tJ
- - SenSOl")' fibe"
. ... 0 10 . nun..u. o( lr ill"min~1 n.
Proprio~I;"'e fibers
P"'r.liY"'p"lho!lic: fobc-rs
-- - - - Sympalhelicfobc-rs
oi
i Ii I

SpillllllraCI and spIna l

nucle us of l" gemi~1I1 n.
{r~in and t(>m pcr3Iu.e)
5h"'l , 111"'1 M

Anle.IOI' ethmoidal n.

- ,i-s., 1a"d mlern;ll o~S<l1 (rTlI:'dilll

.. lami ) brtm(h~ of ,Interior
,.
~\.u ill;lry Move

_ ;.m"!lcolerTll~,,1 o .
.: ~orna1lct>fxi;rl n

- - ' - - Facial (VIII n .

.... ronic hrJnch~ ""d ," -~- Chorda
- . ; opalil'ine II"nWion
~ .If alveolar hr,"lndleS

0:>"01'. middle, post{'lior)

,finn.lOmltai n.

,... branOieS (j'ust .....ior

;I [aler"l, n;u.opalatille
::o;.":'l"lOrsuperior lTIed~11)

~. (. of pterygOid canal

"i'rJrY"8t'.11 manch

Inferior d(:.'fltll '~"s, >' Ungu'ill n.

" lil'lh;./of del'l13l ~M: '--"\~.;:.
J~p
6'f 011DN
sfnr<rllJn.) """ ,.........::

RE 11.68: TRI GEM INAL NERVE (V) _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ __

-~geminaJ nerve (CN V) carries sensory infor main sensory nucleus and the rostral descendin g
n from th e face, the sinu ses, the teeth, and nucl eus of eNv. Those axons processing pain and
an terior portion of the o ral cavity. It has 3 sub temperature se nsation (protopath ic sensation)
ns: (I) op hthalmi c- se nso ry innervation, (11 ) termin ate in th e ca udal descencling (spinal) nucl eus
ary- sensory innervation, and (III) mandibu of CN V. The trigeminal nerve also carries proprio
: nsory innerva tio n. and motor innerva tion cepti ve in formation from muscle spindles in the
-eo mastica tory and tensor tympani muscles. masticatory and extraocular mu scles. The primary
:ary sensory axons from trigeminal (semilunar, sensory cell bodies found in the mesenceph(llic
-' rian) ganglion cells that process fine discrimi nucleus of CN V are the o nly primary sensory neu
e tou ch (epi criti c sensation) terminate in the rons to reside in the CNS .
161
REGIONAL NEUROSCIENCE Brain Stem and Cerebellum

Motor fibers Grea ter petrosal n. Carotid plexus Ion internal carotid a,1
Sensory fibers Deep petrosal n. Ge niculate ga ngli on
Parasympathetic fibers facial (VII) ncrve
lesser petrosal n.
Sympathetic fibers Molor 100\ of facial n.
Nerve o f pterygoid canal
Internal acoustic meatus
O tic ga nglion Nervus inlcrmedius fOI fadal n.)
Pterygopal atine ganglion nucleus of fa cial n.
Superi or salivator), nucleus
Facial muscles ,.,..".,." ':>:-.:~:
Nucleus of the solitary Iratt
OCCipitofron tali s m.
(frontal belly) - - - -, \
Orbicu laris oculi m.
Corrugator supercili; m.
Zygomaticus major m.
Zygomaticus minor m. ~\ j
Depressor
supereil,; m.
Procerus m.
Levator labi;
sup eri oris m.
Levator labii superioris
alaeque nasi m, _ I-
LevaTor anguli
orism.

Nasalis 111 .

Occipito
Depr e~sor
frontalis m .
septim. (ocapi tal belly)
D<:dpilal branch
O rbf<:ularis
of pos teri or
oris m. J
auri cu lar n.
D epressor labii
Branches to
inferiori s m. - - --C
auricu lar musd es
Mentalism. Posterior auricuhu n.
Transvers us Nerve to stapedius m.
menti m.

Ri.o rius m.

Buccinato r m.
Tympanic n.
Platysma

Submandibular g,mglion
Lin gual n.
II I Glossopharyngeal (IX) n.

Posterior belly of digastric m.

Stylohyo id m.
~~~ CIIQN
:\":.":::
Chocda tympa ni Caroti cotympa nic n.
FIGURE 11.69: FACIAL NERVE (VII ) _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

The faci al nerve (CN VI I) has motor, parasympa
thetic, and se nsory co mponents. The motor fib ers
distribute to the mu scles of facial expression; the
scalp, auricle, buccinato r, stapediu s, and stylohyoid
muscles; and the posterior belly of the digastric
muscl e. Oamage results in ipsilateral paralysis of
facial expression, in clud ing the forehead (Bell's
palsy). Activation of th e stapediu s dampens th e
ossicles in the presence of sustained lo ud noise;
damage to CN VII also results in hyperacusis.
Parasympathetic nerve fibers from the superi or
162
salivatory nucleus distribute to the pterygopalatint:
ganglion, which innervates the lacrimal glands, a"'"
to the submand ibular ganglion, whi ch jnnervat~
Ihe submandibular and sublingual salivary gland!'
Special sensory taste fibers from the anterior tv. a-
thirds o f the tongue (via the chorda tympani) a M
the soft palate (via th e greater petrosal nerve),
axons of the geniculate ganglion, co nvey informa
tion to the rostral portion o f the nucleus soli ta ri ~
in the medulla.
.lIe - Stem and Cerebellum REGIONAL NEUROSCIENCE

~ib ulocochlear (VIII) Nerve

G reater petro5a1 n.
Spiral ganglion of cochlea
Geniculate ganglion of facial n.
Cochlea r part of
\"esli bulocochlea r n. Facial cana l
ur Tool of facial n. of malleus

"'o('fV US inte rmedius

___ Ty m,,,nil c cavity

,.... ---- -,
.'!:lllihliocochiear (VIII)
,~ _ _ _ Incus

;;,!.c-- - Chorda tympani

loblongata
- - -- :~~;'~i',"~u~o':f:superior
duct
Ampulla of la teral
semicircular duct
Utricle

Ampulla of posteri or
semicircular du ct
Saccule

Superior diViSion}of vestibular pari of

Inferior division vestibulochlear n.

'Dc"~I} Cochle" nudei " ' ' '' ibu'' " ganglion

VestilxJlar pa rt oi ves tibulocochlca r n.

Inferior cerebellar

peduncle (to cerebellum)

'lGU RE 11.70: VESTIBULOCOCHlEAR NERVE (VIII) _ _ _ _ _ _ _ _ _ _ _ _ _ __

- -e vestibu locochl ear nerve (eN VII I) ari ses fro m
""'volar prim ary sensory neurons in th e vestibu lar
Sc::arpa' s) ganglion and the spiral (cochl ea r) gan
s. _ Dn. The periphera l process of th e vestibu lar gan
[) ::ion neurons innervates hair cells in the utricle and
d ""'""'"e saccule that respond to linear acceleration
; "avity) and in the ampullae of the semicircular
.....ets that respond to angular acceleration (move
- ent). The utricle, the saccule, and the semicircular
::.... cts provide neu ral signals for coordination and
~ ui libr ation of position, and moveme nts of the
head and the neck. The ce ntral pro cesses of
vestibul ar gangli on cells terminate in th e medial,
lateral, superior, o r inferior vestibul ar nuclei in
the medulla and th e pons and in the cerebellum.
Th e periph eral processes of spiral gangli on cells
innervate hair cells that lie along th e cochlear duct
in the o rgan o f Corti. They convey hea ring in forma
tio n into the do rsal and ventral cochlear nuclei via
the central axonal processes. A lesion to CN VIII
resu lts in ipsi latera l deafness and vertigo and loss
of equi librium.
163
REGIONAL NEUROSCIENCE Brain Stem and Cer e b ~ 1

-- S~n sory rohe l'5 c.,Jl;';ula~f\ g;lJlglion 01lacial n.

- - MOlu, fibe,s

....... P3r.uympa lhelic

-~.

m~rs

>.
N"rvp. o f pte rygoid (JnJ~ ~
,j
Vi
~
l Intenor 5 all v~ to ry
nu cklu 5
Ptern;opdlalm e ~~ n ~II ()n ~

MJncllbul.v Solj13l~ Ir./l~' nucJ.. u~

O liL ,lnd
,I)
AUIlCulolempo'JI
I '~.o"ci nerv<:'
Nucleus ambi guu s

~'Glossophal)'ngeal (I X) nerve
, Com m unication 10 aur iCular

'\C"..om.....m'cation to faCIal "

S u~ri(lr cerviCdlwmpathetic 8anlliion

,
r-- V~B u l (XI n.

%1 ! Ph.:u yl1 gcal b'and> of va gus n.

'"

,
~l
I
C~rDlid $Inu s bra nch of ti cmOpMr)'l1 I'1H1n
Inte rn al carotid a.

;f. ~p
Ph.:l, )'nllt'",I. lun< ~I",r. ,md ~ogual
b,.:mche. ( ) i glrmopOJ'Y"gealll.. ', : Comn1of> caro,id J , ~

FIGURE 11.71 : GLOSSOPHARYNGEAL NERVE (IX) _ _ _ _ _ _ _ _ _ __ __ __
The glo ssop haryngeal nerve (e N IX) is a mixed
nerve w ith motor, parasympatheti c, and sensory
compo nents. M o tor fibers fro m the nucleus
ambiguus supply the stylopharyngeus muscle and
may assist in innervation of pharyngeal muscles, for
sw allow in g. Preganglionic axon s fro m th e inferior
salivatory nucl eus travel with e N IX to the otic gan
glion, whose neurons innervate th e parotid gland
and the mu co us glands. Special se nsory axons
from the petrosal (i nferior) ganglion convey infor
mation from taste buds on the posterior third of the
tongue and on parI o f the soft palate. These axons
terminate in the rostral portion of th e nu cleus 501i
tarius. Axons from additional primary sensory neu
rons in the in ferior ganglion carry general sensation
164
c .~ -
from the posteri or third of the tongue and from .
pharynx, the fauces. the to nsil, the tympanic C3\
the eustachian tube, and the mastoid cells. The
central axon branches terminate in the descenru"'.
(spinal) nucleu s o f eNv. The general sensory fibe
fro m the pharynx provid e th e afferent limb of the
gag retlex. Additional primary sensory nem o ns
inn ervate the carotid body (chemorecepti on oi
C02) and the caro tid sinus (baroceptors) and
co nvey the central axons to the caudal nucleus
solitarius. Primary sensory neurons in the superiOf
ganglio n innervate a small regio n behind the ear
and convey general se nsa tio n into the desce ndir_
nucl eus of eN v.
Nucl eus so litarius = Solitary tract nucleu s.
 Stem and Cerebellum REGIONAL NEUROSCIENCE

\')gus (X) Nerve Gloosopharyngeal <IX) n.

/ ' Oo ",It,,,.,,,,, nucleu~ 01X
M eninge,,1br1lncn o f ,'a8u~n,
Auricu la r branu. of ~illlLls n. C:---c-' '::","Y, ""'" nud"us
Spi nal J r~(:1 <Inri ~pin,,1
Auditory I Eu~<Jchi~n) tub<!
nud e l' s of tn geminal n.
Levator veli p~lalln1 m.
.' I. i
Sa lringo-.pha rvngc-u\ m,
PalJ togto'lus
C'~nia l r!"llli oi accessory n,

Superio r constJldnl m.
af pharynx
Stylophar ynl)""-'S m.

Mid dle
oi
can~::"~;'~''~m:,:::::::~~Tt
pha.yn~ :
'
Inle.io . ganglion o i vagus n.

Phar~'n gea l
branch 01 vagus n. (mOlar to mu,d+'s 0 1
Infe"or ~on STri(:tor m. of pha.yn~ pharynx and .,a~t ~; sen so ry 10 lower p h,lrYJlX)
CriCO lhyroid m. \ C V'S"b ,,",h to <.:arct id sinu s hranc h oi glossoph aryn gea l n.

L""" ",,,,'I pl exus

Su pe-r;or l.Jryn gea l n.
RighI recu.ren, laryoge.ll n. Int"ffi.ll branch (Hmsory)
u(\em.:d branch (mota. 10 c"llh~' foid m.)
- - Mololfo~

- Afferenl fibers

..... Pill,lS~'mp;'lthe lk libers

Hep <ll ic L'dnch of amerior

ve ga l trunk lin I ~ss.c r omentum)

Ce liac br~n<,;h~ s (from an ter ior

Jnd posterior VJ g~1 trunk~
10 celiac plexus l CMdidc pleXUi

Esophageal ple~\Is

~. C"". b",""h,",' "'~'",""..' l .un k

Vag,,1bo--aoches lparasymp<lt hetk mmnr.
!.e<;r"lamOlo r a r)d ,;ensory tII~>e.s) dccomj)ilny
wperior m~nl..,.ir ,,,Icry .md it:; brand>eS to
smaIl1nte5 line, cec um. appendix .md colon,
"ftl!fl a~ (1)1 as left co li c i; plenlCj ftexure

Sma llinies linl'l

:CU RE 11.72: VAGUS NERVE (X)

_ _ _ _ __ __ _ _ _ _ _ _ _ _ _ _ _ __

\ agus nerve (eN X) is a mixed nerve w ith
'or, pa rasympathetic, and sensory components.
'\, axons from the nucleus ambiguus in the
\C ulla supp ly muscles of the soft palate, the phar
and th e larynx, w hi ch control speakin g and
allo win g. A les ion results in hoarse ness, dys
2gJa, and decreased gag refl ex (efferent limb).
;-ganglionic para sympathetic axons from neurons
me dorsal (motor) nucleus o f CN X in the
-!'"Jf'dulla distribute to intramural ganglia associated
-n thoracic and abdominal viscera to supply
:onomic innervation to the hea rt, the lung. and
--e GI tract to the descending colon. Special
sensory axons fro m the nodose (inferior) ganglion,
w hich ca rry information from taste buds in the pos
terior pharynx (fou nd mainly in children), send cen
tral bran ches to terminate in th e rostral nucleus
solitarius. Primary senso ry axons from the inferior
gangli on, whi ch co nvey general sensat io n from the
larynx, the pharynx, and the th oracic and abdomi
nal viscera, terminate mainly in the ca udal nucleus
soli tarius. Primary sensory axons from the superio r
(jugular) ganglion, w hich convey general sensatio n
from the ex ternal auditory meatus, terminate in the
descending (spina l) nucleus of CN V.

165
REGIONAL NEUROSCIENCE Brain Stem and Cerebellu ~

Accessory (XI) Nerve Crania! roOl oi accessory n.

(jo ins v~gu s n. to supply
muscles of larynx - excepl
Nucleus cricothyroid - via recurrent Jugular foramen
laryngeal n.)
,/~ Superior ganglion of vagus n.

Vagus (X) n. L - - Accessory (XI) nerve

: . . . - - - Interna l branch of accesrory n.
accessory
f-- - - 1 nferior ganglion of vagus n.

Foramen r - lstspinaln.(C1)

~ _ _ _ _ 2nd spinal n. (e2)

L-_ _ _ _ External branch of

accessory n. (to

sternocleidomastoid
and trapezius mm.l

: - - - - - - SternoCleidomaS10id m.

spinal n. {O )

~ _ __ _ 41h spinal n. (4)

Trapezius m.

- Moto r fibers
- Proprioceptive fibers 4' ~:*;-
<f ,,@.\
~ ...; -

FIGURE 11.73: ACCESSORY N ERVE (XI ) _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ __

The accessory nerve (eN XI) is a mo tor nerve w ith
cranial and spinal po rtions. The cranial portio n
ari ses from LMN s at the caud al end of th e nucleus
ambiguus; the axons travel through an internal
bra nch that distribu tes w ith the pharyngeal and
laryn geal bran ches of the vagus nerve (CN X) and
th e nerves to th e soft palate, These axons are often
considered to be part of CN X. The spinal portion
arises from LMNs in the lateral part of the upper
four or five se gments of the cervical spi nal cord.
166
The axons then emerge as rootlets from the lai e-~
margin of the spin al cord, ascend behind the de-
ticulate li gaments, and coalesce as a singl e ner. e
Thi s nerve then ascends through the foramen rna:.
num and joins the vagus nerve to exit thro ugh cr
jugul ar foramen. The spin al accessory LMNs su
the sternocleid omastoid mu scle and the upp er
two-thirds of the trapezius muscle. D amage to .
division results in weakness of head rotation ar .::
shoul der elevation.
 Stem and Cerebellum REGIONAL NEUROSCIENCE

Styloglossus m. Hypoglmsal
Meningeal
branch nucleus

1tfinsic muscu lalure of tongue

_ _ _ A' -_ _ _,
Hypoglossal (XII) nerve
(in hypoglossal canal)

Occipita l condyle

Infef iOf ganglion of vagus n.

Ventral rami of C l. C2 and C)

forming cef'llical plexus

_ _ _ Superior ceNical sympathetic

trunk ganglion

Superior root ldescendens

hypoglossi) of ans.a cervical is

~Jossus rn. >.c;.-;~:";~:'2 fJ'--'L----=~_ Inlernal carotid a.

Ceniohyoid m.

Inferior root (descendens

Hyoglossus m.
hypoglossi) of ansa cervica l is
Thyrohyoid m.

~_ _ _ Ansa cervical is (ansa hypogloss i)

Omohyoid m.
(supefior hell y) - - - - Inlernal jugular v.
Stern othyroid m . _~- '
_ _ _ Omohyoid m. (inferior belly)
Sternohyoid m.

Motor fibers

- - Semory fi bers

U RE 11.74: HYPOGLOSSAL NERVE (XII ) _ _ __ _ __ __ _ _ __ _ __ _ _

hypoglossa l nerve (e N XII) is a moto r nerve.
'5 in the hypoglossa l nucleus of the caudal
t"dulla exit fro m the ventral surface of the medulla
iIle preoliva ry sulcus {between the medullary
'("amid and the inferior olive) to innervate the
",nsic hyoglossus, styloglossus, chondroglossus,
-.d genioglossus tongue mu scl es and the intrinsic
superio r and inferior lo ngitudinal, transverse, and
vertical lingual tongue muscles. Damage to this
nerve leads to weakness of the ipsilateral to ngue
muscles; the protruded tongue d eviates toward
the weak side because of th e un oppo sed acti on of
the innervated contralateral geni oglossus muscle.

167
REGIONAL NEUROSCIENCE Brain Stem and Cerebellu

lIary V1)
wpcriOi ,,1I'eoliu
Trigeminal nerve (V)
Glossopharyngeal nerve (I X)
====
W/",0;
nerve ~
Vagus nerve (Xl '%:/'//
h
ganglion ,J(la greater Fa cial ne rve (V II )
and feS5f:f pall/ine
nerves
Facial (VII) (intermediate nerve)

plus gener.]1 senS<lUon via lingu,)! bt:mches

Trigemin.;a[ ( V) (mui1lary Vll
Via pharyngeal /]Janeh of
pterygopala tine sangJion
Facial (VII ) (i ntermediate nerve)
Via Slealer pelfoS<J1 ~rvl! and
pterygopalatine gang/ion

Vagus (X) Glossopharyngeal (IX)

Trigeminal (V)
Vi,) internal branch or Via pharyn t:4~Ai pJexus l:1 :"_"''l (m~x illary V2)

- Via p terygo
superior larynge,li nI;fVf' Via tonsillaf brandle5

Ta ste plusgeneral palaline gatlglion

sensatjon via iJnd greater and

lillgual bran, lesser palatine
,

Via inferior
alveolar nef~
1}0::

Facial (VII ) (i ntermediate nerve)

- - TaSI~ viol cilOldd Iympani and /j~ua' nerve ~~p . !,~

FIGURE 11.75: AFFERENT INNERVATION OF THE MOUTH AND THE PHARYNX _ _ _ _ __

Primary sensory axons of the maxillary and
mandibu lar d ivisions of eN v (trigemina l ganglion)
provide general sensation to the mouth, the tee th,
the gums, the sinuses, and the anterior two-thirds of
the tongue. General sensation to the posterio r third
of the tongue, the pha rynx, and the larynx is pro
vided by primary sensory axons of eN IX (inferior
or petrosa l ganglion) and, to a lesser extent, by
eN x (inferior or nodose ganglion) for the phan ....
and the larynx. Chemosensatio n from taste buds
conveyed by eN VII (genicu late ganglion ) for Ih.
anterior two-thirds of the tongue, eN IX (petrosa
ganglion) for the posterior third of the tongue, a""
eN x (nodose ganglion) for the posterior pharvn\.

168
 Stem and Cerebellum REGIONAL NEUROSCIENCE

Do,P.IT"""'skoe" groups

~ffiia n reticulilr 10m"",," -----"

~>R,,'h. nudei
~:el al l e ticula r formation -
and nudei

"'edlal reticular ----,,L'

formation

' ,jK:~r--~ Major noradrenergic and

~ adrenergic cell groups

Raphe ""'''.'. - -

Raphe nudei
1. Obscurus. pa1!idus
2. Magnus
3. Pontis
4. DOl'5alis. centralis superior

E 11.76: RETICULAR FORMATION: GENERAL PATTERN OF NUCLEI IN THE BRAIN STEM

_
'F..JCular formation (RF), the neuro nal core of
_:.Il stem, consists of neurons w ith character
~e ndritic morphology. The RF extends from
_<:fa l sp inal co rd through the hypothalamus
"'ie se ptal region. RF neurons are large cells
Z\Onal arboriza tions that terminate at a dis
TOm their ce ll bodies and dendritic tree; they
~ intern eurons. The major nuclei are found in
zone (sensory functions), a medial zone
'unctions), and a co lumn of raphe nuclei
(serotonergic neurons). The sero to nergic neuro ns
exert modulatory influences on their targets. The
catechoJaminergic neurons (locus coeruleus,
tegmental noradrenergic and adrenergic gro ups) in
several regions of the RF have w idespread projec
tions and exert mainly modulatory influen ces on
their targets. The dopaminergic neurons of the mid
brain are included in this illu strati o n, altho ugh some
experts question whether they are RF neurons.
169
REG IONAL NEUROSCIENCE Brain Stem and CerebeIL.:...

\
, ;'

A. Thalamus and

-
hYf)Othalamus

.J

l~
"'"{ Thala nlUS:

'-#\, .' 7 . "

.J.\ ;_....,.,;(
In rralamin':l1 nuclei
Reticular nucleusuf

\
~ .
t. .
. .

Gu~f'
'/' .
I -J
)
Ii thalamus
MiOli", oct",

Late ral hypothalamic arE'~

'v j ~. -r"( . .~ . ' throogh septalnuclei

.'\... J./
_ -- - - Pcriaqueduclal gray mailer
l atera l ~ F of the mid brain _ _ __ , ~
~
7' R.Jphe n udei (dorsal, cenllal sureriOl l
B. Mid brain Sub~lanlia nigra - - - - ' - - _ - Ventral tegmental nucleus
Interpp.l'Iunwlal nucleus --\"'.--- ---41,/

,/'''' .'\.
_____'_ ~ /l P'lrahrachial nucleI.JS
Loc,us COe ruleus
AS ,< f. 4;--,c-c, -- - Pa.aporui nc RF (Ialeral gal.e Center)
......( _. ..

C. Pons Raphe n uclei (pontis) " . ii Po ntine RF (pon ti~, c3u d.a lis, ovalis)
L3Ieral R: F ~

A-/"
O. Medulla
,..",'" 0 '
Medullary RF (gigimlocellularJ ." '.b
Z ,('
,J"'I.fo.'~ A2

Raphe nudei (obscul us.
pallidus, milgnus)
"
~ R~~,"0'Y"""';
~~ Rostral Vl medulla (RVLM)
". ~ A:~Al
~ ~
~ : LUe lal felKtRill nucleus

,-
--'
E. Spinal cord-medullary junc tion . ") lamina 7 - caudill RF
J. Perkins
MS, MFA

'w.... "

FIGURE 11,77. RETI CULAR FORMATION : NUCLEI AND AREAS IN THE BRAIN STEM AND
DI ENC EPHALON ___________________________________________

Many of the named nudei of the RF are present in medulla. Catecho lam inergic neurons are found ,
the medulla, the pons, and the midbrain. Impo rtant the locus coeruleus and the tegmental groups.
medial RF groups include th e medull ary (giganto Raphe nuclei are fo und in th e midlin e and in thE
ce llular) and po ntine (caudal and rostral) RF, whi ch wi ngs of cells that extend laterally. Th e core 01u
are in volved in reticulospin al regul atio n of spinal RF co ntinues rostrall y fro m th e lateral regions 0-"
cord LMNs; and the parapo ntine RF (PPRF), the the brain stem into the lateral hypo thalamic ar
hori zontal (lateral) gaz e center. l ateral RF areas and extends through the hypothalamus to the
and nuclei (such as the lateral reticular nucleus) are sep tal nuclei. The intralaminar, midline, and rel,
involved in polymodal sensory functi o ns. RF respi lar th alami c nuclei are classified as part of th e R;
ratory and ca rdiovascular neurons are found in the
170
 -------~ ~---------
olem and Cerebellum
Oladory input
E 11.78: MAJO R AFFERENT CONNECTIONS TO THE RETICULAR FORMATION _
~ e sensory information {particularly nocicep
-om spinal cord somatosensory sources and
all brain stem sensory modalities is sent to
..Li?ral regions of th e RF. Olfactory input arrives
5Jl olfactory tract projections int o forebrain
~ . Many li mbic and hypothalamic structures
Corte~ ...:::::;""+-
A,ou ul
Tha lamus
Inlral.amlr'l.l( nucleus
SI.l atu m _ _ --'
I
ARAS
Iteticul~r _ _ _:j;>
formallon
Spinal cord
mOlor reSioo.
"ia renruto;p,f\al
u at:(s
~. Fron1 i.1rteral ,tt.w!;" n ~,
Nudeus '''ticui.1rns t~ e.1~ po<1 li5..
p." ,ln.edi)M relicui.1rr tM;I""s.
locul o:letvle~ ,apl>e n"";";
.: !) (-rom locus coerule...s, raplr.. nudel.
wIW~llegme<ll.'I.'u
RE 11 _79 : MAJOR EFFERENT CONNECTIONS OF THE RETICULAR FORMATION
_ _ __
ascending reticu lar ac tivatin g system (ARAS) of
is responsible for consciousness and
...sal. It projec ts through nonspecific nuclei of
.,. ~a lamu s to the cortex; lesions in this area lead
roma. The RF sends extensive axonal projec
r-! to sensory, moto r, and autonomic regions of
soinal cord that modu late nociceptive input,
.:.an glionic autonomic outflow, and LMN ou t
respectively. The RF sends extensive connee
-rs to brain stem nuclei (such as the nucleus
REGIONAL NEUROSCIENCE
A,.in l1(m
Trijtemin." nvcicw, vnllb..JIi, nuclt'IJ~
(<xhI~~, nucie,,!. 0Ihe< "udljO<~ nUdel. I. Pe,~i ns
nuclo!'U! 01 the sol~arl U-iCI, ,",w,
.,~
~1Qr coikullll idee!) Livers)
_ __
provide input to the RF, particu larly for visceral and
autonomic regulatory fun ctio ns. The cerebral cor
tex/ the glob us paliidus, and the cerebe llum also
provide input to th e RF medial zones invo lved in
motor regulation.
H)'pOlhalamU5, ~ptu m limbic fortb~in ~rU'
1 Hrp pocamp.llrormal'on,
a ~wg'Wo , d nu cieu;. N'ci eu.
lcr.~ml _s. oliac lor, lubercle.
d n~~ l a'~. pre irool~ l. in.u l ~r C.()tle.'
_ _ _ _--{(j',
. _ _ _ Br~in , tern ~ ut(}f'L<>mic cenlers .};;
'-. / Nud eu. of the , oljt~ry tract
-----;(i ' Mulliple brain,tern regions
for l elpoint modulalion
Spinal cord ..,n,ory r~l\ion,
outon<>mk re&io~s Oors.1 oorn !or
T1Ll ;"t",rnedi<JI~ let~1 ,,,II U>iurnn, noc,rer<,ve
S2$a int.,,-m~~l;'''e ~r;,l' modlli<>,;""
G
t Vi" ~i'r\ ioreb[. i~ bu ndl~. (5) 11X1l.tding ventrol.,,,.-.I and vcntromcd,.1
~llongitL!(\",.lll''''''OJlu!, - !egnrenl1Jm Df caudal br. ,n ,tem
h~nulop-edl.t~cul tr.ct. :~'; Intri-fet,cd", connoection ,
rnamm,II01~"""n t.lI !racl J. P.. rkin
(4) From odf~n...--g;c. n()f ><I0-...,e<~' 'ep/))e r\t~1 ,rtd low. coerulem), ""MtA
~ ,1I!f)!
- serot~ (cJoph~t nu cle i
tractus solitarius) and to autonomic regulatory ce n
ters and nuclei for modulation of visceral func ti ons.
Efferent RF projections to the hypothalamu s, the
septal nuclei, and the li mbic forebrai n areas help to
modulate viscera l autonomic func tions, neuro
endocrine outflow, and emotional responsiveness
and behavior. Efferent RF projections 10 the cere
bellum and the basal ganglia participate in modulat
ing UMN control of LM Ns.
171
REG IONAL NEUROSCIENCE Brain Stem and Cerebelhr

(cholinerglcs)

area

Interleukins; To pin eJ I
other ~(}(}d-borne
(melato nin)
substances

lei

~~?
Areas assoCiated

M
'."

). p",;"
wi th arou sal
Areas assoCiated Wilh
QlmN the induction 01 ,Ieep
Spinal co rd !::li Sympat hetic ch ain gangli a

FI G URE 11.80: SLEEP- W AKEFULNESS CONTROL _ _ _ _ __ __ _ _ _ _ _ _ __

Sleep is a normal physiological state involving a
cyclic temporary loss of consciousness; it is readily
reversed by appropriate sensory stimuli. Sleep is an
active process initiated by activity in several chemi
cally specific collections of neurons of the brain:
locus coeruleus of the pons (noradrenergic); raphe
nuclei of the medulla and pons (serotonergic);
nucleus solitarius of the medulla; cholinergic neu
rons of the brain stem tegmentum; lateral RG par
ticularly in the pons; anterior, posterior, and
preoptic areas of the hypothalamus; and reticular
nucleus of the thalamus. Many of these regions
actively inhibit the lateral (sensory) portion of the
RF, which is responsible for maintaining a waking
state and consciousness. Circulating substances
172
such as interleukin (Il)-1 ~ can act on key sites i'"
the hypothalamus and the brain stem to influll(
components of sleep. Illness behavior involves
enhanced slow-wave sleep induced by IL-1 ~ an.::
other intlammatory mediators. Non-REM, or slcr.
wave, sleep, initiated by hypothalamic neurons
and other regions, is accompanied by decreasec.
activity of the locus coeruleus and the cholinerg:::
tegmental neurons. During REM sleep, activity if;
the noradrenergic locus coeruleus neurons and _.
serotonergic raphe neurons diminishes, whic h ~
vents the cerebral cortex from attending to exte""
stimuli. Dreams are likely the result of the corte\.
attending to internal stimuli from stored mem ore.
 Stem and Cerebellum REGIONAL NEUROSCIENCE

LinguLl--, "Unlolded" s<hemalic o i (crebel1um

demomtr.lling body map Aw.b

~u:!ra l hC!misph('fc
I Re gions I I Lobes I

P()~ l e rior
lobe

Flocn,lonodula r
lobe

Unfolded schematic vf t:erebel lum d emons traling regi ou~ and lobes

" RE 11.81: CEREBELLAR ORGAN IZATION: LOBES AND REGIONS _ _ _ _ _ _ _ __

ce-rebellum is organi zed ana tomically into globose and embo li form, and dentate, respectively)
-or lobes: anterior, posterior, and flocculo that help to regulate the ac tivity o f the re ticulo-
ar. Di stin ct sy ndromes are associated with spinal and vestibulospinal tracts, the rubrospinal
.....age to each lobe. The functional organi za tion tract, and the co rti cos pinal tract, respectively. At
~e cerebellum follows a vertical arrangement of
least 3 rep resentatio ns of the body are mapped
~p heres: (1) ve rmi s (m idline), (2) paravermis,
onto th e cerebellar cortex. The cerebell ar cortex
31 lateral hemisphere. Each fun ctiona l regio n
has multiple, ord erly, small info ldings, o r co nvolu
~ ciated w ith specific deep nuclei (fastigial. tions, called folia.

173
REGIONAL NEUROSCIENCE Brain Stem and Cerebe....

Posterior view id Habenu lal trigone

Pulv;nCl! .-- Later.)} Geniculate
" !neal gland _ _ Medial bodies
Sup Cfior collicul us
IIi
Trochlear (IV) nerve
Superior medullary Velum---j
Sup er;

CerebeliM "ddl

peduncl e~ { "'II e~.

Inferiot

Lateral recess ~
Dentate nucleu s
Tlmia o f 4th ventricif.

l. - lingul.. TUB - luber

( l -" (cnrrallo bule PYR - pyramid

CUl = ( ulmen UV" uvula

0 '" ded ive TON'" tonsil

Cerebellar cortex
FOl - JuUu m

fasciculus Woldlis
J~~
Mttdian sagiltal section
Bodv of forn ix ~ H.J\)e{lul.}r co mmissure ~r,~
Choroid pl",xu) of 3rd ventricle ~ _____ { Pineal gland
,~. ; . -
Inter ventricular fo ram en . _ !iplenium of corp us callosum
( OrMOnrO I ~~
Thabm u~ ~
!=erebral aqueduct (of Sylviu s)
A nt. co mmissure---...,:
". LingulCi
Post. commissure - - ' L~ Cen[rall obule
Ma mmillOlry body~ '*ihor--- Vermis
i,
O<:vlommor (!Ill
Stlpe,ior (ollic ulus
,11'- Sup. medullary velum
.. medullary velum
Mediallongitutlinal fasciculu ~ -
Choroid plexu~ of
41h veolride
4th venlrid e
,II

Median apertu re {o r MagendieJ - "\- 7

,C
I'ti"'""'
p""m'd }
Uvu la Vel mls
'"'-- - - Nodule
Central canal of spinal cord

FIGURE 11,82: CEREBELLAR ANATOMY: LOBULES _ _ _ _ _ _ _ _ _ _ _ _ _ __

The 10 lobu les of th e ce rebellar cortex are seen in
both midlin e and cross-sectional views. Inputs to
the 3 layers of the cerebell ar co rtex arrive mainly as
mossy fibers; the inferi or oliva ry nucleus sends
climbin g fibers to end on Purkinje cell dendrites,
and the locus coeruleus sends diffuse varicose
inputs to many regio ns o r I'h e cerebellar cortex. The
deep nuclei provide the "coarse adjustment" upon
w hich is superimposed the " fine adjustment" from
174
the cerebell ar cortex. The cerebellar cortex se--C.
its output via inhibitory Purkinje cell projecti ons
(using CABAl to deep nuclei, which in turn p" .~
to UMNs. Affe rents and efferents pass throug.'"
cerebellar peduncles, w hich con nect the cere!Je
lurn w ith the brain stem and the d iencephal o'L
table lists the majo r afferent and efferent pro!C[
tions through the 3 cerebellar peduncles.
 - tern and Cerebellum REG IONAL NEUROSCIENCE

Pedunde
Spinocerebellar T"~'"",I~
Inferior 00=1 Fastigiobulbar. ar>d r~ncUiaI1
(restiform body) Rostral Undnate fasdcu lus nucle;
)
Cuneocerebe!lar
OIive<CTebellar
ReliculoceceUellat Direct cercbellovestibular
Trigeminocercbcllar (10 lVNj
Raphe<erebellar
Vestibulospinal
lux tarestiform body
(primary, secorn:lary)

Middle
Pontocerebellar
(brachium pontis)
Superior Ventral Spinocerebellar Den tatoth alamic
(Brachium Trigeminocerebellar Den tato rubrJI
l evel of section conjunct ivumJ Te<:loCerebeliar Dentatoreticular ;
Superior colliculus
Interior col liculus
lnlerpositusruhral connectionS II
(gtobose, embolifOrml ~
Coeruleo-cerebellar

nucleus

Fourth
ventricl e

CN VI

--::=-~-:---'%~---------t-
-
Medi,'II
lo ngi tudinal
f,)sciculus

Te ctospinJ t trac I

"""- - - Nucleus eN VII

0'
'3(ial

"
Ponline nuclei

~L R E 11.83: CEREBElLAR ANATOMY: DEEP NUCLEI AND CEREBElLAR PEDUNCLES _ _ __

deep cerebellar nuclei are found at lhe roof of
~o urt h ventricle in a cross-sectiona l view of the
r:;, at th e leve l of cranial motor nucl ei for eNs VI
~ V II. The fastigial nucleus receives input from
, erm is and se nds projections to reti cular and
:ibular nuclei, the cell s of origin for the reticulo
~.al and vestibulospinal tracts. Some vermal and
culonodular Purkinje cells project directly to the
? al vestibular nuclei, w hich some authors con
sider a fifth deep cerebellar nucleus. The globose
and emboliform nuclei receive input from the par
avermi s and project to the red nucleu s (rubrospinal
tract). The dentate nucleus receives input from the
lateral hemispheres and projects to the ventrolat
eral and ventral anterior nuclei of th e thal amus,
which then project to the cells of origin for the co r
ticospinal and corticobu lbar tracts. The 3 cerebellar
peduncles are seen in this cross section.
175
REGIONAL NEUROSCIENCE D ieneeph...

Thalamocortica l

radIation.

Central sulcus

Th almic n udei
CM
Centromedian
LD late ral dorsal

lP
l ateral posterior
MD Medial dors.lJ

VA
VentfaJ an/erior
VI
Ventral inlermedial
VL Ventral laleral
VPl Ventral po~'efola'eral
VPM Ventral postero medial r 1ntetnal medullary lamina
r Intraraminar nuclei
An te ri or nudei O ther medi ill nuclei
r M idline (median) nuclei

~fp
CII!!N
Pulvinar :',';.':::

/ Medial geniculilte body

From globus pallidus

and subslanliil nigra

Lat~al genkvlate body

Irael
Someslne lic from body

Someslhetic from head (lrigeminal nel lie)

(spinothalamic Ifac! and medial lemniscuS)

FIGURE 11.84: THALAMIC ANATOMY AND INTERCONNECTIONS WITH THE

CEREBRALCORTEX _________________________________________

The thalamus conveys extensive sensory, mo tor,
and auto no mi c information from the brain stem
and the spinal cord to the cortex. Th alami c nu clei
are reci procally interco nn ected wit h regions of the
cortex. Specifi c thalamic nu clei project to circum
scri bed regio ns o f th e co rtex; these nu clei in clude
( 1) sensory proj ecti o n nuclei (VPl- somatosensory;
V PM- trigeminal; lateral geniculate body- visual;
medial geniculate body-a ud itory>. (2) mo tor
related nuclei (VL and VI---<:erebellum; VA and VL
basal ganglia), (3) autonomic and Jimbic-related
176
nuclei (anterior and LO- cingulate cortex; MD
fro ntal and cin gul ate cortices), and (4) nuclei
related to association areas and l P- parietal c o'- ~
N onspecific th alam ic nuclei (j ntralamin ar nu d .
[CM , parafascieularl and medial VA) send d i ffu ~
con nectio ns to w idespread regio ns of the cere
cortex and to other thalam ic nuclei. The reticui.r
nucleus of the thalamus helps to regulate the
excitabili ty o f thalamic projection nuclei. Some :
amic lesio ns can lead to excruciatin g neuropalt
pain, referred to as thalam ic syndrome.
 on REGIONAL NEUROSCIENCE

Fornix

,,~potha la mic sulcus

.;.ruerior commissure

Paravenlricular

.1
Ve ntromedial
ArcUolte -----l~
-"""'=
(in fundibular)
Mil mmillary ------r , ',
,.
,
tracl

In fund ibulum (pituitary """., ~

Dorsa l longitud ina l
Hypophysis (pituitary gland)
fasci c ulu s a nd Olher
descending pathways

Lamina terminalis - ' Hypothalamic

sukus
~-a,'cntricular hypothalamic '", ', "' - - ,
Supraoptic hypo"""n" "",01,",, - - -:
Supraopti cohypophyseal lracl - --"--'7--:-'
Tuberohypophyseal ""' _ _
Mammillary
Hypotha lamohypophysea i " '" ~ body

Infundibulum {pituitary sta lk}

'-A""",, (infundibular) nucleus

Median eminence
.aenohypophysis Fibrous trabecula _ _ __ of tuber cin ~ reum
lmerior lobe Of Neurohypophysi s
Diluilary gland) Pars i"",,~edl " _ _ __ Infundibu l.:l ' (p 05te rior lobe
stem o i pitu i lar~' gland)
Pars diSlalis - _ __
Infundibul,u
process
Cleft

L RE 11.85: THE HYPOTHALAMUS AND THE PITUITARY GLAND _ _ _ _ _ _ _ _ __

-'F hypothalamus is the major region of the eNS
~ ve d in neuroendocrine regulation and control
. isceral functions, such as te mperature regula
food and appetite regulation, thirst and w ater
~ nce, reproduction and sexual behavior, parturj.
and control of lactation, respiratory and cardi o
.3Scular regulation, gastrointestinal regu latio n,
~ ' ess responses, and reparative states. It is sub
- .ided into rostralto-caudal zon es (preoptic,
.JI1 terior or supraoptic, tuberal, and mammillary
Of posterior) as w ell as medial-ta-Iateral zones
(periventricular, medial, lateral). These zones
contain some discrete nudei and more diffu se
"centers" or areas. The neuroendo crine portion of
the hypothalamus consists o f (1) magnoceUular
portions o f the paraventricular nucl eus (PVN) and
the supraoptic nucleu s (SON), which send axons
directly to the posterior pituitary, (2) releasing
factor and inhibitory-factor neuron s, which proj ect
axon s to the hypophyseal portal vasculature in the
median eminence, and (3) the tuberoinfundibular
system.

177
REGIONAL NEURO SCIE NC E Diencepa llill

.m:~
".' '_ ''J'';'~~Cor pus callosum
oc ___ : .: "h:IJ.....

,':Mecuan
forebrain
1

I ,al ,
MammIllary
.
Optic (II) complex
nerve
Oculomotor (III)

~hySe.a llfact ,f' ,f$

Anterior I
o f pituitary
Posteri or lobe
of pituitary 4~'~oN
..:' -,"': : ~

FI G URE 11.86: SCHEMATIC OF HYPOTHALAMI C NU CLEI _ _ _ _ _ _ _ _ _ _ __ _

Hypothalam ic nuclei and areas are associated with
visceral and neuroendocrine functions. The magno
cellular neurons supraoptic nucleus of the SON
and PVN release oxytoc in and vaso pressin into the
posterior pituitary general circulatio n. PVN parvo
cellular neuron s containin g CRH project to th e
hypophyseal portal system in the median eminence
and induce the release of AOH. Descending
axons of th e PVN project to brain ste m and spinal
cord preganglionic neurons and related nudei to
regulate outflow from the ANS. The anteri or and
178
posterior areas coordin ate parasympathetic and
sympathetic outtlow, respectively. Nudei OM arw4
VM, and the lateral hypothalamic area regu late
appetitive, drinkin & an d re prod uctive behavio r.
The preoptic area regulates cyclic n e uroe nd oc ri~
behavior and thermoregula tion. The suprachias
malic nucleus receives visual inputs from the cp'
tract and regulates circadian rhythm s. Several
hypothalamic regio ns are involved in the regula e
of sleep.
 alon REGIONAL NEUROSCIENCE

CN V

Uncus
COrliCOsp rnallract fibers
Basilar artery

.. ".--~

Level of section
(mid pons)
';.",8
~.;.\ . :;;;;":wx,

i.,,;,

\ ~;J!/I
. ~1
.~~
~\.~~. ..
\-.~"
\ ..... \~ . .
)

lateral
,. ~.

Superio r
cerebellar cerebellar
bemisphere peduncle

Medial lemniscus Pontine legmenltlm

Cerebellar vermis Fourth ven tri cle

RE 11.87: HORIZONTAL SECTIONS THROUGH THE FORE8RAIN

_ _ _ _ _ _ _ __

~ plates co mpare anatomical sections, MR descendin g corticospinal, corticorubral, and

-"c"',,;, and ( T scans. These are cut in th e tru e hor
~J plan e, nol in the older, 25 0 tilt. Th e most
~ a nt anatomical relationships in th ese sec
center arou nd the internal capsule (Ie). The
Mi oj the ca udate nucleus, medial to the anterior
of the Ie, forms the lateral margin of the
r-~I po le of the lateral ventricle. The thalamus is
" 'al to the posterior limb of the Ie. The globus
-Idus and th e putamen are lateral to the wedge
~.i.iO e d Ie. The posterior limb carries the major
corticoreticu lar fibers, and the ascendin g se nsory
fibers for the so matosensory and trigemin al sys
tems. The mos t posterior portion s also ca rry the
auditory and visual projections to th eir cortices.
The genu carries the corticobulbar fibers. Th e ante
rior limb carries cortical projections to the striatum
and the po ntine nuclei (pontocerebellar system).
"The MR images for the horizontal sectio ns are
T2-wei ghted, with high signal intensi ty, so the ven
trical s (cerebrospinal tluid) appears bright."
179
REGIONAL NEUROSCIENCE Teleneephalcr

Amygdala
C()(licospina! tract fi bf' r~

\ ~,.

\ ~,

... ~\
i ~,
h
level of section '\
""',

L"
"

(rostra l pons)
.: ~fl J
''f ~

t~-." , /)';,.

.' .~I\".
'r
~
': }
,: '
~... . '~
)i"'~".
~.
..'

,.
..
,~ . . \
'.\

I
/." -

I },
I
~
\
"
Superior
cerebellar

Foul1h
:1,

veolride
M """
lem ~

it ' . .,.'
Lateral cerebellar
! ~j. ", c. ' . ' ).
.....

,\
'~'.
\: .. '

J ....~ .:-I
J.''''
.. 'Ii Vermis of cerebellum

'~ --. ~ ~.

0_
........c~
"....m
1

Rostral Pons

180
 ph.lon REGIONAL NEUROSCIENCE

Posterior cerebral
arteries

Cerebral peduncle

Amygdala

Tail o r caudate lobe

nucleus --~ )~
\ , \' \
:.~
\\ '\
level of section
(mid brain)
/
HiPpoca mpa~~I_ _ _""::"'~_ _ _ fl
formation -

Inferior horn of

lateral ventricle --~~-~'-~:_ _---.J,.--~

Entorhinal cortex j!(~Q~

Decussation of
Cerebral aqueduct
superior cerebellar
Occipilallobe
pedunde

Cerebellar verm is

Miilbra/n

181
 l8L

5tWJ i)/\
aqolll.'l!dp:>()
Ji!IIOIqaJaJ p npanbl'
Il'Kj<lJil:> snln:l!llo:'l
JO!Jadns

(!lnWl'Il!410d)' 4 pue
U!l'Jq P !W !I!JlSOJ)
UO!I~S' JO lilA""
VO!l l' WJOJ
Il?dwC:)odd!H

sn<> pnu
,{jail\,
illejn:>!ua8 Il'JaIFl
l eJq aJ~ :) J01..Iil\Uy

sn<lpnu pa~

DN3DSOlln3N lVNOl9311

ephalon REG IONAL NEUROSCIENCE

O rbitOLrontal cortex An terior limb of internal capsule

Putame n Glo bus pa llidus (internal a nd

external segments)
Claustrum
H ead of caudate nucleus
Insular An teri or
cortex commissure

Colum n~
of forn ix

Posterior limb o{
l evel o( section inte rnal capsule
(anterior commissure
and caudallhalamus)
Th,llamus - -,

Temporal lobe ~ ~

Choroid ple xus -.:~

r ail o f cauda te

nucleus --~...;....- ---.:::::""

Atriu m of' 1 ,~t~'~"~'_.!.:..-3_--

ven lricle

Extreme
capsule

Extern al
C<1.psuie
iormenian

Pulvina r

Occipital lobe commissure

o
.JCIt'IttACRA'i ..
w ::. "

Anterior Commissure and Caudal Tha lamus

183
- REGIONAL NEUROSCIENCE

-~ ......,
Frontal lobe

, Anterior limb oi
! inter nal capsule

Head of caudate nudeus

Claustrum GenuofcOf~
caHosum
Extreme
capsule
./ \\ Genu a
in tern a.
\ '.
\\ ~\~ capsw

level of sect ion ~

(head of c<ludate Insular \;
and mid thalamus) con ex A nt ~
hori'"
Posterior limb of tau?...
internal capsule \'('n:-

Auditory radiations ~
Tail oi caudate nudeus ~
Tempo,," obe ~

~ '4'

Optic
radialion Third
venln~

Temporal pole o f
lateral ve ntricle Globus pallid;....;.

Cho roid p lexus Putamen

\... Spleni um of the
Fimbria ()f fornix
corpu s callo sum
Oeeipilat robe
Pulvinar -"C"4~
o~

Head of Ca udate a nd Midthalamus

184

ph. lon REGIONAL NEUROSCIENCE

, - A" ,,,;m limb o f internal capsule

Putamen
Rostru m of corpus callo su m
Claustn)m
Frontal pole o f laleral
Insular ven tricle

l evel of secti on
sal ga nglia and internal capsule)

latera l and ven tral

thalamic nuclei
Postelior limb o f
inlem al capsule

Pulvinar
Tail of caudate
nucleus -~_
~-,..' .

Temporal po le of
lateral ven tricle

Head of
caudate
nude;

Optic radiatiom
Chofoid plexus '--. Goo, of in Iernal capsule
Body of
Medial thalamic iarnix Splen ium o f corpus
nuclei - - -- ' ollosum

O ccipilallobe

Basal Ganglia and In ternal Capsule

185
REGIONAL NEUROSCIENCE

(ingulate cor tex

Genu of corpus ca ll ~

Frontal lobe
Body or caudate
nucleus

level of section
(dorsal caudate, splenium and genu
of corpus callosum )

,!/~ '
Insular cortex ~ ' ":i' t/;;:;l~
Parietal lobe - -

.'\;:
....,,,,.::.
~ -.
, ~" ~~
~

Frontal ~
lateral \ e"""
Occipital
lobe Septum pellu cid...

Spjenium
of corpus
callosum
..IC1HNA.C:~
C ~!N I

Do rsal Caudate, Splenium, and Genu of Corpus Callosum

186
 halon REGIONAL NEUROSCIENCE

Antr jO( cin gulate

cortex Fro nta l lobe

Centrum se miovale
Body of caud'lI
nucleus

l evel of section
{body of corpus callosum)

,, ~
-----1\~.--
Parietal lobe

,"
'h '

" ,.....,..
~

Occipita/ lobe Body of lateral venlride

Booy of corpus callosum

Body of Corpus Callosum

187
REGIONAL NEUROSCIENCE
Telencephalo<

<. 'y ;;"f~'\:.

,q,._.
"
, .L
\ . ~.

Fronlal lobe
Cingulate gyrus

)t., 4< -

.~~
?'l
.. 1
-- f ""
l evel of section
(centrum semiovale) ~ /i
I
.-;;,
t
( >~\W
<1' .
/~;
!, -12tW)),
~"'.
'<
',:'. ''.,X '

]. ~.'
.~j
" ~.
M
~
'.'
.j_ ,I ,~~t
.
"
. J' ~ ~
.", .10>
.

J U . ~~,
;

(\ "
~W~
'0; - :

Occipilal
lobe

-"'.....
__ I
<...,.~

..... .""

Centrum Semiovale

188
 ephalon REGIONAL NEUROSCIENCE

Superior Iron lal

gy rus Cingulate gyrus

Middle fro nlal

gyrus
Genu o f corpus callo sum

Inferior fronla!
g)' rU 5 Fron tal pol e of
l evel of section la te ra l ve nt ri cle
(gen u of corpu s callosu m)

Subcallosal
gyM
I............
Lateral

~
iissure ~

Tempot"a\
pole

Genu of Corp us Ca ll osum

RE 11.88: CORONAL SECTIO NS THROUGH THE FOREBRAIN _ _ _ __ _ _ _ __

- -.a,e pl ates co mpare coron al anatomical sections
\ \R images. Th ey show impo rtant relati onships
.,. Ie. the basal ganglia, and the thalamus. Basa l
"':brain structures (such as the nucleus accum
i, the substantia innominata, a nd the nucle us
..I.SC!lis), thalamic nuclei, and th e important tempo
rallobe structures (amygd aloid nuclei, hippoca m
pa l form ati on) and pathways (forn ix, stri a termi
nalis) are illustrated. The MR images for the coronal
sect io ns are T2weighted, w ith a reversal technique
(FLA IR) thal results in the ventricles (cerebrospinal
nuid) appearin g dark.
189
REGIONAL NEUROSCIENCE

Cingulum Cingulale gyrus

Para oOaCIOry gyrus Body o f corpus calloSl.-

Head or caudale
nudeus - - - , Septum pellucidL~
Nud~J 5 accumbens
Fron tal pole (II
Anleri()( limb o i
latet"al vent~
internal CJpsule

b .:"
l evel of section

(head of caudate nucleus!

nucleus accumbens)

Insuldf COd ex

l a tera l Ihsure

TempOial pole
(

). 'J t
Amygdala
--J.l : (

Oplic nerve b"

".

0
-'CHN A.CIU.I:....c
,.': : .-::

Head of Ca udate Nucleus/ Nucle us Accumbe ns

190
halon REGIONAL NEUROSCIENCE

G ngulate gyrus
Columns 01 fOrnix
Body of corpus callosum
Cingulum

Head of Ciludate nucleus Septum pellucidum

lateral ven tricle

An teri()( limb of iOlem al

capsule - - - -,

level of sedion
(anterior commissure! Globus pallidus
columns of fornix)

Claustrum -~~~"""

Insular cortex --t~"'::::::;;gl...-

Pulam'O -:::-==~;;;;;:;;;:::f---':':;
Lateral fissure

Nucleus basalis --4~~---ll------''---''(4;'

Temporal lobe - ---.__"

Amygdala

Thi(d ventricle Extreme

capsule
Supra optic re<:ess
External
capsule

Anter ior Co mmi ssure/ Co lumn s of Forn ix

REGIONAL NEUROSCIENCE lelencephal

Stria terminalis Cingulale gyrus

and terminal vein

Body of caudate
nudeus

AnTerior limb of
intetnal capsule

Level of section
(amygdala, anterior limb
Claustrum
of internal capsule)

Putamen

Insular cortex -----.!

G lob us pallidus
~ ,~
df,,","tJ
i ; (, .
'.or.
. . './ Ii
,

extern al segmen t
~ .'10 '. ~i}ff..'
Temporal cortex \_
~~/'~'
'?(~. ."
.ti%,~
.". ';$
~::; "il. .'

~",;,~
'~~:~~Z;.' ),
'N",.
'1$?}: ~
'~
Amygdala
Colu mn s o f fo m lX

G lohus p.ll1 idus Third ventricle

inlem al segment
H ypothalamus
O ptic (ract

..I"""
_ .. C.....,...,
I
......:: "

Amygdala, Anterior limb of Inte rnal Capsul e

192
phalon REGIONAL NEUROSCIENCE

Third "en!rid e
Cingulate gyr us
ROSlral th.1Iam u$
Body of corpus callosum
Hypothalamus
Cingulum
Posterior limb o i
internal CClPsulf>
Columns o f fornix
G lobus pdllidus
internal se gment Body of lateral

Putamen

level of section
(mammillary bodies)

Insular corl ex

Lateral fissu r e

( I(lustrum
Inferior horn of
:-:::::~~::::~=t
:I.

lale ral venlobe

Temporal tricle ---==~'W;~:==~r",;r:.

Hippocamp:J1 Ex te rnal
formation capsule
Inte rpedul'lcular iossa '--. ,Vlid,jlece",,,, 1 artery
Mdmmillary body
Basis pontis
CNebral peduncle

in basis pontis

.JOHN ....., , _
CII!JN I
....:,:"

Mammillary Bodies

193
REG IONAL NEUROSCIENCE

, ~~T~~"
Cingulate gyrus
Ongulum - - - - - .
Anterior thalamus
Medial dorsal thalamlJS
Mammillothalamic tract
Body 01 caudate nudeus
Venlfallalefal thalamus
Postelior limb oi
internal capsule ,, ~ \

l evel of Section
(mammi11othalamic tract/substantia nigra,
rostral hippocampus)

Insular cortex

Tempo,, ' pole of

---
''N''
~

t - : : - -- ,
j.,
.,. -(o

i
-;.';

ventricle 7"<;: .-\~

Tail of caudate nucleus _ _" ...
Tempor..1
cortex --'

L Globus pa
external sE' ..-

C lobu s pallidus
Cerebfill peduncle internal segmen

Substantia nigra "--- Basis pOntis

(ort icos pinaltraCI

fi bers in basis pon t ~ "
'-Medullary pyramids

"""'" ACR.......,
011lN
,.......,,: I

Mammillothalamic Tract/Substantia Nigra, Rostra l Hipp oca mpu s

194
 REGIONAL NEUROSCIENCE

Body of corpu s ca llosum

Cingulale (Oriel(

Interventricular Third ventricle

fo ramen of Munro

Cingulum

Suia ter mi na lis

Body of ca udate
nudeus

level of section
(midthalamus)

Cenlromedian thalamus

Ta,1 o f the caudale nucleus

Medial
geniculate
nudeus

Corle:< of Superio r
cefebellum cerebellar peduncle

Ento rhina l Pons (floor o f

cortex - ---' fourth ven tricle)

L_," ,,,,,, cerebellar

peduncle

Medulla

Midtha lamu s

195
REGIONAL NEUROSCIENCE Teleneeph. . .

::\~ I { ~
Cingulate gyrus
Body of corpus callosum

Body of forni)!.

Pulvinar
'>

level of section
(geniculate nuclei)

Medial geniculate nucleus

Lateral geniculate nucleus

Tail of caudate nucleus

Inferior role o f lalera l ventricle .'~.:.

.
Ah,....
~r~
;-':-..::.~;;
04

-----4

comm
Cerebellar
Peri aQueduc.: _
cortex
gray matter
Middle cere bellar
peduncle I

D ecussati on o f superior
cerebellar peduncle M edulla

,JOHN A.CRAC-or.:
0 UllN I
,.,......."

Genicu late Nu cle i

196
 REGIONAL NEUROSCIENCE

Cingulale gyrus Body 0/ corpus callosum

Cingulum
Crus of fornix
Superior colliculus

Pulvinar
Third ventrid e
Sody 0 1caudate
nucleus
Body of lateral
ven tricle

Level of section
c:aMial pulvinar and superior colliculus)

Tail of caudate nucleus

Fimbria o f hippocampal

Inferior
forma tion;;~=====~~~~~;;~~"-.
pole o f
laleral ventride

Hippocampal formation
--j":---~:"'----II"-,~~

lateral
cerebellar
hemi5f)hefe caJliculus

EnlOrh;nal cortex _f--'.r

, .......l.D
\...C,,,.i>ell,, vermis
Middle celebellar
peduncle
Fourth venTricle
Super ~ cerebellar
peduncle

Caudal Pulvinar and Superior Colliculus

197
REGIONAL NEUROSCIENCE Telen eeph

Splenium of corpus
(ingulate gyrus callosum

Pineal

Trigoneof lalera
venlricle
Crus o f fornix
ChorOid p'
Pukinar
Inferior p.
ofl a te '~
ven 'nc ~

level of section
(splenium of corpus callosum )

Tail of caudate
nuculeus _____

Optic radiations _____

Hippocam pal & . ---.,_~ ~

formation . ' .. I...... ' 4
Tempo ral ii$:.

lobe \"

V.
"-.J

L.u eral

cerebellar

hemisphere

Denlat e nucleus

Cerebellar vermis

Fourth ve ntride ....... A.CI!A.IC.-.c

' OlIN
,~,w"
I

Sple nium of Co rpus Callosum

198
"'.-c..cl aIon
,
, , ,,
,"
,
,
RE 11 .89 : LAYERS OF THE CEREBRAL CORTEX _ _ _ _ _ _ _ _ _ _ _ _ _ __
ns of the cerebral co rtex with specific func
'-d ro les, such as the so matosenso ry cortex and
~~o tor cortex, demonstrate histological charac
- cs that reflect their func tio n. TIle sensory
;;::::\ has large granule cell layers (granular cortex)
"pceiving extensive input, w hereas the motor
re, has sparse granule cell layers and extensive
REG IONAL NEUROSCIENCE
, ,
","
.. ,
pyramidal ce ll layers, refl ectin g exte nsive output.
Specific and nonspecific afferents terminate differ
entially in these structu rally uniqu e regions of the
cortex. Monoamine inpu ts terminate more diffusely
than do the specific inputs, reflecting the role of
monoamines as modulators and enhancers of the
activity of o ther neuro nal systems.
199
REGIONAL NEUROSCIENCE

I~
Key for Abbreviations
3 Horizon tal cell
" b Cell of Marlinotti
c Chandelier cell
d Aspiny granu le cell
e Spiny granule cell
f Stellate (granu le) eel
g Small pyramida l ce ll
o f layers II. III
h Small pyramidal
association cell
Small pyra midal
asso ciation <J nd prOjK'l
ce lls of layer V
La rge pyramida l
projection cell (Bel z (;

, ~
v ~-v~-

Cortical interneurons Cortical association Effe re nt

neuro ns neuron

~~p
cen bodies and
Black {
dendrites
Bro wn {
axo ns of in te rneuro ns
and association neurons
Rod { axons of efferen t
neurons
OIlQN
IN".' : ~

FIGURE 11.90: CORTICAL NEURONAL CEll TYPES _ _ _ _ _ _ _ _ _ _ _ _ _ __

The cerebral cor tex has many anato mically unique dendritic arborizatio ns that run perpendicular ::

cell types, with characteristic cell bod ies, dendritic the cortical surface and arbo rize in upper layes.

arbo rizati on s, and axo nal distribution. G ra nul e ce lls The axons of pyramidal cells, w hich function as

are local ci rcuit neuro ns w ith small cell bodies, projection neuro ns (e.g., corticospinal tract ne.:

lo calized dendriti c trees, and axons that distri bute ro ns), leave the cortex an d may extend for up :

locally. Granu le ce ll s function as recei ving neuro ns meter before synapsing on target neurons. Theso

for thalamic and other input, and they mod ulate unique anatom ical characteristics give rise to b

the excitability of oth er cortica l neuron s. Pyramidal co ncep t that neuronal structure exp lains neu r

cell s po ssess more vari ed cell bodies, with large fu nction.

basolateral dendritic branching patterns and apical

200
 REGIONAL NEUROSCIENCE

Vertical co lumns (O.5 -1.Q mm wide)

Small .

' - - - Corticocorlical olfferent --- 1 ----------

rticaf projection (efferent). _ _ _ 1______- - - -
' _ _ _<C:~o: ..
to subcortical struc tures

' - - - - - - - Thalam ocortical afferent ----1----------

l. Perkins Corticocortical eflerenlS
....._ _ _ _ _ _ _ _ (."m' 10 adjacent or nearby _ _.L___________
"ti, .... r...
c BN. vertical columns)

E 11.91: V ERTI CAL COLUMNS: FUNCTIONAL UN ITS OF THE CEREBRAL CORTEX _ _ __

~n tal studies of sensory regio ns of the cere
cent::, provided anatomical and physiolo gical
.a:'lCe that discrete info rmation from a specifi c
_ ' r o r conveying specific fun ction al characteris
. orocessed in a cylindrical vertical zo ne o f
in the co rtex that sp an s aU6 layers of the
",.-.;:s
e'\. The se vertical units va ry (rom O. S to
""1 in diamete r. The dia meter corresponds to
~ horizo ntal e'pall se of a large r pyram ida l
~:. unit. Bo th thala m ic and cortical afferen ts
arborize in the verti cal column and synapse on
both stell ate (granule) cells and pyramid al neuro n
dendrites. Information from a vertical column
can b e sent to an adjacent or nearby column via
corticocortical efferents, or it can be sent to distant
structures by commissural fibers (cortex on th e
o ther side) or p rojec tion fibers (s ubcortical stru c
tures). The minimal elem ents o f the vertical unit
are sh own.
201

REGIONAL NEUROSCIENCE Teleneeph.

'-t I
"
...
- ------1
,, t ""
,.
Association Fibers
l ong - 10 distant regio ns o f
_______ .,..:'
I I
~~
i~ilaterill hemisphe re 4# : 1.

Y, } ..
r-~"',""
\, .
Short - to nearby regions o f _______ ),. ~ \ , j
ipsilate ral hemisp ht're I

T / \
Comm;,," ,,1 Fib",
To cor ti ca l regio ns of
(on tr<l iater<ll hemisphere
<~_
""
.
..
:7
r \. .: . 1l-
"
-

Projection Fibers ~ nucleus 1

Corti cos pinal tract ~:: j \ _L
Cortlcobulbar tract . ~... t"". Putametl
~
;
.
Corticorubrospinal system

Corticoreticulospinili system \ 1 Globus J

Cortlco bulbospinal system

T ...
.
. I pallidus ;,:. ,
(po lysyn ilptic) III Venwce;
Corti coteetat fibe rs
Car lica pontine fibers
(to cereb ellum)
,,# <;: H ippocampus
,,:,~po'h,' ,m~
1"
.I
' '-p
",,-- '"
~.
--. .).
'.

"-.
C orticostriale fiber<>

(to ba!Hl1~ngliiJ )

Coniconigral and
conicosublnalamic fi bers
,I '1~
La te ral venui~
(la teral po le
Corticonud eiJf fibers (to seco nda ry ~ensory nuclei)

Corticothalamic projec1ions

Corticohypotha lamic a nd corlicoaulo nomic fibefs ' . PMa

Conicoo liva ry fibers "".

.""
,-,
Co rticoli mbic fibe rs (in subcortical ro renra in )

FIGURE 11.92: EFFERENT CONNEOIONS OF THE CEREBRAL CORTEX _ _ _ _ _ _ _ __

Neuro ns of the cerebral co rtex send efferent co n
nedio ns to 3 majo r regions. Short o r long associa
tion fibers are sen t to o ther cortical regio ns of the
same hemisphere by sho rt associatio n fibers
(nearby) or lo ng association fibers (at a distance),
Commissural fibers are sent to co rtical regions of
the other hemisphere thro ugh the corpus callos.
or the anterior commi ssure. Pro jectio n fibers a"E
sent to numerous subcortica l structures in the
telencephalo n, the diencephalon, the brain st ~
and th e spinal cord , Th e major si tes of termi na:'!
of th ese co nnectio ns are li sted in th e diagram.

202
 ...ha lon REGIONAL NEUROSCIENCE

Small
pyramid.,11
,ell

Subcortical projectio ns (mainly) Corticocortical axons Cortico cortical Corticothalamic axo ns

Some corticocorlical axons Commissural axons axons Some corlicoco rlical clXons
Some commissural axon s
Some ptojection <Ixons
J. Perkins lod.:austru m
-U, IoI''''
e~

E [1. 93: NEURONAL ORIGINS OF EFFERENT CONNECTIONS OF THE

CEREBRAL CORTEX _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

ICia tion fibers destin ed for cortical regions of
;":1e hemisphere arise mainly from smaller
.-.idal cells in cortical layers II and 111 and from
~_ d pyramidal cells in layer VI. Commissural
destined for cortica l regio ns of the opposite
"'Onere arise mainly from small pyramidal cells
in co rtical layer III and from some modified pyram
idal cells in layer VI. Projection fibers arise from
larger pyramidal cells in layer V and also fro m
smaller pyramidal cells in layers V and VI. O nly a
small number of projection fibers arise (rom the
giant Betz cells in layer V.
203
REGIONAL NEUROSCIENCE

MS I
Motor-sensory
{ Ms II

PremO lar; nrien tati on;

eye arld head movements

Visual I

Preironlal;
inhibitory conlrol

of be-havior;

higher intdligence

l anguage: reading;
speech
MOior conlrol
of speech --~I

5,, 1 } Sensory-motor
MO lnr-sen ~fy
III
PremOlor Temporocingulate and
Prefrontal;
parie tocingu lilte pathwilY
inh ibitory control

of behavior;
, Visual II I
highe r intelligence
,Visual II

~ ~f:~
{I@\.
,./

Fronto

cingulilte ,.
pathwa y _ _- ,I

Cingulate gyrus ~?L Corpus callosum

(emotional behavior) Hippocampal commi ssu re
il nd ci ngulum OilaclOfY '----Ante rior commissure

FIGURE 11.94: CORTICAL ASSOCIATION PATHWAYS _ _ _ _ _ _ _ _ _ _ _ _ _ _~

Cerebral cortex neurons have extensive connec
fions with other regio ns o f the brain (projectio n
neurons), w ith the opposite hemisphere (commis
sura/ neuronst and w ith other regions o f the ipsi
lateral hemi sp here (associatio n fibers). The cortical
association fibers may con nect a primary sensory
cortex wi th adjacent associa tio n areas (e.g., visual
cortex, somatosensory cortex), or link multiple
regions of cortex into complex associatio n areas
204
(e.g., polysensory analysis regio ns), or interl.i1"
importan t areas involved in language functicJo.
cognitive function, and emotiona l behavio r
analySiS. Damage to these pathways and a 5S
ated co rtical regi ons can result in loss of spec.
sensory and motor capab iliti es, aphasias (I
disorders), agnosias (failure of recogni tion
apraxias (performance deficits).
 alon REGIONAL NEUROSCI ENCE

:;'.t)E'rior occipitofronta 'i

faSciculu~ -
-----=7"""~:::r:::::~7- .....,
. J...~JP"'..

'"deulu,1~
looghud'o. ~~~III"~
ierior
;mntal --jf--"''''''"",f' '-_ -.3
~culus

",nClnate _ _;';',-;~
~ cul us

~{?
-
I ~

~~

Superior
occipito(ront a l - - - ,
fasciculus
I
...
"::'Itu d i nal ---::-i~--"
Superior
;.a SC iClI llI s ~".
q .
nudC'Us . Claustrum\ . ,~
, I /~ Lateral
~utamen
Thalamus ..,),. ~. fis sure

Inferior I Globus
1' 1
oc:pl.ofro nta l~ paJlidus

-~~
fa sciculus - til"
UnCIn ate
lilSClcuius
r,"c, Hypothalamus" - o "" I
\."
~.
/
_ _'"
~.
;~) I.MP5,..""
o~
..... f ..

URE 11 .95: MAJOR CORTICAL ASSOCIATION BUNDLES _ _ _ _ _ _ _ _ _ _ __

rsociation fibers co nn ecting cortica l regions in
I f hem isphere with adjacent or distant regions of
S<l me hem isphere are categori zed as short asso
o n fibers (a rcuate fibers) o r lo ng association
f 5. The lo ng assoc iati o n fib ers often are recog
- ed anatom ically as specific association bund les
r1d may have numero us fiber syste ms entering.
:ing, and traversing them . Impo rtant named
bund les include the uncinate fascicu lus, the
su perior lo ngitudi nal fasciculus, the superior
and inferior occip itofro ntal fasciculi, and the
cingulum. Th e cingulum is a bundl e through w hich
th e major mo noa mines (dopamine, norepineph.
rine, sero tonin) and par t o f the cholinergic projec
tions travel to their w idespread target sites.

205
REG IONAL NEUROSCIENCE
TempO f<l l lobe
AI.
FIGURE 11.96: NORADRENERGIC PATHWAYS _ _ _ _ _ _ __ _ _ _ _ _ __ __
Noradrenergic neurons in the brain stem project
to widespread areas of the eNS. The neurons are
found in the locus coeruleus (group A6) and in
several cell gro ups in the RF (tegmentum) of the
medulla and the po ns (groups A 1, A2, AS, A7l.
Axo nal projectio ns of the locu s coeruleu5 branch
to the cerebral cortex, the hippoc,ampu s, the hypo
thalamus, th e ce rebe llum, brain stem nuclei, and
the spinal cord. The locus coeruleus acts as a mod
ulator of the exci tability of other projec tion systems
(including nociception), such as the glutamate sys
tem, and helps to regulate attention, alertness, and
sleep-wake cycles. The RF groups are interco n
nected with sp inal co rd, brain stem, hypothalamic,
206
Telencephalr
\
and limbic regions involved in neuroendocri ne
tro l, visceral fun ctio ns (temperature regul ation
feeding and drinking behavior, reproductive be-1.
ior, autonomic regulation)' and emotional be
A sparse set of epinephrine-containing neuron5.
th e medu llary RF are similarly interconnect ed \\.
each other. The RF gro ups can work in co ncer:
with th e locus coerul eus during challenge or in
response to a stressor to coordinate aler tn ess ,:-t
appro priate neuroendocrine and autonom ic
responsiveness. The central noradrenergic ana
adrenergic neurons and th eir receptors are the'
gets of many pharm acological agents, inclu d in~
those for depression, analgesia, and hyp ertens

Temporal lobe
Raphe d~y" ;'
Raphe pontis centralis ",,~'i'''/
Raphe ~'"" ;< , /
Raphe pallidus and 0",,",,,, -
RE 11.97: SEROTONERGIC PATHWAYS _ _ _ _ _ _ _ _ _ __ _ __ _ _ __
nergic neuro ns (5-hydroxytryptamine
' -~ I. found in raphe nuclei of the brain stem and
en l w ings of cells, have widesp read proj ec
hat innervate every major subdivision of the
.:.. '-=euro ns in the nyf le \..! s raph_~.Qp~!'l!i s and
"'ltJcleus cenlralis superior projec t rostrally 10
ate the cere-brarcoriex, many limbic fore-
structures (hippoca mpus, amygdala ), the
gangli a, many hypothalami c nuclei and areas,
so me thalamic regio ns. N euro ns in the raphe
': us, pontis, pallidu5, and obscurus project
r e cauda lly to innervate many brain stem
_ ns, th e cerebellum, and th e spinal corcL The
eclions of the nucleus raphe magnus to th e
3d1 horn of the spinal cord influence opia te
~ ~ es i a and pain processing. The ascending
REG IONAL NEUROSCIENCE
\
\
seroto nergic systems are involved in regulatio n of
emotional behavior and a wide range of hypothala
mic functions (neuroen docrine, visceral/auto
nom ic). Serotonergic neurons are involved in
sleep-wake cycles and stop firing during REM sleep .
Projections to the cerebral cortex modulate the
processing of afferent inputs (e.g., from the visual
co rtex). The descending se ratanergi c neuron s
enhan ce th e effects of ana lgesia an d are essen ti al
for opiate analgesia. They also modulate pregan
glionic auto no mic neurona l exci tability and
enhance th e excitability of LMNs. Many pharmaco
logica! agents (in cl udin g drugs for treating depres
sion, other cognitive and emotio nal behaviora l
states, headaches, and pain) target serotonergic
neuro ns and their receptors.
207
REGIONAL NEUROSCIENCE
Ventra l tegmental Jre~
\
ci8!N
FI GURE 11.98: DOPAMINERG IC PATHWAYS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
Dopaminergic (DA) neurons are found in the mid
brain and the hypotha lamus. In the m idbrain, neu
ro ns in the substantia nigra p<!rs compacta project
axons mainly to the s.tria_tu~ (caudate nucleus,
putamen) and to the globus pa lli~u s and the sub
tha lamus. The nigrostriatal projection is involved in
basal ganglia ci rc uitry that aids in the planning and
execution of cortical activities, especially in the
motor system. Damage to the nigrostriatal system
results in Parkinson's disease. The antiparkin so nian
drugs, such as L-dopa, target thi s syste m and its
rece ptors. DA neuron s in the ~entra l tegme~!a l
area and the mesencephali c RF send mesolimbi c
p-rojections to the nucl eus accumbens, th e amyg
dala, and the hippocampus and meso co rti ca l pro
j~ctions to the frontal cortex and some cortica l
association areas. Th e mesolimbic pathway to the
nucleus accumbens is invo lved in motivation,
reward, biological drives, and addictive behaviors,
208
Telencep h.,:;
~Jf'
j.""",",,
,-,-,...."
particularly substance abu se. Projectio ns to Ii,......,
struc tures can induce stereotyped, repetitive
behaviors and activities. The mesocortical pro;e.
tio ns influence cognitive func tions in the plam
and carrying-out of frontal cortical adivities al'lQ
in attentio n mechanisms. The mesolimbic and
mesocortical systems and their receptors are -
targets of neuroleptic and antipsycho tic agents
influence behaviors in schizophrenia, obsess" -
co mpulsive disorder (OeD), attention deficit
hyperactivity disorder (AOHO), and other
behavioral states. DA neuron s in the hypotha
form the tuberoinfundibular dopamin e path \" iJ'"
which projects from th e arcuate nu cle us to til:
co ntact zone of the medi an emin ence, where
dopamine acts as a prolactin inhibitory factor.
Intrahypothalami c OA neuron s also influ ence
other neuroendocrin e and visceral/auto nomic
hypothalamic function s.
 ph.lon
(01 Meyl'left)
Bra,n, lem legmentJ!
cholinergic Woup
RE 11.99: CENTRAL CHOLINERGIC PATHWAYS _ _ _ _ _ _ _ _ _ _ _ _ _ __
al cholin ergic neuron s are found mainly in the
~ s basalis (of Meynert) and in septal nuclei.
deus basa lis neurons project cholinergic axo ns
Ie cerebral cortex, and septal cholin ergic neu
_ project cho linergic axons to the hippOCilmpa l
lion. These cho linergic projecti o ns are
. ed in corti cal activation and memory func
parti cul arly conso lidation of sho rt-term
r-.lOry. They often appear to be damaged in
-eimer's di sease. Drugs that enhance choliner
- nction are used for improvem ent of memory.
-.eor chol in ergic neurons, found in th e brain stem
REGIONAL NEUROSCIENCE
teg mentum, project to structu res in the thalam us,
the brain stem, and the cerebellum. The proiec tio ns
to the thalamus mod ulate arousal and the s!eep
wa ke cycle and appear to be import.ant in the initia
tion of REM sleep. Cho linergic interneuro ns are
present in the striatu m and may participate in basal
gangli a co ntra! of tone, posture, and initiation of
movement Or selectio n of wan ted pattern s of activ
ity. Acety!choline is the principal neurot ransmitter
in aUprega nglio ni c au tonomi c neuron s and LMNs
in th e spin a! co rd and the brain stem.
209
REGIONAL NEUROSCIENCE

Internal nasal branch (la t~ral ra mus) }0, anll!n.OI O lfactory bulb
Exlerna l nasa l I!lhmoidill n. O lfactory IraCI
branch _ _ _ ~Olbctory nerves
\

M axillary n.

Pterygopalatine
ga nglion and branches
'" Nelve o f pterygoid canal
Grea ter petrosal n.
Oeep pe\rosal n.

- - - - Pharyngeal branch
~ Naroopa latinc n.
Cpa s ~ng to !oqllum)

PosteriOI inferio r nasal branch

l esser (mi no r] palatine nn.

Cribri form 01
O lfac torv .
, bulb
Olfactory nerves
liil(['

(nlerna l n.lsal bra nch (medial

ra mus) o f anterio r ethmoid,,] n . --_ _ __

Nasopalatine n. _ _ _ _ __ _-."

Incisive cana! __

N~I
~Ium

~~p
, CIIQ'
....... ;.:

FIGURE 11.100: OLFACTORY NERVE AND NERVES OF THE NOSE _ _ _ _ _ _ _ __ __

The o lfactory nerves and their projectio ns into th e
eN S are an important compo nent of forebrain
function. Bipo lar cells found in the o lfacto ry epithe
lium are the primary sensory neurons. The peri ph
eral axon, a chemosensory transd uce r, responds to
the uniqu e chem ica l stimuli of airborne molecu les
entering the nose. The central axon s of bipolar neu
rons aggregate into approximately 20 olfactory
nerves that traverse the cribriform plate and end in
glo meruli o f the ipSilateral olfac tory bulb. These
nerves are vulnerable to tearing, resulting in anos
mia. Unlike neurons in o ther sensory systems,
210
these bi po lar neuro ns ca n proli ferate and reg;::
ate. After processing in the olfactory bul b, miL-a
neurons and tufted neurons project via the altar
tory tract directly and indirectly to limbic for~
stru ctures, in cl udin g septal nuclei and amygd
nu clei, an d thereby infiuence the hypothalam...s.
and its regulatio n of neuroendocrine and viSCer:3.
autonomic function. The olfactory sys tem is
involved in territorial recognition and defensE'
and wa ter acquisition, social behavio r, reproc _
tive behavio r, and signaling o f danger.
 III. SYSTEMIC NEUROSCIENCE
... A. SENSORY SYSTEMS
SO 'lATOSENSORY SYSTEMS

- -=- re 111.1 : Somatosensory Afferents to th e Spinal Cord

- =- re 111.2: Soma tosenso ry System: Spinocerebellar Pathways

=-SU re 111.3: Somatosensory System: The Dorsal Column System and

Epicritic Modalities
~ re 111.4: So matose nso ry Syste m: The Spinothalami C and
Spinoreti cul ar Systems and Protopat hic M oda liti es

:-_ re 111.5: Spinothalamic and Spinoreticular Nociceptive Processin g

in the Spinal Cord

- .:oure 111.6: Mechanisms of N europa thic Pain and Sympathetically

Maintained Pain

- ..:.ure 111.7: Descendin g Co ntrol of Ascending Somatosensory

Systems

TRIG EMINAL SENSORY SYSTEM

- u re 111.8: Trigeminal Sensory and Associated Sensory Systems
ENSO RY SYSTEM FOR TASTE

- ure 111.9: Anatomy o f Taste Buds and Their Receptors

-=igure 111.10: Taste Pathways

~ \UDITORY SYSTEM

~ ure 111.11: Periphera l Pathways for Sound Receptio n

Ggure 111.12 : Bony and Membranous Labyrinths

=- ure 111.13: V III Nerve Innervation of Hai r Cells o f the

O rgan of Corti

figure 111.14: Coch lear Receptors

figure 111.15: Afferent Auditory Pathways

f igure 111.1 6: Ce ntrifu gal (Effe rent) Audito ry Pathways

w. VESTIBULAR SYSTEM
Figure 111.17: Vestibu lar Receptors

Figure 111.18: Vestibular Pathways

Figu re 111.1 9: Nystagm us

6. VISUALSYSTEM
Figure 111.20: A nato my of the Eye
Fi gure 1IJ.21: Anterio r and Posterior Cha mbers of the Eye
Figure 111.22: The Retina and the Pho toreceptors
Fi gure m.23: Arteries and Veins o f the Eye
Figure 111 .24: Visual Pathways: Retinal Projectio ns to the Thalamus and the Brain
St e m
Figure 111.25: Visual Pathways: The Retino-Geniculo-Calcarine Pathway

Figure 111.26: Visua l Pathways in th e Parietal and Temp oral Lobes

211
SECTION B. MOTOR SYSTEMS
B.l. LOWER MOTO R NEURONS (LMNs )
Figure 111.27: Lower M otor Neuron D istribution in th e Spin al Cord and the Bra in Stem
Fi gure 111.28: Alpha and Gamma Lower Motor Neurons
B.2. UPPER MOTO R NEURONS (UMNs)
Figure 111.29: Cortical Efferent Pathways
Figure 111.30: Corli cobul ba r Tract
Fi gure 111.31: Corticospinal Trac t

Figure 111 .32: Corticospinal Tract Terminations in the Spinal Cord

Figure 111.33: Rubrosp inal Tract

Figure 111.34: Vestib ulospin al Tra cts

Figure 111.35: Reticulos pi nal and Co rti co reticular Pathways

Figure 111.36: Tectospinal Tract and Interstiliospinal Trac t

Figure 111.37: Spinal Cord Te (minations of Majo r Descending Uppe r Mo tor Ne uro n Tracts
Figure 111.38: Ce ntral Contro l of Eye M ovements

Figure 111.39: Centra l Con trol of Respi ration

B.3. CEREBELLUM
Figu re 111.40: Func tional Subdivisions of th e Cerebellum

Figure 111.41: Cerebe llar N euronal Circui try

Figure 111.42: Circuit D iagram s of Affe rent Co nnections in the Ce rebell um

Figure 111.43: A fferent Pa thways fa the Cerebellum

Figure 111.44: Cerebellar Efferen t Pathways

Figure 111 .45: Vestibu loce rebe ll a r a nd Cerebe lloves tibula r Pa thways
Figure 111.46: Schematic Di agram s of Efferent Pathways Fro m th e Cerebellum to Upper Motor
Ne uronal Systems
B.4. BASAL GANGLIA
Figure 111.47: Conn ectio ns of th e Basal Ganglia

Figure 111 .48: Basic Basa l Gan glia C ircuitry and N eurotransmitters

Figure 111.49: Parall el Loops o f Circu itry Through the Basal Ganglia

Figure 111.50 : Connections of the Nucleus Accumbens

SECTION C. AUTONOM IC. HYPOTHALAM IC, AND LIMB IC SYSTEMS

C.T. AUTONOMIC NERVO US SYSTEM
Figure 111.51: General O rganizatio n o f the Auto no mic N ervou s System
C2. HYPOTHALAMUS AND PITUITARY
Figure 111.52:
Gen eral Anatomy of th e Hypothalam us

Fi gure 111.53:
Sectio ns Th rough the Hyp othalamus: Preoptic and Supraop tic Zones

Figure 111.54:
Sections Through the Hypothalamus: Tuberal Zone

Figure 111.55:
Section s Through th e Hypothalamus: M ammi ll ary Zon e

Figure 111.56:
Sc hem ati c Recon stru ctio n of the Hypoth alamus

Figure 111.57:
Fo rebrain Regions Associated W ith the H ypothalam us

Figure 111.58:
Afferent and Efferent Pathways Associated W ith the H ypo thalam us

Figure 111.59:
Schem atic Diagram of Major Hypoth alamic Afferen t Pathways

Figure 111.60:
Sc hem atic D iagram of Major Hypo thalami c Efferent Pathways

Figure 111.61:
Summary of General Hypothalamic Con nections

Figure 111.62:
Paraventricular N ucleus of the H ypo thalamu s: Regulati on o f Pituitary

Neurohormonal O u tflow, Auto no mic Pregan glioni c Outflow, and limbi c Acti\'i:'
Figure 111.63: Mechani sm s of Cytokin e In fl uen ces o n the Hypothalamu s and Oth er Bra in
Reg ion s and o n Beh avior

212
 111.64: Circumventricular O rgans
111.65: Hypo physeal Portal Vasculature
111.66: Regulatio n o f Anterior Pituitary Hormone Secretio n
e 111.67: Posterio r Pituitary (Neu rohypo physeal) Ho rmones: O xytocin and Vasopressin
e 111.68: Vasopressin (Antidiureti c Ho rm one) Regulation o f Wa ler Balance and Fluid
Osmo lali ty

... e 111.69: Th e Hypothalamu s and Th erm oregulati on

==-e 111.70: ShorHerm Regulation of Blood Pressure

re 111.71: Lo ng-term Regulation of Bl oo d Press ure

~ :- e 111.72: Neural Control of Appetite and Hunger

;. e 111.73: Neural and Neuroendocrin e Rol es in th e Fi ght-o r-Fli ght Respon se

_'.Ire 111. 74: Neuroimmuno mo dulatio n

J \IBIC SYSTEM
~""'!lI re 111.75 : A nato my of the Limbic Forebrain
=-""!Ure 111.76: Hippocamp al Formati o n: General A nato my
~~re 111,77: Neuronal Connections in the Hippocampal Formatio n
- .:;ure 111 .78: Maj or Afferent an d Efferent Co nn ecti o ns of the Hi ppocam pal Form ati o n
:-~ure 111.79: Majo r Afferent Connections of the A mygdala
:-~ re 111.80: Maj o r Efferent Co nn ectio ns of the A mygdala
- ~ure 111.81 : Summary o f Major Afferents, Efferenls, and Interconnectio ns o f the A mygdala
:-- ure 111.82: Major Affere nt and Efferent Connections of the Septal Nuclei
=-.,:)ure 111.83: Major Connections of the Cin gulate Cortex
OLFACTORY SYSTEM
"0
ure 111.84: O lfactory Receptors
~ ur e 111.85: O lfacto ry Pathways

213
SYSTEM IC NEUROSCIENCE Sensory Systems- Somatosen sor

Proprioception

Touch and
pressure
. -----...
Pain (nociceplion) { I --....
and temperature T --....... -- ' --- -~,

c".'- '''.~
'

Dorsal spinocerebellar
tract
Ii

Spinothalamic and ---f<

; -- "'
spinoreticular tracts i\

J. PerbF
MS ..,..
To skeletal ( ...-'" c ~.

muscle 0( -----

FIGURE 111.1: SOMATOSENSORY AFFERENTS TO THE SPINAL CORD _ _ __ _ __ __

Unmyelinated IUNM) and small myelinated 1M) originate. M axons also project into fasciculi gra:i

axons that convey nociception and temperature and cuneatus for lemniscal pathways destined

sensation terminate in laminae I and V (origin for conscious interpretation. M proprioceptive ax;

spinothalamic tract). Other UNM axons terminate terminate directly on lower motor neurons (l "'"

in the dorsal horn, from which neurons for poly (via la afferents) and the la interneuronal poo;

synaptic reflexes and for the spinoreticular system and also terminate on neurons of origin for tr

originate. M axons for touch and pressure termi spinocerebellar tracts.

nate in the dorsal horn, where additional reflex M = myelinated; UNM = unmyelinated.

connections and spinothalamic projections

214
 Systems-Somatosensory SYSTEMIC NEUROSCIENCE

/'

-....... "'-,,,,,,,...
',,
~<

..." ~,,-
" ",. <"" "',':'
Cerebe llum '\;Y"~
.\ 'c '\d
'7 1.,); ~_J:.:.)?1
"i / ' ! J ,./
""- . >{ (*->.;/
r~ \'-,..1...,,,,,.' Cuneoce rebellar trtlct
: :;J . ;-
; .'
_~

: ': " Inferior cerebellar peduncle

i~}.-I La te ral (accessory) cuneate nudeus
~-r~\
' ~l~~ Rostral spinocereb ellar tract (RS(T)
/
L-l
Upper Body / ( \
(above T6 )

J! .. ' __
.L
- - - - 1-"'.:.JJJt-c-""--4-.-
-- .
; .\~~l ~
-
_ la (to cune(lce rebellar tract)
-~-- Ib (to RSCT)
Ventral spinocerebellar tract (Vscn -J, <1>- - Do.-sal spinocerebellar tract (DScn
(1' i If\
"7~:';,~";'~ ) r).1 !"~ j .~
--.___.U~\\ .1__-. . - Ia(10 oscn ,. Perkins
~---.~- ................~. ~\ ::to.'~---'-"'-"-----'~_ _ Ib (to VSCT) MS,M FA
OllON
Anterior w hite commissure '--'-"

RE 111.2: SOMATOSENSORY SYSTEM: SPINOCEREBELLAR PATHWAYS _ _ _ _ _ _ __

ocep tive pnmary soma tosen sory axons
- jo ints, tendons, and ligaments (represented
afferents from Golgi ten don organs) term i
.R o n neurons of o rigin (border cells, dorsal
.,., neurons) for the ventral spinocerebellar
- ~ and the ros tral spinocerebellar tra ct from the
-er and upper body, respectively (16 is cutoff).
-nprio ceptive prim ary somatosen so ry axons from
""'.... e spi ndles (Ia afferents) terminate on neurons
of origin (Clarke's nucleu s, laterall external]
cuneate nucleus of the medull a) for th e dorsal
spinocerebellar tract and the cuneocerebellar
tract from the low er and upper body, respectively
(16 is cuto ff). The do rsal spinocerebellar, rostral
spinocerebellar, and cuneocerebellar trac ts remain
ipsilateral. The ventral spinocerebellar tract crosses
twice, once in the anterio r white commissure and
again in the cerebellum.
215
SYSTEMIC NEUROSCIENCE Sensory Systems- Somatosens.

I
Cerebrum
"
\
I '/ L1 ' .j

e.'-.-~ h
~ "d~,
CerC'h ral (Ort el( _ p
po;tcentr~1 gyru s J
Po,"""' hmbor
"" "" I""pro _ _-'l.._~_ __
Ventral p<lslelOlateral
(VPl) nucleus of
t
'--"'"

\:"
///-'r\ f
/
~
'Y ~
J J:..-J
J
'''.'',m,,,,

thalolmus "w" "

..... )
~
' ' .J I ...
,If.
I..,
:d'<c~

.' .,.
1;
~,~.;'
""""" 1
J. Pe rkins
.~
M~ MM,
1;'1,. ,~

"" Midbrain
. 1"'
1-=~' Medial lem"iscus

. 1
Pons
.
~' II~ ~,
./

" -"'ia"\! nlni sc.u s ~l

, "~ "'
'.

.\
'.L trr l
''''y.'Ir'p~v
I' ,.
/

I
Gracile nucleus
~ Cu[)a te nucl e us

~m"i"",1
L we, medun::. 'j; , IS

o .. , lor",,1 ,"co " ,,'" door

~
. (decu ssa tion of the me

Fas.clculus
fJscicu vae , ,s.
l u~ c.uneatus~ '\ ~
'" . II LateralcervICal nuc Ieus (C I -C 2 0nlyl

t
I'" '

c"""r' p""r ",", )f"',

/ \. ~ ,\ J j. .J~
_ .., p,"p~o,~: ~~,;,po~ilion
To", ,p i n "b""p"

~ 1~ " ....:: " ",

J' ' \
lumbar spinal cordi L
'
....-h~..-~'t~
'!!... __~.~~
I \

_.3"-_ --
.~ -~
""OI"'''''P''OO, '"bSit loll n
'''''"' pressure, VI rallO
- T-"h

FIGURE 111.3: SOMATOSENSORY SYSTEM: THE DORSAL COLUMN SYSTEM AND

EPICRITIC MODALITIES _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Prima ry somatosensory M axo ns that co nvey fin e nates. Th is tract crOsses in th e medulla and pr
discriminative to uch, pressure, vibratory sensation, to the ventral posterolateral (VPL) nucleus o f tho<
and co nsciou s joint positi o n se nse project directly thalamus. The VPl nucleus terminates topogra'
into th e dorsal column system, w here they are ca ll y in the prim ary se nso ry co rt ex. The spin o
topographically organi zed (fasciculus gracilis for cervical system contributes polysynaptic
lower body [below Tbl, fasciculus cuneatus for mechanoreceptive informatio n to supplement
upper body [T6 and above)). These axons termi that in the do rsal column system.
nate, respectively, in nucleus gracilis and nucl eus M = myelinated.
cunea tus, from w hich the medial lemniscus o rigl

216
 Systems-Somatosensory SYSTEMIC NEUROSCIE NCE

;,
\ !
Cer~brum ~

'
C~r~br al corle!C:
poslcenllal gyru'- r ~
~
'{
PO~l enornmb of
( -~""
' .i ( "-.-J
.J~. - NOMpecific
mt erna l caps ule ~ . - - --'-:- --,jl-----'-- thalam ic nuclei
\ :.j (ce ntrorn edianl
',..,.
1\ . , 4

JJ
(Vr u nuc1eus of .
th.llamus \ .... , .......

\.:. ".; J I I ~

~
,....)
J. Perkins
.~"
, Deep layers of
Midbrain ,. superior colliculus and

r~.... ",,'''q'''''''''''Y

Pons

lower medulla

Spmothalamlc/sPIl10r~!l cular
\'R
...~l "..;
u teral leticular formation

sys tem (irom all spllla lleve!sl ----'

Ce rvical spinal cord :I~~F'~

Aowk" ".i" <om"';"" , -r.l';I!':Ij=::C :s::::::::==

-.
Pain. tempe rature
Polin

lumbar s.p inal cord ,

A01erior while commiS5Ure - - ;h ; = ,~:::::== Pain, tempera ture

"", _. - Pain

" RE 111.4: SOMATOSENSORY SYSTEM: THE SPINOTHALAMIC AND SPINORETICULAR

SYSTEMS AND PROTOPATHIC MODALITIES _ _ _ _ _ _ _ _ __ _ __
~ r y somatosensory UNM (C fibers) and small
; 8 fibers) axons that convey nociceptive infor
-.' n (fast, locali zed pai n), temperature sensa
and light, movin g touch termi nate on neuro ns
minae I and V. These neurons send crossed
DOS into th e spin othalami c tract projectin g to
VPL nu cle us of th e th alamus. These th alami c
...rons project to the secondary somatosensory
":ex (5 11 ), as well as to the primary sensory cor
Prim ary se nso ry C fibers co ntribute to a large,
.2SCading network in the dorsa l horn resu lting in
bilateral projectio ns into the spinoreticular tract.
This system ends mainly in the reticular formation
(R F), fro m w hi ch po lysynaptic proj ections lead to
no nspeCifi c, medi al dorsal. and anterio r th alamic
nuclei. Some spinoreticular fibers termi na te in the
deeper layers of the superior co!li culus (spinotecta
pathway) and pe ri aquedu ctal gray. Co rti ca l regio"'$
such as the cingul ate, in sular, and prefron tal
regio ns then process and interpret noc icepti\ e
info rmatio n related to slow, excrucia ting pain.
M = myelin ated; UNM = unmyelin ated .
SYSTEM IC NEUROSCIENCE Sensory Systems-Somatosen"'

Gating Mechanism Spinal Mechanisms of Nociceptive Processing

DOrs.)I column aiierent

...
r'
O --- '~ ' ,.~--
,---
'.
\ I
!
f
!
/
/~

"'\
.~
,'.,
I.
'j
~
I j

i I

,j,
/ ,
J
\i',
iI ~.
\ \ /
.
!

,.
i
~,
Spj nolhc1 l am~
spi norelicu:";'
tract
"
,
!
\( ,
"
,. Perkins
._".- pYi'=
MS, MfA
0 1lllN
"" .~"
'-.. -."-..;, .. .q.r(J '
~ ~ ~ ~ ~

FIGURE 11 1.5: SPINOTHALAM IC AND SPINORETICULAR NOCICEPTIVE PROCESSING
IN THE SPINAL CORD
Unmyelinated primary afferents (c, AcS fibers) th at
convey "fast, localized pain" and temperature
sensation terminate in laminae I and V, from w hich
the crossed spinothalam ic axons originate. UNM
primary afferents (C fibers) also termil'late on neu
rons in the dorsa l horn, from whi ch a cascading sys
tem of recruitment, convergen ce, and po lysyn aptic
interconnectio ns o riginates. Th is system con
tributes to the spinoreticular trac t (ma inly crossed),
218
~All'A'A.
i,
.AlTA. ..__... _-
- --------------- - - --i
which projects into the RF and th en project~
synaptically to nonspecific, medial dorsal, arc
an terior thalamic nuclei. It contributes to perce"
tio n of excruciating pain and its emoti onal Cor-'
tatfon via cortica l regions such as the cingula:?
insular, and prefron tal cortices. The gatin g mea.
ni sm, shown o n th e left allows prim ary dorsc.:
column axon col!aterals to dampen pain proa!!l
ing in the dorsal horn.
 Systems Somatosensory SYSTEM IC NEU ROSCIENCE

~s of Neuropathic Pain Cenlral seroto ni n .md

- ... S 01 sympathetic postganglionic nerve fibel'> on 10 affe renl ending~ and 1 ~ sensory ce ll hodies_ cenllal norepineph rine
:t-d ~ hre5hold for firing of C fibers (hype resthesia) and AS fibef"S (allad yn ia). p ath ways
ore-atio n of a-<ld renergi c receplorS on 1 sensory affere nt e ndings and 1 senSory cell bodies.
oI:JIoe ep haptic afferent activati on .
Q 0
o
............-::: nt hyperaclivation of wide dynamic range neurons.
ooi&r".Jte excilotoxic cell death of inhibitory neurons (gluromate storms).
3Cy of central de~endin s serotonin, no repinephrine, opioid
-.cP. palhw.lY$ to conlrol nociceplion.
ZJtion by pain decreases ga ting of nociceptive input.limiling
-K..1I therapy to ini tiate gating.
-'":8 of C fibel""S in spinal cord.
~tOn of interneuron dendrite s into Dorsal root
~ I spinal co rd laminae.

Preganglionic
sympathetic fib er

Sym pathetic J. Perkins

.vIS, MfA
chain ganglion Q~

~ RE 111.6: MECHANISMS OF NEUROPATH IC PAIN AND SYM PATHETICAllY

MAINTA INED PAIN _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ __

- cascading dorsal horn system-which receives
-..ary afferent C fibers of nociceptive origin
- projects into the spi noreticular system for the
ious interpretation of excruciating and neuro
- ic. pain-is illustrated. The sympathetic connec-
-s in ad dition to secretin g norepineph rine into
oIood and activatin g the secretion of epineph
can synapse with terminals and cell bodies of
primary nociceptive neurons in neuropathi c pain
syndromes. Descendin g ce ntral noradrenergic and
serotonergic projectio ns are also shown. Specific
mechanisms relevant to neuropathic pain, particu
larly complex regional pain syndrome (reflex sym
pathetic dystrophy), are described in the numbered
sites.

219
SYSTEM IC NEUROSCIENCE Sensory Systems-Somatosen-

From cer ebral from hypori1al<Jmus Stimuli from higher From

cortE'X and (i ncludes beta- center, (P'iydlO- sensory
limbic forebrain endorphin axonal logical, pLacebo cort ex
~ projec.tions] effect, etc.)

~ -t~ Eo'eph, "o ooo,,'o'o, ",''co

.
~
'
.. , '\ ." - Periilqued uda l gr'IY ma il er

>.,

" "', ) M'db,,'o

. LOM <oe~ ." , , , ,f'

Anee,", pa'o m~" t
\ IF ~-'K.
tngen1 u1al nerve _________-.. . .. . ...
. '~. L" ,"-,"-
ru,e \\
~
In -

~_~
Sp,"llogemm"
",""m,,',
,m, oed ",",00 OU'I'USY'"~
' 11 IstJ" ,' \ ' ;
-I
'.
'
I ..
,
J Llte'" cetlcul" lo,m,,'oo

~
I .
I
,,,,,;tem I ' ( ., . '. & Medu'"
..
norJdrene' egmental
'goc " II gmups
\" ..!"
,;
:-N
'

~",
Descending no repi nephrine
Serotonin
palhWay~
palhway~
t I Corliconudear fi ber~
<D----- Spinoreticular pathway

) ! D ecussa tio n o f the pyra mids

.......:t;

Pos,e, o'a
l era l lu n iculu~

I
~
/
,
.
,.' ,
~:
.r.--.~
1 , ;;' I
L An terolatera l fu niculus

h -n-containing neuron
'okep " '
/
d' /
/ / "..;
4 f.,
"I
" t \ 5 1
.. -.: -~ ~
Afferent pain neuron - / -
I
o f dorsa roo ,gan~ho)
n .'
/ '! Il~ lI \. "" \ Sp'oa' wd
'I,I) " ".

~ ; ')
/" /'~ 7~'-""
/ 4"''' ' ' -- ~..... :-....~,~,
, _
~ ,l
:.
t ~".L . .-. .~ - ~'} ~'" ~~~~
. 'i,iiJtFJ'. --~ - ,..-...~ , - . . ~".;;;. ,
. .Y-"-'''''''''''''~~".~~"'~ I."";'"
".~I .
tI ,I.~
Spino rE'ti cular neuro n

FIGURE 111.7: DESC END ING CONTROL OF ASCENDING SOMATOSENSORY SYSTEMS _ __

Descending connection s from the cerebral co rtex, neurons (nucleus raphe magnus) modulate the
the limbic forebrain structures, the hYPolhalamus processing of nociceptive information in the c:ic:J
(the paraventricular nucleus and the periarcuate horn of the spinal cord. The central descendirr;
beta-endorphin neurons), the periaqueductal gray noradrenergic and seroto nergic pathways, inilu
(PAG), the RF of the brain stem, central noradrener enced by the PAG and other higher centers. 2-e
gic neurons (of the locus coeruleus and other brain important for endogenous (and exogenous) r
stem tegmental groups), and sero tonergic (SHn lation of pain.
220
SYSTEMI C NEUROSCIENCE Sensory Systems- Somat

From cef"ebral r-mn1 hypothalamus Stimuli fr om higher From

cortex and {indude!i beta cen t~r s {ps~'ch o ~enso r y
limbic forebrain endorphin axonal logk'<ll, placebo (;(lrt ex
~ projections) effect, etc.!

~. -~'~
1S *
-/
,
/ E"'.ph,I;"<0"" ;";"8 "'"'''''

. t :..._..c
:>'~<-_ _
, . . '\ I>eriaqueductal gray m<ltlC'f

., Midbra in

locus coeruleus

~f(e~ent pain ribers \_. !" ......

'" t".'m; ~" "en', ~ '\"':"!~

SPinallrisp.nlimd7 -- ' i"l

!rJet and nucleus :
-,
.,/

j
_' Latera l ret icular (orm atioc.
I

~ Medulla
Enkephalin-coru aining neuron

Brainst emtegment al~

.
h ~i
- ':
,
no radrenergic cell groups

Descending norepine Phrin.e pathw,ly

\.'
~

a "" ,r,
' Coniconudear fibers
Serotonm p.lthway

Posterola teral funiClJlus d

7

!=E Spinore lic ular pathway

De c u~sa ljoll

~ An l erolaler,,1
of the pyraf"1;

funiculus

Enkephalin<on tainif'lg neu ro n .? 1~''''' II! t~ { "

'"
Afferen t pain neuron f- "" Spinal c(
o f oorsal foot ga n ~ ion ----r-
\

--
'
.. ---_ _L:.:.;~"'"-
---f-'<r~~
/ ""'.
A

..
... -;tJi/#Jt
,, ~~--..,
-" ....~
.,....------' '" - ,
.\,.
'-._ ; \
tf ~
".
Spinoreticular neuron 't

FIGU RE 111.7: DESCEND ING CONTRO L OF ASCENDING SOMATOSENSORY SYSTEMS _

Descendin g connections from the cerebral cortex, neurons (nucleus raphe magnus) modulat
the limbi c forebrain stru ctures, the hypothalamus processin g of nociceptive information in II
(the pa raven tricu lar nucleus and the periarcuate horn of th e spinal cord. Th e ce ntral descel
beta-end orphin neuro ns), the periaqueducta) gray noradrenergic and seroto nergic path wa~ ~
(PAG), the RF of th e brain stem, central noradrener enced by the PAC and other higher cente
gic neuron s (of the locus coeruleus and other brain important for endogenous (an d exogenol!
stem tegmental groups), and serotonergic (SHT) lation of pain.
220
-
~ns
- ory System s- Tri geminal SYSTEMIC NEUROSCIENCE

,
Cerebr,J1cortex.

(ntem.ll capsule
1-- (:",,~ ,"j. nudc~ (i nlr;,t;.,minilr)

f--- v,,,,,, po slerolTlff:iial N PM )

,,,-,de us ()/ th;JJ;u ,"IUS

Midbrain

I mesenceph,J1rc nudeus
Dorsal tli ~mina J ~mn"j'K~~_---=~~~~
~
(dors~l l.i8c'ninolh,)l,)mi( If.

Venrr"llri~emin.'1
. ,
(venfldl trfgen,inOlh,..l.lmic (.act!
Pontine tl!mpl!. "IUrf - -
Propfioctptio n ' f,om mu sd.. . pi nd l... -

,~.
<~~~s;:~'
,./ . .'". ~.
~~2~~'
~
. \ .oot oi m.lndibul.!.r n.

r~i'~<~"'~"~~~~;:~~~~~~~
Later,,1 r I
Medi~ lr

Medull"ry
Venlral trigemin.J11
Spin,,1

(ngcmin..1

Spin. 1 ,

mgemin,,1 nucleus

I .r I I I

Ccrvi.cal spinal cnrd

9G URE 111.8: TRI GEMINAL SENSORY AND ASSO CIATED SENSORY SYSTEMS
_ _ _ _ _ __
- IJ)ns of primary sensory neuro ns enter the brain and to in tralaminar thalamic nuclei, w hich are asso
-:::I"n, travel in the descending (spinal) (ract of V ciated w ith nociceptive processing. The caudal
rid terminate in {he descendin g (spinal) nucleus of descendin g nucleus of V also sends bilateral
-\xons of the trigeminal ganglio n (V) supply the prOjections to the RF to process excruciating pai n
e, the anterior ora l cavity, th e teeth, an d th e (similar to th e spinoreticular system). Primary sen
_...ms; axons of the geniculate ganglion (VII) and so ry axon s that carry fine discriminative modalities

-
e ju gu lar ga nglion (X) supply a small zone of th e
~'e rna l ear; and axons of th e petrosal gangli o n (IX)
oply general sensa tion to the posterior oral cavity
.-xl fhe pharynx. Axons from the descending
cleus of V project into the crossed trigeminal
miscus (ventral trigeminothalamic tract 1VTT11 "
"1ich termi nates in the ventral posteromedial
.. PM ) nucleus of the thalamu s. The V PM nucleus
ojects to the lateral primary sensory cortex (51)
from V (simi lar to th e dorsal col umn) te rminate
in the rostral descendi ng nucleus of V and the main
sensory nucleus of V, w hich contribu te to the
VTTI. A po rtion o f the main sensory nucleus also
projects ipsilaterally to the VPM nucleus. The
mesencephalic nucleus o f V, a primary nucleus
inside the (NS, mediates muscle spindle reflexes
for masticato ry and extraocular muscles.
221
SYSTEMIC NEUROSCIENCE Sensory Systems-Taste
-SenSi

B. Sccti(Jn through
vallate papilla
Foliate
papillae

--.... T,,~t .. buds

Duct of gusta tory

(Ebner's) gland

M"""
and M
Fun giform C. Tule bud
papillae
t of Irigi

Va llate pa pillae:
~EPithelium
Basement membra ne

M icrovilli ~"_. Nerve p lexus

Taste pore ," # I),h T:--- Nerve fibers emerging

itom taste buds

LMge nerve
fiber
Intercellular

Fibroblast

~ Small nerve fiber

largE! nerve fiber

low"
De!tmosomes Collagen oblofl;!
Schwanncell
C ranules _ _
Basement membrane Pee
Epithelium ----.j o i~

D. Detail of taste pore E. Detail of base receptor cells

~~p

OIIl!N : ......... 1I

f iGURE 111.9: ANATO MY O f TASTE BUDS AN D THEIR RECEPTORS - - - - - - -- - IGURE III

Taste buds are chemosensory tran sducers that and posterior regions of the tongue and, less fre- 'rimary sen
consist of bundles of columnar cell s that lie within quently, on the palate and the epiglottis. Nerve anglion (\ /j
the epith elium. They translate individual molecular fibers for taste show complex responses to electr~ odose (infE
configurations o r combinations of molecul es for cal activity across populations of many nerve fibers. le anterior
salty, sweet, sour, and bitter sensations into action The integrative interpretatio n of taste takes place -'lird of the I
potentials of both small and large primary sensory the e NS. ~spect ivelv
axons. The taste buds are found on the anterior
.'
irt of th e n.
,et), which
222

 ry Systems-Taste SYSTEMIC NEUROSCIENCE

venlral poste romedial (VPM)

nucleus of thala mus
Sensory cortex (just below

face area)

Lateral hypothalamic area

Amygdala

Ponti ne taste area (parabrac hial nucleus)

Trigeminal (V) n.
Trigemi nal (semilunarl ganglion
;e;;encephalic nucleus
_ ''Iotor nucleus
- .,geminal n.

~~'%l~~M;n
~' ldibular n,
Pons - ---"-""-
Pterygop alatine gan8iion

Greater petrosal n. = = = =+ - -Lf.:P"j?fll f

Geniculate ganglion Ne~e ofote'",oiid canal

Facia l (VII ) n.
and Nerv us intermedius

ROSlral part of nucJeus

of solitary tract -~-
Fungiform papillae
Glossopharyngeal (IXl n. - - - --f,

-
---+\.(-- -~ Foliate papillae

Lower parI of merlulla "

oblongata ",-, " "Ihi '" papillae

"""
Petrosal (inferior) ganglion

of glossopharyngeal n.

Nodose (inferior) ganglion

of va gus n. - - -_ _/
Vagus (X) n.

'I CU RE 111.10: TASTE PATHWAYS _ _ __ _ _ _ _ _ _ __ __ _ _ _ _ _ __

:>rimary sensory axons of neurons of the geniculate
.,anglio n (VII ), the petrosal ganglion (I X), and the
~o dose (inferior) ganglion (X) supply taste buds in
Ie anterior two-thirds of the to ngue, the posterior
mird of the tongue, and th e ep iglo ttis and palate,
espectively. These axons terminate in th e rostral
part of the nucleus solitarius (nucleus of the solitary
'ract), which sends ipsil ateral projections mainly to
the parabrachial nucleus in the po ns. The
parabrachial nucleus pro jects to the VPM nucleus
of the thalamus, the hypothalamus (lateral hypo
thalamic area, paraventricular nucleus), and the
amygdaloid nuclei. These no n thalamic projectio ns
are associated with th e emotio nal, motivational,
and behavio ral aspects of taste and food intake.
223
SYSTEMIC NEUROSCIENCE
Sensory 5ystems-Audiror~'

Umhs ,,{ c bn _
front al section
Prominence of lateral semicircular CJncll
of stapes in oval (vestibular) wi ndow
Incus
Tegmen
Semicircular duCtS, ampullae, ulricle and saccule
Epitympanic Facial nerve (VII) (cut)
Vestibu lar nerve

Auricle
I acou stic meatus
Veslibuloco chlear
nerve (V III)

j'.

"
Tv
R,
\~ l

to
External
acoustic meatus Scala vestibuli "
Cochlear duel Cochlea
containin g spiral

,
I
organ (COrti)
Scala tympani ~~p
CIIl!N
I~.W;~
Pharyngotympanic (auditory) tube N o te: Ar rows inDICate course of sound wav~s

FIGURE 111.11 : PERIPHERAL PATHWAYS FOR SOUND RECEPTION

_ _ _ _ _ _ _ _ __ Fie
The sound transd uction process involves move axons of neuron s of th e cochl ear (sp iral) ganglion. Th e
ment of mech an ical so und through the externa l The basi lar membrane in the cochl ea shows maxi Yes !
ea r and the external acoustic meatus. across the mal displacement spatially accord ing to the can
tympanic membrane, leveraged as a mechanical frequency of impinging to nes. with low frequen roUi
force by the bones of th e middle ear (ossic1es) via cies stimulating the apex (heli cotrem a) and high incu
the oval window to produce a fluid wave in the frequencies stimulating the base. The eustachian mer
cochlear duct This fluid wave ca uses differential (pharyngotympanic) tube permits pressure
mOvement of the basilar membrane, stimulating equilibration between the middle ear and the
hairs on the apical portio n of hair cells to release outside world .
neurotransmitters that stimulate primary sensory
224
""6lsory Syste ms-Audito ry SYSTEM IC NEUROSCIENCE

Bony a nd mem branous labyrint hs: sch e ma

Ampullae

Anteri or semici rcular canal and duct

~o?I'io r
semicircular
,...,=~- Dur,l mJter

- cln d duct

Endolymph atic sac

-" lon body and Endolymphatic duct in

.e-:branou s limbs ve stibular aqueduct

Utricle

al semicircular

and duct _ _ ~ Saccule

capsule _ _ __

"'!PeS in oval Ductus reuniens

~,:, !lula r ) window

-panic cavity _ _ _ _ _-.:::~

-".'nal acouslic meatu s -

oo ------------~

- "'panic membrane
Cochlear Jqueducl
md (cochleClr)
"<fow (ciosed

:occondtlry

-,J<tnic membrane)

AG URE 111.12: BONY AND MEMBRANOUS LABYR INTHS _ _ _ _ __ _ _ _ _ _ __

--~ relati onship between the cochlea and the
3tibular apparatu s (utricle, saccu le, semicircu lar
-.a ls and du cts) and th e bony labyrinth that sur-
" ds th em is illu strated. The ossicles (malleus,
"IICU S, stap es) leverage move ment of the tympanic
l1brane to produ ce movement of the oval
window. M ovemen t of the oval window causes th e
tluid wave to move through the scala vestibuli and
th e sca la tympani of the coch lea and ricochet onto
th e round window. The 3 semicircul ar can als are
located at 90 angles to each other, represe ntin g
tilted X, Y, and Z axes.

225
SYSTEMIC NEUROSCIENCE Sensory Systems-Audit ~

Seelion through turn of cochlea

ves tibuli
(perilymph w eakly +)
Osseous
spira l lamina Vestiblilar (Reiss n.,...-,.
membrane

Nerve fibers
(endolymph
+ 80 mV)

Spiral
Modiolus of ligame.-;
cochleus
'ic.
membra-.

$pirJlga nglion _ _
Spiral Qrg-,,-
(Coni)

Scala tympani - _ _- '

(perilymph 0 mY)

Cochlear nNve Osseous cochl ea

~{p

Inner hair cell

OIl!!N :;:.... : 1

FIGURE 111.13 : VIII NERVE INNERVATION OF HA IR CEllS O F THE ORGAN OF CORTI _ _ _ _ FIG l
Primary sensory axons of th e sp iral (coch lear) gan forces from movement of the fluid basilar mem Fluid
glion innervate inner and outer hair cells of the brane in re lation to the ri gid tectorial membran e. the h
organ of Corti, located on th e basilar membrane. This represents the complex transd uction process panll
Th e axons are activated by release of neurotrans of the converting external sound waves to action orgar
mitters from the hair celis, wh ich occurs when the potentials in spiral ganglion axo ns. MOVE
hairs o n the apical surface are moved by shearing ial mE
resul t:
226
~m5-AuditOliensory Systems-Auditory SYSTEMIC NEUROSCIENCE

Distort Reissner's
5. Short waves (high membrane and basilar
1. Sound waves frequency, high pitch) membrane of cochlear
impinge o n
ear drum,
,.K'''! base of cochlea duct and its contained
organ of corti, thus
.al rRej ~sner's) causing it stimulating hair
M 4. Sou nd waves
vibrate transmitted Long waves (low cells which are ill
up scala frequ ency, low pitch) contact with the
ochlear ducl 3. Stapes ves tibuli in at apex of cochlea tectoral memberall e.
"'Idolymph moves in medium or Impulses thell pass
~ mV) and out up cochlear Ilerve.
its contained
of oval perilymph
-- Spiral 2.0ssicles
vi brate as
ligament ,
- Tectorial
membrane

Spiralorgall
Corli)

"-'ilar
!embrane

cochleJ

8. Impact of wave
7. 6. Wave transmitted
on membrane of
scala tympani ac ross cochlear duel
round window
in medium of its in medium of endolymph.
causes it to move from scala vesti bul i to
contained
in and out at scala tympani. (Note:
perilymph
round window in wav~ may also travel
l:oN
";.',':: opposite phase to
around helicotrema
oval wi ndow
at apex of cochlea.)

- FIGU RE 111.14: COCHLEAR RECEPTORS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

,. Fluid movement through the scala vestibuli, around

the helicotrema, and ba ck through the scala tym
55 pani moves the basilar membrane on w hich the
organ of Corti and its hair cells reside differentially.
v\ovement of hairs by shearing forces of the tector
ial membrane on the apical portion of the hair cells
'esults in their depolarization and in the release of
neurotransmitters. This release stimulates action
potentials in the primary afferent axons of spiral
ganglion cells. Efferent axons from the olivo
cochlear bundle that ca n modulate the sensory
transduction process and the excitability of hair
cells are controlled by descending central auditory
pathways.
227
SYSTEMIC NEUROSCIENCE Sensory Systems- Auditor y

/ AcouSlic ilrea oi
lemporal lobe corlex

Medial geniculate body

Brachium of
inferior collkulus

Inienor colliculus--ff1.f
Midbrain

a nd acoustic a rea
oj corlex:
_ low tones
"~' ,~ ,, ~
L..' lerillremniSCi ~ _ Middle tones
High tones

Medulla oblongata ~

Superior o livary
comp~x
Dorsal cochlear nucle us
lnle rior cerebellar peduncle
,- - - Vemral COChle<l f nucleus
II / Cochlear division of
veslibulocochlear nerve

,/
'/ Intermediate
acoustic slria
I
Trape zoid body Iventral acoustic stria) .Ii
FIGURE 111.15: AFFERENT AUDITORY PATHWAYS _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
Ce ntral axon projections of th e sp iral ganglio n
neurons terminate in dorsal and ventral cochl ear
nuclei in several to notopic maps (shown in th e
cochlea in colors). These cochlear nuclei projec t
into the lateral lemniscus (LL), via acoustic stria,
but many projections remain ipsilateral. The LL ter
minates in the nucleus o f the inferior co lliculus. The
inferior colliculus, in turn, projects via its brachium
to the medial geniculate body (MGB) of the thala,
mus. The thalamus sends to notopic projectio ns to
228
~!p
I-!elicular
forma tion
Hai, ce lls
"':',':!UN
...'::
th e pri mary auditory co rtex on the transverse
gyru s of Hesch!. Th e accesso ry audito ry brain stem
nuclei (superior oli vary nucleus for lateral sound
localization, nuclei of the trapezoid body Ino t
shown I and the LL) send crossed and uncrossed
proj ections through the Ll. Because sound is repre
sented bilaterally, a unilateral lesio n in the LL, the
auditory thalamus, audito ry radiatio ns, or audi tory
cortex results in diminution in hearing and auditory
neglect contralateral to simultaneous stimulation.
,ditory Se nsory Systems-Auditory SYSTEMIC NEUROSCIENCE

Excitatory e ndings - -
Inhibitory e ndings -+ _ -+
Intermediate endings - "" ~

Temporal cortex Fib~(5 may be

'nf" ' ;"rcoH;cul,, , } eXCltdtory or
l inhibitory

",

Brachium
o f inferior
co llicu lus
Inferi o r
colliculus

Tympa ni c
memb rane

Nuclei of

lateral .~~~_ _ _--;:~ .

lemnisci - - - -"" -
Stapes Middle ear

(-l~~S~~~==t===T;:r~~:n:;n" ,oe"" fibers

n ucleus of tri geminal n.
::::::::::",,?J-;?'~l:- I nerve fibers
~-C'o ,,;, ' cochlear nucleu s
<.-.\-_-",""",1 cochlear nucleus
/ ':ff"'enl ol ivocochlear fibers (inhibit hair cell s
and afferent nerve term inals via coch lear
division of vestibulocochlear nerve)

)
fOfmalion
Tr,'lpezoid body
N
". " Superior olivary complex A"erenl nerve fibers

~IG URE 111.16: CENTRIFUGAL (EFFERENT) AUD ITORY PATHWAYS _ _ _ _ _ _ _ _ _ __

:Jescending pathways travel from the aud itory
iem cortex, the MGB of the thalamus, the inferior
j colliculus, and accessory auditory nuclei of the
'"train stem to caudal structures in the pathway to
j ~e r mj n a te in the cochl ea r nuclei and the superio r
pre olivary nu cleus. These centrifugal co nn ections
e oe rmit descending contro l of in comin g auditory
lr\ ,ionn atio n. Th e olivocochlea r bund le from the
Dr\
n.
superior olivary nuclei projects back to the hair
cells in the orga n of Corti to modulate the trans
duction process between the hair cells and the
primary afferent axons. The moto r nuclei of V and
V II send LMN axonal projec tions to the tensor
tympani and stapedius muscles, respectively, to
dampen refl exes of the ossicles in response to sus
tained loud noise.
229

SYSTEMIC NEUROSCIE NCE Sensory Syslems- Vestibu icr

A. Me mbrano us labyrinth B. Position within base of sku ll

Canals
Superior superior Po~tl!rior
Vestibular Plane o!
semicircular horiZOflld1
GllliJl a nd
V'fide
Ve stib ula r and
cochlear divisio ns
H ori zontal
o f ve5tibulo ' -' . ~---::-~
/....
cochlear n. ._~'-~
P1~nc
Cochlea r duct of sacc ule
, Plane of ' Svperior
superior canal ;C"!~'
Horh:onlal

("-..1 '

C. Section of crista
ampullae semicircular
canal
PlMleof
r
po51erio r canal
Po~tel i or

Opposite wall ~~
o f amp ulla _ _ _ _ E. Structure and innervation
of hair cells Excitation
Gelatinous cupula- - -

Hair tufts t I
Hair cell s::---
)1.1) iii,
Nerve fibers

Hair cell (type I)

~

l!2 .

. 1& ;lb:~ .Ulr.~

L
H air cell (rype II)

Supporting cells

O. Section of macula ~ -......---Efferen t nerve

Affelefll nerve
, endin g

~ Affe(ent nerve
Effe renl nerve

Basemenl
""'"
. /M yelin sheadl

~~!% CIIiiI\I
; ......; : :

FIGURE 11 1.17: VESTIBULAR RE CEPTO RS _

_ _ _ __ _ _ _ _ _ _ _ _ _ _ __

Th e ves tibular receptors includ e hair cell s in th e
crista e ampuliaris of the utricl e (lin ea r acceleration
or gravity) and saccule (low-frequency vibratio n)
and in th e maculae of the o rthogo nall y orien ted
semici rcular canals (angular acceleratio n or move
ment o f the head). Hair tufts fro m the cristae
ampullaris and the maculae are embedded in a
gelatinous substance, w hich is moved by gravity
(utricle) exerting force on the calcium ca rbonate
crystals (oto liths) resting on top of the hairs, or by
fluid in a se mici rcul ar canal (h ead movement).
Ben ding of the kinocilium in the hair tufts depolar
izes th e hair cell, causing the release of neurotrans
mitters that stimulate actio n po tentials of primary
sensory axons o f the vestibular (Scarpa's) ganglio n.
Additio nal efferent projections fro m the e NS mod
ulate this transduction process.

230
estibular Sensory Systems- Vestibular SYSTEM IC NEU ROSC IENCE

Vestibulospinal Tracts
7-+'----f'<~-;I
~ Su~~pe'~
: I';;O'}I Vt'stibular nudei
Excitatory endings - - - -
Inh ibi tor y endings _ _
Ros tral
Ascetldln8 'Ibers in medial
lo ngitud in al fascicul i _ _ _ _L-<1'
Tmnk

~
Ve ntral

Caudal
Somatolo jlica.1pa tte rn
In la teral vestibula, nucleus

.,

Motor neuron
(oontfOliing .,,' m"'"<"'''Jt:~
Med ia l ve stibulosp inal
fi bers in media l Lat era l Fibers irom c ristae
(fOt,ltional ~Ii mu lj)
Vffi..i bulot p inOiI

"'''

interneuron
Inhibitory
interneuron F iber~ from maculae
(gravitatio nal ~limull )

Lower ~== To flexor muscl es

e ll) - To eXlensor muscl es

Inhibitory ending
ells To axia l mu sdes - - -<1===:::3' To axial muscles
I il
""";',,,o ,yending
ending
Lumbar part of spinal co rd
"--l"'...' vestibu\ospi J'l all r.lcI
\

~=:: To flexof musd es

- To exten!oOf ml1Slei
J{p
tT 01lliN
, .: .. : 1

- FIGU RE 111.18: VESTIB ULAR PATHWAYS _ __ _ _ _ _ _ _ _ __

_ __ _ _ __
by Primary afferent vestibular axons fro m the vestib
ular ganglio n terminate in the superio r, inferior,
lar medial, and lateral vestibular nuclei and directly
ans in the cerebellum (deep nudei and cortex).
ry Descendi ng axo ns are sent fro m th e medial
Ion. vestibulospin al tract (medi al nucleus) to spinal cord
od LMN s to regul ate head and neck movements.
Descendin g axons are se nt from the lateral vestibu
lospinal tract (l ateral nucl eus) to all levels of spinal
cord LM Ns to activa te extensor mOvements.
M ultiple vestibular nuclei project to the cerebellum
to modulate and coordinate muscle activity for
basic to ne and posture (shown in figure 111045) and
to extraocular lMNs via the medial longitudinal
fascicu lus to coordinate eye movements w ith head
an d neck move ments . Some ascendin g axons
from the vestibular nucl ei may reach th e th alamu s
(nea r VPM and posteri or nucle i), with projection s
to th e lateral postcentral gyru s (area 2, mo ti o n per
ceptio n and spa tial orientati o n) and th e in sular and
temporo parietal co rti ces.
23 1

SYSTEMIC NEUROSCIENCE Sensory Systems-Vestibular

I' Slow phase " , Rapid phase (saccadic movement-----

~ ~
mainT;Jin~,~~e~~~~.~Ie'ation ~ .~~ maintain~~~I~raTiun ~.#
Ho, j2ontai
semicircular
:
/ _-
'~
~ ,~
\ Hori~onlal
.,_ semi(.ircuJar
Horizoolal
~('mK I(c.ular
vi \.. ;rP
H".ri~()nl"
_,c,' ("t-l. tJ.. ' "; seomlclrculJ"
(a na l excited canal dej)l'e~sW unil.' InpUT , r 1 1"-. -\. canal depres~e,c
conhnues,
but is
opposed by
Medial rectU $ inhibition Medial rectus motor
Medial r~I US nlOlor
moto r n~~;~~s ~V neurons depressed imm sat:oOO:: \ neuro ns exCited
exdted -------""
~
Abducens imernudeil' neuron
ce nle,
Medial
"
Ascending Abd ucen<
tr.:ad of _ _ _ _ In/1ibi torv Il Ucltc'Ui
Deirer5 intemeu(on~ depressed

Medi..!! Vestibular
nud ei bun,
and lateral
nudeo s depressed interneu ron
vestibu lar
nudei, e~cited exdted by saccadic :,t I
cen teo' ,~ I - Excitol tOI\
bur,;
Abdu cens Abduc ens interneuron
nucleus
depressed "'"''''''''--'''''''''''''f''' \-- Abducen~
Parapontine
(VI)
nerve
nucleus
e};cited by Oculomotor
reticular fo rmation saccadic (III) nerv~
(PPRF)
(III) Laler"
La teral
rectu,
recws
muscle
Lateral muscle Laler al
rectus
~
Eyes move in opposite direction to head; tend to prt!~t!rve visua l
iixiltion: rille determin ed by degr ee oi hOri20ntal canal exci tation
rec lus
muscle
---
Eyes sna p bac k in same direction as head
--- ~~p CJ I@N
:'<\"',:;

FIGURE 111.19: NYSTAGMUS

Nystagmus, the alternating back-and-forth move
ments of the eye, requi res central coordination
of extraocular LM Ns and eye movements.
O pto kinetic nys ta gmu s~ the v isually activated
movement o f the eyes via tracking mechanisms,
allows the eyes to return 10 a forw ard positio n by
visual association cortex projections through the
superior col liculus to extraocular LMNs. Vestibular
nystagmus involves vestibular projections via the
medial lo ngitudinal fasciculus to extraocular nuclei
(LMNs). The slow phase (or drih) of vestibular
nystagmus results in the provocation of eye move
ments as if the head were turning. It is caused by
asymmetrical input fro m receptors of the semi
circular canals o r from damage to vestibular nuclei
o r the vestibular cerebellum. The fast phase
(saccadic movement) is the provoked return to
a forward pOSitio n.

232
:stibular ilenso ry Systems- Visual SYSTEMIC NEUROSCIENCE

Ho rizontal sect ion

Scleralvenous sinus ~"""" of lens
(Schlemm's /Cmo"
c hamber

,C'' '''I processes

--,
Ciliary parI o' ,e."",_,
-eldon o f lateral
orizonlaJ ,Te" do, of
rectus muscle
r.icircula; media l rectus
epressed muscle

IS motor
e.~ ciled

duc:en~
nu cleu~
lre.sed

_bilOry
burst
~e l.Jr O I1

ta ror)'
b ur~ 1
~ur OI1

O ptic (visual)
..ller,,1 part
V itr eous body
re<:lu<
Choroid
'lUsde
Hyaloid canal
Perichoroidal m,,-p/

Sclera ' -_ _ Lamina cribrosa of sclera

i!jN
.... ,' fascial sheath of eyeball ~Vp"i, nerve (II)
(Tenon's ,;,,,,,'el/
/ ':e,",,' relinal artery and vein
Fovea centralis "0 m" u';
O uter sheath o i optic oe,ve -"--
Subarachno id '~' CP'/

FI GURE 111 .20: ANATOMY OF THE EYE _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

clei --"ere are 3 major layers of the eye. The outer layer, epithelium of the posterior iris. The retina contains
;:he fibro us tunic, consists of the protective cornea the photoreceptors for transduction of photon
\ e '"ansparentJ and sclera (opaque). The middle layer, en ergy into neuronal activity. Aqueous humor is
...., ~ vascu lar tunic (uveal tract), cons ists of the secreted fro m blood vessels of th e iris into the
"'(Koid the Ciliary body, and the iris. The trans- posterior chamber and flows through the pupil into
.p ' biconvexlens surroundedbya capsule of the dnterior chambet; where it is absorbed into
P::' /5 suspended!/om tile Ciliary process the trabecularmeshwork //}/O Scillemm s canalat
~. me 1/717er layer, Me laterlldl tile Ii7dOcornealangie. Claucoma results when
"'t:> neuroretli7a, the nonpigment absorption ofaqueous humor is blocked Vitreous
a..an body, and the pigment humor fills the interior of the eyeball.
233
SYSTEMIC NEUROSCIENCE Sensory Systems-Visual

.Posterior ~miling I~min a (Dcscemel's membrane)

TrOlbewlar meshwork ;mo spaces Endothelium of an terior cham ber

o f iridOCOIfleal an gle IFonlanaJ Cornea
St;ieral vt'nou~ ~inus (5chlemm's I
Sc.Ieral spur

Pectinate ligamen t,
........ jnr arte rial circle of

"nlerior ciliary I'ei~

~. Anlerior chamber
~FOldsof iriS

M inor <llterial
d lcle of iris

Ciliary
pr0ce5S
Merid iona l Circ ul ar
fibers fibers

-
.... ./
~

( ili.ny muscle
Nucleus of len$

,.
Zonular fibers
Pigment epithelium
Capsule of l en~
G liill)' body
~I ichnroinal spiKe Ug.tment of le nSI iiridial part o f retinol) Sphincter muscle of pupil

Notl:!: Fur daril),. Ollly single plane of zo nular fiben sh own; aau.l l ~. fibcf"s surround entire cirOJmference of lenr;

FIGURE 111.21 : ANTER IOR AND POSTERIOR CHAMBERS OF THE EYE _ _ _ _ _ _ _ __ I

The ciliary mu scle and the pupillary co nstri ctor (crossed axons through the po sterior co mmissure)
mu scle are supplied by parasympatheti c post to the Edinger-Westphal nucleus. Thi s nucl eus
ganglionic M nerve fibers from the cili ary gangli on projects to the ciliary gangli on via e N III (effe rent
(preganglionics in the Edin gerWestphalnucieus, limb), whi ch res ults in both direct (ipsilateral) and a
axons in CN III). Ciliary muscle contractio n reduces consensual (contralateral) pupillary constriction. a
the tension on zonular fibers and ca uses the lens to The pupillary dilator muscle is supplied by sympa o
curve, w hich induces accommodation for near thet;c postganglio nic UNM nerve fibers from the n
vision. In the pupillary light reflex, light shone in superio r cervical ganglion (preganglio nics in Tl C
one eye en ters the CNS via the optic nerve (CN II) and T2). a
(afferent limb) and terminates in th e pretectum . M = myelinated; UNM = unmye linated. a
euro ns of the pretectum project bilaterally If

234
Visual Se nsory Systems-Visual SYSTEMIC NEUROSCIENCE

Re tinal lay!1"5 Cells

Visual R ~ce ptors
Nerve Iber l r Jnner limiting
A.. Eyeball me mbratl('
~uriJce oi

Inner plexiform
w:niculale body
Llyer

Inner nuclear
layer
II I cell "
i5uppor lill!; ~d i a l ce ll )
IJipo),,.. ce ll
Amelre ine cell
Oul~r plexiiOlm
fa\"ef

OUler nvdeal
layer i I U'il~
choroId
PhOIOIKcplo r
I,,.,
Pijtll)Cf1 t B. Section
, ep'lhO!liu m th rOug h retina

Rod Photoreceptor Cone photoreceptor

I Outer
plexiiorm
layer

O. Rod in liSh'

Nuc levs

hiller
segmef1 1

!CURE 111.22: THE RETI NA AND THE PHOTORECEPTORS

_ _ __ _ _ _ _ _ _ __ _
-~e retina, a tissue-thin piece of eNS containing
-.e photoreceptors, is attached to the vascular
c at th e ora se rrata. The o uter segments (rods
cones) of th e ph oto receptors are embedded in
. ment epithelium to prevent backscatter in the
~ er part of the retina. The rods and cones con
l synap tica lly with bipolar cells, which in turn
nect with the ganglion ce ll s of the retin a (equ iv
t of secondary sensory nuclei). Horizon tal and
;.acrine cells provide ho rizontal interco nnections
....e retina. In the macul a {3 mm in diameter), th e
fovea centralis (0.4 mm in diameter), the central
point for visual focusing, consists of cones for color
visio n (p hotop ic), w hich project, with very little
conve rgence, to ganglion ce lls. The periphera l
retinal pho to receptors, mainly rods for night vision
(scoto pic), project, \-vith great convergence, to
ganglio n ce ll s. Rod light tran sdu ction invo lves
co nve rsio n of II -cis-retinal (fro m rhodopsin) to
an all-trans form, provoking calcium influx w ith
hyperpolari za tion and a decrease in sodium
co nductance.
235
SYSTEMIC NEUROSCIENCE
Corn i I ~----""
A nterior chamber ~ ..
Scleral venous sinus
ISchlemm's 1.:<1( 011)
Iridocorncal angle
IlenS
Vitreous t:h,Jmhel
J r Lo ng I)Osterior ciliary artery
p t - Shorl posletiof cili ary arll!ries
Central refinal ar{~
({Ii~ .
"'-'--''-- OPl ic ner ve (II)
Superior tempor...1retinal
arte rio le and venule - -
Macula and iovea centralis
Inferior temporal (etinal ____
Jf\eriole and venule - - - -
Right retin al vessels: ophthalmoscopic vi ew
FIG U RE 111 .23: ARTERIES AND VEINS OF THE EYE _ _ _ _ _ _ _ _ _ _ _ _ _ __
The ce ntra! retina! artery and its branches supply
blood to the retina. Thi s arterial system, derived
from the ophthalmic artery (the first branch o ff the
interna l carotid artery), is often the site w here
ischemic or embolic events (transient ischemic
attacks) herald the presence of serious vascular
disease. Cili ary arteries supply the middle vascular
236
Sensory Systems- Visual
Minor anerial circle of iris
Major arlE'fial circle of iris
Blood vessels of ciliilry body
Bulbar conjunctiva and conjuncliv<ll vessels
Anteriur c iliary artery <lnd vein
iii
Ora serrata
MUKular artcry Clnd vein
Ex trin~ic eye- musd~
~
,. Long po sterior ciliary ar tery
H--- Vorticose vein
, - - - . Episclera l artery a nd vein
" Re linal a rlery and vein
and vein / Superior macular arteriole
. venule
Superior n3sill ret inal
ar teriole and venule
Optic d isc
~ I rl ferior rl;lSJJretinal
ar teriole and venule
~J~
Inferior macular
arteriole and venule
n...::
1!!iN1
tuni c, w hich also contribu tes blood to the retina.
Blood supply to the retina can be disrupted if the
retina becomes detached. Vessels enter and exit
the retina at the optic disk (nerve head), loca ted
nasally and slightly inferiorly fro m the geometric
midpo int of the eyeball. The macula is located terr C(
porally and slightty inferiorly from this midpoint. kl
rl<
~ .
<
-,
~

ls-Visu a ..ory Systems- Visual SYSTEMIC NEUROSCIENCE

To visual cOflex
From visua l cortex
_ _- To Vi5Uili cortex

Supra chiasmatic _ _ Pulvin ar

nucleus - - -____

sUp r ior colliculus

Nucleus of accessory
optic tract

l a le ral geniculate

body _ _ _- '

Pontine

tegmen tal reticul,ir

nucleus

Infeliorolive -------~l_

To pleg<lngJionic

sympathetic neurons

(ll - T2) that project to

the superior cervical ganglion
and regula te melalonii""~"'C
from the p ineal gland -
~:,e~l~iO: ,p
"_ _ _ _ _

!;N
; . ::

a.
FI GU RE 111.24: VISUAL PATHWAYS: RETI NAL PROJECTIONS TO THE THALAMUS AND
THE BRAIN STEM _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

'e ::eti nal projections travel through th e optic nerve,
it 21iasm, and tract and terminate in several regions.
j
-;he lateral geniculate body mediates conscious
nterpretation of visual input. The superior
(olliculu s, a seco nd pathway through the pu lvinar
m th e associa ti ve visual cortex, provid es loca li zing
n iormation for visu al stimu li. It also provid es
desce nding co ntralatera l connections {tectospina l
'~ac t} to cervical LMNs to mediate reflex visual
erieds o n head and neck movements. The
pretectum mediates th e pupillary light reflex. The
suprachiasmatic nucleus of the hypoth alamu s inte
grates light flux and regulates circadian rhyth ms
and diurnal cycles. The nucleus of the inferio r
accesso ry op ti c tract may help to mediate brain
stem responses for visual tracking and may inter
co nn ect with sympath eti c preganglio ni c neurons
in T1 and T2 (regu latin g the superi or cervical
gangli on).
237
SYSTEMIC NEUROSCIENCE Sensory Systems- Visu al

Centra l darker eirde

represents macular .:co ne

O verlapping Lighte51shades repr~n l

vi'S...,,[ fi elds , monocular fields

Each qu adrant
a different co lor

Projec tion on Projection on

leli l f lin.:! right re tina

Projection on left Projection on right

dmsa[ lateral do rsalla!"",,1
M eyer's Meyer's geniculate nucleus
Ipsila gemculate nucleus Ipsilateral
, lehll

loop loop

: --::- (;:.-"- --,

:.; + s ,, "
'
,,--------- ' ' ,, """ " " ~
' .40
, L-~ _
6'"
,..",.
_"'_. ;"::,,"i,", .
I, :,jr. 4 '
,, .... .. I " ."
t ::. .: .... :;
, !. _____ . : 2 , ,
L ;I . .
,,,
, 2
1 .. -,.-; I
-) ';: ~
, "
, '

."_,,,"l Contralateral Co ntralateral

,
Projection on 1011
occipita l Projec tio n on right
lube
.,. - occipital
lobe
Calcarine

FIGURE 111.25: VISUAL PATHWAYS: THE RET INOGENICULOCALCARINE PATHWAY

_ _ __
Th is visual pathw ay co nveys info rmatio n fo r fi ne.
grained conscious visual analys is of the o utside
world. It is organi zed topographically (retinotopic)
th rough its course to the calcarine cortex in the
occipital lobe. The nasal hemiretinal gangli o n cell
axons cro ss the midline in the optic chiasm,
whereas the temporal hemiretinal ganglion cell
axons remain ipSilateral. Thus, each optic tract con
veys information from the contralateral visual field,
and damage produ ~es a contralateral hemian o pia.
238
The op tic tracts terminate in the lateral gen iculate
body o r nucleus, w hich is organized in 6 layers.
The optic radiatio ns project to the calcarine (striate
cortex (area 17). The portion of the optiC radiations
that loops throu gh the tempo ral lobe (Meye(s
loop) can be damaged by a tumor o r mass; such
damage results in a contralateral upper quadran
tanopia (quadrant defici t). Bilateral convergence
from the right and left retinas takes place first in the
primary visual co rtex, area 17.
 )elso ry Systems-Visual SYSTEMIC NEUROSCIENCE

Parietal lobe Spa tia l vis ua l pathway: p<lsitional

relationship between objects in
vis ual scene, analysis oi motion

Fron tal lobe

, I
direction
motion responsive

Occipital
lobe

shape and color

percep tion
Temporallobe

,
Obje<:1 recognilion p athway:
high r~olu tion and form

' IGURE 111.26: VISUAL PATHWAYS IN THE PAR IETA L AND TEMPORAL LOBES
_ _ __ _ __
' eurons in the primary visual cortex (V1, area 17)
send axons to the association visual cortex (V2 and
.'3, areas 18 and 19). V2 and V3 also receive input
~'o m the superior callieulus via the pulvinar. Vl ,
'2, and V 3 project to th e middle temp oral area
.and V4. Middle temporal neurons are directi on
~ electiv e and motion respo nsive and furth er project
.:lto the parietal lobe for spatial visual processing.
Th e parietal neurons provide analysis of motion
and of positio nal relatio nships between objects
in the visual field. V4 neurons are involved in
shape and color perceptio n. V4 projects in to the
temporal lobe, in wh ich neurons provide hi gh
reso lu tion object recognition, inclu ding faces,
an imate objects, and classification and o rientation
of objects.

239

SYSTEMIC NEUROSCIENCE Motor Systems-lower Molor Neurons (LM

Oculomolor \III) n.

gji
Q: .;I Red nucleus

" ,
~ QculomolOr n ucleu~

:"-'- - - Trochlear nut .<-:.

'--- - -- - -- Trochlear (IV f"

Tri geminal M n. %;, _ Trigemin.-.' (V "

and ganglion and jjaflgl ion
~ MOlor n ucleL;

Facial (V II) n.
..

-- trigeminal n.
----:;;:-- Abducens nu
;;:
Faci<ll (VII) n,

~ Fada l nucleus

Clossophar\
Glossopharyngeal (IX) n. ----i !IX) n.

Vagus (X) n.
Vagus (Xl n. - - - - '
'-----'-:::.- - ACf;essOty (XI) n.

"Hypoglossal (X II) nucleus

~ Spinal nucleus of accessory n.

i - - -- Spinal cord vel1 tral horn (at all spina l levels)

FIGURE 11 1.27: LOWER MOTOR NEURON DISTRIBUTION IN THE SPINAL CORD

AND THE BRAIN STEM _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Lower motor neurons (l MNs) are found in the
ventral (an terior) horn at all levels of the spinal cord
and in motor cranial nerve nuclei in the medulla,
pons, and the midbrain. These l MNs send axons
into the ventral root and then in to spinal nerves
from the spinal cord ventral horn, and into cranial
nerves from the brain stem. The LMN axons ter
minate as neuromuscu lar junctions on skeletal
muscle fibers.

240
 :o r Systems-lower Motor Neurons (lMNs) SYSTEMIC NEUROSCIENCE

1<1 <liferent
synilpse
on cr -LM N

:>erkins
06, ' IF'"

Iri':i

Ne uro musculdr
junctions on ex!r aiu ~~ 1
ske letill m uscle fibt-rs

Trail and plate endings

on contractile elements
(intraiusal fibers) oi
muscl e spindles

la afferent
Skeletal muscle fibers
(extrafusal) Nuclea r bd~ '"'''-+ --ffl" nerve

Nuclear ch<l in , ,, ,,-tlii jl~

Muscle spindle

FlGURE 111.28: ALPHA AND GAMMA LOWER MOTOR N EURONS _ _ _ _ _ _ _ _ __

~ LMN groups, except the facial nerve nucleus
"'13t supplies the muscles of facial expression, con
tim both alpha motor neurons (a~LMNs) that sup~
- , th e skeletal muscle fibers (extrafusal fibers) and
:-2lll ma motor neurons (y-LMNs) that supply the
;;;m all contractile elements of muscle spindles (intra
J5al fibers). The muscles of facial expression do
not have muscle spindles and are not supplied by
"tLMNs. The a-LMNs regulate contraction of the
skeletal muscles to produce movement. The
"tLMNs regulate the sensitivity of the muscle
spindles for la and group II afferent modulation
of a-LMN excitability.
241

SYSTEM IC NEUROSCIENCE Motor Systems- Upper Motor Neurons (UMNs

Cerebral Cortex: Effere nt Pathways

From {mntal cortex to

tha laml)s, ba ~a l g~ngli a.

pontine nucl ei and

reticular form<l lion

Corticobulbar, cortioorub.al,

cortlconuciear, an d co rt icos pinal

parietal (orte~ to thalamus, pon lin e

pathways --~"""'r nuclei an d reticular iormation

c..uda le nucleus .- -- -~
-.;::
r ~i - Posterior limb o f irllernal capsule
A.nte-rior limb of internal ca psule - - - ,
~~i'!' From occipital eye fi elds to ~uperior co lli c ulu~
Lentiform J Glob us pa lli du s
nucleus
';111 1_ From audilory cortex to in ferior colliculus

From fron tal eye liekl~ 10 ?~

~
." .
Sup",nor coJliculi

in terstitial nucleus of Cajal ~,t -.,.,___ In~erstilial nucleus of Caial

Inlerior co lli CLJ lu~
Ce rebral peduncle
Red nucleus

CorticuS{)inal
Corticonucleaf formation

For pontocerebellar connections

Facial nerve nucleus of oppoSIt e ambiguLJs

side for lower fa ce

Upp er (MI l o f medu lla oblongata I nudeus

(ormation
? yramkl s

ile nucleus
Lower p.l rt o f medu lla oblonttala
Re ticular formation
Derussation o f py.-amids
r- l ateral (cro ssed) cortic ospin allract

Anterior (d irect) co rticosp inal tract

~t$-t ~ Posterior (dorsal) hom

Spinal cord
Anteri or (ven tral) horn s
~~ft%
lrnN",'."."/' l

FIGURE 111.29: CORTICAL EFFERENT PATHWAYS _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Neurons in the mo tor cortex (area 4) and the sup
plemen tal and premOlar cortices (area 6) send
axons to the basal gangli a (caudate nucleu s and
putamen), the thalamus (ventroa nterior IVAl and
ventrolateral [VLJ nucleO, the red nucleus, pontine
nuclei, moto r cranial nerve nucl ei o n both sides,
and th e spina l cord ventral horn mainly on the con
tralateral side. These axons form the corticospinal
tract, the corticobulbar tract, corticostriate projec
tions, corticopontine proj ection s, corticoth alamic
projections, and cortical co nnectio ns to upper
mo tor neuro ns (UM Ns) o f the bra in stem (reticular
formation IRFJ, red IlUdeus, superior co11iculu sl.
242
Neurons of the sensory co rtex (areas 3, ' ,2) send
axons mainly to secondary sen so ry nuclei to reg u
late in coming lem niscal sensory projections des-
tined for conscious interpretatio n. Neurons in the
frontal eye fields (area 8) project to the superior
co lli culus, the horizontal and vertica l gaze centers
of the brain stem, and th e intersti tia! nucleus of
Caja/ to coordinate volun tary eye movements and
associated head moveme nts. Other region s of the
sensory cortex project axo ns to th alam ic and brain
stem structures that regu late incoming lemniscal
sensory info rmatio n.
 to r Systems-Upper Motor Neurons (UMNs) SYSTEMIC NEUROSCIENCE

f>rimary mOlor
co rteK (Are... 4)

Lateral ilspecl oi ceteb,a l cortex 10 show

topogtaphic proJec1io n oi motor (enU~fS
o n precentral gyfUS and puomulor and
supple mental molor cortex

larynx ';;;t

Midbrain ' \
/ til

IV )0 III
111 Upsilateral and conjralilteral)

\)si, . IV IV l ip~ ila l c ra l Jnd contralateral)

VI \1 VI Iip<;ilMeral and oonlralateral)

Pons V V V(ipsilJ lefal and amtralateral)

U""".
lower
VII 10 uppt'! lace (ip~larernl a nd contralatera l)
VII 10 lower iace (contr<llateJa l only)
VII VII

XII (ipsil ate ral a nd contralateral)

IX X. and XI ~~p
..
(psilateral and contralateral)
IX,X,XI LX. X, XI eiil
FIGURE 111-30: CORTICOBUlBAR TRACT _ __ _ _ _ _ __ _ _ _ __ __ __

o The corticobul bar tract (CBT) arises mainly from
the lateral porti on of the primary motor co rtex
area 4). CBT axons project through th e gen u of the
internal capsule into the cerebral peduncle, the
basis ponti s, and the medullary pyramids o n the
ipsilateral side. The axons distribute to mo tor
cranial nerve nuclei on the ipsilateral and contra
j
, lateral sides, e)(cept for the portion of the facial
nerve (eN VII) nucleus that supplies the muscles of
facia l expressio n for the lower face, w hich receive
contralateral prOjections exclu sively. CST projec
tion s to th e hypoglossal nucl eus are mainly
contralateral; projections to th e spinal accessory
nucl eus are mainly ipsilateral. In con tra st to Bell's
palsy (CN VII palsy), in which th e entire ipsilateral
face is pa ralyzed, CST lesions result mainly in a
contrala teral drooping low er (ace that is paretic to
volun tary co mmands (central facia l palsy).

243

SYSTEMIC NEUROSCIENCE Motor Systems-Upper Motor Neurons (UM" ~

Prima ry motor
cor1ex (Area 4)

l ateral aspect 0 1cerebral (Orlex showi n.

topograph ic lucali zation o t" mOlo r een le",
on prece ntral gyrus arid premo tor .l r :,
Posterior
'~ >~
~ w
supplemental moto r ((Jfle-.
limb
"%.,
Intt'rnal
Po. lerior
~..,....... ~.
ca psul e /
Anl erior ~ ,~
' __I Visual .md audito:.
lim b
'- ,.~
__
~
Temp oropontU1e
,:?{' ~:;" Ho rizontal set:tion Sensory
Midbrain Corl icospmal

"-/fA:~
Itnough internal capsule
show ing location of
Co rlicobulba r
principal palhw av5
f rnnloponline
; ~~)
.-.-. ,~ 1' Fron tolna lamic
Pons
) Anterior

Ve ntrilJ aspe-c -
brai n stem shon .,...:;;
decuSS31io r
pyl a !T1~

Ir
'
tI\"' ~ l _""~ """

~ "-i'!.~
fiE- ,d Decussation of pyramids
(approxitTlalety 80% of CST fibers)

Spinal .1" -, \ Lateral (crow~dJ

J ~t;~
cord
",. . ", corticospin al tracl tT l ,
,,;:..
~~:,~/ Anle.i()f (direct) corlicospinal lr,}ct .:>~

FIG URE 111.31 : CO RTI COS PI NAL TRACT _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Th e motor portion of th e co rticosp inal tract (CST) pyramid on the ipSilateral side. Most CST axom
originate s fro m neurons of many sizes mainly from decussate in the decussation of the pyramids a:
the primary motor cortex (a rea 4) and the supple meduUary-spinal cord junctio n and descend in
mental and premo tor cortices (area 6). The primary lateral CST to synapse o n a-LMNs and y.LM i\ 's
sensory cortex (areas 3, 1, 2) contributes axons to both directly and indirectly through interneur()(i""
the CST tha t terminate mainly in secondary sensory CST axons th at do not decussate continue as . ~
nucle i to regulate in comin g lemnisca l sensory infor anteri or CST and th en decu ssa te at the appropn&:
mation processing. The CST travels through the level to terminate direc tly and indirectly o~ L , ',
posterior limb of the intern al cap sule, th e cerebral contralateral to the cell s of o ri gin.
ped uncle, th e basi s po ntis, and the medullary
244
IU MNs)
-'olor Systems-Upper Motor Neurons (UMNs) SYSTEMIC NEUROSCIENCE

I.-. ing
fllNS
rand
!lfll."\
....... eral
. : "llc05pina l - - - - - : - - -
'"><,

'i

Minor component of
1""""",- - - - - ---,1-- """,;OC corticospinJI
tract (ipsilatera l)

Anterior An terior J. Perkins

white corti cospinal MS.MF'\

-=dIal muscles commissure tracl .~

- URE 111.32: CORTICOSPINAL TRACT TERMINATIONS IN THE SPINAL CORD

_ _ _ _ __
~ ~ed CST axons in the lateral CST, intermixed
:- axons of the rubrospi nal tract, travel in the
ear j'unicul us. Most of these CST axons termi
2 di rectly and indirectly on LMNs associated
- distal musculature, especially for skilled hand
ringer move ments. The uncrossed anterior CST
~ decussate predominantly in the anterior
:e commissure and terminate directly and indi
mainly o n LMNs that supply medial muscula
- .; small number of anterior CST axons
termina te ipsilateral to the cortical cells of origin.
An isolated lesion of the CST in the medullary pyra
mids results in weakness of contralateral fine hand
and finger movements. All lesions involving the
CST intermixed w ith other descending mo to r sys
tems (internal capsule, cerebral peduncle, po ns)
produce contralateral spastic hemiplegia w ith
hypertonia, hyperreflexia, and a plantar extensor
response. Lesio ns of the lateral CST produce similar
symptoms ipsilaterally below the level of the lesio n.
245
SYSTEMIC NEUROSCIENCE Motor Systems-Upper Motor Neurons (UMNs)

"b",p'",1 T"
Pnmary rnocttor
cortex (area 4)
<4f~~~ ,
} ~ ~ ~~!il- I, , 7 ""' ...
'm,IIpy,.m'd, ' oell, ~\ ~
) . '~
Giant pyrarnidil l ceils V;c - r

Fibers from glo bose and

emboli/arm, ~ m e from

dentate dee p nvdei via

,"pe"~co,,bell"pe""oc~ ~

Red nuclei ~'.. . ~ Helps 10 hold flexor actions of

RST on UE in check
Helps \0 drive Bexor actions of
Crossed robrospinal and RSTfOl l E
rubromoou1lary fibers - ~
Mid brain

To pyramid

Fildal nerve nudeus

nI,... Ul"ICrossed rubr om~u ll ary
Irubr obulbar) fi bers

Medu ll a oblongate
Lateral reticular nucleus
M edullary reticul ;lr
formation

Ru brospinal lraC! -----------.Ji Inferior olivary rlUcieU5

Inhibitory Interneuron '~_~ Cervical part of spinal cord

\
;-.. l ,l(eral (crosse d )
ExcitaTo ry i nterneuron corticospinal lraC I
Rubros pinai lraci

Pos terior (dor sal) ho rn

!Iltemeurons controlling
afferent input 10 spinal cord
~

-,
To extensor ,_ . lUmbar pall of spinal cord
mu scles
~ ,J ,\ -I"hibi ,o", iote,~",on
~~p
To flexor
muscles
, _O? 1( ) Excil.atory interneuron
01lDl\l
;,',,-:;;

FIGURE 111.33: RUBROSPINAL TRACT _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ __

Th e red nucleus receives ipsil atera l topograph ical
input from the motor cortex and fun ctio ns as an
indirect corticospinal system. Axons of the
rubrospinal tract (RST) decussate in the ventral
tegmental decussa tion and descend in the lateral
brain stem and the lateral funiculus of the spinal
cord, w here they are intermixed ex tensive ly with
axons of the lateral CST. The RST terminates
directly and indirectly on a,-LMNs and y-lMNs in
the spinal co rd, particularly those associated with
flexor movements of the extremities. Th e RST helps
246
to drive fl exor movements fo r th e lowe r extremity
but helps to hold flexor move men ts in check for
the upper extremity, RST lesions usually occur in
conjunction w ith the CST in the spinal cord; corti
corubral lesions also occur in conjunctio n w ith
the CST in the internal capsule and the cerebral
peduncle, These lesions result in contralateral
spastic hemiplegia. Brain stem lesions ca udal to
the red nucleus result in decerebration (extensor
spasticity).
 SYSTEMIC NEUROSCIENCE
-
MNs) Molor Systems-Upper Motor Neurons (UMNs)

h:citatory
endings - - - V~s tibu lolr nuclei
Inhi bitory
~ ndings ..... _ Rostral

Upper lirl1, ;X Trunk

Ve ntr.,1 ./:" Oors ,,1

cerebellum t; l ower limb

Caudal
Soma totopic..'
pattern in lateral
ve stibu lar nucleus
!\ Ves tibu lar

MOlo<"e~l~~'"

:, ganglion
,"d
(cont rolling _ (\
neck muscles) ~

, Fib ers from cristae

(rotatio nal sti muli )
tract

Excita tory
endings 10
Interneu ron Fibers from macula e
(gravitati onal stimuli)

l ower pari of
cervica l spinal ~=:: T;:O l1ex(I( muscles
:... ~)(tensor muscles
cord

t~-l~~:'T;:O::;Uial mu~cles
,,,'''''''' ending
Lum b<lr p ari o f '+-'''"''''1 vestibul O$ pinal tract
spinal co rd
-........,., Inhibitory iniern eulOn
{

~--'TO
' - --
H
exorm
uscles
To .,," "'" muscles

F CU RE 111.3 4: VESTIBULOSPINAL TRACTS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

-"e la teral vestibulospinal tract (LVST) arises from
e lateral vestibular nucleus (LVN ) and terminates
ectly and indirectly o n ipsilateral a-LMNs and
LMNs associated w ith extensor musculature,
especially proximal mu scu lature. If thi s powerful
antigravity extenso r system were not kept in check
by descendin g co nn ecti ons fro m the red nucl eus
and the cerebellum, it wo uld prod uce a co nstant
state o f extensor hypertonia. The medial vestibu
lospinal tract (MVST) arises from the medial
vestibular nucleus (MVN ) to provide inhibiti on of
the a-LMN s and the y-LM N s that control neck and
axial musculature. The MVST terminates mainly o n
interneuro ns in the cervical spinal cord ventral
ho rn. Th ese 2 ves tibulospinal tracts stabilize and
coo rdin ate th e pos iti o n of the head, th e neck, and
th e body and provide important refl ex and brain
ste m co ntrol ove r tone and posture. The vestibu
lospinal tracts work w ith the reticulospinal tra cts to
con trol tone and posture.

247
SYSTEMIC NEUROSCIENCE
Re ticulospinal and
Corlico~tjcuJar Pathways
Ex citato ry
endings
---
Inhibitory
endings
--
MecJial pon tine
relicular fo rma tion ________ f.
pon s ~
Receives input
from multiple
,en$ury S~' stCIllS
via laleral RF
Medial
medullary
relicul
form ation -
Medulla oblonga(J
l ateral reticulo-spina l
Ira c t (pa rt ially ClOSSed );
excites ~nd inhibits
axial (neck and back)
motor neurons and
moclula l ~ affe rent
inpu t 10 sp ina l cord
M otor neurrlnS
(alpha and g<l mma)
FIGURE 111.35 : RETICULOSPINAl AND CORTICORETICUlAR PATHWAYS _ _ _ _ _ _ __
The pontine reticulospinal tract (RetST) arises from
neurons of th e medial pontine RF (nu clei po nti s
caudalis an d orali s). Axon s descend as th e po ntine
(medial) RetST mainly ipSi latera lly and terminate
directly and indirectly o n a-LMNs and y-LMNs at all
levels. This tract has a distinct extensor bias w ith
axial musculature, and it rein forces the action of
th e lVST. Th e ce rebral cortex exerts minimal influ
ence on the pontine RetST; it is driven by po ly
sensory input from trigeminal and somatosensory
248
Motor Systems- Upper Motor Neurons (UMNs)
p,';e,,1
T nickness o f blu~ l ine.
indi ca! i!s deo~ty ot
Frontal ____/" .1:....'"
~~
r
cortica l proj ec tion { O rb ito
ironlal
Tri geminal mOlor nucleus
. 1
,. "'-----"
Receive excitatory
fibers irom pontine,
inhibito ry fi bers
from medullary
reticular formation
~
k I
,
""'"
F"cial nerve nucl eus
Exerts slro ng d rive
over medullary RST
"it ":r~~l M ecJiill (anterio r)
ret iculospinal
-.. trac ti I)roduces
@'* direct t:.'(ci ta tion
.., ~ 1r:% oi motor neurons
-,-- - - Ce-rvical part
oi spinal cord
-:>--- Pos te-rim (dorsa l)
horn inte-rneurons
re-gulating se nsory
inpu t to spinal cord
( I _____ Excitatory
~ inte-rneuron
};':. I , Inh hibito ry
~~p
Interne-uron
Lumb.,r part of
spinal corr!
OII9N
\\'0:::
sources. The medullary RetST o riginates from the
medial RF (nu cleus gigantoce llularisl and is heavily
driven by corti cal input, especially from the motor
and pre molor/supplementa l motor cortices. Axons
o f the medullary (lateral) RetST terminate bilaterally,
directly and indirectly, o n a-LMNs and y-LMNs at
all levels. The medullary RetST exe rts a flexor bias,
reiniorcing the CST and the RST. The reti cu lospi nal
tracts are important regulato rs of basic tone and
posture. They are no t organi zed somatotopically.
 '-----
'UMNs) Motor Syslems-Upper Motor Neurons (UMNs) SYSTEM IC NEUROSCIENCE

Superior colliculus

Midbrain
;f

"'II.
------
!
Interstitial nucleus of Cajal

11\ ./'
Pons
Itt
1------- Mediallo ngiludillal fasciculus

"
".,..,..- '"
I\. :p .. - ~

/ .,...- v,.. ...,~

Upp" """,'
spmal cord ) /
(~f \ :"\
j
I ~....'-\ '\ To axial muscles
~. . . ---.-
To neck _ _ _;;_:::::::;;;;::::::;.::.::
--- ._ _ .::~_ _ ;:-:::_
.ptf
,:::::.~~;:;.
.. -:~._
_ ~_.l'
.::::;::;:;::-::
__ .~;::; ::. ::
...~__.,
..:......_ _ _
of the trunk fo r
rotation al
musculature ,;. movement

~~~
J. Perkins
~"- M,M.FA

'\ 0 10N
." :::::

'\ \
iii\!
. .'. :; . =__ ,! ......-. . ...
.~,,:: _:::~:::~;;
.--~
- _.~:;:~
. _:;.~.:....
J. \.
........
___ To axial muscles

of the trunk for

ro tational
movemen t

e
FIGURE 111.36: TECTOSPINAL TRACT AND INTERSTITIOSPINAl TRACT _ _ _ _ _ _ _ __
The tectos pinal tract (r ST) arises from neuron s in
.ilv deep layers of th e superior colli cui us, decussates in
or the dorsal tegmenta! decussation, descends con
}n s tralaterally near the midline, and terminates directly
aliI'. and indirectly o n a -LMNs and y-LMN s in the cervi
cal sp in al co rd associated with head and neck
s, movements. This pathway mediates reflex and
.al visual tracking infiuences for positi oni ng the head
with regard to visual input. The interstiti ospinal
tract (1ST) arises fro m the interstitial nucleus of
Cajal, a region that helps to coordinate eye move
ments and gaze centers. The 1ST descends ip si
laterally in the medial lo ngitudin al fascicu lu s (MLF)
and terminates directly and indirectly on a-LM Ns
and y-LMNs associated with axial muscu lature of
the trunk involved in rotatio nal movement.

249
SYSTEMIC NEUROSCIENCE Motor Systems-Upper Motor N eurons ( UM I\ ~

A. Corticospinal tracts B. Rubrospinal tract

.-- -.-~-", -:- r""",

l ate ra l (crossed)
cortico~pinal !r<let

Anterior (d irect)
corlicospinaltracl
'C<.1!:G;. is
~.

;;;J ~
~

;~:f.:" Fibers from leit mOlOr COf1ex

;:::;::;:~ Fibers I'rom left red nucleus
:;i;ii~;: Fibers from lell seOSOfY cortex

C. Reticulospina llracts
". .
.... .. j:" , j j
I
'l1i'::. ' . A
/~
C;;' "N ,
l ateral / v,": . f l"
J. .~I
~ {~
reticulospinal . I \ v",
tract 1, f-~l.IJ;:!).
t vill.
.& .~ o II!IN
:', ';": ::

Medial
ves tibulospinal fibe rs l ateral vestibulospinal tract
Medial re ticulo spinaJ tract
.:~;:;:;, F i be~ from lefl pontine reticular formaiion ;~:~,:, f ibers (rom lett lateral (Deiters') nucleus
:;iiiii~i Fibers h om lef! medullary relicular formation ;;iiiiW f ibers from leit medicI! and inferior nude;
(only 10 cervical and Ihora<ic levels)

FIGURE 111.37, SPINAL CORD TERMINATIONS OF MAIOR DESCENDING UPPER MOTOR f

NEURON TRACTS __________________________________________
C
The lateral C ST and RST terminatio ns are directed tracts, and the vestibulospina l tra cts are directed
mainly toward LMN s associated w ith di sta l limb m ainly toward LMN s associat ed w ith m o re proxi
mu sculature. The anterior CST, the retic ulospinal mal and axial mu sculature. C
P<
~
"

250
(UMI\, or Systems-Upper Motor Neurons (UMNs) SYSTEMIC NEUROSCIENCE

b ci.alo r)' endings --.. - - - -- _ _ _ Frorllal eye iields

lnhibitoryending5 - (Brodmann ared 8)
Occipital
eye fields
,. IBrod mann
areas 17, 18, 19)
~ :e fS!il i a l

...., cleusOf----fl~~
,.
(.aja! '"'f""l

- , - - + i " - M"dlj , ,( longitudinal

fascicu lus
Superior
~ucem oblique m.

~"'ur on
Superior
r;Xlu S m .

lateral
Medial re1: lus

" uci eus

~---jc-:i-- Modli" longi!udin;:tl

iasciculi

'l.ledialloo,,"odlo,,' _"=:::=
~-Jt"~~1- __ Ascending Irael
ofOielers
'asciculi

SOP",m}
Medial Ve'itibulaf i
Lateral nuclei obliq ue m. rect us m.
Inierior

. GURE 111.38: CENTRAL CONTROL OF EYE MOVEMENTS _ _ _ _ __ _ _ _ _ _ __

-
Ce ntral control of eye movements is achi eved
- 'ough the coo rdination of extrao cu lar motor
-ucJei for cranial nerve eN III (oculomo tor),
C'J IV (trochlear), and eN VI (abducens). The
~ra pontjne reticular fo rmation (PPRF) (horizontal
;aze center) receives input from the vestibular
- .uclei, the deep layers of the superior colliculus
1pUt from Vl, V2, V3), the cerebral co rtex (frontal
,?>. e field s), and the interstitial nucl eus of Cajal
nput from th e vestibular nuclei and th e frontal eye
fields). Th e PPRF supplies the ip sil ateral eN VI
nucl eus and the co ntralateral eN III nucleus (via
intern eurons in the eN VI nucl eus) for th e medial
rectu s, thu s coordinating horizontal eye move
men ts. The interstitial nucleus o f Caja l helps to
coordinate vertical and oblique eye mOvements.
Secondary sensory vestibular projections also ter
minate in the extraocular motor cranial nerve
nuclei. Axons interconnectin g the extraocular
motor cranial nerve nuclei travel thro ugh the MLF.
251
SYSTEMIC NEUROSCIE NCE Motor Systems- Upper M oto r Neuron s (UMt\ s

Descending Coolro! from higher centers

Superior ce,ebt:!:II<lf Cerebral corte"

peduncle -...........

Mf;'dia l pa' olJbrachiaJ

.............

Pilcemaker
nueIeus
...'
'\ r RelSu lil le~ pan:maker

funclion

Ventral ~ Oor&ll

respira tory respiratory

nudeus <> ffi nucleus
/'" Dor~1 mOlor nucleus

~
oiCN X
Pcms
nucleus
____ D o rsal resp irvl Ory nudeus
nude us solitariu!>}
{ventrolateral nucleus solitarius}

Mutual inhibition

eN IX
chemorecep tor chemoreceptor
zone zone {Pc02 1
neaf ior<lm en
Ven tr al
o f luschka
respir atory

nll cieu. Nucleu s

ambigulis
tral respiratory nucleus
bo dy (,elfOambiguus nucleus)
Ca wtid body
/ '- lower mo to r neurons
oi phrenic nucleus
Tu lower mOll'lr
neUf()n s for
expi'<ltory r~. ~ 1._ _........

muscle ~ ,:z. 'iIJdI..P""""

folysyna phc Spinal cord
conne<..1ions
To lower motor cb ( 0 , (4 ,(5)
neurons iar lower mo tor neurons for
in~l,irJl or y inlercostal dnd ilCCesSOlY
l ~muscles oi respiration
muscles

,.-J.-,-,
Sp inal cord
tfJ'

(T6 - 12)

FIGURE 111.39: CEN TRAL CONTROL OF RES PIR ATI ON _ _ _ _ _ _ _ _ _ _ _ __ _

f
Inspiration and expi ra lion are regulated by nuclei mu sculature associated w ith expiration. Th e DR(\..
of the RF. The dorsal respiratory nucleus (DRN) (la t receives input from the carotid body chemosensl)t'3
eral nucleus solitarius) sends crossed axons to ter (via CN IX) and the aortic body chemosensors (\ ia
minate on cervical spinal cord lMNs of the phrenic CN X) and from the central chemoreceptive zon'5
nucleus and on thoracic spinal cord lMN5 that of the lateral medulla. The DRN and the VRN
supply intercostals and accessory respirato ry mutually inhibit each other. The medial
musculature associated with inspiration. The
ventral respiratory nucleus (VRN) (nucleu s retro
parab rachial nucleus (P BN ) acts as a resp iratory
pacemaker to regulate the DRN and the V RN. Tht
.,.
ambiguus) se nds crossed axo ns to terminate on medial PBN recei ves input from higher centers
thoracic spinal co rd lMNs that supply accessory such as the amygdala and the cerebral cortex.
252
J M Ns)
Motor Systems-Cerebellum SYSTEMIC NEUROSCIENCE

Lingula --~

Vermis

Lateral h",,,,,h,ce

PrimJry
fissure
"Unfolded" schematic oi
cerebellum demOnSlrJling
body map areas

Poslerlor
lobe

Flocculoflodular
;v----.,0,;,}.'---__J lobe

"Untoldcd" schema lie of cerebellum demonstrating rH1.iou ;. and lobes

:'C.lW_\\\e. 0\
\.rll':me'i\ca\
"unioldln'1,"
niccereoeliaf
sUr/an: In
derivation or
above clr'awam

"'G URE 111.40: FUNCTIONAL SUBDIVISIONS OF THE CEREBE LLUM _ _ _ _ _ _ _ __ _

- ....e cerebellum is classically subdivided into ante
'lOr. middle (posterior), and flocculonodular (FN)
lobes. Damage to each lobe is associated with ipsi
Jaie ral syndromes such as stiff legged gate (anterior
Dbe), loss of coordination with dysmetria, action
"'e mor, hypotonia, ataxia, decomposition of move
--en t (middle lobe), and truncal ataxia (FN lobel.
n e cerebellum is also classified according to a
-ongitudinal scheme, based on cerebellar cortical
regions that project to deep cerebellar !luclei,
which in turn project to and coordinate the acti\ it'
of UMN cel! groups. This scheme includes the
vermis and the FN lobe (projecting to the fastigial
nucleus and the LVNl, the paraverrnis (projectin
to globose and emboliforrn nuclei), and the lareral
hemispheres (projecting to the dentate Ilu cl eus.
Each cerebellar subdivision is interlinked w ith
circuitry related to specific UMI"i system).

2 53

SYSTEMIC NEUROSCIENCE Motor Systems-Cerebellum

Cerebellar Cortex
Exrit.1lory endi ngs - GoIgi (inner stella It cell (inhibitory)
Inhibilory endings _ _ G ranule cells (excitato ry)

Parallel fibers (a)(Ons of gra nule cell

Parallel fibe r, ICU!)

PurkinJ e ce ll s (inhibito ry)

Dendri le~ 01 Pu' kinje ce ll ..........

011.
G ranular
Basket cell layer

White
matter

:limbing filler
(excitatory)

Gl om eru li

, r.bers (e)(citatory)
Verkose axon of locus coeruleus (nOfadrenergic )
...... Purbni., cell axon

~~f!%
...... I"~N
'::

FIGURE 111.41 : CEREBElLAR NEURONAL CIRCUITRY _ _ _ _ _ _ _ _ __ __ __ FIG

The ce rebellum is organized into an o uter
3-l ayered cortex, w hi te matter, deep ce re be ll ar
nuclei. an d cerebe ll ar ped uncles th at co nnect with
the spina l cord, the brain stem, and the thalamus.
In the cortex, the Purkinje cells (the Inajor output
neurons) have their d end ri tic trees in the molecular
layer (a rranged in parall el "p lates"), th eir ce ll bo dies
in th e Purkinje cell layer, and their axo ns in th e
granu lar layer and the deeper w hite matter. Inputs
to the cerebellar cortex arrive as climbing fibers
(from the inferior oliva ry nucleus) or mossy fibers
(all other inputs). The mossy fibers synapse o n Aff,
granule cells, whose axons form an array of paralle l d im
fibers that extend through the dendritic trees of fibe,
several hundred Purkinje cells. Additio nal inter and
neurons modulate interconnectio ns in the molecu arOl
lar layer (o uter stell ate ce ll s), at the Purkinje cell drer:
body (bas ket cells), and at granule cell and molecu cell I
lar layer associations (Go igi cells). Noradrenergic
axo ns of the locu s coeruleus neuro ns te rminate
in all 3 layers and modu late the excitabili ty o f o ther
cerebellar connectivity.
 or Systems-Cerebellum SYSTEMIC NEUROSCIENCE
~ Gene ral Scheme B. Deep Nuclei Relationship With Afferents

_- + Cerebell ar
COlle>.:

, - _ Deep nuclei
~
~
Uppe r m OTor
Purkinje cell

neurons Granule
cell
II
l ower mOTo r D,,,,
neurons nucleus

Afferents Ske letal muscle Mossy fib er Climbing fib er

C. Circuitry of Cerebellar Neurons M o~sy Fibers D. Cirruitry oj Cerebellar Neurons - Climbing Fibe rs

Gran ule Deep

Goigi cell
cell nuc;leu s

Mossy llber Clim bing fiber

'IG URE 111.42: CIRCUIT DIAG RAMS OF AFFERENT CO NN ECTIONS IN THE CEREBElLUM _ _
"erents to the cerebellum include mossy fibers,
dimbing fibers, and locus coeruleus no radrenergic
':'..bers. The mossy fibers synapse in the deep nuclei
.snd on granule celis. The climbing fi bers intertwine
o!: ound the Purkinj e ce ll dendritic tree . Th e nora
drenergic locus coeruleus axons terminate with all
:ell types in th e cere bellar cortex. Th e loops and
J. Perlo:ins
M~ . MfA
I!) lmN
,',-: . ,,,
circuits in C and 0 show interneuronal modulati on
of ah'erent co nnections and Purkinje cell outflow.
The entire circuitry of the cerebellar cortex pro
vides fine-tuning of the original processing in the
deep ce rebel lar nuclei. The en tire Purkinje cell out
put to th e deep nuclei is medi ated by inhibiti on,
usin g GABA as the neurotransmitter.

255
 SYSTEMIC NEUROSCIENCE Motor Systems-Cerebellum

~ uperior cerebellar ped uncle

Cortical inpul ____ <,..... ~
. _~
Nucleus reticular.s
,l
" ..\
legmenl! pon tis con tr<lt ate, al cerebell ar cor te I'(

Ponline nuclei
(contral
~Prim(lry fi~u, e

,Ii,

Upper part of
meduna oblongata,

Spinal

Ves tibular nerve

Inferior ce rebellar pednu de
and gan gli

tract
tract

Spinal input __ (relilY lor cutaneous

informati on)
"'External cune;!te nucleus
(relay for propfioceplive
;nro rmalioo)

Ro str al
spinoce rebellar trac t
~
\
:":
~ ,.
_". k)
_~. ,-
IFrom skin (Io uch <lnd pre ssure)

' . y Frommusde Ispindles

Spinal borde r cells____ '" and golgi tendon organs)

MOlo. in terneuron ~~From skin and deep tissues

11 - ' (pain and Golg; tendon organs)
Lumb ar

~~p
skin (tooch and pressure)
.:and ['o m musee (spindles and
Golg; tendon o rgans)
c ~N
Ventral spinocerebellar tr ..et
\Oocsal spinocerebella r trac t

FIGURE 111.43; AFFERENT PATHWAYS TO THE CEREBELLUM _ _ _ _ _ _ _ _ _ _ __ H

Afferents to the cerebellum terminate in topo dorsal spinocerebellar tract and cun eo cerebellar EfJ
graphically organi zed zones in both the deep tract derive mainly from mu scle spindl e output, nu
nu clei and the cerebellar cortex. The body is repre whereas the ventral and ros tral spinocerebellar m,
sented in the cerebellar cor tex in at least 3 separate tracts derive mainly from golgi tendon o rgan o ut ter
regions, Afferents traveling through the inferior put. The middle ce rebellar peduncle co nveys ve~

cerebe llar pedun cle include spinoce rebellar path
ways (dorsa l and rostra l spin ocerebellar tracts,
cuneocerebellar tfacO, the inferior olivary in pu~ RF
input (rom the lateral reticular nucleus and other
regions, vestibular input from th e ganglion and
vestibular nuclei, and so me tri geminal input. Th e
256
mainly pontoce rebellar axo ns that carry crossed re fl
cortico pontocerebeliar inputs. A fferents traveling retj
thro ugh the su perior cerebellar ped uncle include ane
the ventral spinocerebellar tract, visual and audi ally
tory tectocerebell ar input, some trigemin al input, bell
and noradren ergic locus coeru leus inpu t.
 . to r Systems-Cerebellum SYSTEMIC NEUROSCIENCE

--
Excit;atory endings

Inhibitory
cerebral
e ndings of

Purkinje cells

~-""" ,"'l capsule

Ve ntral ante rio r and ventral

late r..l nuclei o/thalamus
r
Mesenc.ephillic retiCU!ilr formalion - - - - -fi II I 1_

Reu nucleus
Descending fi ber~ iwm
supe rior cerebellil r p ed unde~
Fasti!,;ial nucleus

Hook bundle oi Russell

Emboli/orm nucle us

Dentate '"' I" ",

Section A - B
Cerebellar co rtex
viewed irom
below

Section B - C
viewed from
above
Vestib ular nuclei

Inferior cerebellar peduncle

In(erior o live

f'on to m<.><iullary relicu!,ll fOr m,ltio n

- C URE 111.44: CEREBELLAR EFFERENT PATHWAYS _ _ _ _ _ _ _ __ _ _ _ __ _

~-ere nt s from the cerebellum derive from the deep
oJC!ei. Projec tions rrom the fastigial nucleus exit
.linly thro ugh the inferior cerebellar peduncle and
Fminate mainly ipsilaterally in the LVN and other
:tibular nuclei, and in pontine and medullary
ocular nuclei, regulating the vestibulospinal and
nKulospinal tracts. Proj ections from the globose
.n:i embol ifo nn nuclei project mai nly contra later
. through the decussatio n of th e su perior cere
ellar peduncle to the red nu cleus, with a smal!er
con tributio n to the VL nucleus of the thalamus, and
mainly modulate activity of the RST. Projectio ns
~rom the dentate nucleus project mainly con tralat
era lly through the decussa tio n of the superior cere
bellar peduncle to the Vl nucleus and, to a lesser
extent, the VA nucleus of the thalamus, and mainly
modulate activity of the CST. A small projec tio n
from the dentate nucleus also projects to the con
tralateral red nucleus and to brain stem reticu lar
formation.

25 7
SYSTEMIC NEUROSCIENCE Motor Systems-Cerebellum

Cerebellovestibulllr pathways Vesti bulocerebellar pathways

/Fastigial nucleus
Excitatory endings - - - - --
Inhibitory endings __ - / Globose nuclei
/Emboliform nucleu~

/Dentale nucleus

~~p
CIIDN ~\ '.',".'; :

Vesti~ular
nudel
{ L"
Medial- - -
Inferior----- /~;hn.< from cristae

Nodule Vestibular nerve Fibers from maculae

FIGURE 111.45: VESTIBULOCEREBELLAR AND CEREBElLOVESTlBULAR PATHWAYS _ _ _ __ fl

Primary sensory vestibular inputs terminate in the nuclei and to the pontine and medullary medial RF.
4 vestibular nuclei and in the fastigial nucleus and Thus, primary and secondary vestibular neuron s n
the cerebellar co rtex of th e vermis and th e FN lobe. project to the fasti gial nucleu s and the cerebellar th
The vestibul ar nuclei also proj ect to the cerebellar cortex, and both the cerebellar cortex and the th,
cortex of the vermis and the FN lobe. Purkinje cells deep nuclei project back to th e vestibular nuclei. of
in the vermis and the FN lobe, in turn, projec t back This extensive reciprocal vestibulocerebellar cir SU i
to the vestibular nuclei and the fastigial nucleus. cuitry regulates basic spatial pOSition and body be
The fastigia/ nucleus proj ects to the vestibular tone and posture. en
0';
th,
rna
258

 tor Systems- Ce rebellum SYSTEMIC NEU ROSCIENCE

....._ ; , ~"), -r--_..-' ': -- -_ ~ "
/ .. .; J . IF Cerebra l
co rtel(
:~
. :)' ,
",.
r
.,
r~
1 /
7

.
~

..,
~, ~

./

l ateral
,/
..,',..,- ! ;

Cerebellar (
~/ h,m;'Ph~ I
/ y',\ ; , j
"
,j
.

corte"
""--- '.-
.' ~"'._./
Dentate nucleus

P,,,"~; :,;=~c-;"ob~""'::::'"~d=~
~ ~~~ ;:::~======::::::~-
embolifOfm Pontine
J " , nuclei Red nucleus ., nu cleus

Skeletal

muscles
l ower motOl"
~,~

/ ~/ .......

','~-"/'

:;.,..--C
'-."

Cerebellar
cortel(

r= - . Aocc ulu~

~~
TO
r T1OCCUlo-ooduiar lobe T lowl'f motor
neurons

Spinocerebellar
tracts
Vestibular afferen ts
and secondary sensory
~. ~, ..~t
~-
t 1:1.'
- Skeletal J. Perkins
loiS, MfA

projections Ii ~ -.of I: muscles CI ~

FIGURE 111.46: SCHEMATIC DIAGRAMS Of EffERENT PATHWAYS FROM THE

CEREBElLUM TO UPPER MOTOR NEURO NAL SYSTEMS _ _ _ _ _ _ _ __
- ne lateral cerebellar hemisphere connects thro ugh
-1? dentate nucleu s with VA and VL nucl ei of the
-'al amus, the major tha lamic inputs to th e cells
.; origin of the CST in th e motor and premotor!
supplemental motor cortices. The pa raverm al cere-
oellar co rtex connects through the globose and
: boliform nuclei w ith the red nucleus, cells of
origin for the RST. The cerebellar connections to
-ne cell s of origin for the CST and the RST are
"'1ainly crossed, and th ese UMN systems cross
again before terminating on LMNs. Thus, the cere
bellum is associated with the ipsilateral LM Ns
through 2 crossings. Th e vermi s and the FN lobe
connect with the fasti gial nucleus and lateral
vestibula r nuclei. The fastigial nucleus projects
mainly ipsilaterally to cells of origin for the vestibu
lospinal and reticu lospinal tracts, exerting mainly an
ipsilateral influence o n spinal cord LM Ns through
these UMN system s.
2 59
SYSTEMIC NEUROSCIENCE Motor Systems-Basal Gan glia

Connedions of Basal Ganglia

Fro ntal

Precentral cerebr al cortex

Caudate {Head
nucleus Body
Tail Postcentral
cerebral corle).:

Cla ustrum

A&

,l>"
Temporal cone'

,
"m
,"

Lenti for r-
External segmen t} Globus nucl eu~
}
Internal segmeOl paliidus

'Fasciculus lenticulari s
Sub thala mus
Ansa lentic ularis
Hypothalamus

Pars compacta
~- Corticorubral, conicobul bar
Substantia and cort icospinal fibers
{ Pars reti c ularis

---
nigra

-
nudei from upper pons and midb rain tshown separiltely)
~----- -' ~
~
-'
.
Projections back 10 cortex a nd basa l ga nglia Projections from cortex and basal ganglia
---..., Dopa minergic projection of
substantia nigra
Thalamic and subthalamic
===:'
.. Cortical pro jection
Corticoslrial.ll projection
'" Slriatal projection
projections =
----,~ Pall idal projection

~~~
OIIl!N

:;.: -":;:

FIGURE 111.47: CONNECTIONS OF THE BASA L GAN GLIA

_ _ _ _ _ _ _ _ _ _ _ __ FI<

The basal ganglia consist of the striatum (caudate the STN projects mai nly to the globus p allidus.
Inr
to
nucleus and puta men) and th e globu s pa llidu s The striatum projects to the GP" The GP in ternal
SIr;,
(GP), The substantia nigra (SN) and the su bthalamic segment projects to the thalam us (VA, VL, and
inh
nucleus (STN), w hich are reciprocally connected centrom ed ian nuclei), and the external segment
exc
with the basal ganglia, are often included as part of projects to the STN. Thal am ic VA and VL nu clei
driv
the basal ganglia. Inputs to the basal ganglia from provide input to the cells of origin for the CST.
the cerebral cortex, the thalamus (intralami nar D amage to basal ganglia com pon ents often r esu lt~ cosl
tory
nuclei), the SN pa rs compacta (dopamin ergic in movement disorde rs. Damage to the dopam in e
baSe
input), and the rostral raphe nuclei (sero to nergic neuro ns in the SN pars compacta results in
input) are directed mainly toward the striatum, and Parkinson'S d isease.
segr

260
~anglia
Motor Systems-Basal Ganglia
"OrtE'X
:ortE'~
:rforrr
em
SN - Sub5tantia nig ra
STN - Subthalami<: nucleu5
GLUT .. Glu tama te
GABA .. Gamm a aminoburyric acid
DA .. Dopamine
FI GURE 111.48: BASIC BASAL GANGLIA CIRCUITRY AND NEUROTRANSMITTERS
_ _ __ _
Inputs from the cerebral cortex and the thalam us
to the striatum are excitatory (glutamate). Th e
stria to pallidal and pallidothalamic connedions are
inhibi tory (C ABA). The combination of th ese
excitatory and inhibitory influ en ces resu lts in a net
drive over the thalamocortical (and resultant corti
cospinal) output. Extensive inhibitory and excita
tory circuitry that produce complex modulation of
basal gan glia output also derive from the in ternal
segment of the GP, the SN, and the STN. Th e
SYSTEMIC NEUROSCIE NCE
3HT - 5-Hydroxytr yplamine (serotonin)
PC '" Pa r~ compacta
PR =- Par~ reti cutala
ACh '" Ac etylcholine
G P = Glob~ pallidus
SUB P '" Subs tan ce P
dopaminergic nigrostriatal con nection s can e\ ert
bo th inhibi to ry and excitatory effec ts on ' :1'
striatum. In Parkinson's disease, the lo ~! o ~ nigro
stri atal dopamin e axons aJlows bot;'" .... '2 ~.? {i ve symp
to ms (bradykinesia) and posit;\ , ~\'"'1ptom 5 (resting
tremor, muscular ri gidity, posture "' ~ ~abil i ty ) .
Ad ditional intern eurons such a3 .:--e e"\citatory
cho linergic interneurons in e ':;rl.3:tum are foune
in so me basal ganglia s tr uctJ~e--
SYSTEMIC NEUROSCIENCE Motor Systems-Basal Ganglia

b. limbic loop

SfN

c. Cognitive loop d. Oculomotor loop

Cortex Cortex
--~
-----11~,Onlai ~"~

SN = Substantia n;grd Vl .. Ventfolateral nucleus MD '" Medial dorsal nucleus

PC .. Polr5 compacta VA = Ventral anterior nuclcu5 p.arvo .. Parvocel'ular

PR .. Pa rs relicuta la eM .. Cel'womcdian nucleus magno .. Magnocellular

5TH " Subthalamic nucleus C PI .. Globus pallidus !nlemal segment CII!!N
w..;, :

FIGURE 111.49: PARALLEL LOOPS OF CI RCUITRY THROUGH THE BASAL GANGLIA
Th e co rticostriatal, striatopallidal, and pallidothala
mic con nections form parallel loops for motor, lim
bic, cognitive, and oculomotor circu itry. The motor
circuitry is processed through the putamen, the
limbic circuitry through th e ventral pallidum and
the nucleus accumbens, the cognitive circuitry
through the head of the ca udate nucleus, and the
oculo mo tor circuitry through the body of the
caudate nucleus. Connections through the GP and
262
FlGUI
the SN pars reticulata or ventra l tegmental area The nu
then project to appropri ate regions of the thalam u ~ end of
to link back to the cortical neurons of ori gin for the rostral
initial corticostriatal proj ections. These parallel structu/
loops through th e basal ganglia and the co rtex nUcleus
modulate specific subroutin es of cortical activity tegmen
distinct to the appropriate fun ctio n. The SN pars The nu(
compacta may act as the principal interconnec
tions among these parallel loops.
Is-Basal Gan,btor Systems- Basal Ganglia
.
_ SYSTEMIC NEUROSCIENCE

Hippocampal
ionnation Precommi S5urai forn ix

Ven tral amygdalo rugal

pat hway OA median

- FIGU RE 111 .50: CO NN ECTIONS O F THE NUCLEUS ACC UMBE NS _ _ __ _ _ _ _ _ __

Th e nucl eus accumbens is located at th e anterior
nus end of th e striatum in th e int erior of th e ventra! and
the rostral forebrain. Inputs are derived fro m limbic
structures (amygdala, hippoca mpal fo rmatio n, bed
nucleus o f the stria terminal is), and the ventral
tegm ental area, via a ri ch dopaminergic projectio n.
The nucl eus accum be ns is central to mo tivational
states and addictive be havio rs. [t also appears to be
a prin cipa l region in brain reward circuits associ
ated w ith joy, pleasure, an d gratification . The
involvem ent o f the nucleu s accumbens w ith a
specitic limbic basal ganglia loop helps to provide
m o to r ex pression o f emo tional responses and
acco mp anying gestures and behaviors.

263

 Autonomic, Hypothalamic, and Limbic Systems- Autonomic

SYSTEMIC NEUROSCIENCE Nervous System

Input ' 10 i!u tOll om ic pr!;!ga nglionlc

Inpu t, to dut onomic
neururlS ui sympathetic Jnd
pr e-g ~n 8Iionic neuron,
p,)rasyrnpathl'tic nerv o us sys tems:
limbic iorl'brain ~tructures
PreS<l tl l\lionic
So me corticJ I .egions
- - - - - sy",p~t het i c
~Iypolh.:lli)mic .egiuns
POSI&~ nglioJlic Brain St!:'m ce nte rs
+ - -- .. _- .. s\'mp.:lIhel ic Jnd nuclei
Sensory inputs
Preg.lnslionic

<- - - - . pari)Fynlpathelic

Po~tsansliOrliC

"" " "'" ~ parasvmpathe tic

Ed in ge r.Wes tphdl

Ciliary g.J np,lion

PupilJa.v (on~trlctor m . .... . . ~~ ___ _ ~ __ ~ ~
CiIr"lIym. . Supe,io.
.. Ii
Plelygop.:lL:it int' ~ns~on OWomotor III I! n. \
nucleU5
Wcttmalgland s"' __ ___ ,
Glands Ot nasill mUCOSil " "\ Infel lor
Submandib~lar SJ ngl lOll Faoal tV II ) fl . i sJ.liv~ t p r )'

Sall.,ary
[S"bm""",,,
S ubltngu~ 1
."d....__ .......... .
gland \:: GlossophJ rvngeJI (IXI n.
~
~::.
!
\
"" ""

Dorsa l mo tor
gl~n(b otlC ga~g
l' O n '-' ..~ . (au\ onom i
Parot'dgla nd ...--- .-!~ ____ V X
agusl)n
..-#.~3.:
~ \ nucil;u> of X

Smoot h m, c~ .d ia( m., ~ec.eto.\' ... -;!' .~it ---" L..lIer al ho rn

gl.mo~ in heart lung, ,';see' a,
(intermt'(\iol,lterJI cell column)
G t l.aCllu de~cendjn g coJon
ganglid

To ~"SUlldr smooth muscle 111 ....\ Th OfJcic

5kin and mus<:lc~. "'flector .... --
sp in ~ 1 co rd
p ill 111m" 1\\,<;.11 glands in limb s White Gray rJmu s fOO l
(Tl L2 J
U)mmunic.1tls communi ca ns
Se cretion oi

e pine phrrne and

o.
no replnephrrnr'
gJnglia
Into blood

~' ~ .

"""O''''"Od;.:::~"o.,y'''m __ " '~'--""""" ~ ,f$ \- ~"~:~>

.r./ "
To "'"'"
. m.. ,moolhm
. II., '
"",'.- .......- .. j

L _______ ,j

IIi"". ''''. ,.11'0

"boh " " .
"o,h,
. _ 4r~ ,'" Sacral
spinal cord
'I',er'"~ < Intramural
,'.~'O"' I ~1)l,
~~'
S (S2S4)
'wi!
mooth m., <;ccre
lower G( Ir"'c ,
10",
, ,Je '~l" >
. " &~ngliJ
" '"
PohK 00.
. -.,
. _ ""'"
.

FIGURE 111 .51 : GENERAL ORGANIZATION OF THE AUTONOM IC NERVOUS SYSTEM _ __ _

The auto no mi c nervous system is a 2-neuron chain
connecting preganglionic neurons Ihrough ganglia
to visceral target tissues (cardiac muscle, smoo th
mu scle, secretory glands, metabolic cells, cells of
th e immune system ). Th e sympath etic division
(sympathetic nervous system rSNSJ) is a thora
co lumbar (T1-12 ) system arisin g from the inter
med io late ral cell column of th e lateral horn, actin g
thro ugh chain ganglia an d collateral ganglia; it is a
system for fight-o r-flight, designed for reactions in
an emergency. The parasympathetic division
(parasym pathetic nelv ous system IPsNSJ) is a
264
craniosacral system arising fro m brain stem nud e
associated w ith cranial nerves (CNs) III, VII, IX, a-....
X and from the intermediate gray in the 52-54
spinal cord. Connections from eNs III, VII, and 1\
act through cranial nerve ganglia; connecti ons ircr
eNX and 52-54 act through intramural gan gli a in
or near the target tissue. The PsN5 is a hom eosta
reparative system. Central connections from the
limbic forebrain, the hypo tha lamus, and the brain
stem regulating 5NS and PsNS o utflow to the bcx::t.
act mainly through connections to vagal and s~ rr~
pathetic preganglionic neurons.
Jl onom ic -\ utonom ic, H ypo th alami c, and Limbi c Systems-H ypothalamus
and Pitui tary SYSTEMIC NEUROSCI ENCE
iotlic
, """
t('fTIS;
lIJres
""",

'"",

r!lers
udei
PUTS

~I

"
,

Preoplic nudei
2 Poraventri aJlJ.r nucleus
J Anterior hypothalamic area
4 SupraoPllc nucleus
5 Lat eral hypothalamic J.rea \
f) Dorsal hypolnalamic area
7 D orsomed ial nucleu.
"
B VenlromediJ.1nucleus

9 Postt'l'iof hypo th alamic area

10 Mafllmlllary!xxh'

TT Opticchi,ul1l 18 Intel'Pf'"
..oro region
12 Lamina terminalis 19 Thala.~

13 An terio!" commisWle 20 Tube!

14 Hypotl\;llamlC wkus 21 Opoe "'I!l"O

I S Intel thalamic adhe~ 22 Infu"","_

1& Fol nix 23 Ante...

17 Seplum pellocidtlm 24 POSIerG'

-
;:>i
FIGURE 111.52 : GENERAL ANATOMY OF THE H
The hypo th alam us, a collection of nuclei and fiber
....'IUS _ _ _ _ _ __ _ _ __

"'"'-Cmmillary or posterior. From the medial

pd tracts in the ventral diencephalon. regulates vis
ce ral autonomi c functi ons and neuroendocrin e
fun cti on, parti cularly from the anterio r and poste
Jm rior pituitary. Many nuclei are fou nd between the
posteri or bound ary (mamm illary bodies) and the
:ic anteri or bo un dary (lam in a terminalis, anterior com
mi ss ure) o f the hypothalamu s; these nuclei are
subdivided into 4 general hypot halam ic zones:
(1 ) preopti c, {2) anteri or or supraoptic, (3) tuberal,
at th e eN III \ entricle to the lateral
the nuclei are s-_ ooi\ ided into 3 general
perive ntricular ] medial, and (3) lateral.
...:..-lfy gland is attacfled at the base of the
;..amus via the media ""' em inence and the
m (p ituita!) sta, .. . The med ian emi
M1 important zone of neuroe ndocrine
n.

265

 Autonomic, Hypotha lamic, and Limbic Systems-Hypothalamus

SYSTEMIC NEUROSCIENCE a nd Pituitary

Sections Through Hypothalamus I

1 COI'PlJS <:allow m
Plane 1

,
~~~~~~~~~~~~~~~~.i'~~~sePtum Anteri o r horn o f lateral ventricle
Head ofpellucidum
caudate nucleus
-,;." Septal area
!Ii. Column of fornix
AnteriOT limb of inl ernal capsule
:i'J,

~;~==:==~===~~~'.~~~~?~~
PUTamen

'"
~i . 3rd ventp<lllidu$
Globus ride

,~~~~~~!t~=]~==::~~~~~~~~~~ l Medial commis ~Jle

Lateral preoptic ilrea
pleoplic are.t
Amerior
'-.~_ _~:;;;".._ _-\;;--_ __ Anlygdala
"" . Substantia innomina la

~~p
Cl:oN
~ ...: : :
Optic chiasm

Plane 2
~,/
Interventricular foramen (of Monro)

lrd ...cnllide

Column oi iornix

Perivennicul,)! nucleus

An$,llenllcularis

~ , _ Paraventriculilf nucleus

- _ mfer;or thal".mic peduncle

JI'_"~~~~;:::~=::::::::::=,~==
!' e )
'- .. La leral hypoth.,l.,mic .lIea

-===-- ~J Ante rior hypothala mic area

S"p"opHm" "" ISON)

. .-.,~./ r ..
7
~
OptiC tract
O pfic chiasm

FIGURE 111.53: SECTIONS THRO UG H THE HYPOTHALAMUS: PREOPTIC AND

SUPRAOPTI C ZON ES _ _ _ _ _ _ __ _ _ _ _ __ _ _ _ _ __

The major nuclei in the preop tic zone include the
medial and lateral preoptic areas. The organum vas
culosum o f the lamina terminalis (OVLT). a circum
ventricular organ, is presen t in this hypothalamic
area. The major nudei in the supraoptic (anterior)
zone include the supraoptic nucleus (SON ), the
paraventricular nucleus (PVN ), the suprachiasmalic
266
nucleus, the anterior hypo thalamic area, and the
lateral hypo thalamic area (LHA). Some nuclei, such
as the PVN, have many subregions (magnocellular
and parvocellular) that contain collections of chem
ica lly specific neuro ns w ith disc rete projectio ns and
functio ns. These gro ups o f neurons sometimes
in termingle w ithin o ne subregio n o f the nucleus.
lothalamus Autonomic, Hypothalamic, and limbic Systems- Hypot halamus
and Pituitary SYSTEM IC NEUROSCl E1\C
IS I Sections Through Hypothalamu s II

~~~~~~~~~~~~~~;
Plane 3
, of caudate nucleus
callosum
_horn of laleral ventr icle
Septum pellucidum
" - - - - _- -- - - - -- ---i, - Body of fornix
-r-~------__..---t-- Thalamus
Mamilio tha lamic twCl
3rd ventricle
POSl- lim b 0 1 internal capsule
Pul.amen
pallid us
Dorsal hypoth al amic area
i I nu cleus
of fornix
Ansa lenticularis
nucleus
l aleral hypothalamic area
I i hypothalamic area
Supraoplic nucleus

Pementricular arcuale nucleus

Plane 4
Thalamu s

Mamill(')fh"lamic tract
Post. limb of IIlIE'rnal cap~ule

f'u tamen
Zona incerta and fields of Forel
Fasciculu s lenticularis
Dorsa l hypo thalamiC area
Perjvenlricuiar nu cleus
Dorsomt.'dial nucleus
Aberrant pallidoiugal fibers
l atelal hypothalamic area
Ven tromedi al nucleus
Oplic Iract
SupraoptiC nucl eus

<:.::::::::::;:.-~__ Tub",ral nucl ei

'" Periventr icul,)f arcua te nucleus

FIGURE 111.5 4: SECTIONS THROUGH THE H YPOTHALAMUS: TU BERAL ZONE _ _ _ _ __

Ie
The major nuclei in the tuberal zone include the
dorsomedial (OM) nucleus. the ventromedial (VM)
iuch nucleus, the PVN, the arcuate nucleus, the peri
Jlar arcuate area (~ndorphin cells), the tuberal nuclei,
lelll th e dorsal hypothalamic area, and the LHA. Some
and nuclei from the supraoptic zone (PVN, SON, LHA)
extend cauda l~ into the tuberal zone, The median
eminence extends (rom this region, and axo ns
from releasing..factor and inhibitory-factor neurons
that control th e release of anterior pitui tary hor
mones funnel down to th e co ntact zon e, v,,'here
they release th ese factors into the hypophysia l
portal system, whi ch balhes the cells of the anterio
pituitalY,

26'
 Autonomic, Hypothalamic, and limbic Systems- Hypot hal amu s
SYSTE MIC NEUROSCIENCE and Pituitary

Sections Through Hypothalamus fit

r: ;
) '\
Corpu s ca llosum
Plane 5

lail of caudal~ nucleus

f Anl_ horn of taler,,-I venllicle
Body oi forni:-

'I
3rd vO;: r1 lrid~
Th.Jla rn us
Post. limb of i nl~rna l capsule
Field H, of Forel
i/I Id H, of forel
'.: Marnmillotll.llamfc Ifact
.... PUlamen
Globu~ pallidus
Post. hypothalami!..: ;lIea
$uulhalJmic nucleus
(aprulupechmcvlar Iransilion zone
l ,l\e ral nypoth,JlCln lic area
Opli<:tract
p,uts o f medial Column o f tOn'll);
mamm il1 Jry nucleus j nterc ~ l a tus

~~~...

i
l ale ral m<lmmillary nude u~

ClliiN
', .-:, :
~

i
Plane 6
Thalamus
3rd \'entr icle

Field H, of Forel
Field t-L~ oi Forel
Putamen

G lohus pallidus

MammilioThalamic trac l

PO$L hYllothaJaOl ic ':lIea

Subthalamic nud E-US

~
_c_" ~ !" ~ In{. norn of laleral IIl!'ntricie
_ ~ Laleral hvpolhalilmic are.,)
"ii-"
~ "" . Amygdala

ma mml'" ary b - Jv1.amm iliary peduncle

n ucleus Supramammillary decussa\lon

FIGURE 111.55: SECTIONS THROU GH THE HYPOTHALAMUS: MAMMILLARY ZONE _ _ __

The major nuclei in the mammillary zone include extends thro ughout most of the length o f the hypo
the medial and lateral mammillary nuclei, the pos thalamus and shom neuronal characteri stics seen
terior hypo th alam ic area, and the LHA. The LHA in the brai n stem reticul ar formatio n.

268

Ilam ~
. ...... o nomic, Hypothalamic, and Limbic Systems- Hypothalamus
Pituitary SYSTEMIC NEUROSCIENCE

1';.~~~~~~:".\: lamillothalamic t;:ao

..,,~- p,,,,,,,;o, area .
. I ~
Jr----""W- nucleus
Perrven\ncu ar

'\!uclcus
iotcrcalatus

Ventromedial \'1.!mm,ILlr.
nucleus comple\' /
nerve Tuberohypophyseal [rad

OculomOlOr ! I!! no:> r. 0:>

h

(irom
Irac t Pons

-~- Posterior lobe

o! pituitary

FIGU RE 111 .56: SCHE MATIC RECONSTRUCTIO N OF THE HYPOTHALAMUS _ _ __ _ __

-\ schem atic 30 recon stru ctio n of the hypotha la
mus in sagitta l section shows the nuclei, th e areas,
and the zo nes that occupy this small region of the
diencephalo n. Many pathways are represented,
including the fornix, the mammiliothaiamic tract,
the median forebrain bundle, the supraopticohy
pophysial tract, the tu berohypop hysial (tuberoin
fundibular l trac t, and bra in stem co nnections w ith
th e hypothalamu s. wh ich include th e dorsallongitu
din al fasciculus, th e descendin g med ian forebrain
bundle, the mammillotegmental tract, and descend
ing connections irom the PVN to preganglionic
autonomic nuclei.
269

 Autonomi c Hypothalamic, and limbic Systems-Hypothalamus

f

SYSTEMIC NEUROSCIENCE and Pituitary

Supplementary mOtor
(premo!Of) ;lrea
Molor area .......... ( ingula te gyrus

som"o"oro,y,,,, ~ Fornix

Corpus C;;Iliosum ' - / / ,J..........

, .;,' ' ".

Hyp<r
I

Orbital cortex
Amygdala
Hippocampal formation

~~p
Parahip pocamp;'ll gyrus

lION :\".':;: 1

FIGURE 1i1.5 7: FOREBRAIN REGIONS ASSOCIATED WITH THE HYPOTHALAMU S _ _ _ __

A number of forebrain region s are intimately co n peria mygdaloid co rtex. Important subcortical
nected with th e hypothalamu s, some through regions of th e limbic forebrain include the hippo
d irect fiber projections and some through indirect campus (a three-layered co rtex), the amygdaloid r
co nnections. Important cerebral cortex region s nuclei, and the septal nu clei. Important thalamic j
include the prefron tal cortex, the orbitofrontal con nectio ns include the medial dorsal and anterio r 5
(orbital) co rtex, the cingulate cortex, the insular nudei. Important olfactory connections inclu de the ~
cortex, th e parahip pocampal cortex, an d the o lfactory tract, nu cle i, and cortex. h
tl
a
c
P
B
d
270
la _ onomic, Hypothalamic, and limb ic Systems- Hypothalamus
-:) Pituitary SYSTEM IC NEUROSCIENCE
- II i
( inlwlille)o:\ rus
Cor pus cilllo~um

I
~>(;:::::~'
~
,:~I dorsal
: nucleiJnd
01thalamu)
Septal nuclei

From s~pta" subtallosal,

preoptic.. and
frolllotempCll3! Jr~JS

Hypothalamic nu clei

S"iJ.
,.,-mif'1a lis ii~~~~~ Olfactory bu lb
Orbiloj, ontJI cortex prOJl"Ctlon )
Hippocampal formation

:;-~tic ular H'pophvsls

\ ent,~1 am\ gdaloiugal pathway

Oors.ll Iongit ucinal 1.1 ;.c ICU:U'

" ,Me.li," to",b",'" " " ",,,Ie rdeKenclm1;l

- ,vled",""y C.lrdio vascular cente~
, ,,,",,, IX) nc,,'(:

FIG URE 111.58: AFFERENT AND EFFERENT PATHWAYS ASSOCIATED WITH THE
HYPOTHAlAMUS __________________________________________

-\pot hal amic co nnections are numerous and com
ex. Some regions of th e ce reb ral cortex (pre
-onta l, orb itofr ontal) and th e thalamus (anterior)
x nd axo nal projections directly to the hypo
,....,alamus. Diverse afferent pathways arise from the
- ppocampal formation and the sub icu lum (fornix),
':'1e amygdaloid nudei (stria term inalis, ventral
o71ygdalofugal pathway), and the habenula Ifasc>
cul us retroflexus). The retina sends direct retinohy
pothalam ic fibers to the suprachiasmatic nucleus.
Srain stem projectio ns, some compact and some
diffuse, ascend to the hypothalamus th rough multi
pie pathways (not shown here). Efferent connec
tion s fro m th e hypoth alamu s include those to the
med ian emin ence (from multiple nuciei), the poste
rior pituitary (suprao pticohypophys ial tract), the
septal nucl ei and the anterior perforated substance
(median forebrain bundle); those to the thalamus
(mammillothalami c tract); and those to many brain
stem and some spinal co rd sites (dorsal longitudinal
fascicu lus, median forebrain bundl e, mammil
lotegmenfal tract, direct co nnections from the PVN
to preganglio nic neuro ns, and o thers).
2 71
 Autonomic, Hypothalamic, a nd limbic Systems- Hypoth ala mus
SYSTEMIC NEUROSCIENCE and Pituitary

Brain Stem and BloodBorne Dien cl!'ph~ron and Telencephalon

I Inpuls

L-----------+j l 1lof11lOr'leS,
(glucose. Na.-, cyloi.:ines,
others)
no SeN) I H:etino hyp olh al..ulic Ihlihway

Assoc. RF
polysenSOl)'
information

Mammillary peduncle I (to lateral

(to LHA. lateral prl'oplic .Jll!a)) I MD m idline nuclei
~_.::.:::..::=:::::,",,:::oo:. ____...J
Mam millary nucleus)

d Dlf
. I Ito medial areas) Hypothalamus MF8 E
'h"" ~

"'rB
(to PVN)
1 ....;A::::"t"orioO,t"<o;",8"":'o"~<oertc":..J

I1\\ I
(lo lHA )
(t~ lHA, Lateral
po-eoptlC .Jrea.. other)
I. Posterio r orbito(ronlal corre)!,

Olfactory bulb,
rel ated region s
1-+-+---1'I (to PVN, lHAJ I

\\ 7I I (to PVN ) (to ~d i a l preoptic area,

AHA, SON)
I ,
Stri a termina lis

(via MFR, ST, VAFP) I, O lfactory-re lated projection

(to preoptic, medial areas) I, V6r'ltralam ygdalo fugal pillhwily

Dors.al and ventral NA bundles (to widespread an:as)

(via MfB, Olfr
\to mediill milffiITlill.uy nucleus) I_ Postoommissurill fornix

_"
(to .... ] zones)
I ,
Pre<ommissuul iOlni)!:

L----------'!"""-,.JI {IO widespread iI/east

OlF .. Dorsal longitod ina l lasciculus PVN .. Polraveotricular nucleus SON - Supraop tic nucleus
KF - Retic ular for ma tion CA .. catechobm ine Sf - Su ia (erm in-Ollis
lHA .. Lu e ra! hypo thillamic area NA .. "'Ofadl~ic VAf.P = Ventral amygdalofug.l1pa lhwoly
MFB = Median foreb ra in bu nd le 5HT .. :;Hy~oxytrypt amine. ser()(onin MO = Medial dorsal f1udelJS 01 thalam us ClmN
H':", :
NTS "' NucieuSlla ctUS50li(ariu s AHA - An tetio4" hypoth alamic area

FIGU RE 111 .59: SCHEMATIC DIAGRAM O F MAJOR HYPOTHALAMI C AFFERENT PATHWAYS _

The hypothalamus receives extensive input from
many regions of the eNS. Descending inputs arrive
from limbic forebrain structures (hippocampal for~
mati o n, subicu lum, amygdala), the cerebral cortex
(anterior (ingu late, orbitofrontal, prefrontal), and
the thalamus (medial dorsal). Ascending inputs
arrive from extensive areas of the autonomic brain
stem (tegmental nuclei, periaqueductal gray,
parabrachial nuclei, nucleus tractus solitarius
[NTS]); the locus coe ruleu s and tegmental ca te
cho!amine nucl ei; raphe serotonergic nuclei; and
the brain stem reticular formation. Th e retina sene:
input directly to the suprachiasmatic nucleus, a ~
lion of the hypothalamus that modulates diurna
rhythms. Bloodbo rne substances (cytokines, ho,"
mones, glucose, Na+, o thers ) influence the hvpc
thalamus via numero us ro utes and mechanism::..

272
amu s .\utonomic, Hypothalamic, and limbic System s-Hypothalamus

- an d Pituitary

Brain Ste m Efferent Pathways

SYSTEMIC NEUROSCIENCE
Diencephalon, Telencephalon,
and Pituitary Efferen t Pathwa ys

Irom Ol agnoccllular
PVN, SON)
Anterior pill.litarv

" om RF and IF neuron s)

(t rom arCLlil[C,
pt.:. '1\ enlriClJI<Jf nuclei)

_ (irom LHA )

H) pothalamus

ill I,
'" !.lI"O 1":'"<e63J m.lmmHlar y nuclei)

(irOn! medial "orr, O!"". efl\ncular nucleus) f - - - - - -

periventriculaf area )

iirom !oeptal nuclei.

p reopti c Mea)

(from periarCUilli: ,1 fC,]

(from p.uvo<eliular PVN

<lnn ;.on le scattered ,]J('<l S
I<lter;il and dorsal)

RF - Releasin g faClor
I F " Inhibitory facto r
PVN ., P,uaventricu!;u nud\.lS
SON - SUPfilOplic nucleus
MfB " Media n foreb rai n bundle
SHT : 5-Hvd rox~ lr\p!amin" ~""' o'.){1-'l M Teg T " Mammilioleg merllal tract
SNS - Sympa thetic nt'J' 01.1."" ,:"''" FilSc. RelrOO exu s " F,lsci cu lu s Re troflexus
Dlf " Dor.<,,)1 longi tudinal ra ,,,-ocu ,, ~ (HilbenuloPf>duncu lar Tract)
ILC - Int ermed:oI,H",ral ce'! c(>.'-"" S1 " Stri;) termirldlis
DM N of X : OmSill malar tautOnomic nue;,,:.!, of \ CA - Ca tt'cholaminf>

FI GURE 111.60: SCHEMATIC DIAGRAM OF MAJOR HYPOTH ALAMIC EFFERENT PATHWAYS _ _

Th e hypothalamus gives rise to ex tensive effe rent
prOjectio ns. Ascending efferents are sent to the
limbic forebrain structures (amygdala, septal mlele',
anteri or pe rforated substan ce), the cerebral co rtex
(orbitofrontal cortex and tem pora l associ atio n
cortex), and the thalamus (medial dorsal, anterior).
Extensive projections are sent to the median emi
nence (releaSing and inhibitory factors for control
of anterior pituitary ho rmones; dopamin e projec
tions from the arcuate nucleus and the PVN) and
to th e posteri or pituitary. O ther efferent projections
are sent directlv and indirectly to the preganglionic
neuro ns a t the SNS and the PSNS (median fore
brai n bu ndl e, dorsal lon gitudinal fasciculus,
mammill otegmenta l tract, d irec t project ions from
the PVN), to \.videspread autonomic and visceral
nuclei (noradrenergic neurons, serotonergic
neurons, parabrachial nu clei, NTS, per iaq ueductal
gray, tegmental nuclei, in terpedu ncul ar nucleus),
and to the brain stem reticular formatio n.
273
 Autonomic, Hypoth ala mic, and Limbic Sys te m s-H yp o th a l am u~
SYSTEMIC NEUROSCIENCE and Pituitary

(orIel(

Papez circuit

limbic {()(ebra;n Midbrain te8rnenturn

structures Brain stem autonomic structures

MD Thalamus Brain stem

autonomic struClures
Amerior thal amus

Hypophysea! porlal system _ _ _.,

An terior Spinal cord

Posterior pituit,uy SNS pregangllonics

SupraopticohYDo phY5t'al traCI - - - -

Dl F .. eorsallonglludinal i a$Cicu lus MO - Medial dor5al nucleus of thalamus

M f B .. Median forebrain bundle f'X - Forni"
ST '"' Stria termin alis MTT .. Mammilk)(ha lamic ' r;let
VAFP .. Venval amygdalofugal pathWolY SNS .. Sympathetic nervous system

C ~.~N

FIGURE 111.61 : SUMMARY OF GENERAL HYPOTHALAMIC CONNECTIONS _ _ _ _ __ _

The lateral, medi al, and periventri cuiar zo nes of th e
hypotha lamus have specific connectio ns with
the cerebra l cortex, the limbic fo rebrain structures,
the thalamus, and w idespread areas of the brain
stem. Extensive efferent projections of the hyper
thalamus are directed toward regulation of pre-
ganglio nic sympathetic and parasympathelic
neuro ns and toward release and regulation o f
hormo nes of the anterio r and posterior pituitary_
The anterior pituitary hormones regulate hormone
secretio n and functional activities of many target
structures throughout the body_

274
 .\ utonomic, Hypothalamic, and limbic Systems-Hypothalamus

- -:- nd Pituitary SYSTEMIC NEUROSC IENCE

-,
\J
loc~s coe ruleus

Parabrachial nuclei

Hypophyse:a l porla l , },stem

in th e median eminence

Antefio,

/
pituitary

Nudeu! ,:,c' ~::::~~l

SOhLlrius
i
, \ Do!';al mo tor

To rntramura l gan glia ---,v;;:;;-;-:;Jiiiiii~:".!'A:+-+--\i--

Va gu> \ - \
nucleus oi X
comp lex
(au tonomic)

Interm Nliolater 31
cell column In
laleral hom

To sy mpaThetic chain ganglia, _ _ --::==:---_~

colla te ral gan gli a, ad renal medu lla Too racic

spinal cord

(lH2)

FIG URE 111.62: PARAVEN TRI CULAR NUClEUS OF THE HYPOTHALAMUS : REGULATION OF
PITUITARY N EU RO H ORMONA L OUTFLOW, AUTONO MI C PREGANGLI ON IC
O UTF LOW, AN D LI MBIC ACT IVITY _ _ _ _ _ _ __ _ _ __ _ __

The PVN has man y projectio ns that help to coordi
na te pituitary neuro ho rmo nal o utflow, auto nomic
pregangli o niC outflow, and limbic activity. Magno
cell ular neurons send axon s to the posterior pitu
itary, rel eas in g oxytocin and vasopressin into the
general circulation. Cortico tropin-releasing factor
(CRF) neurons and some vasopressin neurons send
axons to the median eminence; these axons
release thei r hormon es into th e hypophyseal portal
system, influencing the release of ACTH , PVN
parvocellular neurons send direct descending pro
jections to preganglio nic neurons o( the parasym
path etics (d orsal moto r nucleus of CN X) and the
sym pathetics (interm ediolateral cell co lumn in the
Tl -L2 lateral horn) and to the NTS. These neurons
also send axons to several important limbic-related
struc tures, such as the amygdala, the parabrachial
nuclei, and the locus coeruleus.
2 75
 Autonomic, Hypothalamic, and Limbic Systems- Hypothalamus
SYSTEMIC NEUROSCIENCE and Pituitary

Behaviors Iniluenc:ed by Cytokines; ~" y -""~,. .,>'<

Illness hr.havior
Affective b ehavior
Cognitive beh avior
..,.-~ - "- ,.
Autonomic an d neuroendocrine
regulation

bloocJ.brain oo.mer (BBB)

~~
O rg.lnum vil K ulafum of the lamina

VasCl,jlature to hypothalamu s

,
~~~
G = Inlerle ukin-l !3 {ll-1~l e
Other cytokine~ actin g on brain:
IL 6 {inlerleu kin-6)
TNF-o. (tumor nt'cr()!;is faClor-o )
IL-2 (inlerJeukin-2)
Se""""
g;eog!ion or X
~ h----l+- Nudeus tf"dCTus
solil.1r'us
Vag.l l arierenlS ".'Ujl
irIviscera y- _ _ _ _... ~ 1-' ,, \ Dorsal mOto r
[a utonomic)
,I ellerents to nudeusorx
intramural ganglia

associated with Somalic Periphera l

vagal aife.-enls nl?rve

Sympathetic
Spinal cord
. ganglion

e - ~ -"'0
\;II'Target

CD Cytokin es tr~ n s ported directl)' acrm s th e 888. 's' Cy tokin e activation oi o ther afferents tha t modu late
a> C)'to kincs cros&ing IntO Ctlrebmsp in,d fluid at OVlT. Of .lC lingon cells
\V dorsa l horn sensory plOce!...~ng tv m.:my site'>..
thai release PG E2 01 neurons thai projeCt to visct'fa~aulonomic Wuelvre'>- C~'lokine modulation o f norepinephrir'le release from
sy mpatheti c: !'lerv e termilla l.
C!l Cylo kine-stimulated r~l eas(> o f small mol ecu les (su ch as NO) Ihat directly
cross in (() th e brain and act as mediators. o Cytokin e modu lation of ne uro transmitt e r intracellul Jr
Signalin g in t.arse t cells.

Cytoki ne stimulalion 0 1 vagal afferetlts (t hr o ugh p.>raoeurons] thai

@ modulate aCllvity in nucleus Iradus ~Iitariv:o. iniluendn!! the mvlliple

C~tokine mo dulation o i p ituitary hOlmonc release.
<lctivi!ies of Ihe para l'en tflw1ar nudeus .l,xl man y olh er sites _

FIGURE 111.63: MECHANISMS OF CYTOK INE INFLUE NCES ON THE HYPOTHALAMUS AND
OTHER BRAIN REGIO NS AND O N BEHAVIOR _ _ _ _ _ _ _ _ _ _ __
Cytokin es, including interleukin (IL)- '~, IL-6, tumor
necrosis factor-ex, and IL-2, ca n influ ence central
neuronal activity and behavior. This diagram illus
P
trates IL-l access to the brain: (1) directly cross ing
the bl ood-brain barrier into the brain, (2) acting on
circumventricular organs (OVLT), releasing small
mediato rs such as PGE2, (3) acting o n vascular
endoth elial cells, releasing nitric oxid e (NO), which
acts in th e CNS, (4) activatin g vaga l afferents into
th e NTS via paragan gli on cells, and (5) activating
o ther afferent nerve fibers. IL-l can evoke illnessp
behavior (fever, indu ction of slow-wave slee p,
decreased appetite, lethargy, classic illn ess symp
toms), influence autonomic and neuroendocrine
regulation, and influence both affective and cogni
tive function s and behavior.

276

~mus Autonomic, Hypothalamic, and Limbi c S) s:lems- H\-pothaJamu s

- and Pituitary SYSTEMIC NEUROSCIENCE

) Pineal gland

<,,.,;~
"'\
Organ um vascularum
of the lamina terminalis
(O VU ) /

Median eminence

~ .
. <~.~ \
Area
postrema

-
~
~ /
\ ,,
.--
"'" I

FIGURE 111.64: CIRCUMVENTRICULAR 0 G.-

Circumventri cular organs are "windows o n <he
brain" that are devoid of the usual tight-juncOCr
endothelial appositions and instead have leoes-
trated vasculature. Thus, the circumventricu;ar
organs have no blood-brain barrier. Som e oj-
organs- th e OVLT, the subfornical organ, a nd ' ~
area postrema-have associated neurons \\ ; ...
projec tions to hypothalamic and other viscera.
structures. They also have cells that can release
small molecules, such as PGE2, into the CSF
affectin g target structures at a distance. The
~.::. ~ i te of axonal release (from PVN
~~ '"1()cel kdar neuro ns) of oxytoci n
a'iOpressin into the general circula
~'1 eminence is a zone of neuro
"ra1Sdu ctio n for the secretio n of
~ and inhibitory factors into the
- oonal vasc ulature; th ese facto rs influ
-~.' or anterior pituitary hormones.
::;and synthesizes and releases the ho r
n,
277

 Autonomic, Hypothalamic, and limbic Systems-Hypothalamu-s

SYSTEMIC NEUROSCIENCE and Pituitary

Blood Supply of Hypothalamus and Pituitary Gland

"

". . ,
"J>- J Hypothalamic \'esseis K

nucleus

~
lans h ypophysea l Anterior brOl.nch} Superior

portal veins \1k

Posle.'"ioo branch hypophy>eal
Short hypophy>eal
'"ery
portal veins ~

Artery of trabecul~
Trabe<ukl

Efferent vein to
_ caverno us >lnu>
,~

tr;;.~ Anterior lobe

Simil ar >ecl ion through
Secondary pl ex us of
supraopti c nucleu>
hypophyseal port al ,ystem

l ate ral branch """"" Efferent vein to

,cd caverl10us sinu~

M ed ial branch /

cf (.wernOU 5 si nu s

Infer ior hypophysea l Internal ca ru tid arter" ~

Superior hypophyseal artery

Portalv,,;n >
Lateral hypophyseal vein ~ /
Inferior hypophyseal ~!p
~05te rior lor
Q '~N
FIGURE 111-65: HYPOPHYSEAL PORTAL VASCULATURE _ __ _ _ _ __ _ _ _ _ __

The hypophyseal portal vascula r system derives
from arterioles that enter the med ian em in ence at
the base of the hypothalamus. The primary capi l
lary plexus is a site where releas ing and inhibitory
factors that influence the secretion of anterior pitu
itary hormones are released from axons w hose
neuron s reside in the hypothalamu s and other Ci\. ~
sites. These releaSing and inhibitory factors th en
travel th rough venu les into th e seco ndary caprl lan
plexus in very hi gh concentrations and act directl\ .
on anterior pituitary celi s, w hich synthesize and c
secrete th e hormones of the anterior pi tuitary.

~
o
278
un >\utonomic, Hypothalamic, and limbic Systems- Hyp othalamus
a nd Pituitary
-
Emotional and extt'TOCepliv/:!
j;,iluences via afferen t nerves
\0 hypothalamus .:.-.:.-~~-~~
Blomi-borne
mol ~cutar
mnu ences on
CRF neurons
Adrenocor tica l
Thyroid hormones
hormones
FIGURE 111.66: REGULATION OF ANTERIOR PITUITARY HORMONE SECRETION _ _ _ __ _
Neurons that synthesize releasing and inhibito ry
iactors for control of anterior pituitary hormones
send axons th at termi nate on the primary capillary
plexus of the hypophysea l portal system (the zo ne
of neuroendocrine transduction) and release these
factors into th e hypop hyseal portal blood. Th ese
factors then flow into the seco ndary hypophyseal
portal plexus and regulate the release of anterior
pituitary hormones. These hormo nes ac t o n target
organs to affec t the release of target organ
SYSTEM IC NEUROSCIENCE
nucleus
. : ,.-..:
Specific secretory cell s of
ante riOI lohe (adenohypo physis)
influenced by neuroseoetion5
from hypothalamus
Skin
(melanocytes)
Es trogen
Testosterone Progestero ne
hormones or to intluence metabo lic and functiona l
ac tivities. For example, CRF neurons release CRF
into the hypophyseal portal blood, regulating the
release of ACTH, which in turn regu lates th e
release of cortisol from th e adrenal cortex.
M agnocell ular neuron s of the PVN and th e SON
send axons directly to the posterior pituitary; th ese
axons release oxytocin and arginine vasopressin
directly into the systemic circulation.
279

 Autonom ic, Hypothalamic, and limbic Systems-Hypothalaml.i ,~

SYSTEMIC NEUROSCIENCE and Pituita ry

Ne uros.ecrelory En ding (pos te rior pituitary)

"'' ' Jll ' t'(

Pi tuicvte processes

., ,z~t
'!'fa "I Iiit Axon

~
:>.:
'rf.,jji~ t '/~z'1' /d"
"''HI
( l' . . /1 ~;o~

t %ll
,
"" " '.~ :> .. '<"~ ~ ~
I: "tr,~
I "

1;
"f" ! ' -, ,- ,
I,

[}Ii ~ :,,,~'t to'! ~

'~I, i d -
I" I: '"

i~~!"\!f;)-~-~~d'~..\
" ' ,: l t,-, {I,'
:."
t\ I.,
iiw'
'0/- I',
". \ ~. .. ,\ -1" l'

---. .' ~,11. ~I'"

J-,; BrWFl -s\cm t .,11 \ .-"~\ ,,('''''.

\i
~ '. 2\ ;J~ ~ "'M
.,-
patnways I
V :'
i?: ~i .~ .. ~ ~-~
."J.,~-<;~-', "".. ::'''~
M
"'" "'r- --".,',' ''
fZ....

~rti
.
C.~pillJ.ry-
-'/0 .,- . .?O: '~~

-... - ;it:~I<,st
Neurosecretory
Endothelium ~es id(-'S

M;l51<:ell Collagen space

~<l5ern e l1l memb ran e

:.ulsr
Arlerial O rigin uf Y;lsopf"('ssin
tohvpCl
Supriloplic nucl eus Cell of
Neurolw p() p h y~ eJ I sli praop lil
II<lCI ,/'" nucleus

He rri,\ S bod le , _ __________.---1(

I,
Axonal
tran ' f10 rt I'"ij
~
Fenestrated
Anterior

.j
P05lerior lobe
Ineumoypophy!>i s)

/~! e of~'J " ,j

Po,terlor
of se(re l olY

produf l JJ
' .~
c.lpillary

lobe - ..

~~~
O'Il!N ~\:.: :I
vi

FIGURE 111.67: POSTERIOR PITUITARY (NEURO HYPOPHYSEAL) HORMONES,

F
OXYTOCIN AND VASOPRESSIN
_ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Magn ocellular neuron s in the PVN and the SON neurons possess extensive protein synthetic capac \
send axons directly through the infundibular region ity and transport the vesicles in w hich their hor b
and th e pituitary stalk, to tenTlinate on th e vascula mo nes are packaged to the axon terminals with b
tu re in the posterior pituitary. Neurons from borh very fast axoplasmic transport The hormo nes are u
nuclei syntheSize and release oxytocin and arginine released fro m the termi nals and di ffuse through the it
vasopress in in to the systemic vasculature. Brain fenestra ted capillari es directfy into the systemic v a~ al
stem and forebrain pathways terminate on the culature (inset: Neurosecretory Efferent Endings u
magnocellular neurons and regulate th eir secretion From Magnocellular Neurons in PVN an d SO N). n,
of oxytoci n and vasopress in. Th ese ma gnocellul ar
280
l am ~. ~utonomic, Hypothalamic, and Limbic Systems-Hypothala mu s
SYSTEMIC NEUROSClE' CE
- and Pituitary

MKHANISM OF ANTIDIURETIC HORMONI IN IItc.llIAIINC. tJlIlNI V{IIIJM! ANI> (ON( ~NJRAJION

0;m01Jr.~~j'~"di:: ! ~;\]i~~~~j~:~
ADH IS produced In >upraoplic and I'Jravenln<.;u.

lar nuel .. i of hypolha lamu~ and d"""end~ along ; n I J~e 1

0r,,1 or I)
nerve fibers 10 neuruh~1K)l'hy,,;,
Blood
""chJnS,,\\Jth . : 1'
whe r.. it i to red
,ec!emJ lI0 55 " J gut I ' i
for sub-5eq uent
loss Into bod\ cJ\ltie s IJ scites, efiu
rele.J.>e
...ont or loss o!'.lernall\ Ihcmm
m,lge s"!"!ai

,
hypoth.llamic ~more
(eplOl$ and by low.
blood \'oIume aifec!ing
loor.>(;( and c~ r otid
volume recep tors; low
osmo l<llily ~nd high !.>lao<!
"''''''3j
volume in hibit AOH rel ease

In presen ce of ADH, blood flow

to rena l med ulla is dimi"i~ h ed,
th us augmentin g hypertonici!)' oi
medull ary il1lers!ilium by minimi z
ing depletion oi solutes via bl ood-

ADH cause, \vall s 01 co ll ectin g duc ts to become

mOle permeabl e 10 "'01(1; ' ond thus permits 0"
mo lal equil iblation arid Jbsorption of walei' into
the hypertonic intersti tium: a Ima MvoIunle o i
hishl~ CQnCCf1lra ted Ur~le is excreled

o --.J
I
270
/ ,
290
,
310 - 30 - 20 -1 0 0 10 20
Plasma o~mol ali.y (mO~m ;kg Hp) % ( h;lnge in blood volume o r pteu' e

FIGURE 111.68: VASO PRESSIN (ANTIDIURETIC HORMON E) REGU LATION OF WATER BALANC E
AND FLUID OSMOLALITY _ _ _ _ _ _ _ _ __ _ _ _ _ __ __

Vasopress in regu lates the vo lume of water secreted
by the kidneys. Vasopressin secretion is regulated
by the osmola li ty of body fluids and by blood vol
um e and pressu re, Changes in body fluid osmolal
ity of a few percent are suffi cient to significantly
alter vasopressin secretio n. Decreases in blood vol
ume and pressure of 10% to 15% or more are
needed to affect va sopressin secretio n. The blood
vo lume and pressure sensors are found in the pul
monary vessels, ca rotid sinu s, and the ao rti c arch.
These baroreceptors respond to stretch of the ves
sel wall, which is dependent on blood vo lum e and
pressure. The diagram shows th e mechanisms of
action of vasopressin o n th e kidney, with resultant
effects on urin e vo lu me and concentration.

281
 _.......,.....
~ ._ ...
........,,~.:-....,..-~=~~-=~~.-.?,S::_""'-~_._~.~"'.,:'''.;;,;,;.~....s.........._
.

Conservation a-f!
regulation loss
,.,
production oihe~-

o _J:f~.'N_
.' . -j{.
.-.k/<;0-;...;,;..
""""&Q.0_
. ;.:
C:';.<~.,~,~" 0::

Inflammatory cylokines,

Neurohumcral mechanism fot- increasin8 ~

thyrotropic activity of anteliOf lobe "

fO elevate metabolism '(
~!and

, t--- R e~pil a tion

t
37C (98.6 f )
regulation

l
Cutaneous
blood vessel
constricti on
(dilation)

~~?
o.._. -
~

Perspiration t
M"'''

FIGURE 111.69: THE HYPOTHALAMUS AND THERMOREGULATION

_ _ _ _ _ _ _ _ __
The preoptic area of the hypothalamus contains
wa rmth-sensitive neurons. The posterior hypothala
mic area (PH A) contains cold-sensiti ve neurons.
The preoptic area an d the anterio r hypothalamic
area initi ate neuronal responses for heat dissipation
(parasympatheti c); the PHA initi ates neuronal
respon ses for heat generation (sympathetic).
Neu ro nal pathways arisi ng (rom the brain stem and
the limbic forebrain areas can modulate the activity
of these thermoregulatory systems. The preoptic
area is responsive to pyrogens and the inflamma
tory cytokine IL-l p; thi s area can generate an
increased setiJoint for temperature regulation,
thu s initi ating a disease-associated fever. Extensive
hypo th alam ic connectio ns with the brain stem and
th e sp in al cord are used to initiate ap propriate
heat-d issipa tion or heat-generati o n responses.
App ropriate behavioral responses also are initiated
to op timize thermoregu latio n (e.g., going to a
warmer or cooler location).

282
=\utonomic, Hypothalamic, and Limbi c Systems-H\ pothalamus
and Pituitary SYSTEM IC NEUROSCIE',CE

PARASYMPATH ETIC SYMPATHETIC

SmaU aneries;
and "'le<io~

Spinal
(ord II Change in posturE'
(sitling to s tandi~

I
+
Venou~ return

0 '-r-_ _ _~~~-~...",!
o 1 00 200
.\-IA P (n1m rig)

'P. . .asyrn~theli( I
efferent QUtpul \
I MAP

, Firing rate of

cs;:;;;;;;:
- -~
barOfE'Ceplor
.me.en! fi~

t .HE'aft rail'

-
t Qudiac output tMAP . L.

FIGURE 111. 70: SHORT-TERM REGULATIO N OF BLOOD PRESSUR E _ _ _ _ _ _ _ _ __

Both the sympathetic and parasymp athetic divi short-term bl ood pressure reg ulatio n. Th e hypo

sions of the autonomic nervous system are thalamu s. and the NTS are the key sites integrating

involved in maintaining blood pressure on a limbic forebrain dnd cortica l inf1uences over th e

seco nd-by-second basi s. Many descending path brain stem regions that regulate blood pressure.

ways from the brain stem (including the NTS, These brain stem sites have extensive interco n

tegmental catecholamin e nucl ei, the locus nections w ith each other. Th e examp le of blood

coeruleus, raphe nuclei, the rostral ventrolateral pressure regulalion in this plate is based on a

medulla, other medullary reticular regions, chan ge in posture. (ACh == acetylcholine,

parabrachial nuclei, angi otensin II-containing E = epinephrine, NE = norepinephrine,

neurons) and the hypothalamus regulate the auto MAP = mean arterial pressure.)

nom ic preganglionic neurons associated "',fith

283

 Autono mic, Hypoth alamic, and Limbic Systems- Hypothalamus

SYSTEMIC NEUROSCIENCE and Pituitary

Rt'S ponse 10 Decre a sed Blood Volume and Pres~utt RI,'~ponse to Incn'Olu d Blood Vo lume and P' l'ss u re

S~'mp~ thetic I SylTlpilthetie

t Ile'VI' ;Klivity , nerve activity _ ~ ADH
I - - - - - ADH ------.,

eN IX. X eN IX, X

HeJrt

Angiotensin
,lnd

lung,
,it"
"
~Ild lungs

ANt>
A"
.' r~
~
} -"'..;:'
Liver
-":.1_,::,/' ,---~
Adrenal; ,\drenals
I
Ao,'o,"o,'oo,,, TAO"O'' ' 'O , I
, AldoslerollC

I Renin
, .md
Renin
angiofen$in n

r.; ! im(,I;Jle~ renin

L - ;'':.l0

...~"'c":'cec'o
'cC::.) Aldosterone L
~I'u{'rion
:m(/ IdeCleas(>s waler
decreAse! NaCi {'.>:netion l Ki dneys
excretIon)
I

J. Perkins
!.IS, MfA
t
Blood v()Jum~'
,Ifld prt'Sstlrl'
l inneMed I f~O
inrdkt:j
..

,
I Bloud \ " Iun ...
and pr('~~u,,'

o.~

FIGURE 111.71: LONGTERM REG U LATION OF BLOOD PRESSU RE _ _ _ _ _ _ _ _ __

When blood volume and blood pressure change, thelic aCfivatio n, no repinephrine and epinephrine
the kidneys respond by retaining or excretin g NaCl secretion from sympathetic nerve terminals and the
and water in order to resto re blood volum e to its adrenal medulla in crease in the ci rculation and act
normal homeostatic state. With increased sympa- on the kidney to red uce NaCI excretion.

2B4

 M onomic, Hypothalamic, and limbi c S\ ste ms- H\ pothalamu s

ill d Pituitary SYSTEMIC EUROSCIE CE

c::::;';,lJ~ Ca lc.:Inne
II:;W""

Taste .:
QfiQod

Heilring (sounds 01
food pr~par-ation,
call to dinner,
bell,etc.) O ('Cl" a...~
oiCo.. ClP
and Jep:,.,
Thoracic
Dorsal rool

0., .,# .
~"'H ~
Deplt'riOll of body rnJtri~nl Slore5
,",10d" QI
inducin,!t
hun!,,,,

<EV
- Symp,llhelic efier~ls
- Pat"iympa l hel ic~.f ferrnts
- - ....fie1enh land CNS connec:tion~1 r
H unger ( 011111< .,...... - ,.
- - - - lodell"il" palhs

FIGURE 111 .72: NEURAL CONTROL OF APPETITE A" D H l " GE R _ _ _ _ __ _ _ _ __

Th e sensa ti ons of hunger and sa ti ety are comple,
and include multiple neural pathways and cirCldal
ing hormo nes_ Depicted here are pathways
invo lved in the sensatio n of hunger. Altho ugh o ur
understandin g is incomplete, the hypothalamus is
known to pl aya critica l rol e in co ntrollin g appe tite
and food intake_ When food is ingested, chol ecys
tokinin and glucagon-like peplid e-l are released
irom neuroendocrine cells in the intestine. These
hormones :::uppress ap petite and give the sensation
oi ~at j el;\. In the absence of food, the level s of
the::' hormones are low. Long-term regulatio n of
iood intake il1\ olves the ho rmone leptin, w hich is
produced b~ fat cells, \lVhen fat stores are hi gh,
leptin is re leased and appears to act on the hypo
thalamus to sup press appeti te. When body nutrient
stOres are depleted, leptin levels are lowe red.
285
 Autonomic, Hypothalamic, and limbic Systems-Hypothalamus
SYSTEMIC NEUROSCIENCE and Pituitary

Ne ural, Neuroendocrine and Systemic Components of Rage Reaction

paMrn released ~nd
cortex and

Or!);t"ffon,all-on"~ ~-"\';' 11 ",,~i

Median iO<dJ<dh . l'- """C""l -

buo-.dIe ~

"~Ii

"T ,root Id"f il iWi

""

"

Pe!. ic""""fl....-.- '

~~p '
'~N

FIGURE 111.73: RESPONSE

NEURAL AND NEUROENDOCRINE ROLES IN THE FIGHT-OR-FLIGHT
____________________________________________

The classic sympathetic fight-or-fligh t response,
shown here CI S a rage response, invo lves the secre
tion of neuroendocrine "stress hormon es," whi ch
include cortisol fro m the hypothalamcrpituitary
adrenal (HPA) axis and norepinephrine and epi
nephrine from symp ath etic nerve terminals and the
adrenal medulla. Sympathetic connectio ns with
the viscera ini tiate physiological changes to support
th e in tegrated fight-or-flight response. Th ese
changes include diversion of blood (rom th e
viscera and the skin to the muscles, increased hea rt
286
rate and cardiac o utput and contractili ty, bra n
chodilation, pupillary dilation, decreased gastroin
testin al activation, decreased renal activity, and
glyco genolysis from th e live r. Inputs from th e limbic
forebrain, the co rtex, and the brain stem regulale
th e hypothalamic co ntro l of neuroen docrin e an d
autonomic outflow and are key in initiating the
classic fight-or-flight response_ In the fight-or-fli ght
respon se, the brain stem parasympathetic neurons
are inhibited.
 omic, Hypoth a lamic, and limbic Systems-H ypotha lamus
Pi tuitary SYSTEM IC MUROSClE'JCE

( - , - - - --limbjc forebra,n J .,.2 ;

(cingu la!e (0."1"',
anly gd J I~

Cytokin .. a nd inflam matory

mediator ieedb Kk to th"
bra in and
Bra in

Va o;c ular de live ry of e minenn~ nud el

nellfOEndoc:rine hormone~ R"'!':.J.;;ing a"d
"irY' btllflg factor s

, I
Thymus t r~ctw;

.'- ------ ---.~ ,, >olitariu,

,,
Balle ,, \
rn ~rrow
, Vagu~ (,'\) n.
... "0'''' motor
l ~lJ!onomk:)
nu de lJ ~ 01X

---------- ..
Pulmoll ar y
MALT

c
"' -
Spleen

=-----,~ Pr("l!~nglio"ic
>m ph nooes
;\mp ,11 hetic
f ,., ,
tw .................
', ............ =>-~
~
G J 1J _~s()( i ateci
. "Tl pnoid li$sue
(GALT) .... -_ .
~~?
.I Collateral sympathetic

Skin
lym phoid
tissue
ganglion
~, ;;,.-- - ---- ------ --------- ------------- -. --
~ ,<.u..

~ ~~

- IGU RE 111.74: NEUROIMMUNOMODULATIO N _ _ _ _ __ _ _ _ _ __ _ __ _

,... '1 nections from the cerebral cortex, the limbic direct/\ innervate virtually all organs of the immune
;-t"brain, th e hypo thalamus, and the brain stem system. Vagal postganglionics innervate pulmonary
~rt extensive modulation over autono mic pre and gut-assoc iated lymphoid tissue. Pituitary hor
.......:;:glionic outflow and neuroendocrine outflow. mones in the circula tion, and their ta rget organ
?J rohormones and neuro transmitters fro m this
hormones. modulate immune reactivity in all
...d ow target lymphoid orga ns and cel ls of 'he
lymp hoid organs. Co rtisol, norepinephrin e, and
'"'llune system. Thi s ci rcuitry provides th e sub epinephrin e are partic ularly important in mediating
n.te for behavi or, emotional responsiveness,
chronic stress respo nses related to immun e
..... ro nic stressors, and positive complem entary
reactivity. Circul atin g and local cyrokin es and
erven tion s to influ ence immune resp onses. inflammatory mediators act on the brai n and th e
~ m pathetic postgan glionic noradrenergic fibers pituita ry to provide immune-neural signaling.
28 7
 SYSTEMIC NEUROSCIENCE Autonomic, HypothaJamic, and Limbic Systems- Limbic

Major limbic Forebrain Structures

(inguLu{' g~'fUS Intef"ventn c;ul.lr fO fJmen

Anterior nucleus of thalamus
Indu, ium gri,(':um
In tertha lam ic adhesion
CorpU$ callo,um

Septum pellucirlum

Precommissur,ll iornl:ot"

Seopt,ll nuclei Stria

medu llarl5
Subca llosa l
H\,pot l1alamus

Paraterminai gyrus .............

l amina termin alis Calcarine

sulcus

------~
~
med i3l st fia -, (fissure)
O lfacto ry laterill stria .. ~"
tr~ct ~~
~
"
~ ,I _
, -,.~:(
{
---..<ill
bulb:---- ;i Gyrus J;N#H':..::;,..
. fasciolari>
Anterior perforated
substJnce -
De nIa lI'
OptiC chiasm :? gyru s
Pus\commissural fornix \ ',

Mammillary body and

imbriaof
hippocampus ,p~p

mammillotnalamic tract tT 0 llliN

'~ '.'.': ;~

Medial forebrain bondle ---1 Pa. ahippocampa l gyrus

A m~'gdaloid body Descending conne(tions to
(nuclei) /
rt'ticular and tegmental nuclei
Uncus of braIn ste m (dorsallongilUdi nal
Inlerpedun(.u/<;)r nucleus iasciculus)
F.:lsciOJlus rel rollexu>

FIGURE 111.75: ANATOMY OF THE LIMBI C FOR EBR AIN

_ _ _ _ _ _ _ _ _ _ _ _ __
FI
Struc tures of the limbic forebrain are found in a via the stria meduUaris thalami. The cin gulate,
ri ng (limbus) that encircles the diencephalon. Two prefro ntal, orbitofrontal, ento rhinal, and periamyg
n
g)
major temporal lobe structures, the hippocampal daloid areas of the co rtex interconnect with sub
formation w ith its fornix and the amygdala w ith its
Ie
cortica l and hippoca mpal components of th e
ar
stria terminalis, send (-shaped axonal projections limbi c forebra in and are often co nsidered part of
er
.....J-hrou gh the cortex, around the diencephalo n, the li mbic system. Th e limbic system is thought to
~

and into the hypothalamus and the septal region.

sh
be a major substrate for reglilation of emotional
th,
The amygdala also sends a more direct pathway responsiveness and behavior, for individu alized
th,
(vent ral amygdalofugal pathway) into the hypothal reactivity to se nsory stimuli, and for integrated
pc
am us. The septa l nuclei sit just rostral to the hypo memo ry tasks.
da
thalamus and send axons to the habenular nuclei
2BB
Autonomic, H ypothalamic, and limbi c 5 ,,"rc-~~ - SYSTEMIC NELJROSCIE' CE

Body of
caudate
nudeus

'.;'.

Basal ganglia.

Temporal horn
of the lateral
ventrid e

CA regio ns of
hipPocampal
formation
(pyramidal cells)

J. Pericins
MS,MFA

:I '~N

FIGURE 111.76: HIPPO CAMPAL FQRMATI O

The hippocampal forma tion consists of the denU1ie
gyru s, the hippocampus proper (cornu am rnonis
[CAl region s), and th e subiculum. These structures
are in tima tely interconnected w ith the adjacen:
entorhin al cortex. The hippocampus is a seahor*
shaped structure found in the medial portion 0:
the anterior temporal lobe. II bul ges laterally into
the temporal horn of the latem] ventricle. Th e hip
pocampus is divided into several zones of p) ra m:""
dol cells, called CA regio ns (CA1-CA4). Granule
, ----------------
_ ce ntate gyrus. The dentate gyrus
~ pus are three-layered cortica l
- "'""'OOQm pal formation has extensive
;:;=1:1: _ ~.th cortical association areas
~ebr a in structures su ch as the
iiI"C :"l"le ci ngulate gyrus. It is invo lved
:"':::::=:=~ ,.,.- a rt term memory into long
- 'unc tio n with extensive regions

"1 89

SYSTEMIC NEUROSCIENCE Autonomic, Hypothalamic, and limbic Systems-Limb ic

of late ral ventricle

CA3--'-- - -
Mossy fiber ._:'/' / 1.. . ! Schaffer coUaterals

.
d J - ,>---- r

~
'" CAl
, fi . . . \ \ _~ Inputs to subiculum from
~~-'- ;:::::'::::::::;=:;=\;X'::::'=:---:-:-":7.:-::- J
J'
c ingul,lte cortex, amygdala
~. "", '
Subiculu m \',' ;.~ \ \ ...-- ~ Inp uts to ento rhinal corle",
(cingu i,lte cortex, se nsory
,lssociation cortices, baso
lateral dmygdala . in sular
cor tex, olfactory bulb, and
preiro ntaJ cortexi
Perfora n t pathway./""
from en torhin al cortex
to dentate gyrus, CA 1 and
CA3, and the sub iculum

Eniorhina i corlex
J, Perkins
MS. MFA
c~

FIGURE 111.77: NEURONAL CONNECTIONS IN THE HIPPOCAMPAL FORMATION

_ __
The hippocampal formation has an internal cir
cuitry interco nnected with the en torhinal cortex.
Pyramidal neurons of the entorhi nal cortex send
axons to granule cell dendrites in the dentate gyrus.
These granule cell axons (mossy fibers) synapse on
pyramidal cell dendri tes in CA3. Pyra midal cells in
CA3 project to pyramidal cell dendrites in CA 1
(Schaffer collat erals) and CA2. CA 1 pyra mi dal
axons project to pyramidal neuro ns in the subicu
lum. The subicu lum sends axonal projections back
_ _
to the pyramidal neurons of th e entorhinal cortex.
This in formatio n fl ow represents an interna l circuit.
Superimposed o n this circuitry are a host of inter
co nnections with associatio n regions of neocortex
and other limbic forebrain structures. Neurons of
th e subi culu m and CA neuron s of CA 1 and CA3
send axons into the forn ix as efferent projections to
target structures. Th e subicu lu m also sends axons
to the amygdala and the association areas of the
tem poral lobe.

290
.\utonomic, Hypothalamic, and limbic Systems-limbic

-\jjerenl connections-===:
: iierenl connections _

?os1I;ommhsural

o,ecommissural
'omix

(Of pus
MammJlio
callo sum tegmen t;,1
Iract

-\ociiltion
~ r"ilS of _--.0,. Input5 to
"onta l lohe hippocampus
W
_J-;;...-......-::;:- and dentate
5eptal nuclei - -- - - -:;; gyru5:
Raphe nuclei
(SHD
. Loc us coe ru leus
(NE)

Fimb ria

Inputs to entorhinal cortex:

Sensory associatioll cortices
Polysellsory association cortex
Prefrontal cortex
c ortex Insular corte,\
Efferems of suhio:: ulu m Denta te Amygdala
\0 amygdala, association g)' rus Periof.Jnt Olfactory bulb
areas of te mpvralluhe (A rellion_
o.a:h,. a' Inputs to subiculum:
of h ippocaMpI..,
Amygdala

FIGURE 111.78: MAIOR AFFERENT

FORMATION AND EFFERE'T CO , 'ECTIONS OF THE HIPPOCAMPAL
__________________________________________

Pyram idal neurons in the subiculum and the hip
poca mpal regions CA 1 and CA3 give rise to the
efferent fornix. The subiculum projects to hypo
thalamic nuclei (especially mammillary nuclei) and
thalam ic nuclei via the postcommissu ral fornix.
CA 1 and CA3 neuro ns send axons to the septal
nuclei. the nucleus accumbens, the preoptic and
anterior hypo thalamic regio ns. the cingulate cortex.
and the association areas of the fronlal lobe.
Afferent cholinergic axons from sepia I nuclei
traverse the fornix to supply the dentate gyrus and
tr.e ... ppocampal CA regions. Massive inputs arrive
in me hippocampal formation from sensory associ
ation cortices, polysensory associatio n co rtices, the
preironta.l cortex, the insular cortex, amygdaloid
nud ei, and the olfactory bulb via projection s !o
the entorhinal cortex. The ento rhinal cortex is
fully integrated into the internal circuitry of the
hippocampal formation. The subiculum is con
nected reciprocally w ith the amygdala and also
sends axons to cortical associa tion areas of Ihe
lemporallobe.
19 1
SYSTEMIC NEUROSCIENCE Autonomic, Hypothalamic, and limbic Systems-Limbic

TnalJmvs
i I
Bed nucleus of th e Cingulate cortex In lrJI ~m inar M edial dorsal
slri<l t erminahs
Stria lermina hs Brain stem inputs;
M Parabrac hial nuclei
"$. >;.'_ Periaqueduclill gray
Ve ntral tegmental area (DA )
.4#" Raphe nud ei (5Hn
j/ Locus coeruleus (NEI
/.;'J1" Nucleus trac Lus
Corpus callosum
1/
,,-:,j::
solitarius

Anteri o r '\
!/

Sept;!1 '""'~
nudei ' ,_ _ _
.""",
- - -~
~,o_" ,
Prefrontal I ---...
cor tex
. 4."

Hypothalamic inputs:
~{!'
,
l ate r",1hypo thalamic. area . ~-
Ventromedi al nucleus ~

Corticomedial nuclei /'

of the amygdala

Basolateral (hippocampal formati on)

of the amygdala
Sen sor y <l ssociation inpulS:
Te mporal lobe ,enso ry associatio n cortex
En(Qrhin;)1 cortex
Insular ( Ollex
Medial frontal lobe

FIGURE 111.79: MAIOR AfFERENT CONNECTIONS OF THE AMYGDALA _ _ _ _ _ _ __

Th e amygdala is an almond-shaped co ll ection of
nuclei in the medial portion of the anterior tempo
rallobe. It is involved in the emotional interpreta
tion of external sen so ry information and internal
states. It provides individual-specific behavioral and
emotiona l responses, particularly for fear and aver
sion. The amygdal a is subdivided into cortico
medial and basoJateral nuclei and the central
nucleus, w hich mainly provides efferent projections
to the brain stem. Afferen ts to the cortico medial
nuclei arrive mainly fro m subcorti cal limbic
sources, including the o lfactory bulb, septal nuclei,
hypothalamic nuclei (VM, LHA), th e thalamu s
(intralaminar nuclei), the bed nucleu s o f the stria
terminalis, and extensive autonomic monoamine
nuclei of the brain stem. Afferents to the baso
late ral nu d ei arrive mainly from cortica l areas,
including extensive sensory association cortices,
the prefrontal cortex, the cingu late cortex, and the
subicu lum .

292
- Autonomic, Hy pothala mi c, and Limbic Systems-limbic

Bed ("1\.'( .,,""... ; o ~i1f:

SYSTE M IC '\EL.,ROSClE'CE

To ~~
SIre.; .~- ilL , Stria " onio, li, "uC~""j :.~"' ~ ~~
Dorsa l ~ nt.;C..;- _~ r:i ..
Rapre -~ ~.....-
Stri atum (rostral areas)
Cauda te nucleu s
l o cuscO'"l.t"_> '=
Parabf01d1&ai rvoea

Putamen # Periaqueduct.l.l $-7_

Reticul ar forma:iof'

Corpo .
C2~

Fro ntal lobe regions:

Fron\31cortex
Prefro nta l cortex

nuclei

Nudeus - ------- --- __~~'-_'b

acc umben~

SubStolOti.1 inno mina ta (w ith

nucleus w sa1is ACh neuro ns)

Ventral amygd alo fu gal

pathway (VAf P)

earlico rnedial o" lei /

o f th e amygd al a cortex

Baso latera l nuclei

(e.g., visual are as)
of th e amygd a l<l
Central nuc:leus Of t.'""f' a"Tl'o .,;:dala
co rt ex

H - Hypothalamus: Th " Tha la m us:

~ff
Preoptic area M<?dia l dOf'SaI nuc't' us
Anterior hypothalamic area
Ventromedial nucleus
Laleral hypothalamic a rea
Paraven tricular nucleu s
M " M idline tha lamic nucl e i
..-...
~~

FIGURE 111.80: MAJOR EFFERENT CO NN ECTIO\,S OF THE AM YGDALA _ __ _ _ __ _

Efferents fro m the corticomedial nuclei project
through the stri a te rminalis and are directed mainly
tow ard subcortical nu clei such as septal nuclei, the
MD (DM) nucleus of the tha lamu s, hypothalam ic
nuclei, the bed nucleus of the stria te rminalis, the
nucleus accumben s, and the rostral striatum.
Efferents from th e basolateral nuclei project
tnrough the ventral amygdalofugal pathw ay to
=i\t ensive cortical regions, including the frontal
cortex, the ci ngulate cortex, the inferior temporal
cortex, the subiculum, and the en torhinal cortex;
and to subco rti cal limbic regio ns, includin g hypo
thalamic nucl ei, septal nuclei, and the ch olin ergic
nucl eus basa li s in th e substanti a inno min ata. The
central amygdalo id nu cleus receives input mainl y
from internal amygdaloid connections and sends
extensive efferents thro ugh the ventral amygdala
fugal pathw ay to extensive auto no mic nuclei and
monoaminergic nuclei of the brain stem, m idline
thalamic nuclei, the bed nucleus o f the stria
termi nalis, and the cholinergic nucleus basalis.
293
 SYSTEMIC NEUROSCIENCE Autonomic, Hypothalamicl and Limbic System5 -1.im2

Olfactory bulb
Se ptal nLfcie l
Hypo(h~l<lmlc nudei (lH A. VM )
Thalami c nud E' i (inlrillaminar) ----- -1.
I_~?'
-~ I
{.~, Jiit
.
~ ~:~t~IU~~:ll : i Oi ST
- Thala mu. (MD I
\-- - - - - - - - _ H yp ol h.l l am u_~
Red nucleus o( ST PreOI)Lic area
AHA
8 rain stem
Pa rabrolchial nucleu s YM
PeriaquedU<laJ gray
LH A
PYNp
VTA (DA)
Ra phe nud ei (5HT) .Nucleus atTumb~n~
locus seruleus (N EI +---------~-
: Stri a tum (f()~tra ll
NTS Ca udate
Pu tamen

@Fa : Mi~li ne Ih~l amic Olld ei

Kliilf-------- - - 81 aln stem
Red nuclNI$ or ST
Sub~ tanna innomin,lla
I nuci\;" us b,19lis)
NTS
OMNofx
fl.aph~ nud ei (SHn
Lo cu >l:ue rulu s (N EJ
P<trabrachi.lI nucl~i
Periaqu educl al gray
Re ticul ar for matiun

Th ala mu s (MDI ~, Su b.tanti a inn omln " t~

Pre fron tal corl ex
~; (n ucleus ba ~li5J
}; Sept<lll1u c!ei
Cingul ate corte)!; >~~.

Subiculum --------------1 '~'J---------- H ypo thitlamus

;." Frontal cortex
Seno:.ory associ,llion curtex
C b I Temporal lobe sensory assac CO rlex J---------~~ En lorhinJI co rtex
ere fa Enlorhinal corlex \-- - - - - - - _ Subic ulum
corlex { Ensul.:u cortex r------- - (i nRula te (o rlex
Med ial fro nlal cortex
r--------- Infcrio r te mporal (orteJC

DA .. Dopamine M D - Medial Ull rSillllucleus o llha l;;mlU5

5HT - 5Hydroxyn ypl amine (Serotonin) AHA - Anterior h)'poth al amic (If!:'<l
NE - I~ ore pin<:!p hri n e VM - Ventrom edial
5T - St ria terminali s lHA - l ateral hypulnaJamit area
lmN
... " ". VTA .. Ven tral 'e~men t al are a PVNp " f'a ravenlricu lar nu c1eu ~. parvocellular
..." .. NTS = Nud eus tractu s solilMiu s DM N of X '" Dors al moto r (autonomic) nucleus of X

fiGURE 111.81 : SUMMARY Of MAJOR

OFTHEAMYGDALA AffE RENTS, EFFERENTS, AN D INTERCONNECTIO NS
________________________________________

The corticomedial amygdala is connected recipro massive descending efferen ts to extensive au to

cally mainly with subcortical lim bi c forebrain struc
tures, w ith ex tensive additional inputs from brain
stem autonomic and monoaminergic nuclei. Th e
basolateral amygdala is connected reciprocally
with extensive regio ns of limbic and association
cortex, w ith additio nal efferen ts to subcortical
limbi c fo rebrain regions. Both the corticomedial
and the basoJateral nuclei send axons to the central
nucleu s of th e amygdala. The ce ntral nucl eus has
294
nomi c and mon oamin ergic nuclei of the brain stern
as well as to some subcortical limbic forebrai n
regions. These in terconnectio ns provide the inte
grated circuitry that permits analysis of both exter
nal and internal inform ation, w hich provides an
emotional and interpretative context for the initia
tion and control of appro priate behavioral and
emotional responses.
Autonomic, Hypothalamic, and Limbi c S..-s1 '-5. SYSTEMI C NELROSCIE'CE
AFFER ENTS

Major Olfferents fmlll.

HippocamPJI CA pyramidal ct' ll s
A mygciai()id 11ucit'i
Corilcoml!diili nuclE'i \i " stria Termlnalis

Basolatcral nudei via venTr,,1

ilm \, grlil lofugal path way

Vent ra l tegme nta l JreJ

Hypothalamus

Preop tl<- ,lrc')

Anteri or h\pothai amir Mea

P;u,lVE'nlricui<ll nucleus

l ater,,1hypolh;damic <lfea

l ocus cO Nule(ls (NE; no t show n I

e'11rJI I{'gmenlal alea

,., ";,,1111,,,; th alami

EF FERENTS nuclei

Majorc(fe ren'~ roc

Hippocamp JI CA regiOnS } . . .

Via ltlrnlX
DenTilTe g~'ru~ (ACh path)
Habenular nuclei } Via sui., ( ~
Medial dors al nucleus medullaris
(ithe tha lamus tha lami
Vemral tegmental Mea - Via median I ,
H~'Poth alamu s forebrain bUI1{lie
Preop tic area
A nterior hYPOlhillilmic area
Ventromedial nucleus
laleral hypo th" l,l miC are~

(medial dQrsalJ

r - _~ .e

FI G U RE 11 1.82: M AJOR AFFERE NT AND EFFERE TCO"ECfION S OFTHE SEPTAL NUClEI _ _

The septal nuclei are subcortical nudei implicated
by earl y ablatio n and stimulatio n stu di es in the
regulatio n of emotio nal responsiveness (such as:
rage be havior) , In experim ental studies, the septal
nuclei app ear to playa ro le in emotional beha\ iors.
sexual behavior, aggressive behavior, modulation
of auton o mic function s, and aLtention and memor\'
functions (from the cholinergic neuro ns). At1erents
to the septa! nuclei arrive mainly from th e hippo-
carnpo!o the corticomedial and basolateral
aFl1\ goala. the ventral tegmental area in the mid
bra-n and several hypothalamic nudei. Efferents
(rom the septal nucl ei proj ect mainly to th e
hippocampus and the dentate gyrus (via the
iorni"\ , th e habenular nu clei and th e M D (OM)
nucleus 01 the thalamus (via th e stri a medullari s
thalami ), the ven tral tegmental area (via the median
io rebrain bundle), anel several hypotha lamic nuclei.

295
SYSTEMIC NEUROSCIENCE Autonomic, Hypothalamic, and limbic Systems-limbic

Afferent con nections - - - _ ( inguJ3te cortex

Efferent connections - - -_ Association areas o f
parietal cortex
.,,,.
"
,.t/'(,'
"""',>.~".",-.,; .. .Ao
-Ij}.;:t'''-

Corpus callosum
",,;
~
" ,'
-{.

Asr.ocialion
areas of homal
corteK

Mammillary body

~~f'
Amygdala
(basolate ral nudei)

A NT = AnteriOl' nudei OJ the thalamus

o~
.
l ~U" .
~

MD = Medial dorsal nucleus oi the thalamus

FI GU RE 111.83: MAIOR CONNECTIO NS OF THE CINGUlAT E CO RTEX _ _ _ _ __ _ __

The cingulate cortex is lo cated above the corpus
callosum. This cortical region is involved in the reg
ulation of autonomic functions (respiratory, di ges
tive, cardiovascular, papillary), so me somatic
function s (motor tone, ongoing movements), and
emotion al responsiveness and behavior. Lesions of
the cingulate cortex result in indifference to pain
and o ther sensations w ith emo tional connotation
and social indifference. Afferents to the cingu late
cortex arrive fro m association areas of th e frontal,
parietal, and temporal lobes, the subiculum, septal
296
nuclei, and thalamic nuclei (MO, anterior). Efferents
from cingulate cortex neurons pro ject to associa
tion areas of the frontal, parietal, and temporal
lobes and to lim bic forebrain region s such as the
hippoca mpus, th e sub iculum, th e ento rhinal cortex,
the amygdala, and sep tal nuclei. These limbic fore
brai n regions send extensive projectio ns to the
hypothalamus for regulation of the autono mic
and so matic regi ons of the brain stem and the
spinal co rd.
 4utonomic, Hypoth alamic, and limbic Systems-olfactory SYSTE M IC :-'EL.ROSCIE' CE
-

Olfactory Receptors ;;;:~::bulb

~ A. Oi ~t ri bulion oi
oliado r.... epithelium
t b _~c ':;',,"'"

Septum
B. Schema of
section through
~ olf.cto., muw"

... . Cribriform plate

.;-- - -- - - Schwann cell

Ii t
axons
_____ Basement
~ membrane

-
>
- --'- -- - ViUi
.-"

FIGU RE 111.84: O LFACTORY REC EPTO RS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Olfactory receptors are found in a patch of o lfac
tory e pithelium that lines the medial and lateral
w alls of the roof o f the nasal cavity. They are primi
tive specialized bipolar neuro ns w hose nuclei are
in the base of the epithelium. A dendritic process
extends toward the epithelial surface, w idening
into a rod w ith 10 to 30 motile cilia that project
into the mu cous cover. Odorants act on receptors
(G-protein- cQup led) on th ese cilia and bring abou t
a slow depolarizin g generator potentia l. O dorant
in teraction wi th receptors is complex, often req uir
ing odo rant-binding proteins to carry the odorant
through the mucosa. The bipolar neurons of the
oliacto ry epithelium are e NS neurons; they are
unusual beca use they undergo continuous replace
ment and turnover from basal stem cells in the
epilhelium. The UNM olfactory axons cl uster
IOgelher in groups (enwrapped by a Schwann cell)
before passing Ihrough the cribriform plate. Inj uries
to th e cribriform plate can result in anosm ia.

297
SYSTEMIC NEUROSCIENCE Autonomic, Hypothalamic, and Limbic Systems-olfactory

~~
:::~} Efferent r,b ers
' '::::

(; : : ; : : ; ) A"",o, "b",
Granule <::oell Ie.><eited by
Fibers from con tralateral o lfactor y bulb
and inhibitin g 10 miffal I
<lnd tuited cdls! to contralateral olfactory bulb
Mitralc:ell
-;"-' An terior oommis~ure
Re>current plOCeSS .~

..JH.e/ 1>.....-1 Tuited cell ,y..P~"

Med ial olfacto ry stria

I"erig(om / '
er ulMcell i!
,/ Anterior perforated
G'om,,,'"

~
substan ce
O lfactory

'I ~ ""V< fib", I olfactory stria

lateral olfactory
tract nucleus

,
O lfaclory nerve~
, Anterior olfac1ory nucleus
Crib riform plale of ethl'llOid bone

~~p
l~
"'...... n

FIGURE 111.85: OLFACTORY PATHWAYS _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

Primary sensory axons from bipolar neurons pass
through the cribriform plate and synapse in the
olfactory glo meruli in the glomerular layer of the
olfactory bulb. The glomeruli are the functional
units for processing specific odor information.
The olfactory nerve fibers synapse on the dendrites
of the tufted and mitral cells, the secondary
senso ry neurons that give rise to the olfactory tract
projections. Periglomerular cells are interneurons
that in te rconnect the glomeruli. Granule cells
modulate excitability of tufted and mitral cells.
298
Centrifugal co nnections (from serotonergic raphe,
nor adrenergic locus coeruleus) modulate activity in
the glomeruli and o n peri glomerular cells. The
o lfacto ry tract bypasses the thalamu s and projects
to the anteri or olfa ctory nucl eus, the nucl eus
accumben s, the primary o lfactory co rtex, the
amygdala, the peri amygda Jo id co rt ex, and the
lateral entorhinal co rtex. Th e o lfactory cortex has
interco nn ectio ns with th e orbitofrontal co rtex and
th e insular cortex, th e hippoca mpu s, and the lateral
hypothalamus.
tor\'
IN DEX
A de~ e opm m , 75
Associational cortex, : '?'? ~ Cacd
Abdominal nerves, 129
enerentconn ec~,1 9 3
<Jssoc[ation

Abducens nerve, 157. 158, 160.2 51

interconnections. 2Q ..:
Astrocvtes, 7 8, 71

ACA. See Ante rior cerebra l arte ry

Am)'gdaloiugai pa lh\,a... 293
Athe rosclerosis, 47

Accessory nerve, 15 7, 158. 166

AmygdaJo,d nudei, ]O ]93
Auditory cortex, 24

Acetylcholine
AnastolT\O<oeS
Auditory system

enteric nervous system a nd. 132

arte rial ~uppk. 10 br<;irl 50
afferent pathways, 228

synapses. See Cholinergic synapses

veno u" rings around \ er..ebr.:.t'!. 66
bony and membranous labyrinths. 21 ';:

Ac tion poten hals. 16

Anenceph.aJ... 68
centrifugal (efierentl pathways, 229

conduction velocity, 17
Anesthesla_ den--...r()('l" 3- <;1,3
cochlear receptors, 22 7

propagation. 17
Aneurysm. cere!:lra. 5 :
frontal section, 224

Adenohypophysis. 177
Anions. l -J.
ne rve innervatioll of hair cells in, 226

AOH (a ntidiuretic hormone). See

ANS. See AIJl()!'lQl'l""C "E'I"\ O~~~, ~iem
ves tibular recepiors, 230

Vasopressin
An teri()( c er~.;r;:;:""\ .!,?
Auerbach's (myenteric) plexus, 132, 133

Adrenal gland, 135

circle oj \ \ 'iIis and j(
Autonomic channe ls, 91

Adrenergic distribution, 120

occlu sion. 5]
Autonomic nervous system, 116

d
Adrenergic synaps5, 118, 120
separalion,50
cnoJirler gic and adrenergic
Afferent pathways. 294
ID'les suppi'ed 0.. 53
distribution/synapses, ' 19, 120

amygdala, 292
Anterior lobe, cerebel..a: ~ B _ enteric. See Enteric nervouS syste m

audi tory system. 22 8

Anterior whit e cOllL....,~~ = -;. :?\ .l eye. 124

brain stem, 118

spinocere .?J' tr"iId and ~ 15
head and neck. 123

cerebenar pecluncles. ' 74

Antidi uretic hornone. ~ '.2:X)OI'e-;SIn
l ateraJv i~w, 122

cerebellum, 255, 256

Antid ro mic co nduction. 'of>. - ' 9
medial view, 121

ce rebral cortex. 201

Aortic bodv, 25]
innervation. See Innervation, autonomic

cingulate conex, 296

Appetite, 285
orga nization, 264

classification, 18
APs. See Action po:~ "a ' ;
reTle>:; pathways, 118

cranial ne rve nuclei, 15 7

Aqueous humo r, 13]
Autonomic neurons

enteric, 133. 134

Ara chnoid
postganglionic, 71, 86, 264

eye, 124
meninges a nd. -12 preganglionic, 74. 86

heart,. 128
spinal cord. 39. .;0
as basal plate de rivatives, 80

hippocampa l formation, 29 1
Arachnoid cells. , 1
development in brain stem and spinal

hypothalamic, 271, 272

Arachnoid gran ulatioru . .!2
cord,79

intestinal, 131
cerebrospinal l1uid e:"Ct.,...a'q 2'1d ..: ;; o utflow regulation. 275

kidney and upper urete r, 137

ventricular de\elopmen~.:.'1C - Axillary nerve, 103

mouth and pharynx, 168

ARAS (ascendi ng re ocu1ar <=<;""'.,,'-1'13 above e lbow, 104

Olfactory, 298
5~lem J. 171
Axoaxo nic synapse, 4

reproduc tive o rgans

Afea postfema. 277
Axodendritic synapse. 4. 5

femal e, 140
Arousal, brain areas assocated Iqlh, 1-2
Axon hillock, 4

male, 139
Artery!ies)
Axon membrane, bare, 10

septal nudei, 295

to brilio, 50
A..xons, 4, 7

spinal cord, 118

lateral and medial "e".'.~, 5 2
autonomic, 82

somatosensory, 114
carotid,47
central, 10

tracheo bronchial tree. 12 7

cerebral, 48. 49
commissural, 203

urinary bladder and lower ure ter, 138

cervical cord, 64
conicobuJbar tract, 243

vagus ner ve and. 165

communicating, circle oi \ \ ~1Ii ;; and. 51
corticoc:ortical, 203

AICA (anterior inferio r cerebellar eye, 236

corticothalamic, 203

,merY),55 hypophyseal. 56
ensheathment, 9

Alar plate
meningeal. 42, 46
glossopharyngeal nerve and, 1b4

as brain stem deriva tives. 80

~o( st roke, ~ 49
lower motor neuron, 96

in early brain development

pOSterior cranial iossa ' . 55
myelination of, 8

28-day stage, 72
radicul.. r, 63, 65
peripheral. 82

36<1ay stage, 73
skull base fOfamina and. 22
projection, 203

embryo nic vs. adul t comparisons. 78

spinal cord, 64
synaptic morphology and, 1 I

neura l tube development ar'ld, 69

cro.ss-sectional view, 65
terminal. 96

(l-iower motor neurons. 241. 245, 24&. 24 7, diwibulion, 64

unmyelinated vs. myelinated, 214

248,249
loogiludinal view, 63
Axosomatic synapse. 4, 5

Amacrine cells, 235

verte bral,4 7
Axosomatic !ermmal, 11

Amino dcid synapses, 12

ASA (anterior spinal artery ). 63, 64, &5

Amygdala, 275

afferents, 292

Ascending pathways, spinal cord. 41

Ascending reticular activating system, 171

Bare axon membrane, 10

basolateral,294
Association fibers, 204
[3arorece pto r~, 94

corticomedial.29 4
cerebral cortex, 202
Basal gangha, 29

199