Female Pelvic Medicine & Reconstructive Surgery from Journals@Ovid About This Journal

1 March 2014 | Volume 20 | Issue 2 | p 6575

doi: 10.1097/SPV.0000000000000058

Neuroanatomy, Neurophysiology, and Dysfunction of the Female Lower Urinary Tract: A

Review.

Unger, Ccile A. MD, MPH; Tunitsky-Bitton, Elena MD; Muffly, Tyler MD; Barber, Matthew D. MD, MHS

Article Outline

Article Information

From the Center for Urogynecology and Reconstructive Pelvic Surgery, Cleveland Clinic, Cleveland,
OH.

Reprints: Ccile A. Unger, MD, MPH, Department of Obstetrics, Gynecology, and Womens Health
Institute, Cleveland Clinic, 9500 Euclid Ave, A81, Cleveland, OH 44195. E-
mail: ungerc@ccf.org; cecile.a.unger@gmail.com.

The authors have declared they have no conflicts of interest.

Abstract

NEUROANATOMY AND PHYSIOLOGY

These processes involve the coordination of neural events in both the central and peripheral nervous
systems. The central nervous system is composed of the brain and spinal cord, whereas the
peripheral nervous system is composed of the autonomic (visceral, involuntary) and somatic
(voluntary) systems.(3) The 2 types of neurons that are found in these systems are the afferent
(sensory) and efferent (motor) neurons (Table 1). In the lower urinary tract, the autonomic nervous
system is mediated by the sympathetic and parasympathetic nerves, whereas the somatic system is
mediated by the pudendal nerve.8 Parasympathetic nerve fibers emerge from the sacral portions of
the spinal cord, whereas the sympathetic nerves emerge from the thoracic and lumbar portions of the
spinal cord.3

The autonomic nervous system involves a sequential 2-neuron efferent pathway (Fig. 1). A
preganglionic neuron synapses onto a postganglionic neuron before innervating the target organ. The
sympathetic division arises from the thoracolumbar spinal cord (T12-L2) 8 and consists of cell bodies in
the lateral horn of the spinal cord called the intermediolateral cell columns. 7 These cell bodies are
general visceral efferent neurons, and serve as preganglionic neurons that synapse on their
respective postganglionic neurons. These neurons are the paravertebral ganglia of the sympathetic
chain found on either side of the vertebral bodies or the prevertebral ganglia (also known as the pelvic
plexus or the inferior hypogastric plexus), located bilaterally on the walls of the rectum (Fig. 2). The
hypogastric nerve is the postganglionic nerve that travels inferiorly and descends into the pelvis to
form the inferior hypogastric plexus.9 It releases norepinephrine which activates the -adrenergic
inhibitory receptors in the bladder causing detrusor muscle relaxation and the -adrenergic excitatory
receptors in the urethra and bladder neck resulting in the contraction of these muscles. 10

The parasympathetic division arises from cell bodies in the ventral rami of the sacral spinal cord (S2
S4).(8) These cell bodies are the cholinergic preganglionic neurons that exit the spinal cord in the
ventral spinal nerves forming the pelvic nerve.(7) They then synapse with the postganglionic neurons in
either the pelvic plexus or the intramural ganglia found within the bladder.(7) The pelvic plexus is a
paired structure, containing both sympathetic and parasympathetic fibers, with each pair situated on
the side of the rectum and the vagina. The postganglionic nerves release acetylcholine which
activates the excitatory muscarinic (M3) receptors of the bladder, resulting in detrusor muscle
contraction and urine flow.(8) Other postganglionic nerves release nitric oxide which acts on the
smooth muscle of the urethra, causing relaxation and facilitating voiding. (10)

The pudendal nerve arises from the ventral rami of the sacral nerves (S2S4). It passes between the
piriformis and coccygeus muscles and leaves the pelvis through the lower part of the greater sciatic
foramen. It crosses behind the spine of the ischium, and reenters the pelvis into the ischiorectal fossa
through the lesser sciatic foramen. It accompanies the internal pudendal vessels upward and forward
along the lateral wall of the ischiorectal fossa, and it is contained in a sheath of the obturator fascia
termed the pudendal canal (Alcock canal).(9) The pudendal nerve is a somatic cholinergic motor nerve
that activates the striated external urethral sphincter muscle as well as the muscles of the pelvic floor
and external anal sphincter.(8) The pudendal nerve gives off 3 branches in the perineum, namely, the
inferior rectal nerves, the perineal nerve, and the dorsal nerve of the clitoris. (9)

The regulation of micturition is a complex interaction between the autonomic and somatic nervous
systems and the brain, whereas the spinal cord acts as an information transmitter.(7) Sensation of
bladder fullness is conveyed to the spine via the pelvic and hypogastric nerves, whereas sensory
input from the bladder neck and the urethra is carried by the pudendal and hypogastric nerves.
(8,11)
When the bladder fills, these afferent nerves from the bladder and the urethra transmit signals to
respective preganglionic neurons located in the lateral dorsal horns of the spine. (8) Interneurons
project from this area to different regulatory centers in the gray matter of the brain including the
Barrington nucleus in the ponsoften referred to as the pontine micturition centerand the
periaqueductal gray cell bodies of the hypothalamus.(8,12) Of note, there are 2 main types of fibers
carried in the afferent nerves, namely, A- fibers, which contain myelinated axons and transmit low
threshold signals such as bladder filling; and C fibers, which are unmyelinated and transmit noxious
stimuli such as irritation or coldness.(3,7,11) The cell bodies of both fibers are found in the dorsal root
ganglia of the spine at T11-L2 and S2-4. These cell bodies synapse with neurons that transmit
information to the brain involving regulation of bladder function. (8)

When the bladder fills, detrusor muscle activity is thwarted via inhibition of the parasympathetic
nervous system, whereas both the smooth and striated muscles of the urethra are activated. This
controls filling and prevents involuntary emptying and is mainly controlled by spinal reflexes in the
spinal cord that are triggered by the afferent pelvic nerves coming from the bladder. (8) The lateral
region of the pons is also involved and it helps facilitate the activation of these spinal reflexes.
(13)
Interneurons in the spinal cord transmit bladder filling sensory signals to the periaqueductal gray
cell bodies of the hypothalamus, which in turn, signal the pontine micturition center together with the
prefrontal cortex.(14) Activation of the cerebral cortex facilitates awareness of bladder filling and allows
for suppressive signals to be sent to the pons to prevent activation of the micturition center which is
responsible for voluntary voiding.(15)

Excitation of the pontine micturition center through voluntary signals from the cerebral
cortex(3) activates descending spinal pathways leading to urethral relaxation and detrusor muscle
contractility. This results in urine flow and emptying. (8) The descending signals activate the
parasympathetic sacral nerves leading to the release of acetylcholine at the muscarinic receptors of
the bladder. At the same time, there is a release of nitric oxide and removal of adrenergic signals at
the level of the urethra, allowing both smooth and striated muscle relaxation. (8) These mechanisms
facilitate voluntary emptying of the bladder.

Case Report on Hiccup and Lateral

Medullary Syndrome
Mohamed Hamdy Ibrahim1, Alyaa Fadhil2, Sameh Saied Ali3, Salma Fathy
Abdel Kader3,
Mohamed Khalid4, Kiran Kumar4, Shivram Kumar5, Janhavi Sirsat5
1
Department of Neurology, GMCH & RC, Ajman, United Arab Emirates
2
Department of Clinical Pharmacology, GMCH & RC, Ajman, United Arab Emirates
3
Department of Radiology, GMCH & RC, Ajman, United Arab Emirates
4
Department of Internal medicine, GMCH & RC, Ajman, United Arab Emirates
5
Gulf Medical University (GMU), Ajman, United Arab Emirates
Email: mohamedhamdy_neuro2007@yahoo.com
Received 31 March 2015; accepted 6 June 2015; published 9 June 2015

the major swallowing centers of

the nucleus tractus solitaries (NTS) and nucleus ambiguous (NA) and the reticular
formation around them are
located in the dorsolateral medulla oblongata

Differentiate into Schwann Cells in

Fibrin Gel as 3D Culture
Neda Bayat1, Somayeh Ebrahimi-Barough2*, Mohammad Mehdi Mokhtari
Ardakan2,
Jafar Ai1,2*
1
Brain and Spinal Cord Injury Research Center, Neuroscience Institute, Tehran University of
Medical Sciences,
Tehran, Iran
2
Department of Tissue Engineering and Applied Cell Sciences, School of Advanced
Technologies in Medicine,
Tehran University of Medical Sciences, Tehran, Iran
Received 13 November 2015; accepted 21 December 2015; published 24 December
2015

Schwann cells (SCs) have important roles in development, myelination and

regeneration in the peripheral nervous system. Schwann cells are fundamental for
development,
myelination and regeneration in the peripheral nervous system

Lower motor neuron dysfunction in ALS

Mamede de Carvalho, Michael Swash
PII: S1388-2457(16)30006-2
DOI: http://dx.doi.org/10.1016/j.clinph.2016.0
Reference: CLINPH 2007794
To appear in: Clinical Neurophysiology
Accepted Date: 1 March 2016

The final effector component of the lower motor system is the alpha motor neuron (lower
motor neuron - LMN). The axons from these LMNs, known as anterior horn cells in the
spinal cord, project through peripheral nerves to innervate skeletal muscle fibres (Burke,
1981). LMNs are modulated by the corticospinal tracts and also a number of polysynaptic
tracts with different origins, such as pathways projecting from the vestibular, tectal and
rubral nuclei, and from the reticular substance (Burke, 1981; Pierrot-Deseilligny and Burke,
2005). In the spinal cord, local segmental connections to LMNs from direct afferent
sensory input, and from interneurons, modulate peripheral sensory input and descending
projections from higher centres (Jankowska, 1992; Pierrot-Deseilligny and Burke, 2005)
Voluntary and reflex movement programmes are transmitted to muscles through
large, rapidly conducting motor axons from the LMNs - Sir Charles Sherringtons final
common pathway. In the spinal cord the LMNs innervating individual muscles are
organized in columns located in a precise mediolateral topography reflecting their function
and the distal or proximal anatomical location of the particular muscle (Routal and Pal,
51999)

Review
Branched thalamic afferents: What are the messages
that they
relay to the cortex?
R.W. Guillerya,, S. Murray Shermanb
aMarmara University, Istanbul, Turkey
bThe University of Chicago, IL, USA
BRAINRESEARCHREVIEWS66(2011)205219

If the ascending axon that goes to the posterior column nuclei is now looked at from the point of view of its
relationship to the branches that are innervating spinal reflex mechanisms, it has to be regarded as an
efference copy. It carries a reliable representation of the instructions that produce the refle. The medial
lemniscus and the optic tract are sensory pathways bringing messages to the brain about the outside world. For
the ascending spinal pathways considered so far there is no question that the message they carry is relayed to
the cortex. Although the terminals of thalamic afferents that carry messages for relay to the cortex, the
drivers (Sherman and Guillery, 1998), represent only a minority of the afferents in any one thalamic nucleus
(generally less than 10%), they have a characteristic light and electron microscopical appearance in all major
thalamic nuclei and are characterized by comparable patterns of multiple synaptic junctions, with well defined
transmitter and receptor characteristics. (Sherman and Guillery, 1998).

Respiratory Physiology & Neurobiology 169 (2009) 157164

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Respiratory Physiology & Neurobiology

journal homepage: www.elsevier.com/locate/resphysiol

Autonomic function following cervical spinal cord injury

Andrei Krassioukov
International Collaboration on Repair Discoveries (ICORD), Department of Medicine, Division of Physical Medicine and
Rehabilitation, University of British Columbia,
GF Strong Rehabilitation Centre, Vancouver Health Authority, 818 West 10th Av, Vancouver V5Z 1M9, BC, Canada

There are two components within the autonomic nervous system (ANS): sympathetic and parasympathetic
(Krassioukov and Weaver, 1996a,b). The sympathetic and parasympathetic systems interact with each other
within the central nervous system and provide balanced regulation of innervated organs. Autonomic control
occurs via the coordinated function of the central (cortex, hypothalamus, brainstem, and spinal cord) and
peripheral (autonomic ganglia and receptors) autonomic circuits. Although there are obvious functional
differences between the sympathetic and parasympathetic nervous systems, there are also some similarities in
their organisation. In both systems there are two neuronal populations interposed between the central nervous
system and target organs. The first neuron is known as the preganglionic neuron, with the cell body within the
gray matter of the brainstem or spinal cord. Axons of these neurons, called preganglionic fibers, travel within
the ventral roots of the spinal cord or cranial nerves. Both sympathetic and parasympathetic preganglionic
neurons are cholinergic. These fibers release acetylcholine and synapse on the second group of neurons, called
postganglionic neurons. This group of neurons is located within the autonomic ganglia of the peripheral nervous
system. The axons of these neurons are called postganglionic fibers. These fibers have direct connections with
their target organs. Sympathetic postganglionic fibers are predominantly adrenergic and release norepinephrine
(with the exception of cholinergic sympathetic fibers innervating sweat glands and piloerector muscles, Table
1). All parasympathetic ganglionic neurons are cholinergic and release acetylcholine at the target organ
Sympathetic preganglionic neurons reside in the spinal gray matter in the thoracic (T1T12) and upper lumbar
segments (L1L2, Fig. 1) of the spinal cord (Schramm et al., 1993; Krassioukov and Weaver, 1996a,b). The
majority of sympathetic preganglionic neurons are localized within the lateral horns or intermediolateral
nucleus of the spinal cord. A small proportion of sympathetic preganglionic neurons are found near the central
canal of the spinal cord. Axons of the sympathetic preganglionic neurons exit through the ventral roots and
synapse on postganglionic sympathetic neurons located in the spinal paravertebral ganglia (the sympathetic
chain ganglia) and prevertebral ganglia (the celiac, superior, and inferior mesenteric ganglia). The
postganglionic neurons then send their axons, via the peripheral nerves, to innervate the target organs,
including the heart, blood vessels, respiratory tract, sweat glands, sexual organs, and smooth muscles within
the gut and bladder. Parasympathetic preganglionic neurons are located within the nuclei of four cranial nerves
(CN III, VII, IX, X) in the brainstem and within the sacral spinal segments (S2S4). Parasympathetic control of the
cardiovascular and broncho-pulmonary systems and the upper portion of the gastrointestinal tract occur via the
vagus nerve (CN X), which exits from the brain through the base of the skull. There is no parasympathetic
innervation of the peripheral vasculature except in the pelvic organs. Parasympathetic innervation of the
bladder, reproductive organs, and lower portion of the gut is provided by the sacral portion of the spinal cord
(S2S4).

Research Paper
Acute spinal cord injury (SCI) transforms how GABA affects
nociceptive sensitization
Yung-Jen Huang a,, Kuan H. Lee b, Lauren Murphy a, Sandra M. Garraway c, James
W. Grau a
a Behavioral and Cellular Neuroscience, Department of Psychology, Texas A&M University, College Station, TX 77843, USA
b Center for Pain Research, Department of Neurobiology, University of Pittsburgh School of Medicine, Pittsburgh, PA 15213, USA
c Department of Physiology, Emory University School of Medicine, Atlanta, GA 30307, USA

Experimental Neurology 285 (2016) 8295

Aferent fibers that transmit signals related to noxious stimulation project to the dorsal horn of the spinal cord
and engage pain (nociceptive) circuits that are regulated by descending fibers and intraspinal biological
processes (Woolf, 2004). Of particular concern are processes that foster neural excitability within the dorsal
horn, generating a sensitized state that increases behavioral reactivity and enhanced pain. Research has shown
that prolonged activation of unmyelinated (C) fibers can induce a lasting central sensitization that is mediated
by molecular signaling pathways related to those implicated in brain-dependent learning and memory (Ji et al.,
2003; Woolf and Thompson, 1991). Central sensitization can be induced by electrical stimulation of nociceptive
fibers, peripheral injury, or the application of an irritant (e.g., formalin capsaicin). At a behavioral level, the
sensitization of nociceptive circuitsis associated with increased reactivity to mechanical stimulation (Woolf,
2011). At a cellular level, central sensitization within the spinal cord is correlated with increased expression of
the immediate early proto-oncogene c-fos and the phosphorylation of the protein extracellular-signal-regulated
kinase (pERK; Gao and Ji, (2009)).

Complex and interacting influences of the autonomic nervous

system on
cardiac electrophysiology in conscious mice
Heidi L. Lujan, Joshua P. Rivers, Stephen E. DiCarlo
a Wayne State University School of Medicine, Department of Physiology, 540 E. Canfield Ave, Detroit, MI 48201, USA

Autonomic Neuroscience: Basic and Clinical xxx (2016) xxxxxx

Corresponding author.
E-mail addresses: hlujan@med.wayne.edu (H.L. Lujan), jrivers@med.wayne.edu
(J.P. Rivers), sdicarlo@med.wayne.edu (S.E. DiCarlo).
AUTNEU-01877; No of Pages 8
http://dx.doi.org/10.1016/j.autneu.2016.08.017
1566-0702/ 2016 Published by Elsevier B.V.
Contents lists available at ScienceDirect

Autonomic Neuroscience: Basic and Clinical

journal homepage: www.elsevier.com/locate/autneu

Heart rate is slowed by parasympathetic nervous system activity via the muscarinic M2 receptor (Fisher et al.,
2004) and elevated by sympathetic nervous system activity via the beta 1-adrenergic receptor (Rohrer et al.,
1998). In addition, the sympathetic and
parasympathetic divisions of the autonomic nervous system alter heart rate and cardiac electrophysiology
through complex and interacting mechanisms

Central control of visceral pain and urinary tract function

Thelma A. Lovick
PII: S1566-0702(16)30006-6
DOI: doi: 10.1016/j.autneu.2016.02.001
Reference: AUTNEU 1815
To appear in: Autonomic Neuroscience: Basic and Clinical
Received date: 16 February 2015
Revised date: 17 August 2015
Accepted date: 1 February 2016

Afferent input from A and C-fibres innervating the urinary bladder are
processed differently by the brain, and have different roles in signaling bladder
sensation. A fibres relays in the midbrain periaqueductal grey (PAG) and
pontine micturition centre (PMC). Under physiological conditions C-fibre bladder
afferents are normally silent but are activated in inflammatory bladder states
and by intense distending pressure. Following prolonged stimulation visceral
nociceptors sensitise, leading to a lowered threshold and heightened sensitivity.
In addition, sensitization may occur within the central pain processing circuitry,
which outlasts the original nociceptive insult. Visceral nociception may also be
influenced by genetic and environmental influences. Afferent information on
filling status is relayed to the brain by a group of neurons in the lateral funiculus
of the upper sacral cord, just lateral to the dorsal horn.

Sensory and circuit mechanisms mediating lower urinary tract

reflexes
Zachary C. Danziger a,, Warren M. Grill a,b,c,d
a Department of Biomedical Engineering, Duke University, Durham, NC, USA
b Department of Neurobiology, Duke University, Durham, NC, USA
c Department of Surgery, Duke University, Durham, NC, USA
d Department of Electrical and Computer Engineering, Duke University, Durham, NC, USA

Autonomic Neuroscience: Basic and Clinical xxx (2015) xxxxxx

some neurons are activated selectively by different types of mechanical perturbation, while some respond
exclusively to chemical irritation. lectrophysiological studies of unmyelinated afferents located in the dorsal and
ventral roots show that afferent fibers that were unresponsive to mechanical stimulation of the detrusor did
respond to the chemical irritant mustard oil (Hbler et al., 1990), capsaicin (Shea et al., 2000), or cold (Bors and
Blinn, 1957). These afferents are commonly referred to as silent because they are not active under
physiological continence or micturition conditions. Complicating a bipartite distinction, however, is the fact that
some unmyelinated chemosensitive fibers also respond selectively to high bladder pressures (Hbler et al.,
1990), opening the possibility that these fibers could be multi-sensory in nature or that chemosensitive fibers
can locally influence the action of mechanosensitive afferents at high pressures. A mechanistic basis for the
difference in sensitivity between high and low threshold afferents has yet to be established, and several
observations suggest that the development of such an explanation will be challenging. Most notably, there are
a range of thresholds in the high and low threshold fiber populations rather than two clearly separated groups,
and unmyelinated and myelinated fiber types comprise both populations. The afferent signals of the lower
urinary tract provide state information to spinal and supraspinal centers that promote continence and mediate
micturition througha network of reflexes.The reflexes arebroadly organized so as to synergistically contract the
detrusor and relax the urethra to expel urine, and contract the urethra and relax the detrusor to prevent leaking
during storage. Part of the difficulty in developing a
complete theory of neural control of the lower urinary tract is the large number of interdependent reflexes. A
large body of work exists investigating the spinal and supraspinal centers where information is integrated for
individual reflex control.

Autonomic and sensory nerve modulation of peristalsis in the

upper
urinary tract
M.J. Nguyen a, R. Higashi b, K. Ohta b, K.-I. Nakamura b, H. Hashitani c, R.J. Lang a,
a Department of Physiology, School of Biomedical Sciences, Monash University, Clayton, Victoria 3800, Australia
b Department of Anatomy, Kurume University of School of Medicine Kurume, Japan
c Department of Cell Physiology, Nagoya City University Graduate School of Medical Sciences, Nagoya 467-8601, Japan

Autonomic Neuroscience: Basic and Clinical xxx (2015) xxxxxx

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Journal of Behavioral and Brain Science, 2012, 2, 35-47

http://dx.doi.org/10.4236/jbbs.2012.21005 Published Online February 2012
(http://www.SciRP.org/journal/jbbs)

Morphological and Quantitative Study of

Neurons in th
Gracile Nucleus of the Camel Brain Stem
Saleh M. Al-Hussain1*#, Raith A. Al-Saffar1, Sami I. Zaqout2#
1
Anatomy Department, Faculty of Medicine, Jordan University of Science and Technology, Irbid,
Jordan
2
Anatomy Department, Faculty of Medicine, Islamic University of Gaza, Gaza, Palestine
Email: *smbanihani@just.edu.jo
Received August 27, 2011; revised October 13, 2011; accepted November 3, 2011

The dorsal column nuclei (DCN) are among the most throughly studied structures in the
mammalian central nervous system from the anatomical, physiological and behavioral
point of view [1]. The DCN (gracile and cuneate nuclei) are sensory nuclei which are
interposed in the ascending somesthetic pathway from the periphery to the
somatosensory thalamus [2]. Nerve fibers from receptors mediating proprioception,
kinesthesia, stereognosis and vibration enter directly into the dorsal columns of the
spinal cord and ascend ipsilaterally, without synapsing, to the gracile (Gr) and cuneate
(Cu) nuclei. The fibers from the leg (hindlimb) ascend in the more medial portion of the
dorsal column in the fasciculus gracilis, while those fibers from the upper extremity
(forelimb) ascend in the more lateral fasciculus cuneatus [3]. In mammals, hindlimb and
forelimb afferents nerve fibers project to separate bulbar targets, namely the le Gr and
Cu nuclei. This organization allows assay of interacttions between distinct components of
the somatosensory system during the course of development [4]. There is a steady
increase in size of the dorsal fasciculi and their bulbar nuclei throughout the phylogenetic
scale from lower animals to man [5]. The two nuclei make their first definite appearance
in reptiles and reach their highest stage of development in mammals [6]. It is believed
that
the development of the dorsal fasciculi and their nuclei in the mammalian and primate
series is correlated with increasing sensory discrimination in the skin and the increased
development of proprioceptive sense in the limb [5]. The concept that the DCN function
only as relay stations [7] is too simplistic in view of accumulated evidence that the
synaptic passage of sensory information is capable of considerable modulation. Now it is
known that the DCN are major integrating centers in which the processes of presynaptic
and postsynaptic inhibition and facilitation regulate rostral transmission of afferent
information [8-11].

The dorsal fasciculi of the spinal cord are the chief pathways for the conduction of
proprioception (sense of position and movements) and tactile discrimination which reach
the thalamus and ultimately the cortex after a relay at a bulbar level; the Gr and Cu.
These nuclei were particularly attractive targets for studying early stages of sensory
processing in the somatosensory pathways [16]. In all species studied, these nuclei
receive somatotopically arranged primary afferent fibers, contain a representational map
of the trunk and limbs, and give rise to the medial lemniscus, the fiber system that
provides the fastest and more precisely organized somatosensory input to the thalamus
[17-24].

The sympathetic innervation of the heart:

Important new insights
J.H. Coote Press enter key for correspondence information Press enter key to Email the author

R.A. Chauhan

DOI: http://dx.doi.org/10.1016/j.autneu.2016.08.014

3. The organization of the T1T6 sympathetic nuclei

Sympathetic preganglionic neurones lie bilaterally in the intermediate zone of the spinal cord lying most densely
in the grey matter but extend into the white matter on its lateral border and medially across to the central canal
(Coote, 1988, Jnig, 2006). On each side of the spinal cord these sympathetic neurones form clusters arranged
longitudinally in columns (Fig. 1A) with groups of dendrites spreading laterally and medially but most extensively
rostro-caudally for up to 1.5mm to 2.5mm in the cat spinal cord and probably more in human spinal cord. There
is a rich synaptic terminal innervation on these dendrites by both segmental, propriospinal and supraspinal
afferent terminals making synaptic contact via an array of neurotransmitters from excitatory amino acids like
glutamate or inhibitory amino acids like GABA or glycine as well as monoamines and neuropeptides that have a
variety of effects