2 Organization of the Nervous System (pp.
4-5)  columnar hypothesis
Cerebral hemispheres
• The cerebral hemispheres are composed of
four major lobes:
occipital, parietal, temporal and frontal.
 On the medial part of the temporal lobe are
a series of structures that form part of the
limbic system (see Chapter 45).
 cerebral cortex
- outer layer of the cerebral
hemisphere
- and contains neurones that are
organized in both horizontal layers
and vertical columns
- interconnected over long distances
via pathways that run subcortically.
These pathways, together with
those that connect the cerebral
cortex to the spinal cord, brainstem
and nuclei deep within the cerebral
hemisphere, constitute the white
matter of the cerebral hemisphere.
These deep nuclei include
structures such as basal ganglia and
thalamus
10 Organization of the Cerebral Cortex and
Thalamus (pp 20-21)
 Brodmann’s areas
- cyto-architectural maps
- organization of the outer layer of
the cerebral cortex
- which equates to some extent with
the functional organization of this
structure into motor, sensory and
associative areas, as evidenced by
the laminar organization of the
cortex.
- An area of cortex that is
o predominantly sensory in
character has a prominent
layer IV
o cortical motor areas have
a prominent layer V.
- proposes that the ‘column’ of cortex is
the basic unit of cortical processing.
- alternative approach is to view the
cortex as being organized vertically
- vertical organization
Anatomical organization of the cerebral cortex
 neocortex
- consisting of six layers, although in
certain areas of the cerebral cortex
further subdivisions are used, e.g. the
primary visual cortex.
• thalamic afferent fibres
- relaying sensory information
- project to layer IV often with a smaller
input to layer VI
- terminate in discrete patches.
- Input from thalamic afferent fibers
then synapses onto interneurones
within the cortex which in turn project
vertically to neurones in layers II, III and
V, and from there project to other
cortical and subcortical sites,
respectively. Thus, the weight of
synaptic relations within the cerebral
cortex is in the vertical direction. This
arrangement of synaptic connections is
well seen in the somatosensory and
visual
- In many cortical areas with a motor
function, the motor output from that
cortical area is such that it is directed
back at the motor neurones controlling
the muscles that move the sensory
receptors which ultimately project to
that same area of cortex – so called
input–output coupling.
Developmental organization of the cerebral
cortex
Cortical neurons
- In the mammalian CNS the entire
population
- is produced by a process of migration
from the proliferative zones that are
situated around the cavities of the
cerebral ventricles.
Radial glial fibres
- guide and may even give rise to the
migrating neurons
- span the fetal cerebral wall
- direct the neurones to their correct
cortical location in the developing
cortical plate from the ventricular and
subventricular zones
- (see Chapter 1). Thus, developmentally,
the cortex forms in a vertical fashion.
Neurophysiological organization of the
cerebral cortex
Neurophysiologically, if a recording electrode is
passed at right angles through the cortex, it
encounters cells with similar properties.
However, if the electrode is passed tangentially
then cells shift their response characteristics.
This has been shown in many cortical areas
 Columnar organization of the cortex
- ensures that topography can be
maintained and that the
reorganization of the cortex in
the event of a change in the
peripheral input is relatively
straightforward
Functional organization of the cerebral cortex
1. Serial processing models
- The original models proposed that
information processing was performed
in a serial fashion, such that the cortical
cells form a series of hierarchal levels.
Thus, one set of cells performs a
relatively straightforward analysis,
which then converges on another
population of neurones that perform a
more complex analysis
- The ultimate prediction of these
hierarchical models is that one neurone
at the top of the hierarchy will register
the percept – the ‘grandmother’ cell.
2. Parallel processing models
- The discovery of the X, Y and W classes
of ganglion cells in the retina (see
Chapter 24) led to the development of a
competing theory that proposed that
information is analysed by a series of
parallel pathways, with each pathway
analysing one specific aspect of the
sensory stimulus (e.g. colour or motion
with visual stimuli)
- This theory does not exclude
hierarchical processing but relegates it
to the mode of analysis within separate
parallel pathways. In practice, the
cortex employs both modes of analysis.
3. Distributed processing models
- It should be stressed that cortical
columns are not to be viewed as a static
mosaic structure, as one column may be
a member of a number of different
pathways of analysis.
- This organization has been termed the
distributed system theory and
describes the brain as a complex of
widely and reciprocally interconnected
systems, with the dynamic interplay of
neural activity within and between
these systems as the very essence of
brain function.
- Consequently, one column may be a
member of many distributed systems,
because each distributed system is
specific for one feature of a stimulus
and one column may code for several
features of the stimulus
34 Association Cortices: the posterior Parietal • Temporal cortex
and Prefrontal Cortex
- (corresponding to Brodmann’s areas 21,
Association cortices 22, 37 and 41–43).
- involved in audition and language,
- parts of the cerebral cortex
complex visual processing (such as face
- do not have a primary motor or sensory
recognition) and memory
role involved in the higher order
processing of sensory information Neurophysiologically,
necessary for perception and
 area 5
movement initiation.
- contains many units with a complex
These association areas include: sensory input often with a convergence
of different sensory modalities, such as
• Posterior parietal cortex
proprioceptive and cutaneous stimuli.
- (PPC; defined in monkeys as o These units with such a
corresponding to Brodmann’s areas 5 dual input are probably
and 7, and in humans including areas 39 involved in the sensory
and 40); control of posture and
- has developed greatly during evolution movements.
and relates to specific forms of human o Other units with
behavior, such as the extensive use of multiple cutaneous
tools, collaborative strategic planning inputs are probably
and the development of language. more involved in object
recognition.
It has two main subdivisions:
o However, in addition to
1. somatosensory information (centered having these complex
on area 5); sensory inputs, units in
2. visual stimuli (centered on area 7).
this area are often only
• Prefrontal cortex
maximally activated
- (corresponding to Brodmann’s areas 9– when the sensory
12 and 44–47); stimulus is of interest or
- has increased in size with phylogenetic behavioral significance.
development and has its greatest
representation in humans.
- involved in the purposeful behavior of
an organism and thus is intimately
involved in the planning of responses to
stimuli that include a motor
component.
- Within this structure are specialized
cortical areas such as the frontal eye
fields and Broca’s area.
- generally respond to complex sensory
stimuli of behavioural relevance, which
can then be translated into a cue for
movement.