Cerebral Cortex

1. The majority of the human cerebral cortex is devoted

to tasks that transcend encoding primary sensations or
commanding motor actions.

2. Collectively, the association cortices mediate these

cognitive functions of the brainbroadly defined as the
ability to attend to, identify, and act meaningfully in
response to complex external or internal stimuli.

3. Descriptions of patients with cortical lesions, functional

brain imaging of normal subjects, and behavioral and
electrophysiological studies of nonhuman primates
have established the general purpose of the major
association areas.
 Cerebral Cortex
4. Laminar pattern

The standard areas of cortex (isocortex) is characterized as having

six distinct layers. From outside inward:
Molecular layer
External granular layer
External pyramidal layer
Internal granular layer
Internal pyramidal layer
Multiform layer.
The molecular layer I contains few scattered neurons and consists
mainly of extensions of apical dendrites and horizontally oriented
axons, and some Cajal-Retzius and spiny stellate neurons can be
found.
The external granular layer II contains small pyramidal neurons and
numerous stellate neurons.
 Cerebral Cortex
5. The external pyramidal layer III contains predominantly
small and medium sized pyramidal neurons, as well as
non-pyramidal neurons with vertically-oriented
intracortical axons. Layers I through III are the main
target of interhemispheric corticortical afferents, and
layer III is the principal source of corticortical efferents.
6. The internal granular layer IV contains different types of
stellate and pyramidal neurons, and is the main target
of thalamocortical afferents as well as intra-hemispheric
corticocortical afferents.
 Cerebral Cortex
7. The internal pyramidal layer V contains large pyramidal
neurons (as the Betz cells in the primary motor cortex),
as well as interneurons, and it is the principal source of
efferent for all the motor-related subcortical structures.

8. The multiform layer VI contains few large pyramidal

neurons and many small spindle-like pyramidal and
multiform neurons.

9. The layer VI sends efferent fibers to the thalamus

establishing a very precise reciprocal interconnection
between the cortex and the thalamus.
 Cerebral Cortex
10. During early development, there is an additional layer of
neurons present in the future white matter. These are
called subplate neurons and these neurons disappear
during postnatal development.
11. The cortical layers are not simply stacked one over the
other; they develop characteristic connections between
different layers, which define the basic structure of the
cortical columns in the mature cortex (Mountcastle,
1997).
12. There are no actual borders between the layers, and
neurons cross layer boundaries with their dendrites and
axons trees all over. The pyramidal cells (the majority of
the neurons) span at least three layers, and in many
cases all the layers.
The structure of the human neocortex, including the association cortices. (A) A summary
of the cellular composition of the six layers of the neocortex. (B) Based on variations in
the thickness, cell density, and other histological features of the six neocortical laminae,
the human brain can be divided into about 50 cytoarchitectonic areas, in this case those
recognized by the neuroanatomist Korbinian Brodmann in his seminal monograph in
1909. (See Box A for additional detail.)
cerebral cortex
Canonical neocortical circuitry. Green arrows indicate outputs to the major targets of
each of the neocortical layers in humans; white arrow indicates thalamic input (primarily
to layer IV); dark purple arrows indicate input from other cortical areas; and light purple
arrows indicate input from the brainstem modulatory systems to each layer.
 Cerebral Cortex
13. The cerebral cortex sends connections (efferents) and receives
connections (afferents) from many subcortical structures like the
thalamus and basal ganglia. Most of the sensory stimulation arrives
at the cerebral cortex indirectly through different thalamic nuclei.

14. This is the case of touch, vision and sound but not of olfactory
stimulation, which passes to the olfactory bulb and then to the
olfactory (pyriform) cortex. The largest part of the connections
arriving at the cerebral cortex do not come from subcortical
structures however. The main source of cortical stimulation is the
cerebral cortex itself: maybe 99% of the total connections.

.
 Cerebral Cortex
15. Other areas receive impulses from the primary sensory areas and integrate the
information coming in from different types of receptors. These are often called
association areas and make up a great deal of the cortex in all primates, humans
included. Thus, the cortex is commonly described as comprised of the primary
sensory areas, the motor areas and the association area.

16. The motor areas are located in both hemispheres of the cortex. They are shaped like
a pair of headphones stretching from ear to ear. The motor areas are very closely
related to the control of voluntary movements, especially fine fragmented movements
performed by the hand. The right half of the motor area controls the left side of your
body and vice versa.

17. Two areas of the cortex are commonly referred to as motor:

18. Primary motor cortex, which executes voluntary movements

19. Supplementary motor areas and premotor cortex, which select voluntary movements.
 Cerebral Cortex
20. Posterior Parietal Cortex, which guides voluntary movements in
space

21. Dorsolateral Prefrontal Cortex, which decides which voluntary

movements to make according to higher-order instructions, rules,
and self-generated thoughts.

22. Areas that receive that particular information are called sensory
areas. Parts of the cortex that receive sensory inputs from the
thalamus are called primary sensory areas. The senses of vision,
audition and touch are served by the primary visual cortex, primary
auditory cortex and primary somatosensory cortex. In general, the
two hemispheres receive the information from the opposite sides of
the body.
 Cerebral Cortex
23. The right primary somatosensory cortex receives information from the left
limbs and the right visual cortex receives information from the left visual
field. The organisation of sensory maps in the cortex reflects that of the
corresponding sensing organ, in which is known as a topographic map.
Neighbouring points in the primary visual cortex, for example, correspond to
neighbouring points in the retina.

24. Association areas comprise three major groups:

Parietal, temporal, and occipital lobes - all located in the posterior part of the
brain - are involved in producing our perceptions resulting from what our
eyes see, ears hear, and other sensory organs inform us about the position
of different parts of our body and relate them to the position of other objects
in the environment
Frontal lobe - called prefrontal association complex and involved in planning
actions and movement, as well as abstract thought
In humans, the association areas of the left hemisphere, especially the
parietal-temporal-occipital complex, are responsible for our understanding
and use of language
Lateral view of the human brain
Midsagittal view of the brain
 Cerebral Cortex
25. Parietal association cortex is involved in attention and
awareness of the body and the stimuli that act on it;
temporal association cortex is involved in the
recognition and identification of highly processed
sensory information; and frontal association cortex is
involved in guiding complex behavior by planning
responses to ongoing stimulation (or remembered
information), matching such behaviors to the demands
of a particular situation.

26. More than any other brain regions, the association

areas support the mental processes that make us
human.
 Cerebral Cortex
27. The function of the frontal cortex was first suggested by a
dramatic nineteenth century accident in which a tamping
rod was driven through the frontal part of the brain of a
railroad worker named Phineas P. Gage.

28. Remarkably, Gage survived, and his behavioral deficits

stimulated much early thinking about complex brain
functions. The illustration here is a reconstruction of the
trajectory of the rod based on Gage's skull, which is
housed in the Warren Museum at Harvard Medical
School. (Courtesy of H. Damasio.)
 Cerebral Cortex
19. The awareness of physical and social circumstances, the ability to
have thoughts and feelings (emotions), to be sexually attracted to
others, to express these things to our fellow humans by language,
and to store such information in memory certainly rank among the
most intriguing functions of the human brain.

20. Given their importance in our daily livesand for human culture
generallyit is not surprising that much of the human brain is
devoted to these and other complex mental functions.

21. The intrinsic interest of these aspects of human behavior is

unfortunately equaled by the difficultyboth technical and
conceptualinvolved in unraveling their neurobiological
underpinnings
 Cerebral Cortex
22. Clinical evidence from patients with lesions of the association cortex in the
temporal lobe indicate that one of the major functions of this part of the
brain is recognition and identification of stimuli that are attended to,
particularly complex stimuli. Thus, damage to either temporal lobe can
result in difficulty recognizing, identifying, and naming different categories of
objects.

23. These disorders, collectively called agnosias (from the Greek for not
knowing), are quite different from the neglect syndromes. As noted,
patients with right parietal lobe damage often deny awareness of sensory
information in the left visual field (and are less attentive to the left sides of
objects generally), despite the fact that the sensory systems are intact (for
instance, an individual with contralateral neglect syndrome typically
withdraws his left arm in response to a pinprick, even though he may not
admit the arm's existence).

24. Patients with agnosia, on the other hand, acknowledge the presence of a
stimulus, but are unable to report what it is. These latter disorders have both
a lexical aspect and a mnemonic aspect.
 Cerebral Cortex
25. One of the most thoroughly studied agnosias following
damage to the temporal association cortex in humans is
the inability to recognize and identify faces. This
disorder, called prosopagnosia (prosopo-, from the
Greek for face or person).

26. After damage to the temporal lobes, typically the right

temporal lobe, patients are often unable to identify
familiar individuals by their facial characteristics, and in
some cases cannot recognize a face at all.

27. Nonetheless, such individuals are perfectly aware that

some sort of visual stimulus is present and can describe
particular aspects or elements of it without difficulty.
 Cerebral Cortex
28. The functional deficits that result from damage to the human frontal lobe are diverse
and devastating, particularly if both hemispheres are involved. This broad range of
clinical effects stems from the fact that the frontal cortex has a wider repertoire of
functions than any other neocortical region (consistent with the fact that the frontal
lobe in humans and other primates is the largest of the brain's lobes and comprises a
greater number of cytoarchitectonic areas).

29. The particularly devastating nature of the behavioral deficits after frontal lobe damage
reflects the role of this part of the brain in maintaining what is normally thought of as
an individual's personality. The frontal cortex integrates complex perceptual
information from sensory and motor cortices, as well as from the parietal and
temporal association cortices.

30. The result is an appreciation of self in relation to the world that allows behaviors to be
planned and executed normally. When this ability is compromised, the afflicted
individual often has difficulty carrying out complex behaviors that are appropriate to
the circumstances. These deficiencies in the normal ability to match ongoing behavior
to present or future demands are, not surprisingly, interpreted as a change in the
patient's character.
Summary of the overall connectivity of the association cortices
Asymmetry of the right and left human temporal lobes. (A) The superior portion of the brain has been removed as
indicated to reveal the dorsal surface of the temporal lobes in the right-hand diagram (which presents a dorsal
view of the horizontal plane). A region of the surface of the temporal lobe called the planum temporale is
significantly larger in the left hemisphere of most (but far from all) individuals. (B) Measurements of the planum
temporale in 100 adult and infant brains. The mean size of the planum temporale is expressed in arbitrary
planimetric units to get around the difficulty of measuring the curvature of the gyri within the planum. The
asymmetry is evident at birth and persists in adults at roughly the same magnitude (on average, the left planum is
57% larger in infants and 50% larger in adults). (C) An MR image in the frontal plane, showing this asymmetry
(arrows) in a normal adult subject.
The distribution in the human brain of neurons and their projections (arrows) containing
biogenic amine neurotransmitters. Curved arrows along the perimeter of the cortex
indicate the innervation of lateral cortical regions not shown in this midsagittal plane of
section.