The primary somotosensory cortex is located

caudal to the central sulcus in the postcentral gyrus

in the parietal lobe

The somatosensory cortex is subdivided in 4 areas,

each receving a somatotopic arrays of inputs from
the skin; therefore, it contains 4 neural maps of the
surface of the body.
Broadmann area 3 of the primary
somatosensory cortex (SI) receive inputs
from the VPL/VPM thalamic nuclei and
performs the first basic processing of the
somatosensory information (3b: tactile
stimuli; 3a: proprioceptive stimuli).
Neurons in areas 3b and 3a project their
axons to areas 1 and 2 of SI where a
more complex processing of the same
stimuli is performed. Area 2 integrates
tactile and proprioceptive information, i.e.
it combines tactile information with
information concerning limb position to
mediate the tactile recognition of objects.
The four SI areas are extensively
interconnected such that processing of
sensory information at this level involves
both serial and parallel processing.
Strictly speaking the real primary
somatosensory cortex is area 3;
area 2 has features of a higher order , i.e.
of so called unimodal association areas,
that are concerned only (or primarily) with
a single modality and integrate the
different submodalities (e.g. area 5; area
SII)

Somatosensory information is
conveyed in parallel from the
four areas of SI to higher
centers in the cortex, including
the secondary somatosensory
cortex (SII) and the posterior
parietal cortex (areas 5, 7 are
unimodal somatosensory
association areas, while areas
39 and 40 are multimodal
association areas)
Neurons in the primary somatosensory cortex project to neurons in adjacent areas (unimodal
somatosensory association areas; and to primary motor cortex), and the neurons in the unimodal
association areas in turn project to other adjacent higher-order cortical regions (multimodal
association areas in the posterior parietal cortex; also to premotor cortex). The posterior parietal
cortex also receives input from the visual and auditory systems and from the hippocampus;
moreover, it is heavily bidirectionally connected to other multimodal association areas.

The multimodal parietal association areas integrate somatic sensory information

with other sensory modalities to form unified spatial percepts of objects in
extrapersonal or far space.
The multimodal association areas of the cortex are concerned with
integrating sensory modalities.
They appear to be particularly important for two tasks:
(1) the production of a unified perception
(2) the representation of the percept in memory (cf their strong connections
with the hippocampus)
Perception: transformation of sensory stimuli into conscious experience.

Colors, odors, sounds are mental creations which do not exist outside our
brain, but are created on the basis of our interpretation of physical sensory
stimuli (electromagnetic waves of different wavelength, chemical
compounds, pressure waves). This interpretation and hence the
conscious experience (perception) of the external world depend on brain
state, concentration of attention, previous experience (besides the intrinsic
brain structure)

The brain constructs an internal representation of external physical

events after first analyzing various features of those events (e.g. shape,
texture, movement of an object) and then integrating them in in a unified
conscious experience.
How this integration occurs—the binding problem—and how conscious
experience emerges from the brain’s selective attention to incoming
sensory information are two of the most important open questions in
cognitive neuroscience.
The multimodal association areas project to multimodal motor association
areas located rostral to the primary motor cortex in the frontal lobe; these
frontal multimodal association areas transform sensory information into
planned movement and compute the programs for these movements.

Because the multimodal association areas integrate sensory

modalities and link sensory information to the planning of
movement, they are thought to be the anatomical substrates of the
highest brain functions (perception, goal-directed actions, conscious
thought).
Consistent with this, lesions of these association areas result in profound
cognitive deficits.
Lesions in the posterior parietal cortex produce complex defects in
personal or peripersonal spatial perception and selective attention.

Many patients with parietal lesions show a striking

deficit in awareness of one side of their body. For
example, such patients may not dress, undress, or
wash the affected side (personal neglect
syndrome).

In some patients with right parietal lesions the

sensory neglect extends from near space to far
space (contralateral spatial neglect syndrome):
cf inability to copy the left side of a drawing;
cf self-portraits of a german artist after a stroke
that affected the right posterior parietal cortex
The right parietal lobe
controls perception and
attention to both the right
and left parts of the body
and of the extrapersonal
space.
The left parietal lobe
controls mainly the right
part.
Large regions of association cortex are contained within each of the four lobes
and contribute to cognition in distinctive ways (as mainly inferred from clinical
observations in humans and electrophysiological studies in monkeys; recently, from non
invasive imaging techniques: eg fMRI, PET).

The parietal association cortex is critical for sensory guidance of motor behavior
and spatial awarness.

The temporal association cortex is important for recognition of sensory stimuli

and for storage of semantic knowledge.

The frontal association cortex plays a key role in organizing behavior and in
working memory.

The limbic association cortex (which includes the parahippocampal cortex, the
cingulate cortex, the hippocampus and the amygdala) serves complex functions
related to emotion and episodic (autobiographical) memory.

Association areas have much more extensive input and output connections than
do lower-order sensory and motor areas.
All association areas are highly interconnected by a dense network of pathways
within and between the parietal, temporal, frontal, and limbic lobes.
The parietal association cortex (as well as the other association cortices) is
heavily connected to the frontal association cortex

In addition to serial processing,

another principle of cortical
organization is parallel
processing, whereby sensory
information of a given modality
is processed differently in
parallel pathways.

Parallel pathways in each

sensory modality lead to dorsal
association areas (parietal
cortex: dorsal stream) and
ventral association areas
(temporal cortex: ventral stream)
The frontal association cortex consists of the dorsolateral and
ventromedial prefrontal areas.
These areas play a critical role in the executive control of
behavior.

The dorsolateral prefrontal cortex is important for cognitive control

of motor behaviour and for maintaining intention.
In humans injury to this region results in disorganized behavior,
distractibility, incapacity to carry out plans (but normal perceptual and
motor abilities; also tests of intelligence may be normal).
.
The orbital-ventromedial prefrontal cortex (which is linked strongly to
amygdala and hypothalamus and receives input from every sensory system) is
important for emotional control of behavior; it contributes to
motivational states by representing the emotional value of objects that
might become targets of action.
Injury to this cortex in humans (cf P. Cage) results in deficient emotional
control of behavior, in a failure to properly value the expected
consequences of an action (hence inappropriate behavior) and in
abnormal emotional states characterized e.g. by indifference (loss of
appreciation for literature or music, insensitivity to feelings of others, indifference
to the financial consequences of own actions).
All areas of the frontal lobe participate in the control of motor behavior but in
different ways.
Just as in the posterior sensory cortex, frontal areas are connected in series
in a functional hierarchy.

In contrast to the sensory

systems, where information
flows from the periphery into
higher-order areas, in the
motor systems signals flow
from the higher-order areas
of the frontal lobe to the
primary motor cortex.
The orbital-ventromedial prefrontal
Cortex (OF), an area involved in
emotional processes associated with
the executive control of behavior, is
connected to the dorsolateral prefrontal
cortex (DLPFC), which is important for
cognitive control of motor behavior.
DLFPC is connected to the premotor
cortex (PM), which is involved in
generating the motor programs (e.g.
programming the combination of limbs
to be used or the sequence of
movements), and receive major inputs
also from primary and secondary
somatosensory cortices (which send
information about ongoing movements).
PM neurons become active during the
preparation of movement which is then
implemented by activation of neurons in
the primary motor cortex (M1), to which
PM is connected.
L5 pyramidal cells in M1 directly
connect to motoneurons and produce
movement of different parts of the body.
The axons of neurons in layer V of the
primary motor cortex project through the
corticospinal tract to the ventral horn of
the spinal cord. The human corticospinal
tract consists of approximately one
million axons, of which about 40%
originate in the motor cortex. In the
medulla the axons form prominent
protuberances on the ventral surface
called the medullary pyramids, and thus
the entire projection is sometimes called
the pyramidal tract.
Most of the corticospinal fibers cross the
midline in the medulla ( pyramidal
decussation). However, about 10% do
not cross until they reach the level of the
spinal cord at which they will terminate.

The corticospinal axons make

monosynaptic connections
with motoneurons (important for
individual finger movements) and also
with interneurons in the spinal cord
(important for coordinating larger
groups of muscles in behaviors such
as reaching and walking).
The primary motor cortex is organized somatotopically like the somatosensory cortex:
specific regions of the motor cortex influence the activity of specific muscle groups
A continuous stream of tactile, visual, and proprioceptive
information is needed to make voluntary movement both accurate
and properly sequenced.
The motor information carried in the corticospinal tract is
significantly modulated by both sensory information and information
from other motor regions.

The output of the motor cortex is under the substantial influence of

other motor regions of the brain, including the cerebellum and
basal ganglia, structures that are essential for smoothly executed
movements, and are also important for motor learning (the
improvement in motor skills through practice).