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Ganong's Review of Medical Physiology, 23rd Edition

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9<br />

FIGURE 11–3 Brain areas concerned with somatic sensation,<br />

and some <strong>of</strong> the cortical receiving areas for other sensory<br />

modalities in the human brain. The numbers are those <strong>of</strong> Brodmann’s<br />

cortical areas. The primary auditory area is actually located in<br />

the sylvian fissure on the top <strong>of</strong> the superior temporal gyrus and is not<br />

normally visible in a lateral view <strong>of</strong> the cortex.<br />

(Figure 11–3). Not only is there detailed localization <strong>of</strong> the<br />

fibers from the various parts <strong>of</strong> the body in the postcentral<br />

gyrus, but also the size <strong>of</strong> the cortical receiving area for<br />

impulses from a particular part <strong>of</strong> the body is proportionate to<br />

the use <strong>of</strong> the part. The relative sizes <strong>of</strong> the cortical receiving<br />

areas are shown dramatically in Figure 11–4, in which the proportions<br />

<strong>of</strong> the homunculus have been distorted to correspond<br />

to the size <strong>of</strong> the cortical receiving areas for each. Note that the<br />

cortical areas for sensation from the trunk and back are small,<br />

whereas very large areas are concerned with impulses from the<br />

hand and the parts <strong>of</strong> the mouth concerned with speech.<br />

FIGURE 11–4 Sensory homunculus, drawn overlying a<br />

coronal section through the postcentral gyrus. Gen., genitalia.<br />

(Reproduced, with permission, from Penfield W, Rasmussen G: The Cerebral Cortex <strong>of</strong><br />

Man. Macmillan, 1950.)<br />

8 6<br />

Sl<br />

4<br />

Lips<br />

Sll<br />

Nose<br />

Face<br />

Tongue<br />

Thumb<br />

Eye<br />

Upper lip<br />

Ring<br />

Middle<br />

Index<br />

Pharynx<br />

Intraabdominal<br />

Hand<br />

Face<br />

Tongue<br />

Auditory<br />

Little<br />

Lower lip<br />

Teeth, gums, and jaw<br />

Hand<br />

Wrist<br />

Trunk<br />

Forearm<br />

Elbow<br />

3 1 2<br />

Arm<br />

Shoulder<br />

Head<br />

Neck<br />

Trunk<br />

Hip<br />

Leg<br />

Foot<br />

Toes<br />

Gen.<br />

Posterior<br />

parietal<br />

cortex<br />

Visual<br />

CHAPTER 11 Somatosensory Pathways 175<br />

Studies <strong>of</strong> the sensory receiving area emphasize the very discrete<br />

nature <strong>of</strong> the point-for-point localization <strong>of</strong> peripheral<br />

areas in the cortex and provide further evidence for the general<br />

validity <strong>of</strong> the law <strong>of</strong> specific nerve energies (see Chapter 8).<br />

Stimulation <strong>of</strong> the various parts <strong>of</strong> the postcentral gyrus gives<br />

rise to sensations projected to appropriate parts <strong>of</strong> the body.<br />

The sensations produced are usually numbness, tingling, or a<br />

sense <strong>of</strong> movement, but with fine enough electrodes it has<br />

been possible to produce relatively pure sensations <strong>of</strong> touch,<br />

warmth, and cold. The cells in the postcentral gyrus are organized<br />

in vertical columns, like cells in the visual cortex. The<br />

cells in a given column are all activated by afferents from a<br />

given part <strong>of</strong> the body, and all respond to the same sensory<br />

modality.<br />

SII is located in the superior wall <strong>of</strong> the sylvian fissure, the<br />

fissure that separates the temporal from the frontal and parietal<br />

lobes. The head is represented at the inferior end <strong>of</strong> the<br />

postcentral gyrus, and the feet at the bottom <strong>of</strong> the sylvian fissure.<br />

The representation <strong>of</strong> the body parts is not as complete<br />

or detailed as it is in the postcentral gyrus.<br />

Conscious awareness <strong>of</strong> the positions <strong>of</strong> the various parts <strong>of</strong><br />

the body in space depends in part on impulses from sense<br />

organs in and around the joints. The organs involved are<br />

slowly adapting spray endings, structures that resemble Golgi<br />

tendon organs, and probably Pacinian corpuscles in the synovia<br />

and ligaments. Impulses from these organs, touch receptors<br />

in the skin and other tissues, and muscle spindles are<br />

synthesized in the cortex into a conscious picture <strong>of</strong> the position<br />

<strong>of</strong> the body in space. Microelectrode studies indicate that<br />

many <strong>of</strong> the neurons in the sensory cortex respond to particular<br />

movements, not just to touch or static position.<br />

VENTROLATERAL<br />

SPINOTHALAMIC TRACT<br />

Fibers from nociceptors and thermoreceptors synapse on neurons<br />

in the dorsal horn (Figure 11–1). Aδ fibers terminate primarily<br />

on neurons in laminae I and V, whereas the dorsal root<br />

C fibers terminate on neurons in laminae I and II. The synaptic<br />

transmitter secreted by afferent fibers subserving fast mild<br />

pain is glutamate, and the transmitter subserving slow severe<br />

pain is substance P.<br />

The axons from these neurons cross the midline and ascend<br />

in the ventrolateral quadrant <strong>of</strong> the spinal cord, where they<br />

form the ventrolateral spinothalamic tract (Figure 11–2).<br />

Fibers within this tract synapse in the VPL. Other dorsal horn<br />

neurons that receive nociceptive input synapse in the reticular<br />

formation <strong>of</strong> the brain stem (spinoreticular pathway) and<br />

then project to the centrolateral nucleus <strong>of</strong> the thalamus.<br />

Positron emission tomographic (PET) and functional magnetic<br />

resonance imaging (fMRI) studies in normal humans<br />

indicate that pain activates cortical areas SI, SII, and the cingulate<br />

gyrus on the side opposite the stimulus. In addition, the<br />

medi<strong>of</strong>rontal cortex, the insular cortex, and the cerebellum are<br />

activated. These technologies were important in distinguishing

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