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the bridges that permit the adaptation of organismal needs to the external environment This concept of central nervous system function, rst proposed by Broca, was elaborated by Yakovlev and has been adopted more recently by Mesulam and by Benson Such a model retains to a large degree the cytoarchitectural similarities among areas that serve similar functions (ie, the scheme of Brodmann) and also respects the sequence of maturation (myelination) of connecting pathways proposed by Flechsig (see Fig 28-3) The latter corresponds to the functional development of primary, secondary, and tertiary associative cortical regions In this way, localization may be viewed as the product of genetic patterns of structure, which mature during development, and their synaptic formations, which permit the development of complex circuits during lifelong learning and experience From these remarks it follows that subdivision of the cerebrum into frontal, temporal, parietal, and occipital lobes is somewhat of an anachronism of limited psychophysiologic validity, although, in clinical work, it still serves an important practical purpose as a shorthand way of localizing a lesion Some of these delineations were made long before our rst glimmer of knowledge about the function of the cerebrum Even when the neurohistologists began parceling the neocortex, they found that their areas did not fall neatly within zones bounded by sulci and ssures Therefore, when the terms frontal, parietal, temporal, and occipital are used in the text below, it is mainly to provide the clinician with familiar and manageable anatomic landmarks (see Fig 22-7) Following from the issues expressed above, a comment should be made regarding the study of cortical function by functional imaging techniques [positron emission tomography (PET) and functional magnetic resonance imaging (fMRI)] These extraordinary images reveal the brain regions that are activated during various normal behaviors Some techniques use regional cerebral blood ow, regional oxygen content, or diffusion of water as surrogates for local metabolic brain activity and neuronal activation (see reviews by Gore and by Le Bihan) Invariably, an ensemble of areas, a network of the variety mentioned above, is required to perform even seemingly simple tasks such as recalling a name, visualizing or identifying an object, or carrying out a commanded task The fact that multiple areas of the cortex are entrained may seem at odds with the classic view of lesional neurology in which one localized area is assigned to a particular clinical syndrome (as de ned by a behavioral de cit) The discrepancy is in part one of epistemology in that normal function as re ected in functional images does not equate with abnormal function as exposed by a focal lesion Moreover, it appears from these imaging studies that certain regions of the cortex are necessary to fully conduct particular behaviors, but they are not suf cient for their enactment Restated, a number of ancillary regions appear by functional imaging to participate in these behaviors, but they act in a supplementary way A rough analogy can be made with memory function as described in Chap 21, where hippocampal destruction results in complete amnesia but undoubtedly other areas of the brain participate in normal memory Possibly lesions in these supplementary areas may impair function to a small degree, but these areas are not requisite for the behavior and therefore are not demonstrable in the clinical examination These considerations notwithstanding, there is little doubt that functional imaging provides insights into the co-ordinated interaction of spatially distributed brain regions in normal behavior They also promise to illuminate patterns of diseased brain function and recovery from these processes
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General Anatomic and Physiologic Considerations of Cortical Function
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Pertinent to this subject are a number of morphologic and physiologic observations Along strictly histologic lines, Brodmann distinguished 47 different areas of cerebral cortex (Fig 22-1), and von Economo identi ed more than twice that number Although this parceling was criticized by Bailey and von Bonin, it is still used by physiologists and clinicians, who nd that the Brodmann s areas do indeed approximate certain functional zones of the cerebral cortex (Fig 22-2) Also, the cortex has been shown to differ in its various parts by virtue of its particular connections with other areas of the cortex and with the thalamic nuclei and other lower centers Hence, one must regard the cortex as a heterogeneous array of many anatomic systems, each with highly organized intercortical and diencephalic connections Certain general aspects of cerebrocortical anatomy are particularly noteworthy The sheer size of the cortex is remarkable Unfolded, it has a surface extent of about 4000 cm2 about the size of a full sheet of newsprint (right and left pages) Contained in the cortex are many billions of neurons (estimated at 10 to 30 billion) and ve times this number of supporting glial cells The intercellular synaptic connections number in the trillions Since nerve cells look alike and presumably function alike, the remarkable diversity in human intelligence, store of knowledge, and behavior must depend on the potential for in nite variations in neuronal interconnectivity Most of the human cerebral cortex is phylogenetically recent, hence the term neocortex It has also been referred to as isocortex (Vogt) because of its uniform embryogenesis and morphology These latter features distinguish the neocortex from the older and less uniform allocortex ( other cortex ), which comprises mainly the hippocampus and olfactory cortex Concerning the detailed histology of the neocortex, six layers (laminae) can be distinguished; from the pial surface to the underlying white matter they are as follows: the molecular (or plexiform), external granular, external pyramidal, internal granular, ganglionic (or internal pyramidal), and multiform (or fusiform) layers, as illustrated in Fig 22-3 Two cell types relatively large pyramidal cells and smaller, more numerous rounded (granular) cells predominate, and variations in the lamination of the neocortex are largely determined by variations in the size and density of these neuronal types Many variations in lamination have been described by cortical mapmakers, but two main types of neocortex are recognized: (1) the homotypical cortex, in which the six-layered arrangement is readily discerned, and (2) the heterotypical cortex, in which the layers are less distinct The association cortex the large areas (75 percent of the surface) that are not obviously committed to primary motor or sensory functions is generally of this latter type Homotypical areas are characterized by either granular or agranular nerve cells The precentral cortex (Brodmann s areas 4 and 6, mainly motor region) is dominated by pyramidal rather than granular cells, especially in layer V (hence the term agranular) Agranular cortex is distinguished by a high density of large pyramidal neurons In contrast, primary sensory cortex postcentral gyrus (areas 3, 1, 2), banks of the calcarine sulcus (area 17), and the transverse gyri of Heschl (areas 41 and 42) where layers II and IV are strongly developed for the receipt of afferent impulses, has been termed granular cortex because of the marked predominance of granular cells Granular cortex has a preponderance of small neurons (Fig 22-4)
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