The neocortex is divided into multiple areas with specific architecture, molecular identity and pattern of connectivity with the dorsal thalamus. Gradients of transcription factor expression in the cortical primordium regulate molecular regionalization and potentially the patterning of thalamic projections. We show that reduction of Fgf8 levels in hypomorphic mouse mutants shifts early gradients of gene expression rostrally, thereby modifying the molecular identity of rostral cortical progenitors. This shift correlates with a reduction in the size of a molecularly defined rostral neocortical domain and a corresponding rostral expansion of more caudal regions. Despite these molecular changes, the topography of projections between the dorsal thalamus and rostral neocortex in mutant neonates appears the same as the topography of wild-type littermates. Overall, our study demonstrates the role of endogenous Fgf8 in regulating early gradients of transcription factors in cortical progenitor cells and in molecular regionalization of the cortical plate

The functional organization of area 3a, a cortical field proposed to be involved in somato-motor-vestibular integration, has never been described for any primate. In the present investigation, the topographic organization and connections of area 3a were examined in marmosets using electrophysiological recording and anatomical tracing techniques. Multi-unit neuronal activity was recorded at a number of closely spaced sites; receptive fields (RFs) for neurons were determined, and the optimal stimulus was identified. In all cases, neurons in area 3a responded to the stimulation of deep receptors on the contralateral body. The representation of the body in area 3a was from the toes and foot, to the hindlimb, trunk, forelimb, hand and face in a mediolateral progression. In all cases electrophysiological results were related to myeloarchitecture, and the map in area 3a was found to be coextensive with a strip of lightly to moderately myelinated cortex just rostral to the darkly myelinated 3b. To examine the cortical connections of area 3a, injections of anatomical tracers were made into electrophysiologically identified body part representations. Area 3a has dense intrinsic connections and receives substantial inputs from the primary motor cortex (M1), the supplementary motor area (SMA), areas 1 and 2, the second somatosensory area (S2), and areas in posterior parietal cortex (PP). The connections of area 3a indicate that integration of cortical representations of body parts occurs both within area 3a and between area 3a and other somatosensory and motor areas. In addition, there are differential patterns of interconnections between behaviorally relevant body part representations of area 3a, such as the forelimb, compared to other body part representations (hindlimb/ trunk), especially with 'higher order' cortical fields. This suggests that 3a may be an important component in a network that generates a common frame of reference for hand and eye coordinated reaching tasks.

Genetic and neurobiological research is reviewed as related to controversy over the extent to which neocortical organization and associated cognitive functions are genetically constrained or emerge through patterns of developmental experience. An evolutionary framework that accommodates genetic constraint and experiential modification of brain organization and cognitive function is then proposed. The authors argue that 4 forms of modularity and 3 forms of neural and cognitive plasticity define the relation between genetic constraint and the influence of developmental experience. For humans, the result is the ontogenetic emergence of functional modules in the domains of folk psychology, folk biology, and folk physics. The authors present a taxonomy of these modules and review associated research relating to brain and cognitive plasticity in these domains.For several millennia, scholars have debated whether human traits largely result from our biological nature or are a reflection of nurture, specifically our developmental experiences. The debate continues to this day and has recently pervaded the cognitive neurosciences, at least with respect to theoretical models of brain and cognitive evolution (Elman et al

The detailed organization of somatosensory area 3a was examined in macaque monkeys using multiunit electrophysiological recording techniques. By examining topographic relationships, changes in receptive field size, and the type of stimulus that neurons responded to, functional boundaries of area 3a were determined and related to architectonic boundaries. One striking observation was that the location of area 3a varied with respect to the central sulcus. In one-half of the cases area 3a was on the rostral bank and fundus of the central sulcus and in the other half of the cases it was on the caudal bank and fundus of the central sulcus. In terms of topographic organization, we found that area 3a contains a complete representation of deep receptors and musculature of the contralateral body, and that the general organization of body part representations mirrors that of the primary somatosensory area, 3b. These results as well as results from studies of area 3a in ours and other laboratories indicate that area 3a is part of a network involved in proprioception, postural control, and the generation of coordinated movements. Further, comparative analysis of area 3a in a variety of species suggests that its construction is based, to a large extent, on the use of a particular body part rather than on innervation density.

The process of generating functionally distinct neocortical areas requires the formation of an intra-neocortical connectivity map. Here, we explore the early development of murine intra-neocortical projections and find that axons from rostral and caudal neurons remain, respectively, within large rostral and caudal domains of the neonatal neocortex. Despite evidence that thalamic input can regulate neocortical areal properties, we found that the neonatal intra-neocortical projection pattern was not perturbed when thalamic input was absent in Gbx2 mutants. On the contrary, in Fgf8 hypomorphic mutants, the rostral neocortex of which acquires more caudal molecular properties, caudally located neurons ectopically project axons into the rostral cortex. Therefore, neocortical patterning by Fgf8 also contributes to arealization through mediating early development of intra-neocortical connectivity.