Studies in the greater galago have not provided a comprehensive description of the organization of eye-specific retino-geniculate-cortical projections to the recipient layers in V1. Here we demonstrate the overall patterns of ocular dominance domains in layers III, IV, and VI revealed following a monocular injection of the transneuronal tracer wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP). We also correlate these patterns with the array of cytochrome oxidase (CO) blobs in tangential sections through the unfolded and flattened cortex. In layer IV, we observed for the first time that eye-specific domains form an interconnected pattern of bands 200–250 μm wide arranged such that they do not show orientation bias and do not meet the V1 border at right angles, as is the case in macaques. We also observed distinct WGA-HRP labeled patches in layers III and VI. The patches in layer III, likely corresponding to patches of K lateral geniculate nucleus (LGN) input, align with layer IV ocular dominance columns (ODCs) of the same eye dominance and overlap partially with virtually all CO blobs in both hemispheres, implying that CO blobs receive K LGN input from both eyes. We further found that CO blobs straddle the border between layer IV ODCs, such that the distribution of CO staining is approximately equal over ipsilateral and contralateral ODCs. These results, together with studies showing that a high percentage of cells in CO blobs are monocular, suggest that CO blobs consist of ipsilateral and contralateral subregions that are in register with underlying layer IV ODCs of the same eye dominance. In macaques and humans, CO blobs are centered on ODCs in layer IV. Our finding that CO blobs in galago straddle the border of neighboring layer IV ODCs suggests that this novel feature may represent an alternative way by which visual information is processed by eye-specific modular architecture in mammalian V1.

Callosal patches in primary visual cortex of Long Evans rats, normally associated with ocular dominance columns, emerge by postnatal day 10 (P10), but they do not form in rats monocularly enucleated a few days before P10. We investigated whether we could replicate the results of monocular enucleation by using tetrodotoxin (TTX) to block neural activity in one eye, or in primary visual cortex. Animals received daily intravitreal (P6–P9) or intracortical (P7–P9) injections of TTX, and our physiological evaluation of the efficacy of these injections indicated that the blockade induced by a single injection lasted at least 24 h. Four weeks later, the patterns of callosal connections in one hemisphere were revealed after multiple injections of horseradish peroxidase in the other hemisphere. We found that in rats receiving either intravitreal or cortical injections of TTX, the patterns of callosal patches analyzed in tangential sections from the flattened cortex were not significantly different from the pattern in normal rats. Our findings, therefore, suggest that the effects of monocular enucleation on the distribution of callosal connections are not due to the resulting imbalance of afferent ganglion cell activity, and that factors other than neural activity are likely involved.

Posterior parietal cortex has traditionally been considered to be a sensory association area in which higher-order processing and intermodal integration of incoming sensory information occurs. In this paper, evidence from clinical reports and from lesion and behavioral-electrophysiological experiments using monkeys is reviewed and discussed in relation to the overall functional organization of posterior parietal association cortex, and particularly with respect to a proposed posterior parietal mechanism concerned with the initiation and control of certain classes of eye and limb movements. Preliminary data from studies of the effects of posterior parietal lesions on oculomotor control in monkeys are reported.

The behavioral effects of lesions of posterior parietal cortex in monkeys have been found to be similar to those which follow analogous damage of the minor hemisphere in humans, while behavioral-electrophysiological experiments have disclosed classes of neurons in this area which have functional properties closely related to the behavioral acts that are disrupted by lesions of the area. On the basis of current data from these areas of study, it is proposed that the sensory association model of posterior parietal function is inadequate to account for the complexities of the present evidence. Instead, it now appears that many diverse neural mechanisms are located in part in parietal cortex, that some of these mechanisms are involved in sensory processing and perceptual functions, but that others participate in motor control, and that still others are involved in attentional, motivational, or emotional processes. It is further proposed that the elementary units of these various neural mechanisms are distributed within posterior parietal cortex according to the columnar hypothesis of Mountcastle.

We review the evidence for the concept of the “initial” or prototype brain. We outline four possible modes of brain evolution suggested by our new findings on the evolutionary status of the dolphin brain. The four modes involve various forms of deviation from and conformity to the hypothesized initial brain type. These include examples of conservative evolution, progressive evolution, and combinations of the two in which features of one or the other become dominant. The four types of neocortical organization in extant mammals may be the result of selective pressures on sensory/motor systems resulting in divergent patterns of brain phylogenesis. A modular “modification/multiplication” hypothesis is proposed as a mechanism of neocortical evolution in eutherians. Representative models of the initial ancestral group of mammals include not only extant basal Insectivora but also Chiroptera; we have found that dolphins and large whales have also retained many features of the archetypal or initial brain. This group evolved from the initial mammalian stock and returned to the aquatic environment some 50 million years ago. This unique experiment of nature shows the effects of radical changes in environment on brain-body adaptations and specializations. Although the dolphin brain has certain quantitative characteristics of the evolutionary changes seen in the higher terrestrial mammals, it has also retained many of the conservative structural features of the initial brain. Its neocortical organization is accordingly different, largely in a quantitative sense, from that of terrestrial models of the initial brain such as the hedgehog.

Ever since being described by Mountcastle (Mountcastle, 1957), columnar organization of sensory cortical areas has provided key leverage into understanding the functional organization of neocortex. Columnar or clusteredThe term column specifically denotes an organization in which groups of neurons with similar properties are extended perpendicularly to the cortical surface. Clustered organization, as used here, is intended to be similar but more general, where the groups of similar neurons need not have any particular geometry. Because of the limits of recording in the alert monkey, these cannot be distinguished in the present work, and the more inclusive term “clustered” will be used. organization of neurons sharing like properties is now known to be widespread, and probably universal in primary sensory areas. Visual cortex in primates consists of a primary area and a large number of secondary areas, which are organized in a manner both hierarchical and parallel (Felleman & Van Essen, 1991; Young, 1993; Young et al., 1995). One major component in the organization of extrastriate visual cortex appears to be the division into dorsal and ventral “streams” of processing (Ungerleider & Mishkin, 1982), each of which is organized hierarchically. Within each, columnar organization exists at early stages, but becomes less clear at higher levels. Columnar organization has been described at the highest level of the ventral stream, inferotemporal cortex (IT, Saleem et al., 1993; Fujita & Fujita, 1996; Tanaka, 1996), but has not been well characterized at the higher levels of the dorsal stream. Hints of such organization are found in the literature (Saito et al., 1986; Lagae et al., 1994), but systematic measurements are needed. In this paper, I report the existence of clustered organization in the medial superior temporal area (MST) of the dorsal stream, which is arguably the highest dominantly visual area on this pathway. I have measured the selectivity of both single- and multiple-unit activity along oblique electrode penetrations through this area to three different kinds of optic flow stimuli, and find that nearby neurons are more similar in their tuning than are more distant ones. This observation documents the existence of some form of clustered organization and supports the importance of this area in the processing of optic flow information.