Overlapping and nonoverlapping cortical projections to cortex of the superior temporal sulcus in the rhesus monkey: double anterograde tracer studies

To examine how fibers from functionally distinct cortical zones interrelate within their target areas of the superior temporal sulcus (STS) in the rhesus monkey, separate anterograde tracers were injected in two different regions of the same hemisphere known to project to the STS. Paired injections were placed in dorsal prearcuate cortex and the caudal inferior parietal lobule (IPL), interconnected regions that are part of a hypothesized distributed network concerned with visuospatial analysis or directed attention; in a presumed auditory region of the superior temporal gyrus (STG) and in extrastriate visual cortex, the caudal IPL and lower rim of the intraparietal sulcus; and in dorsal prearcuate cortex and the STG. Overlapping and nonoverlapping projections were then examined in STS visual and polysensory areas. Prefrontal and parietal fibers directly overlapped extensively in area MST and all subdivisions of presumed polysensory cortex (areas TPOc, TPOi, and TPOr), although nonoverlapping connections were also found. Although STG and IPL fibers targeted all TPO subdivisions, connections were to nonoverlapping, but often adjacent, columns. Paired prefrontal and STG injections revealed largely nonoverlapping vertical columns of connections but substantial overlap within layers VI and I or areas TPOc and TPOi. The findings suggest that area TPO contains differently connected modules that may maintain at least initial segregation of visual versus auditory inputs. Other modules within area TPO receive directly converging input from the posterior parietal and the prefrontal cortices and may participate in a distributed cortical network concerned with visuospatial functions.     

The TINS Lecture. The parietal association cortex in depth perception and visual control of hand action

Recent neurophysiological studies in alert monkeys have revealed that the parietal association cortex plays a crucial role in depth perception and visually guided hand movement. The following five classes of parietal neurons covering various aspects of these functions have been identified: (1) depth-selective visual-fixation (VF) neurons of the inferior parietal lobule (IPL), representing egocentric distance; (2) depth-movement sensitive (DMS) neurons of V5A and the ventral intraparietal (VIP) area representing direction of linear movement in 3-D space; (3) depth-rotation-sensitive (RS) neurons of V5A and the posterior parietal (PP) area representing direction of rotary movement in space; (4) visually responsive manipulation-related neurons (visual-dominant or visual-and-motor type) of the anterior intraparietal (AIP) area, representing 3-D shape or orientation (or both) of objects for manipulation; and (5) axis-orientation-selective (AOS) and surface-orientation-selective (SOS) neurons in the caudal intraparietal sulcus (cIPS) sensitive to binocular disparity and representing the 3-D orientation of the longitudinal axes and flat surfaces, respectively. Some AOS and SOS neurons are selective in both orientation and shape. Thus the dorsal visual pathway is divided into at least two subsystems, V5A, PP and VIP areas for motion vision and V6, LIP and cIPS areas for coding position and 3-D features. The cIPS sends the signals of 3-D features of objects to the AIP area, which is reciprocally connected to the ventral premotor (F5) area and plays an essential role in matching hand orientation and shaping with 3-D objects for manipulation.     

Organization of the visual reticular thalamic nucleus of the rat

The visual sector of the reticular thalamic nucleus has come under some intense scrutiny over recent years, principally because of the key role that the nucleus plays in the processing of visual information. Despite this scrutiny, we know very little of how the connections between the reticular nucleus and the different areas of visual cortex and the different visual dorsal thalamic nuclei are organized. This study examines the patterns of reticular connections with the visual cortex and the dorsal thalamus in the rat, a species where the visual pathways have been well documented. Biotinylated dextran, an anterograde and retrograde tracer, was injected into different visual cortical areas [17; rostral 18a: presumed area AL: (anterolateral); caudal 18a: presumed area LM (lateromedial); rostral 18b: presumed area AM (anteromedial); caudal 18b: presumed area PM (posteromedial)] and into different visual dorsal thalamic nuclei (posterior thalamic, lateral geniculate nuclei), and the patterns of anterograde and retrograde labelling in the reticular nucleus were examined. From the cortical injections, we find that the visual sector of the reticular nucleus is divided into subsectors that each receive an input from a distinct visual cortical area, with little or no overlap. Further, the resulting pattern of cortical terminations in the reticular nucleus reflects largely the patterns of termination in the dorsal thalamus. That is, each cortical area projects to a largely distinct subsector of the reticular nucleus, as it does to a largely distinct dorsal thalamic nucleus. As with each of the visual cortical areas, each of the visual dorsal thalamic (lateral geniculate, lateral posterior, posterior thalamic) nuclei relate to a separate territory of the reticular nucleus, with little or no overlap. Each of these dorsal thalamic territories within the reticular nucleus receives inputs from one or more of the visual cortical areas. For instance, the region to the reticular nucleus that is labelled after an injection into the lateral geniculate nucleus encompasses the reticular regions which receive afferents from cortical areas 17, rostral 18b and caudal 18b. These results suggest that individual cortical areas may influence the activity of different dorsal thalamic nuclei through their reticular connections.     

A spatiotemporal profile of visual system activation revealed by current source density analysis in the awake macaque

We investigated the spatiotemporal activation pattern, produced by one visual stimulus, across cerebral cortical regions in awake monkeys. Laminar profiles of postsynaptic potentials and action potentials were indexed with current source density (CSD) and multiunit activity profiles respectively. Locally, we found contrasting activation profiles in dorsal and ventral stream areas. The former, like V1 and V2, exhibit a 'feedforward' profile, with excitation beginning at the depth of Lamina 4, followed by activation of the extragranular laminae. The latter often displayed a multilaminar/columnar profile, with initial responses distributed across the laminae and reflecting modulation rather than excitation; CSD components were accompanied by either no changes or by suppression of action potentials. System-wide, response latencies indicated a large dorsal/ventral stream latency advantage, which generalizes across a wide range of methods. This predicts a specific temporal ordering of dorsal and ventral stream components of visual analysis, as well as specific patterns of dorsal-ventral stream interaction. Our findings support a hierarchical model of cortical organization that combines serial and parallel elements. Critical in such a model is the recognition that processing within a location typically entails multiple temporal components or 'waves' of activity, driven by input conveyed over heterogeneous pathways from the retina.     

Cortical connections of the frontoparietal opercular areas in the rhesus monkey

The connections of the frontoparietal opercular areas were studied in rhesus monkeys by using antero- and retrograde tracer techniques. The rostral opercular cortex including the gustatory and proisocortical motor (ProM) areas is connected with precentral areas 3, 1, and 2 as well as with the rostral portion of the opercular area which resembles the second somatosensory type of cortex (area SII) and the ventral portion of area 6. Its distant connections are with the ventral portion of prefrontal areas 46, 11, 12, and 13 as well as with the rostral insula and cingulate motor area (CMAr). The mid opercular region (areas 1 and 2) is connected with pre- and postcentral areas 3, 1, and 2 as well as SII. Additionally, it is connected with the ventral portion of area 6, area 44, area ProM, the gustatory area, and the rostral insula. Its distant connections are with area 4, the ventral portion of area 46, area 7b, and area POa in the intraparietal sulcus (IPS). The rostral parietal opercular region is connected with the postcentral portions of areas 3, 1, and 2; areas 5, 7, and SII; the gustatory area; and the vestibular area. Its other connections are with area 4, area 44, the ventral portion of area 46, area ProM, CMAr, and the supplementary motor area (SMA). The caudal opercular region is connected with the dorsal portion of area 3; areas 2, 5, and 7a; and areas PEa as well as IPd of IPS. It is also connected with area SII, insula, and the superior temporal sulcus. Its distant connections are with area 44; the dorsal portion of area 8 and the ventral portion of area 46; as well as CMAr, SMA, and the supplementary sensory area. This connectivity suggests that the ventral somatosensory areas are involved in sensorimotor activities mainly related to head, neck, and face structures as well as to taste. Additionally, these areas may have a role in frontal (working) and temporal (tactile) memory systems.     

Diversity of connections of the temporal neocortex with amygdaloid nuclei in the dog (Canis familiaris)

Reciprocal connections of amygdaloid nuclei with the temporal neocortex in the dog were investigated. Injections of fluorescent tracers and BDA into particular temporal areas were made in eleven dogs. The topographical arrangement of connections and variations in their density differentiate the temporal neocortex in the dog into a few regions. Among them, the cortex involving the anterior part of the ectosylvian gyrus did not send any amygdalopetal projection. The middle ectosylvian, dorsal zone of the posterior ectosylvian and the anterior part of the Sylvian gyrus were weakly connected with the amygdala. The cortical region involving the ventral zone of the posterior ectosylvian and composite posterior areas, as well as posterior Sylvian gyrus, was characterized by profuse connections with the amygdaloid complex. Cortico-amygdaloid connections originate in the wide cortical area of the auditory cortex of the middle and dorsal part of the posterior ectosylvian gyrus as well as in the auditory association cortex located in the ventral ectosylvian, composite posterior and posterior Sylvian gyri. The connections showed a dorso-ventral gradient of increasing density, in the direction of association fields. The most substantial projection taking rise from the ectosylvian posterior and posterior composite gyri terminated preferentially in the pericapsular sector of the lateral amygdaloid nucleus and, to a lesser degree, in its medial sector. Terminals of connections originating in the Sylvian gyrus occupied preferentially the intermediate part of the lateral nucleus, slightly more medially than that from the ectosylvian and posterior composite areas. Additionally, axonal terminals derived from the composite posterior and Sylvian posterior areas were observed in the basal parvocellular and magnocellular nuclei. Neocortical projections were reciprocated by amygdalofugal connections with two exceptions: the basal magnocellular nucleus was distinguished by a substantial amygdalofugal projection to the temporal neocortex focused on the dorsal Sylvian gyrus, and the central nucleus of the amygdala, in contrast, received an exclusively corticofugal projection.     

Topography of the cortico-cortical connections from the striate cortex in the cat

Injections of radioactive amino acids were placed in regions of the striate cortex of cat representing central, intermediate and peripheral parts of the horizontal meridian and also in regions of lower and upper visual field representations near the vertical meridian. The study of cortico-cortical connections arising from these points revealed several retinotopic arrangements in the distribution of these connections and, it is argued, of the extrastriate cortical recipient areas themselves. On retinotopic distribution exists along the banks of the middle suprasylvian sulcus, in which lower and upper visual fields are rostral and caudal, respectively, and central and peripheral visual fields are ventral (or lateral) and dorsal (or medial), respectively, in the banks. This arrangement is called the lateral suprasylvian area (LS). Another retinotopic distribution exists along the caudal bank of the posterior suprasylvian sulcus. In this region, called the posteroior suprasylvania area(PS), points at, or near, the horizontal meridian are mainly represented, central and peripheral parts of which are located dorsaly and ventrally in the sulcus, respectively. Two other retinotopic distributions of connected exist in visual areas 2(V2) and 3 (V3) of Hubel and Wiesel, in which lower and upper visual fields are situated rostrally and caudally in these areas, respectively. Along the horizontal meridian, central is lateral in V3 and medial in V2, while more peripheral points, (15 degrees, 45 degrees) of V3 and V2 approach each other, in a mirror image fashion, along the coronal plane. However, representations of these peripheral parts of the horizontal meridian are repeated twice again: extensively, caudally along the lateral border of area 18, and more restrictively rostrally, along the lateral border of area 18.     

A visuo-somatomotor pathway through superior parietal cortex in the macaque monkey: cortical connections of areas V6 and V6A

This report addresses the connectivity of the cortex occupying middle to dorsal levels of the anterior bank of the parieto-occipital sulcus in the macaque monkey. We have previously referred to this territory, whose perimeter is roughly circumscribed by the distribution of interhemispheric callosal fibres, as area V6, or the 'V6 complex'. Following injections of wheatgerm agglutinin conjugated to horseradish peroxidase (WGA-HRP) into this region, we examined the laminar organization of labelled cells and axonal terminals to attain indications of relative hierarchical status among the network of connected areas. A notable transition in the laminar patterns of the local, intrinsic connections prompted a sub-designation of the V6 complex itself into two separate areas, V6 and V6A, with area V6A lying dorsal, or dorsomedial to V6 proper. V6 receives ascending input from V2 and V3, ranks equal to V3A and V5, and provides an ascending input to V6A at the level above. V6A is not connected to area V2 and in general is less heavily linked to the earliest visual areas; in other respects, the two parts of the V6 complex share similar spheres of connectivity. These include regions of peripheral representation in prestriate areas V3, V3A and V5, parietal visual areas V5A/MST and 7a, other regions of visuo-somatosensory association cortex within the intraparietal sulcus and on the medial surface of the hemisphere, and the premotor cortex. Subcortical connections include the medial and lateral pulvinar, caudate nucleus, claustrum, middle and deep layers of the superior colliculus and pontine nuclei. From this pattern of connections, it is clear that the V6 complex is heavily engaged in sensory-motor integration. The specific somatotopic locations within sensorimotor cortex that receive this input suggest a role in controlling the trunk and limbs, and outward reaching arm movements. There is a secondary contribution to the brain's complex oculomotor circuitry. That the medial region of the cortex is devoted to tightly interconnected representations of the sensory periphery, both visual and somatotopic-which are routinely stimulated in concert-would appear to be an aspect of the global organization of the cortex which must facilitate multimodal integration.     

Cortical hierarchy reflected in the organization of intrinsic connections in macaque monkey visual cortex

Neuronal response properties vary markedly at increasing levels of the cortical hierarchy. At present it is unclear how these variations are reflected in the organization of the intrinsic cortical circuitry. Here we analyze patterns of intrinsic horizontal connections at different hierarchical levels in the visual cortex of the macaque monkey. The connections were studied in tangential sections of flattened cortices, which were injected with the anterograde tracer biocytin. We directly compared the organization of connections in four cortical areas representing four different levels in the cortical hierarchy. The areas were visual areas 1, 2, 4 and Brodman's area 7a (V1, V2, V4 and 7a, respectively). In all areas studied, injections labeled numerous horizontally coursing axons that formed dense halos around the injection sites. Further away, the fibers tended to form separate clusters. Many fibers could be traced along the way from the injection sites to the target clusters. At progressively higher order areas, there was a striking increase in the spread of intrinsic connections: from a measured distance of 2.1 mm in area V1 to 9.0 mm in area 7a. Average interpatch distance also increased from 0.61 mm in area V1 to 1.56 mm in area 7a. In contrast, patch size changed far less at higher order areas, from an average width of 230 micron(s) in area V1 to 310 micron(s) in area 7a. Analysis of synaptic bouton distribution along axons revealed that average interbouton distance remained constant at 6.4 micron(s) (median) in and out of the clusters and in the different cortical areas. Larger injections resulted in a marked increase in the number of labeled patches but only a minor increase in the spread of connections or in patch size. Thus, in line with the more global computational roles proposed for the higher order visual areas, the spread of intrinsic connections is increased with the hierarchy level. On the other hand, the clustered organization of the connections is preserved at higher order areas. These clusters may reflect the existence of cortical modules having blob-like dimensions throughout macaque monkey visual cortex.     

Perirhinal and postrhinal cortices of the rat: interconnectivity and connections with the entorhinal cortex

The cortical regions dorsally adjacent to the posterior rhinal sulcus in the rat can be divided into a rostral region, the perirhinal cortex, which shares features of the monkey perirhinal cortex, and a caudal region, the postrhinal cortex, which has connectional attributes similar to the monkey parahippocampal cortex. We examined the connectivity among the rat perirhinal (areas 35 and 36), postrhinal, and entorhinal cortices by placing anterograde and retrograde tracers in all three regions. There is a dorsal-to-ventral cascade of connections in the perirhinal and entorhinal cortices. Dorsal area 36 projects strongly to ventral area 36, and ventral area 36 projects strongly to area 35. The return projections are substantially weaker. The cascade continues with the perirhinal to entorhinal connections. Area 35 is more strongly interconnected with the entorhinal cortex, ventral area 36 somewhat less strongly, and dorsal area 36 projects only weakly to the entorhinal cortex. The postrhinal-to-perirhinal connections also follow this general pattern. The postrhinal cortex is more heavily connected with dorsal area 36 than with ventral area 36 and is more heavily connected with area 36 than with area 35. The rostral portion of the postrhinal cortex has the strongest connections with the perirhinal cortex. Like in the monkey, the perirhinal and postrhinal cortices have different patterns of projections to the entorhinal cortex. The perirhinal cortex is preferentially connected with the rostrolateral portion of the entorhinal cortex. The postrhinal cortex projects to a part of this same region but is also connected to caudal and redial portions of the entorhinal cortex. The perirhinal and postrhinal projections to the entorhinal cortex originate in layers III and V and terminate preferentially in layers II and III.     

The processing of first- and second-order motion in human visual cortex assessed by functional magnetic resonance imaging (fMRI)

We have examined the activity levels produced in various areas of the human occipital cortex in response to various motion stimuli using functional magnetic resonance imaging (fMRI) methods. In addition to standard luminance-defined (first-order) motion, three types of second-order motion were used. The areas examined were the motion area V5 (MT) and the following areas that were delineated using retinotopic mapping procedures: V1, V2, V3, VP, V3A, and a new area that we refer to as V3B. Area V5 is strongly activated by second-order as well as by first-order motion. This activation is highly motion-specific. Areas V1 and V2 give good responses to all motion stimuli, but the activity seems to be related primarily to the local spatial and temporal structure in the image rather than to motion processing. Area V3 and its ventral counterpart VP also respond well to all our stimuli and show a slightly greater degree of motion specificity than do V1 and V2. Unlike V1 and V2, the response in V3 and VP is significantly greater for second-order motion than for first-order motion. This trend is evident, but less marked, in V3A and V3B and absent in V5. The results are consistent with the hypothesis that first-order motion sensitivity arises in V1, that second-order motion is first represented explicitly in V3 and VP, and that V5 (and perhaps also V3A and V3B) is involved in further processing of motion information, including the integration of motion signals of the two types.     

Modular organization of occipito-temporal pathways: cortical connections between visual area 4 and visual area 2 and posterior inferotemporal ventral area in macaque monkeys

The modular organization of cortical pathways linking visual area 4 (V4) with occipital visual area 2 (V2) and inferotemporal posterior inferotemporal ventral area (PITv) was investigated through an analysis of the patterns of retrogradely labeled cell bodies after injections of tracers into V4 and PITv. Although cytochrome oxidase or other stains have failed to yield reliable independent anatomical markers for cortical modules beyond V1 and V2, V4 and PITv seem to have modular compartments with specific patterns of cortico-cortical connectivity. Tracer injections of V4 labeled cells in V2 (1) thin stripes exclusively, (2) interstripes exclusively, or (3) specific combinations of interstripe and thin stripe subcompartments. These labeling patterns suggest (1) that there is a complicated organization of inputs to V4, (2) that projections from V2 to V4 display a submodular selectivity, and (3) that projections from V2 to V4 display some degree of cross-stream convergence. Consistent with this framework, extensive regions of PITv provide feedback projections to interstripe-recipient portions of V4, whereas more restricted portions of PITv provide feedback to thin stripe-recipient portions of V4. Similarly, the feedforward projection from V4 to PITv often arose from multiple cell clusters across a wide expanse of V4. When distinguishable fluorescent tracers were injected into two PITv sites separated by 3-5 mm, a variety of projection patterns was observed in V4. In most cases, labeled cells were found in multiple, interdigitating, nonoverlapping clusters of 1-3 mm width, whereas in other cases the two labeled fields were highly intermixed. These results suggest that V4 and PITv contain functional modules that can be characterized by the specific patterns of segregated and convergent projections they receive from lower cortical areas. These specific patterns of intercortical input, in conjunction with intrinsic cortical circuitry, may endow extrastriate cortical neurons with new and more complex receptive field properties.     

Safety and immunogenicity of HIV recombinant envelope vaccines in HIV-infected infants and children. National Institutes of Health-sponsored Pediatric AIDS Clinical Trials Group (ACTG-218)

We applied structural equation modeling to positron emission tomography data in humans to examine functional interactions between the right medial temporal lobe (MTL) and selected right neocortical regions in relation to visual recognition memory. Using a priori knowledge about anatomical connections between these regions as a guiding constraint, we modeled the pattern of interactions [i.e. covariances in regional cerebral blood flow (rCBF)] associated with episodic memory retrieval of spatial location and compared it with the pattern for retrieval of object identity. We also compared these patterns with those associated with perceptual matching of spatial location and object identity. Although displaying no difference in average rCBF across tasks, the right MTL showed domain-specific qualitative differences in interactions with posterior dorsal (parieto-occipital sulcus, supramarginal gyrus) and ventral regions (fusiform gyrus, superior temporal sulcus) but not with a prefrontal region. MTL interactions involving dorsal regions were positive in the spatial retrieval task but negative for object retrieval. Interactions involving ventral regions showed the reverse pattern. No comparable changes were observed during perceptual matching. Using control models, we demonstrated the neuroanatomical specificity of these results. Our results provide support for the notion that the nature of interactions between the MTL and posterior neocortex depends on the domain of information to-be-recovered.     

Efferent and afferent connections of the nucleus lateralis posterior thalami ("pulvinar") in the albino rats (author's transl)

The efferent and afferent connections of the lateral posterior nucleus (LP) of the albino rat were investigated light microscopically with the silver-degeneration-methods and the HRP-methods as well. The results are: 1. The main projection region of the LP is the area of 18a of the peristriate visual cortex. Most degenerating axons terminate in layer IV. A few fibers pass layers III and II and terminate in layer I. It is not sure if there are also terminating fibers in layer IV. We could not find a topistic relation between LP and area 18 a. 2. We observed a small number of degenerating fibers in area 17, too. 3. A part of the degenerating fibers runs to the temporal cortex end enters area 20. 4. There is no evidence for a projection of the LP to both the subcortical regions and to the superior colliculus. 5. The majority of the LP's afferent fibers originates - on the subcortical level - from the superior colliculus. Especially the lamina III (Str. opticum) of the ipsilateral and of the contralateral side is here the source of fibers terminating in the LP. 6. Other subcortical sources of fibers terminating in the LP are: the pretectal region, the ventral part of the LGN, the Zona incerta, the thalamic reticular formation, and the dorsal raphe nucleus. 7. There exists a fiber projection of the area 17 to the LP. The axons originate mainly from pyramidal cells in layer V. It is discussed whether the area-17-fibers terminating in the LP are collaterals of the fibers terminating in the superior colliculus. The projection of the area 18a to the LP is of greater importance. The axons of this area originate mainly from cells of the layer VI. It becomes obvious that the thalamic relay-station of the second visual pathway seems to project nearly exclusively to the neocortex. In contrast to the dorsal LGN, however, the LP is not only a simple relay-station for visual information as also non-visual information arrives here. The morphological basis for these inputs has not yet been clarified completely. We have to take into consideration as well as the connections with the superior colliculus and the pretectal region and the cortical connections. It is remarkable that there exists also a projection of LP-fibers to a region outside the classical visual cortex. In mammals of higher evolution that kind of projection extends increasingly. It is discussed if - under comparative-anatomical aspect - the morphological changes in the pulvinar region are an expression of the neocorticalization, whereas the morphological changes in the dorsal LGN reflect mainly the functional specialization of the visual system.     

Cortical feedback improves discrimination between figure and background by V1, V2 and V3 neurons

A single visual stimulus activates neurons in many different cortical areas. A major challenge in cortical physiology is to understand how the neural activity in these numerous active zones leads to a unified percept of the visual scene. The anatomical basis for these interactions is the dense network of connections that link the visual areas. Within this network, feedforward connections transmit signals from lower-order areas such as V1 or V2 to higher-order areas. In addition, there is a dense web of feedback connections which, despite their anatomical prominence, remain functionally mysterious. Here we show, using reversible inactivation of a higher-order area (monkey area V5/MT), that feedback connections serve to amplify and focus activity of neurons in lower-order areas, and that they are important in the differentiation of figure from ground, particularly in the case of stimuli of low visibility. More specifically, we show that feedback connections facilitate responses to objects moving within the classical receptive field; enhance suppression evoked by background stimuli in the surrounding region; and have the strongest effects for stimuli of low salience.     

The dorsomedial cortical visual area: a third tier area in the occipital lobe of the owl monkey (Aotus trivirgatus)

In the owl monkey, microelectrode mapping of Brodmann's area 19 indicates that this region contains part or all of at least 5 separate representations of the visual field, each of which adjoins the anterior border of V II and collectively are termed the third tier of cortical visual areas (V I is the first tier; V II is the second tier). Described in detail in this report is one of the third tier areas which is located on the dorsal surface and the adjacent medial wall of the occipital lobe and corresponds to a densely myelinated zone of cortex. In this dorsomedial area (DM), the representation of the horizontal meridian is partially split, and thus, like V II (see ref. 4) and the dorsolateral crescent5, DM is a second order transformation of the visual hemifield. In one abnormal owl monkey, a portion of the upper quadrant was represented twice in DM. This abnormal case may provide some clues as to how the normal pattern of visuotopic organization is established in the developing brain.     

Ventral intraparietal area of the macaque: anatomic location and visual response properties

1. The middle temporal area (MT) projects to the intraparietal sulcus in the macaque monkey. We describe here a discrete area in the depths of the intraparietal sulcus containing neurons with response properties similar to those reported for area MT. We call this area the physiologically defined ventral intraparietal area, or VIP. In the present study we recorded from single neurons in VIP of alert monkeys and studied their visual and oculomotor response properties. 2. Area VIP has a high degree of selectivity for the direction of a moving stimulus. In our sample 72/88 (80%) neurons responded at least twice as well to a stimulus moving in the preferred direction compared with a stimulus moving in the null direction. The average response to stimuli moving in the preferred direction was 9.5 times as strong as the response to stimuli moving in the opposite direction, as compared with 10.9 times as strong for neurons in area MT. 3. Many neurons were also selective for speed of stimulus motion. Quantitative data from 25 neurons indicated that the distribution of preferred speeds ranged from 10 to 320 degrees/s. The degree of speed tuning was on average twice as broad as that reported for area MT. 4. Some neurons (22/41) were selective for the distance at which a stimulus was presented, preferring a stimulus of equivalent visual angle and luminance presented near (within 20 cm) or very near (within 5 cm) the face. These neurons maintained their preference for near stimuli when tested monocularly, suggesting that visual cues other than disparity can support this response. These neurons typically could not be driven by small spots presented on the tangent screen (at 57 cm). 5. Some VIP neurons responded best to a stimulus moving toward the animal. The absolute direction of visual motion was not as important for these cells as the trajectory of the stimulus: the best stimulus was one moving toward a particular point on the face from any direction. 6. VIP neurons were not active in relation to saccadic eye movements. Some neurons (10/17) were active during smooth pursuit of a small target. 7. The predominance of direction and speed selectivity in area VIP suggests that it, like other visual areas in the dorsal stream, may be involved in the analysis of visual motion.     

Cortical afferents of the perirhinal, postrhinal, and entorhinal cortices of the rat

We have divided the cortical regions surrounding the rat hippocampus into three cytoarchitectonically discrete cortical regions, the perirhinal, the postrhinal, and the entorhinal cortices. These regions appear to be homologous to the monkey perirhinal, parahippocampal, and entorhinal cortices, respectively. The origin of cortical afferents to these regions is well-documented in the monkey but less is known about them in the rat. The present study investigated the origins of cortical input to the rat perirhinal (areas 35 and 36) and postrhinal cortices and the lateral and medial subdivisions of the entorhinal cortex (LEA and MEA) by placing injections of retrograde tracers at several locations within each region. For each experiment, the total numbers of retrogradely labeled cells (and cell densities) were estimated for 34 cortical regions. We found that the complement of cortical inputs differs for each of the five regions. Area 35 receives its heaviest input from entorhinal, piriform, and insular areas. Area 36 receives its heaviest projections from other temporal cortical regions such as ventral temporal association cortex. Area 36 also receives substantial input from insular and entorhinal areas. Whereas area 36 receives similar magnitudes of input from cortices subserving all sensory modalities, the heaviest projections to the postrhinal cortex originate in visual associational cortex and visuospatial areas such as the posterior parietal cortex. The cortical projections to the LEA are heavier than to the MEA and differ in origin. The LEA is primarily innervated by the perirhinal, insular, piriform, and postrhinal cortices. The MEA is primarily innervated by the piriform and postrhinal cortices, but also receives minor projections from retrosplenial, posterior parietal, and visual association areas.     

Responses of neurons in primary and inferior temporal visual cortices to natural scenes

The primary visual cortex (V1) is the first cortical area to receive visual input, and inferior temporal (IT) areas are among the last along the ventral visual pathway. We recorded, in area V1 of anaesthetized cats and area IT of awake macaque monkeys, responses of neurons to videos of natural scenes. Responses were analysed to test various hypotheses concerning the nature of neural coding in these two regions. A variety of spike-train statistics were measured including spike-count distributions, interspike interval distributions, coefficients of variation, power spectra, Fano factors and different sparseness measures. All statistics showed non-Poisson characteristics and several revealed self-similarity of the spike trains. Spike-count distributions were approximately exponential in both visual areas for eight different videos and for counting windows ranging from 50 ms to 5 seconds. The results suggest that the neurons maximize their information carrying capacity while maintaining a fixed long-term-average firing rate, or equivalently, minimize their average firing rate for a fixed information carrying capacity.     

Corticothalamic connections of extrastriate visual areas in rhesus monkeys

Corticothalamic connections of extrastriate visual areas were studied by using the autoradiographic anterograde tracing technique. The results show that the medial extrastriate region above the calcarine sulcus projects mainly to the lateral pulvinar (PL), medial pulvinar (PM), and lateral posterior (LP) nuclei. In addition, the dorsal portion of the medial region has connections to the lateral dorsal (LD) as well as to intralaminar nuclei. The dorsolateral extrastriate region projects strongly to the PL and LP nuclei, to the PM and inferior pulvinar (PI) nuclei, and to the LD and intralaminar nuclei. The lateral extrastriate region above the inferior occipital sulcus (IOS) has strong connections to both the PL and PI nuclei and has minor projections to the PM and oral pulvinar nuclei. The ventrolateral extrastriate region below the IOS projects mainly to the PI nucleus and to the caudal portion of the PL nucleus and has some projections to the PM nucleus. The ventromedial extrastriate region medial to the occipitotemporal sulcus has strong connections with the ventral and medial sectors of the PI nucleus. This region also projects to the caudal portion of the PL nucleus and has minor connections to the LP nucleus. Finally, the annectant gyrus projects to the PL nucleus and to the rostral portion of the PI nucleus and has minor connections to the PM nucleus. Thus, the medial and dorsolateral extrastriate regions are related mainly to the PL and LP nuclei as well as to intralaminar nuclei. In contrast, the ventrolateral and ventromedial regions are connected strongly with the PI nucleus. This connectional organization appears to reflect functional differentiation at the cortical level.