Prestriate Areas Research Articles

Sub-cortical areas process physical size but not numerical value.

A robust finding in the numerical cognition literature is that the physical size and the numerical value of two to-be-compared digits interact, resulting in a size congruity effect (SiCE). The current study focuses on the possible role of prestriate areas in a digit comparison Stroop-like task. In the visual pathway, prestriate areas refer to regions from the retina up to the primary visual cortex (V1). I hypothesized that processing of physical size, but not numerical value, begins already in prestriate areas. This is because physical size is a basic visual feature while the numerical value of a symbol is a learned convention that should be retrieved from long-term memory. Adult participants compared the size or the numerical value of two digits. Without participants' awareness, I projected the digits either to the same eye, or each digit to a different eye. The latter type of presentation prevents prestriate areas from taking part in comparing the digits. Therefore, slower aresponse time under this condition hints atthe involvement of prestriate areas. Evidence confirmed the initial hypothesis, demonstrating slower performance when the stimuli are segregated between the eyes but only for physical size comparisons. This finding suggests that at least the initial processing of physical size, when relevant, is done before, and by different neural substrate than numerical value. The implications of the study and future directions are discussed.

The influence of visual and phonological features on the hemispheric processing of hierarchical Navon letters

The possibility of allocating attentional resources to the “global” shape or to the “local” details of pictorial stimuli helps visual processing. Investigations with hierarchical Navon letters, that are large “global” letters made up of small “local” ones, consistently demonstrate a right hemisphere advantage for global processing and a left hemisphere advantage for local processing. Here we investigated how the visual and phonological features of the global and local components of Navon letters influence these hemispheric advantages. In a first study in healthy participants, we contrasted the hemispheric processing of hierarchical letters with global and local items competing for response selection, to the processing of hierarchical letters in which a letter, a false-letter conveying no phonological information or a geometrical shape presented at the unattended level did not compete for response selection. In a second study, we investigated the hemispheric processing of hierarchical stimuli in which global and local letters were both visually and phonologically congruent (e.g. large uppercase G made of smaller uppercase G), visually incongruent and phonologically congruent (e.g. large uppercase G made of small lowercase g) or visually incongruent and phonologically incongruent (e.g. large uppercase G made of small lowercase or uppercase M). In a third study, we administered the same tasks to a right brain damaged patient with a lesion involving pre-striate areas engaged by global processing. The results of the first two experiments showed that the global abilities of the left hemisphere are limited because of its strong susceptibility to interference from local letters even when these are irrelevant to the task. Phonological features played a crucial role in this interference because the interference was entirely maintained also when letters at the global and local level were presented in different uppercase vs. lowercase formats. In contrast, when local features conveyed no phonological information, the left hemisphere showed preserved global processing abilities. These findings were supported by the study of the right brain damaged patient. These results offer a new look at the hemispheric dominance in the attentional processing of the global and local levels of hierarchical stimuli.

The relationship between visuospatial disorders in Parkinson’s disease and thickness of retina and cortex

This paper presents the current data on the mechanisms of formation of visuospatial disorders in Parkinson’s disease (PD). Optical coherence tomography method was used to study the retina, using the protocols designed to assess the thickness of ganglionic layer, retinal nerve fiber layer, and macular map. Central structures were assessed using MRI of the brain followed by evaluation of the cortical thickness. Patients underwent neuropsychological tests, including tests designed to assess the severity of the disease, visuospatial perception, and cognitive functions. The results revealed the retinal and prestriate areas with the most significant changes in patients with different stages of PD, and allowed us to hypothesize the role of changes in the peripheral and central structures of the visual analyzer in formation of symptom complex of visuospatial disorders in PD.

Color experience in blindsight?

Blindsight, the ability to blindly discriminate wavelength and other aspects of stimuli in a blind field, sometimes occurs in people with lesions to striate (V1) cortex. There is currently no consensus on whether qualitative color information of the sort that is normally computed by double opponent cells in striate cortex is indeed computed in blindsight but doesn’t reach awareness, perhaps owing to abnormal neuron responsiveness in striate or extra-striate cortical areas, or is not computed at all. The existence of primesight, the experience of colored afterimages in blindsight, has been taken to suggest that qualitative color information is computed either in pre-striate or striate cortical areas but is not broadcast to working memory. I argue here that a recent study in which color phosphenes were induced in a blindsighter using bilateral transcranial magnetic stimulation indicates that computations necessary for conscious color vision are lost in blindsight. Owing to this loss, the neural responsiveness in extrastriate cortical areas is abnormal and hence is unable to give rise to color awareness. Blindsight is thus degraded vision in which the computations necessary for conscious color vision have been lost.

What shall we do with the preattentive processing stage: Use it or lose it?

Since Neisser (1967) and Treisman & Gelade (1980) many models of selective visual attention have included a stage of processing. Some researchers have suggested that the term preattentivehas outlived its usefulness. While there are uses of preattentivethat should be avoided, we argue against abandoning the concept altogether. If there is selective visual attention and if there is processing of visual stimuli prior to selection, then there is processing. We hold that a goal of this processing is to abstract a representation from the stimulus that can be used to guide attention. In that representation, features are coarsely coded in parallel. It is a mistake to think of this representation as self-sufficient visionwhose output can directly control behavior. While some forms of perceptual grouping and segregation may be accomplished preattentively, it does not seem to be possible to go directly to a motor response without engaging attention- demanding processes. In particular, while there may be some conjoining of features, the accurate explicit binding that appears to be required for object recognition entails selective attention to specific objects. Building on the Reverse Hierarchy Theory of Hochstein & Ahissar and on re-entrant ideas of DiLollo, we suggest that features are initially encoded through a feed-forward pathway from V1 to higher visual areas in both ventral and dorsal pathways. Preattentive guidance of focused attention can then be modeled as re-entry from anatomical- ly high-level loci onto earlier loci. Parietal areas appear to guide spatial selection and prestriate areas may guide feature-based selection. In this framework, no part of the brain is exclusively preattentive . The same anatomical areas may participate in the feed-forward production of a representation and then in the more detailed, selective processing modulated by attention through subsequent reentry. Acknowledgment: Grant support from NEI, NIMH, AFOSR

Neurons in Area V4 of the Macaque Translate Attended Visual Features into Behaviorally Relevant Categories

Neural processing at most stages of the primate visual system is modulated by selective attention, such that behaviorally relevant information is emphasized at the expenses of irrelevant, potentially distracting information. The form of attention best understood at the cellular level is when stimuli at a given location in the visual field must be selected (space-based attention). In contrast, fewer single-unit recording studies have so far explored the cellular mechanisms of attention operating on individual stimulus features, specifically when one feature (e.g., color) of an object must guide behavioral responses while a second feature (e.g., shape) of the same object is potentially interfering and therefore must be ignored. Here we show that activity of neurons in macaque area V4 can underlie the selection of elemental object features and their "translation" into a categorical format that can directly contribute to the control of the animal's behavior.

Distribution of alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionate-type glutamate receptor subunits (GluR2/3) along the ventral visual pathway in the monkey.

By using immunohistochemical methods, we examined the distribution of cells expressing subunits of alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionate (AMPA)-selective glutamate receptors (GluR2/3) in the cortical areas of the occipitotemporal pathway in monkeys. GluR2/3-immunoreactive (-ir) cells were primarily pyramidal cells; this category, however, also included large stellate cells in layer IVB of the striate cortex (V1) and fusiform cells in layer VI of all the areas examined. GluR2/3 immunoreactivity differed among the areas in laminar distribution and intensity. In V1, GluR2/3-ir cells were identified mainly in layers II, III, IVB, and VI. The prestriate areas V2 and V4 and the inferior temporal areas TEO and TE contained GluR2/3-ir cells in layers II, III, and VI. In the TE, GluR2/3-ir cells were also abundant in layer V. In area 36 of the perirhinal cortex, neurons in layers II, III, V, and VI were labeled in a similar manner to the TE labeling, but with greater staining intensity and numbers, especially in layer V. Thus, GluR2/3 immunoreactivity increased rostrally along the pathway. Within V1 and V2, cells strongly stained for GluR2/3 formed clusters that colocalized with cytochrome oxidase (CO)-rich regions. These distinct laminar and regional distribution patterns of GluR2/3 expression may contribute to the specific physiological properties of neurons within various visual areas and compartments.

Functional organization of macaque V3 for stereoscopic depth.

Recordings were made from single and small groups of cells in prestriate area V3 of the visual cortex of the Cynomolgus macaque (Macaca fascicularis). The majority of cells in V3 were selective for orientation and stereoscopic depth, these cells being segregated into two sets of functionally distinct columns. Orientation columns in V3 have been previously demonstrated; here we show that V3 also contains columns of segregated disparity-selective cells. On the basis of its cellular properties, functional organization, and intra-cortical connections, we propose that V3 contributes to the processing of stereoscopic depth information and that the parietal areas to which it projects use this information for the analysis of object depth and three-dimensional form.

Corticopulvinar connections of areas V5, V4, and V3 in the macaque monkey: a dual model of retinal and cortical topographies.

The connection zones of cortical areas V3, V4, and V5 (MT) with the thalamic pulvinar nucleus in the macaque monkey were identified. A combination of single- and dual-tracer techniques was used to study their topography and to establish whether these zones occupy separate or overlapping pulvinar territories. In each case, the retinotopic distribution of tracer in the pulvinar was charted by reference to its parallel distribution within the maps of cortical areas V1 and V2. Each of the areas V3, V4, and V5 were found to connect with both the 1° and the 2° maps located within the inferior and lateral pulvinar nuclei and to respect the previously identified topographies of these maps. However, V5 connects to a narrow zone lining the rostrolateral margin of the lateral and inferior pulvinar and V4 to a broader zone within the body of these two nuclei, which is adjacent to but separate from the V5 zone; the V3 zone overlaps both. Focal injections into cortex produce columns of pulvinar label whose trajectory defines a line of isorepresentation. The lines of isorepresentation in the 1° and 2° maps are approximately linear and parallel and adopt a rostrolateral to caudomedial axis; in the 1° map, this axis is roughly perpendicular to the facet of the inferior pulvinar that lies adjacent to the lateral geniculate nucleus. The connections of V5 and V4 can be modelled as successive zones along the axis of isorepresentation, with registered visual topographies. The scheme is extended by existing reports that inferotemporal cortex connects to the caudomedial pole of this axis-reflecting an occipitotemporal cortical gradient, in that V1 and other prestriate areas, e.g., V3, connect to the opposite pole. Thus a simple model of the mapped volume in the pulvinar arises, in which a unidimensional cortical topography is represented orthogonally to retinal topography. Adjoining this volume medially, within the inferior and medial pulvinar, is a second, heavier zone of V5 connectivity, which is poorly topographic. Both the medial and the rostrolateral zones of V5 connectivity may overlap with previously identified regions of tectal input to the pulvinar.

Cortical input to the nucleus of the optic tract and dorsal terminal nucleus (NOT-DTN) in macaques: a retrograde tracing study.

Using retrograde tracing methods, we investigated the cortical projection to the nucleus of the optic tract and dorsal terminal nucleus of the accessory optic system (NOT-DTN) in macaque monkeys. Tracer injections at electrophysiologically identified recording sites in the NOT-DTN resulted in retrogradely labelled neurons in layer V of various cortical areas. The strongest projection always arose from the middle temporal area (MT) and the adjoining cortex anterior to MT in the superior temporal sulcus. A less dense projection came from the middle superior temporal area (MST). In addition, retrogradely labelled cells were consistently found in areas V1 and V2 at moderate to high density. Furthermore, sparse to moderate labelling occurred in prestriate area V3. These findings were compared with the label resulting from control injections into the superior colliculus in two additional cases. Our results indicate that the cortical input to the NOT-DTN as the sensorimotor interface for the pathway subserving stabilizing eye movements during the optokinetic reflex and smooth pursuit mainly arises from the motion-sensitive areas MT and MST in the superior temporal sulcus, as well as from areas V1 and V2. Clearly the projection to the NOT-DTN does not arise from a single cortical area.

The cortical connections of area V6: an occipito‐parietal network processing visual information

The aim of this work was to study the cortical connections of area V6 by injecting neuronal tracers into different retinotopic representations of this area. To this purpose, we first functionally recognized V6 by recording from neurons of the parieto-occipital cortex in awake macaque monkeys. Penetrations with recording syringes were performed in the behaving animals in order to inject tracers exactly at the recording sites. The tracers were injected into the central or peripheral field representation of V6 in different hemispheres. Irrespective of whether injections were made in the centre or periphery, area V6 showed reciprocal connections with areas V1, V2, V3, V3A, V4T, the middle temporal area /V5 (MT/V5), the medial superior temporal area (MST), the medial intraparietal area (MIP), the ventral intraparietal area (VIP), the ventral part of the lateral intraparietal area and the ventral part of area V6A (V6AV). No labelled cells or terminals were found in the inferior temporal, mesial and frontal cortices. The connections of V6 with V1, and with all the retinotopically organized prestriate areas, were organized retinotopically. The connection of V6 with MIP suggests a visuotopic organization for this latter. Labelling in V6A and VIP after either central or peripheral V6 injections was very similar in location and extent, as expected on the basis of the nonretinotopic organization of these areas. We suggest that V6 plays a pivotal role in the dorsal visual stream, by distributing the visual information coming from the occipital lobe to the sensorimotor areas of the parietal cortex. Given the functional characteristics of the cells of this network, we suggest that it could perform the fast form and motion analyses needed for the visual guiding of arm movements as well as their coordination with the eyes and the head.

Neural representation of three-dimensional features of manipulation objects with stereopsis.

In the first part of this article, we review our neurophysiological studies of the hand-manipulation-related neurons in the anterior part of the lateral bank of the intraparietal sulcus (area AIP). We describe the properties of visually responsive neurons in area AIP. Object-type visual-dominant neurons responded to the sight of objects and showed selectivity not only for simple geometrical shapes, but also for complex objects such as a knob-in-groove and a plate-in-groove. Some of the object-type visual-dominant neurons showed selectivity for the orientation of the longitudinal axis or the plane (surface) of a plate or a ring. In the second part of this article, we review our study of binocular visual neurons in the caudal part of the lateral bank of the intraparietal sulcus (c-IPS area), in particular, of axis-orientation-selective (AOS) neurons and surface-orientation-selective (SOS) neurons. AOS neurons preferred long and thin stimuli, were sensitive to binocular disparity, and tuned to the axis orientation in three-dimensional (3D) space. SOS neurons preferred broad and flat stimuli and were tuned to the surface orientation in depth. Some SOS neurons responded to a square in a random dot stereogram (RDS) with orientation tuning, suggesting that they encode surface orientation from a disparity gradient. Others responded to solid figure stereograms with orientation disparity and/or width disparity. It was concluded that the c-IPS area is a higher center for stereopsis, which integrates various binocular disparity signals received from the V3 complex and other prestriate areas to represent the neural code for 3D features. It may send projections to the AIP area and contribute to visual adjustment of the shape of the handgrip and/or hand orientation for manipulation and grasping. Neurons of the AIP area may also receive monocular cues of depth from the ventral visual pathway to discriminate the 3D shape of the object of manipulation.

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.

Hemispheric specialization for global and local processing: the effect of stimulus category.

Neuropsychological evidence indicates that the global aspect of complex visual scenes is preferentially processed by the right hemisphere, and local aspects are preferentially processed by the left hemisphere. Using letter-based hierarchical stimuli (Navon figures), we recently demonstrated, in a directed-attention task, lateralized neural activity (assessed by positron emission tomography) in the left prestriate cortex during local processing, and in the right prestriate cortex during global processing. Furthermore, temporal-parietal cortex was critically activated bilaterally in a divided-attention task that involved varying the number of target switches between local and global levels of letter-based hierarchical stimuli. Little is known about whether such stimulus categories influence such hemispheric lateralization. We now present data on brain activity, derived from positron emission tomography, in normal subjects scanned during either local or global processing of object-based hierarchical stimuli. We again observe attentional modulation of neural activity in prestriate cortex. There is now greater right-sided activation for local processing and greater left-sided activation for global processing, which is the opposite of that seen with letter-based stimuli. The results suggest that the relative differential hemispheric activations in the prestriate areas during global and local processing are modified by stimulus category.

Distribution and density of monoamine receptors in the primate visual cortex devoid of retinal input from early embryonic stages

Developmental mechanisms that regulate the areal and laminar distribution of various macromolecules, including neurotransmitter receptors in the cerebral cortex, are not known. In the present study, we examined the development of monoaminergic receptors in the rhesus monkey striate and peristriate visual cortex in the absence of input from the retina. Binocular enucleation was performed between embryonic days E60 and E81, prior to the ingrowth of geniculocortical fibers into the cortical plate and before genesis of the granular and supragranular layers of the visual cortex. The animals were delivered at term (E165) and sacrificed at 2 or 12 months of age, and their brains frozen and the occipital lobes cut at 20 microns in the coronal plane. Cortical binding of 3H-clonidine, 125I-pindolol, 3H-5-HT, 3H-ketanserin, 3H-spiperone, 3H-SCH23390, and 3H-prazosin that label various monoamine receptors were autoradiographically visualized and quantified using a computer imaging system. All radioligands displayed specific laminar patterns in the striate and prestriate areas in both groups of animals. The areal and laminar distribution in the anophthalmic monkeys was similar to that in the controls. Significantly, in all enucleated animals, just as in the controls, a particularly high density of 3H-clonidine and 3H-prazosin was observed in the sublayers of layer IV involved in color vision. The present results show that the monoamine receptors in primate visual cortex can establish and maintain distinct laminar and areal patterns in the absence of activity or molecular cues originated from the retina, and provide new insight into the cortical consequences of secondary congenital anophthalmia.

Comparison of neuronal selectivity for stimulus speed, length, and contrast in the prestriate visual cortical areas V4 and MT of the macaque monkey.

1. Prestriate area V4 and the middle temporal area (MT) compose the first stage in which the ventral and dorsal visual cortical pathways are segregated. To better known the functional dichotomy between the two pathways at this level, we recorded cell responses from V4 and MT using anesthetized, immobilized macaque monkeys and compared the selectivity for speed of stimulus motion and stimulus length and the sensitivity to luminance contrast between the two areas. 2. V4 cells were as selective as MT cells for speed. The sharpness of tuning was not different between the two populations. The optimal speed varied widely in both areas, but both of the two distributions showed peaks at 32 degrees/s. 3. V4 and MT cells were similar in that about one-half of the cells (45% in V4 and 48% in MT) showed inhibition by long (16 degrees) bars. However, V4 cells preferred stimuli whose lengths were distributed around the lengths of the receptive field, whereas an overwhelming majority of MT cells preferred stimuli whose lengths were much shorter than the lengths of the receptive field. 4. The cutoff contrast at which one-half the maximum response was elicited was distributed widely in both areas, and the two distributions considerably overlapped. MT cells as a whole, however, were slightly more sensitive to the luminance contrast than V4 cells. 5. There was a tendency toward local clustering for cells with similar speed preferences in MT but not in V4. Pairs of MT cells recorded within 400 microns had smaller difference in the optimal speed than that of cell pairs taken randomly from the whole sample of MT cells.

Direct temporal-occipital feedback connections to striate cortex (V1) in the macaque monkey.

Although there have been reports of sparse projections from temporal areas TE, TF, and even TH to area V1, it is generally believed that cortical afferents to V1 originate exclusively from prestriate areas. Injections of anterograde tracers in anterior occipital and temporal areas, however, consistently produce labeled terminals in area V1. In order to confirm these results and display the full range of foci projecting to V1, we injected V1 in two monkeys with the retrograde tracer fast blue. Feedback connections were found, as expected, from several prestriate areas (V2, V3, V4, and MT). These originate from neurons in layers 3A and 6. Connections were also found from several more distal regions, namely, areas TEO, TE, TF, TH, and from cortex in the occipitotemporal and superior temporal (STS) sulci. Filled neurons occurred in two small foci in the caudal intraparietal sulcus. These more distal feedback connections tend to originate only from layer 6. An additional injection of the retrograde tracer diamidino yellow in area V2 of one animal revealed a similarly widespread network of feedback connections. In some areas (in the STS and in TEO), 10-15% of fluorescent neurons were double-labeled. These results indicate that feedback connections to early visual cortex derive from a widespread network of areas, including limbic-associated cortices. These connectional patterns testify to the massive recursiveness of anatomical pathways. As there are no reports of projections from V1 to anterior temporal cortices, our results also indicate that some cortical feedback connections may not be strictly reciprocal.

Neural correlates of motion perception.

Most neurons in the prestriate area MT of the macaque monkey exhibit direction selectivity to moving visual stimuli. Deficits in the initiation of pursuit eye movements and those in the discrimination of the direction of moving random dot stimuli have been produced by MT lesion. Also, facilitation of the direction discrimination has been produced by MT microstimulation. These experiments provide evidence of causal relation between activities of MT neurons and motion perception. However, recent studies have shown that MT neurons are also affected by visual cues that are not directly involved in the motion perception. It is suggested that integration of motion-related inputs and other inputs observed in MT neurons is important for the segmentation of visual images into different objects.

Role of the dorsal prestriate cortex in visuospatial configural discrimination by monkeys

Cynomolgus monkeys ( Macaca fascicularis) learned a series of visuospatial configural discriminations in which particular discriminative stimulus objects were rewarded only in particular spatial locations. For example, object X was rewarded if it was on the left but not if it was on the right. After ablation of part of the dorsal prestriate cortex they were impaired in learning discriminations of this kind. The same animals were not impaired in learning visual object discriminations in which spatial position was irrelevant, nor in learning spatial discriminations in which object identity was irrelevant. The results were compared with previously reported results from fornix transection in the same tasks; the deficit following dorsal prestriate ablation in visuospatial configural discrimination learning was similar in severity to that which followed fronix transection. The results show that the dorsal prestriate area has a more general role in visuspatial processing than was known hitherto, and they suggest that it interacts with the hippocampal formation and fornix in visuospatial memory tasks.

Cortical connections of inferior temporal area TEO in macaque monkeys

In macaque monkeys, lesions involving the posterior portion of the inferior temporal cortex, cytoarchitectonic area TEO, produce a severe impairment in visual pattern discrimination. Recently, this area has been shown to contain a complete, though coarse, representation of the contralateral visual field (Boussaoud, Desimone, and Ungerleider: J. Comp. Neurol. 306:554-575, '91). Because the inputs and outputs of area TEO have not yet been fully described, we injected a variety of retrograde and anterograde tracers into 11 physiologically identified sites within TEO of seven rhesus monkeys and analyzed the areal and laminar distribution of its cortical connections. Our results show that TEO receives feedforward, topographically organized inputs from prestriate areas V2, V3, and V4. Additional sparser feedforward inputs arise from areas V3A, V4t, and MT. Each of these inputs is reciprocated by a feedback projection from TEO. TEO was also found to have reciprocal intermediate-type connections with the fundus of the superior temporal area (area FST), cortex in the most posteromedial portion of the superior temporal sulcus (the posterior parietal sulcal zone [area PP]), cortex in the intraparietal sulcus (including the lateral intraparietal area [area LIP]), the frontal eye field, and area TF on the parahippocampal gyrus. The connections with V3A, V4t, and PP were found only after injections in the peripheral field representations of TEO. Finally, TEO was found to project in a feedforward pattern to area TE and to areas anterior to FST on the lateral bank and floor of the superior temporal sulcus (areas TEm, TEa, and IPa, Seltzer and Pandya: Brain Res. 149:1-24, '78), all of which send feedback projections to TEO. Feedback projections also arise from parahippocampal area TH, and areas TG, 36, and possibly 35. These are complemented by only sparse feedforward projections to TG from central field representations in TEO and to TH from peripheral field representations. The results thus indicate that TEO forms an important link in the occipitotemporal pathway for object recognition, sending visual information forward from V1 and prestriate relays in V2-V4 to anterior inferior temporal area TE.