Anatomical evidence for the posterior boundary of area 2 in the macaque monkey

This study examined the architectonic organization of the macaque's primary somatosensory cortex near the tip of the intraparietal sulcus (IPS), using myelin and Nissl stains plus immunohistochemical labeling with the SMI-32 antibody. The surface cortex between the IPS and central sulcus (overlapping area 2) was distinguished from surrounding cortex (areas 1 and 5) by relatively light SMI-32 immunoreactivity. This distinguishing architectonic feature was most evident between the post-central dimple and cortex immediately anterior to the tip of the IPS. Physiological mappings verified that the architectonic transition correlated with a change in receptive field properties, consistent with their marking the boundary between areas 2 and 5. These results suggest that area 2 occupies surface cortex anterior to the IPS, but not within the IPS.     

Crossmodal processing of object features in human anterior intraparietal cortex: an fMRI study implies equivalencies between humans and monkeys

The organization of macaque posterior parietal cortex (PPC) reflects its functional specialization in integrating polymodal sensory information for object recognition and manipulation. Neuropsychological and recent human imaging studies imply equivalencies between human and macaque PPC, and in particular, the cortex buried in the intraparietal sulcus (IPS). Using functional MRI, we tested the hypothesis that an area in human anterior intraparietal cortex is activated when healthy subjects perform a crossmodal visuo-tactile delayed matching-to-sample task with objects. Tactile or visual object presentation (encoding and recognition) both significantly activated anterior intraparietal cortex. As hypothesized, neural activity in this area was further enhanced when subjects transferred object information between modalities (crossmodal matching). Based on both the observed functional properties and the anatomical location, we suggest that this area in anterior IPS is the human equivalent of macaque area AIP.     

Morphological patterns of the intraparietal sulcus and the anterior intermediate parietal sulcus of Jensen in the human brain

Distinct parts of the intraparietal sulcal cortex contribute to sensorimotor integration and visual spatial attentional processing. A detailed examination of the morphological relations of the different segments of the complex intraparietal sulcal region in the human brain in standard stereotaxic space, which is a prerequisite for detailed structure-to-function studies, is not available. This study examined the intraparietal sulcus (IPS) and the related sulcus of Jensen in magnetic resonance imaging brain volumes registered in the Montreal Neurological Institute stereotaxic space. It was demonstrated that the IPS is divided into two branches: the anterior ramus and the posterior ramus of the IPS, often separated by a submerged gyral passage. The sulcus of Jensen emerges between the anterior and posterior rami of the IPS, and its ventral end is positioned between the first and second caudal branches of the superior temporal sulcus. In a small number of brains, the sulcus of Jensen may merge superficially with the first caudal branch of the superior temporal sulcus. The above morphological findings are discussed in relation to previously reported functional neuroimaging findings and provide the basis for future exploration of structure-to-function relations in the posterior parietal region of individual subjects.     

Polymodal motion processing in posterior parietal and premotor cortex: a human fMRI study strongly implies equivalencies between humans and monkeys

In monkeys, posterior parietal and premotor cortex play an important integrative role in polymodal motion processing. In contrast, our understanding of the convergence of senses in humans is only at its beginning. To test for equivalencies between macaque and human polymodal motion processing, we used functional MRI in normals while presenting moving visual, tactile, or auditory stimuli. Increased neural activity evoked by all three stimulus modalities was found in the depth of the intraparietal sulcus (IPS), ventral premotor, and lateral inferior postcentral cortex. The observed activations strongly suggest that polymodal motion processing in humans and monkeys is supported by equivalent areas. The activations in the depth of IPS imply that this area constitutes the human equivalent of macaque area VIP.     

Cortical neurons projecting to the posterior part of the superior temporal sulcus with particular reference to the posterior association area. An HRP study in the monkey

Corticocortical connections from the posterior association area to the posterior part of the superior temporal sulcal cortex (STs area) were studied in the monkey by means of retrograde axonal transport of horseradish peroxidase (HRP) or wheatgerm-agglutinin-conjugated HRP (WGA-HRP). After injecting 0.05-0.2 microliter of 50% HRP or 5% WGA-HRP into the STs area, labeled cells were examined in various cortical regions. The dorsal wall of the STs receives fibers mainly from the inferior parietal lobule (area 7) and superior temporal gyrus (area 22), whereas the ventral wall and floor part of the STs receive fibers from the posterior inferotemporal gyrus (area TEO) and prestriate cortex (areas 18 and 19). The deeper parts of the dorsal wall close to the floor region of the STs area also receive many fibers from the cortical walls surrounding the intraparietal, lunate and lateral sulci. Both the dorsal and ventral cortical walls of the intraparietal sulcus send fibers mainly to the deep dorsal wall of the STs. The ventral wall of the STs, on the other hand, receives fibers only from the ventral wall of the intraparietal sulcus. The medial surface of the prestriate cortex and the parahippocampal region send fibers to both walls of the STs. In the prestriate-STs projections originating from areas around the parieto-occipital sulcus, a topographic correlation is present; area 19 located anterior to the sulcus projects to the dorsal wall, whereas area 18 situated posterior to the sulcus projects to the ventral wall. Only the dorsal wall receives fibers from the cingulate (areas 23 and 24) and subparietal gyri (area 7). The deeper part of the dorsal wall and the ventral wall of the posterior STs area are interconnected with each other, while the upper part of the dorsal wall does not appear to receive fibers from the ventral wall.     

Distinct Parietal and Temporal Pathways to the Homologues of Broca's Area in the Monkey

The homologues of the two distinct architectonic areas 44 and 45 that constitute the anterior language zone (Broca's region) in the human ventrolateral frontal lobe were recently established in the macaque monkey. Although we know that the inferior parietal lobule and the lateral temporal cortical region project to the ventrolateral frontal cortex, we do not know which of the several cortical areas found in those regions project to the homologues of Broca's region in the macaque monkey and by means of which white matter pathways. We have used the autoradiographic method, which permits the establishment of the cortical area from which axons originate (i.e., the site of injection), the precise course of the axons in the white matter, and their termination within particular cortical areas, to examine the parietal and temporal connections to area 44 and the two subdivisions of area 45 (i.e., areas 45A and 45B). The results demonstrated a ventral temporo-frontal stream of fibers that originate from various auditory, multisensory, and visual association cortical areas in the intermediate superolateral temporal region. These axons course via the extreme capsule and target most strongly area 45 with a more modest termination in area 44. By contrast, a dorsal stream of axons that originate from various cortical areas in the inferior parietal lobule and the adjacent caudal superior temporal sulcus was found to target both areas 44 and 45. These axons course in the superior longitudinal fasciculus, with some axons originating from the ventral inferior parietal lobule and the adjacent superior temporal sulcus arching and forming a simple arcuate fasciculus. The cortex of the most rostral part of the inferior parietal lobule is preferentially linked with the ventral premotor cortex (ventral area 6) that controls the orofacial musculature. The cortex of the intermediate part of the inferior parietal lobule is linked with both areas 44 and 45. These findings demonstrate the posterior parietal and temporal connections of the ventrolateral frontal areas, which, in the left hemisphere of the human brain, were adapted for various aspects of language production. These precursor circuits that are found in the nonlinguistic, nonhuman, primate brain also exist in the human brain. The possible reasons why these areas were adapted for language use in the human brain are discussed. The results throw new light on the prelinguistic precursor circuitry of Broca's region and help understand functional interactions between Broca's ventrolateral frontal region and posterior parietal and temporal association areas.     

Cortico-cortical projections from the prefrontal cortex to the superior temporal sulcal area (STs) in the monkey studied by means of HRP method.

Cortico-cortical connections from the prefrontal cortex to the superior temporal sulcal cortex (STs area) were studied in the monkey by means of retrograde axonal transport of horseradish peroxidase (HRP). After injections of 0.15-0.6 microliter of 50% HRP into the STs area, labeled cells were found in various cortical regions. In the prefrontal-STs projections, main features of topographic correlation were revealed; the posterior part of the STs area receives fibers from the superior frontal convexity (areas dorsal to the principal sulcus) and areas 8 and 6, whereas the anterior part of the STs area receives fibers from the inferior frontal convexity (areas ventral to the principal sulcus) and the frontal pole (area FD). The principal sulcus sends fibers to the entire STs area except for its ventral wall of the posterior part. A small cortical area adjacent to the inferior ramus of the arcuate sulcus (area 45 of ref. 41) sends fibers to the entire STs area. In addition, the orbitofrontal cortex projects mainly to the rostral part of the STs area, and the parahippocampal gyrus (areas TF and TH) projects to the deeper part of the entire STs area.     

A Comparison of the Ipsilateral Cortical Projections to the Dorsal and Ventral Subdivisions of the Macaque Premotor Cortex

The cortical connections of the dorsal (PMd) and ventral (PMv) subdivisions of the premotor area (PM, lateral area 6) were studied in four monkeys (Macaca fascicularis) through the use of retrograde tracers. In two animals, tracer was injected ventral to the arcuate sulcus (PMv), in a region from which forelimb movements could be elicited by intracortical microstimulation (ICMS). Tracer injections dorsal to the arcuate sulcus (PMd) were made in two locations. In one animal, tracer was injected caudal to the genu of the arcuate sulcus (in caudal PMd [cPMd], where ICMS was effective in eliciting forelimb movements); in another animal, it was injected rostral to the genu of the arcuate sulcus (in rostral PMd [rPMd], where ICMS was ineffective in eliciting movements). Retrogradely labeled neurons were counted in the ipsilateral hemisphere and located in cytoarchitectonically identified areas of the frontal and parietal lobes. Although both PMv and PMd were found to receive inputs from other motor areas, the prefrontal cortex, and the parietal cortex, there were differences in the topography and the relative strength of projections from these areas.

There were few common inputs to PMv and PMd; only the supplementary eye fields projected to all three areas studied. Interconnections within PMd or PMv appeared to link hindlimb and forelimb representations, and forelimb and face representations; however, connections between PMd and PMv were sparse. Areas cPMd and PMv were found to receive inputs from other motor areas—the primary motor area, the supplementary motor area, and the cingulate motor area—but the topography and strength of projections from these areas varied. Area rPMd was found to receive sparse inputs, if any, from these motor areas. The frontal eye field (area 8a) was found to project to PMv and rPMd, and area 46 was labeled substantially only from rPMd. Parietal projections to PMv were found to originate from a variety of somatosensory and visual areas, including the second somatosensory cortex and related areas in the parietal operculum of the lateral sulcus, as well as areas 5, 7a, and 7b, and the anterior intraparietal area. By contrast, projections to cPMd arose only from area 5. Visual areas 7m and the medial intraparietal area were labeled from rPMd. Relatively more parietal neurons were labeled after tracer injections in PMv than in PMd. Thus, PMv and PMd appear to be parts of separate, parallel networks for movement control.     

Effective Connectivity of Depth-Structure–Selective Patches in the Lateral Bank of the Macaque Intraparietal Sulcus

Extrastriate cortical areas are frequently composed of subpopulations of neurons encoding specific features or stimuli, such as color, disparity, or faces, and patches of neurons encoding similar stimulus properties are typically embedded in interconnected networks, such as the attention or face-processing network. The goal of the current study was to examine the effective connectivity of subsectors of neurons in the same cortical area with highly similar neuronal response properties. We first recorded single- and multi-unit activity to identify two neuronal patches in the anterior part of the macaque intraparietal sulcus (IPS) showing the same depth structure selectivity and then employed electrical microstimulation during functional magnetic resonance imaging in these patches to determine the effective connectivity of these patches. The two IPS subsectors we identified—with the same neuronal response properties and in some cases separated by only 3 mm—were effectively connected to remarkably distinct cortical networks in both dorsal and ventral stream in three macaques. Conversely, the differences in effective connectivity could account for the known visual-to-motor gradient within the anterior IPS. These results clarify the role of the anterior IPS as a pivotal brain region where dorsal and ventral visual stream interact during object analysis. Thus, in addition to the anatomical connectivity of cortical areas and the properties of individual neurons in these areas, the effective connectivity provides novel key insights into the widespread functional networks that support behavior.     

Maps of space in human frontoparietal cortex

Prefrontal cortex (PFC) and posterior parietal cortex (PPC) are neural substrates for spatial cognition. We here review studies in which we tested the hypothesis that human frontoparietal cortex may function as a priority map. According to priority map theory, objects or locations in the visual world are represented by neural activity that is proportional to their attentional priority. Using functional magnetic resonance imaging (fMRI), we first identified topographic maps in PFC and PPC as candidate priority maps of space. We then measured fMRI activity in candidate priority maps during the delay periods of a covert attention task, a spatial working memory task, and a motor planning task to test whether the activity depended on the particular spatial cognition. Our hypothesis was that some, but not all, candidate priority maps in PFC and PPC would be agnostic with regard to what was being prioritized, in that their activity would reflect the location in space across tasks rather than a particular kind of spatial cognition (e.g., covert attention). To test whether patterns of delay period activity were interchangeable during the spatial cognitive tasks, we used multivariate classifiers. We found that decoders trained to predict the locations on one task (e.g., working memory) cross-predicted the locations on the other tasks (e.g., covert attention and motor planning) in superior precentral sulcus (sPCS) and in a region of intraparietal sulcus (IPS2), suggesting that these patterns of maintenance activity may be interchangeable across the tasks. Such properties make sPCS in frontal cortex and IPS2 in parietal cortex viable priority map candidates, and suggest that these areas may be the human homologs of the monkey frontal eye field (FEF) and lateral intraparietal area (LIP).     

The organization and connections of somatosensory cortex in the brush-tailed possum (Trichosurus vulpecula): evidence for multiple, topographically organized and interconnected representations in an Australian marsupial

Microelectrode mapping techniques were used to determine the organization of somatosensory cortex in the Australian brush-tailed possum (Trichosurus vulpecula). The results of electrophysiological mapping were combined with data on the cyto- and myeloarchitecture, and patterns of corticocortical connections, using sections cut tangential to the pial surface. We found evidence for three topographically organized representations of the body surface that were coextensive with architectonic subdivisions. A large, discontinuous cutaneous representation in anterior parietal cortex was termed the primary somatosensory area (SI). Lateral to SI we found evidence for two further areas, the second somatosensory area (SII) and the parietal ventral area (PV). While neurones in all of these areas were responsive to cutaneous stimulation, those of SI were non-habituating, whereas those in SII and PV often habituated to the stimuli. Moreover, neuronal receptive fields in SII and PV were, in general, larger than those in SI. Neurones in cortex adjacent to the rostral and caudal boundaries of SI, including cortex that interdigitated between the discontinuous SI head and body representations, required stimulation of deep receptors in the periphery to elicit responses. Within the region of cortex containing neurones responsive to stimulation of deep receptors, body parts were represented in a mediolateral progression. Injections of anatomical tracers placed in electrophysiologically identified locations in SI revealed ipsilateral connections with other parts of SI, as well as cortex rostral to, caudal to, and interdigitating between, SI. Injections in SI also resulted in labelling in PV, SII, motor cortex, posterior parietal cortex and perirhinal cortex. The patterns of contralateral projections reflected those of ipsilateral projections, although they were relatively less dense. The present findings support recent observations in other marsupials in which multiple representations of the body surface were described, and suggest that multiple interconnected sensory representations may be a common feature of cortical organization and function in marsupials.     

Quantitative analysis of basal dendritic tree of layer III pyramidal neurons in different areas of adult human frontal cortex

Large long projecting (cortico-cortical) layer IIIc pyramidal neurons were recently disclosed to be in the basis of cognitive processing in primates. Therefore, we quantitatively examined the basal dendritic morphology of these neurons by using rapid Golgi and Golgi Cox impregnation methods among three distinct Brodmann areas (BA) of an adult human frontal cortex: the primary motor BA4 and the associative magnopyramidal BA9 from left hemisphere and the Broca's speech BA45 from both hemispheres. There was no statistically significant difference in basal dendritic length or complexity, as dendritic spine number or their density between analyzed BA's. In addition, we analyzed each of these BA's immunocytochemically for distribution of SMI-32, a marker of largest long distance projecting neurons. Within layer IIIc, the highest density of SMI-32 immunopositive pyramidal neurons was observed in associative BA9, while in primary BA4 they were sparse. Taken together, these data suggest that an increase in the complexity of cortico-cortical network within human frontal areas of different functional order may be principally based on the increase in density of large, SMI-32 immunopositive layer IIIc neurons, rather than by further increase in complexity of their dendritic tree and synaptic network.     

Spatio-temporal analysis of cortical activity evoked by gustatory stimulation in humans.

Gustatory activated regions in the cerebral cortex have not been identified precisely in humans. In this study we recorded the magnetic fields from the brain in response to two tastants, 1 M NaCl and 3 mM saccharin. We estimated the location of areas activated sequentially after the onset of stimulation with magnetic source imaging. We investigated the primary gustatory area (area G) precisely, and found it at the transition between the parietal operculum and the insular cortex. The central sulcus was activated less frequently than area G but with almost the same latency in cases of NaCl stimulation. Following area G, we found activation in several cortical regions, e.g. both the frontal operculum and the anterior part of the insula, the hippocampus, the parahippocampal gyrus and the superior temporal sulcus.     

Brain Activation of Identity Switching in Multiple Identity Tracking Task

When different objects switch identities in the multiple identity tracking (MIT) task, viewers need to rebind objects’ identity and location, which requires attention. This rebinding helps people identify the regions targets are in (where they need to focus their attention) and inhibit unimportant regions (where distractors are). This study investigated the processing of attentional tracking after identity switching in an adapted MIT task. This experiment used three identity-switching conditions: a target-switching condition (where the target objects switched identities), a distractor-switching condition (where the distractor objects switched identities), and a no-switching condition. Compared to the distractor-switching condition, the target-switching condition elicited greater activation in the frontal eye fields (FEF), intraparietal sulcus (IPS), and visual cortex. Compared to the no-switching condition, the target-switching condition elicited greater activation in the FEF, inferior frontal gyrus (pars orbitalis) (IFG-Orb), IPS, visual cortex, middle temporal lobule, and anterior cingulate cortex. Finally, the distractor-switching condition showed greater activation in the IFG-Orb compared to the no-switching condition. These results suggest that, in the target-switching condition, the FEF and IPS (the dorsal attention network) might be involved in goal-driven attention to targets during attentional tracking. In addition, in the distractor-switching condition, the activation of the IFG-Orb may indicate salient change that pulls attention away automatically.     

Projections from areas 5 and 7 to subdivisions of the cat sensorimotor cortex

Projections from the parietal cortex (areas 5 and 7) to subdivisions of the sensori-motor cortical region were investigated in cats using axonal degeneration techniques. Differences between the density of distribution of association fibers proceeding from these areas were found within the parietal and sensorimotor cortex. Area 5 projects mainly to the posterolateral portion of the cruciate sulcus (areas 4fu and 4) and to fields 4y, 4sfu, 6iffu, 6aa, and 6ab to a lesser extent. Area 7 is connected mainly to the medial portion of the lower lip of the cruciate sulcus (areas 6iffu, 6aa, and 6ab). Somewhat fewer fibers proceed to areas 4fu and 4. Fewer projections proceed from the parietal cortex to the somatosensory than to the motor region. Only a few single fibers connect the primary somatosensory region (fields 2, 3a, and 3b) with area 5, while area 7 does not project into this area. Neither field 5 nor 7 projects to the secondary somatosensory cortical area.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 20, No. 3, pp. 319–326, May–June, 1988.     

Somatotopic Organization of the Third Somatosensory Area (SIII) in Cats

Multiunit microelectrode recording techniques were used to study the location and organization of the third somatosensory area (SIII) in cats. Representations of all major contralateral body parts were found in a small region of cortex along the lateral wing of the ansate sulcus and between the lateral sulcus and the suprasylvian sulcus. The systematic map of the body surface included forepaw and face regions previously identified as parts of SIII. The forepaw representation was generally buried on the rostral bank of the lateral wing of the ansate sulcus. The representations of the face and mystacial vibrissae were largely exposed on the rostral suprasylvian gyrus, but part of the representation of the face was also buried in the lateral wing of the ansate sulcus. Representations of the trunk and hindlimb extended from the suprasylvian gyrus onto the medial bank of the suprasylvian sulcus. We had expected to find these latter body parts in more medial cortex just caudal to the representation of these parts in the first somatosensory area (SI). Instead, neurons in penetrations in cortex caudal to the SI trunk and hindlimb representations were unresponsive to tactile stimulation. The unexpected location of the hindlimb in SIII led us to determine whether the proposed parts of SIII had similar cortical and thalamic connections. Injected anatomical tracers revealed that the representations of both the forelimb and hindlimb were interconnected with SI and a region of the thalamus just dorsal to the ventroposterior nucleus. Similarities in patterns of connections of forelimb and hindlimb portions of SIII supported the conclusion that SHI as presented here is a functional unit of cortex. We conclude that SIII has a somatotopic organization that does not parallel that in SI, and that SIII is not entirely coextensive with either area 5 or area 5a of Hassler and Muhs-Clement (1964).     

The organization of somatosensory cortex in the short-tailed opossum ( Monodelphis domestica )

The organization of neocortex in the short-tailed opossum ( Monodelphis domestica ) was explored with multiunit microelectrode recordings from middle layers of cortex. Microelectrode maps were subsequently related to the chemoarchitecture of flattened cortical preparations, sectioned parallel to the cortical surface and processed for either cytochrome oxidase (CO) or NADPH-diaphorase (NADPHd) histochemistry. The recordings revealed the presence of at least two systematic representations of the contralateral body surface located in a continuous strip of cortex running from the rhinal sulcus to the medial wall. The primary somatosensory area (S1) was located medially while secondary somatosensory cortex (S2) formed a laterally located mirror image of S1. Auditory cortex was located in lateral cortex at the caudal border of S2, and some electrode penetrations in this area responded to both auditory and somatosensory stimulation. Auditory cortex was outlined by a dark oval visible in flattened brain sections. A large primary visual cortex (V1) was located at the caudal pole of cortex, and also consistently corresponded to a large chemoarchitecturally visible oval. Cortex just rostral and lateral to V1 responded to visual stimulation, while bimodal auditory/visual responses were obtained in an area between V1 and somatosensory cortex. The results are compared with brain organization in other marsupials and with placentals and the evolution of cortical areas in mammals is discussed.     

The organization of somatosensory cortex in the short-tailed opossum (Monodelphis domestica)

The organization of neocortex in the short-tailed opossum (Monodelphis domestica) was explored with multiunit microelectrode recordings from middle layers of cortex. Microelectrode maps were subsequently related to the chemoarchitecture of flattened cortical preparations, sectioned parallel to the cortical surface and processed for either cytochrome oxidase (CO) or NADPH-diaphorase (NADPHd) histochemistry. The recordings revealed the presence of at least two systematic representations of the contralateral body surface located in a continuous strip of cortex running from the rhinal sulcus to the medial wall. The primary somatosensory area (S1) was located medially while secondary somatosensory cortex (S2) formed a laterally located mirror image of S1. Auditory cortex was located in lateral cortex at the caudal border of S2, and some electrode penetrations in this area responded to both auditory and somatosensory stimulation. Auditory cortex was outlined by a dark oval visible in flattened brain sections. A large primary visual cortex (V1) was located at the caudal pole of cortex, and also consistently corresponded to a large chemoarchitecturally visible oval. Cortex just rostral and lateral to V1 responded to visual stimulation, while bimodal auditory/visual responses were obtained in an area between V1 and somatosensory cortex. The results are compared with brain organization in other marsupials and with placentals and the evolution of cortical areas in mammals is discussed.     

Decoding successive computational stages of saliency processing

An important requirement for vision is to identify interesting and relevant regions of the environment for further processing. Some models assume that salient locations from a visual scene are encoded in a dedicated spatial saliency map [1, 2]. Then, a winner-take-all (WTA) mechanism [1, 2] is often believed to threshold the graded saliency representation and identify the most salient position in the visual field. Here we aimed to assess whether neural representations of graded saliency and the subsequent WTA mechanism can be dissociated. We presented images of natural scenes while subjects were in a scanner performing a demanding fixation task, and thus their attention was directed away. Signals in early visual cortex and posterior intraparietal sulcus (IPS) correlated with graded saliency as defined by a computational saliency model. Multivariate pattern classification [3, 4] revealed that the most salient position in the visual field was encoded in anterior IPS and frontal eye fields (FEF), thus reflecting a potential WTA stage. Our results thus confirm that graded saliency and WTA-thresholded saliency are encoded in distinct neural structures. This could provide the neural representation required for rapid and automatic orientation toward salient events in natural environments.     

Autoradiographic evidence of visual cortical projections to the frontal cortex in the cat

In II adult cats, areas 17, 18, 19 as well as the lateral suprasylvian area were separately injected with L-[5-3H] proline and their efferent projections to the frontal cortex were autoradiographically searched. Only area 19 and lateral suprasylvian area showed such projections; terminal sites were localized in the ventral and dorsal banks of the cruciate sulcus and in the adjacent mesial surface of the brain. The possibility that these labeled regions may correspond to the monkey's frontal eye field is discussed.