Visual topography of V1 in the Cebus monkey

The representation of the visual field in the striate cortex (V1) was mapped with multiunit electrodes in the Cebus monkey. Nine Cebus apella, anesthetized with N2O and immobilized with pancuromium bromide were studied in repeated recording sessions. In each hemisphere, V1 contains a continuous representation of the contralateral visual hemifield. The representation of the vertical meridian (VM) forms the external border of V1 except at the anteriormost portion of the calcarine fissure. The representation of the horizontal meridian (HM) divides the area so that the representation of the lower visual field is located dorsally, and that of the upper field ventrally. The convoluted surface of V1 can be only partially unfolded, and no precise "flattened" map can be obtained without introducing surface discontinuities. The visual topography of V1 is presented in a series of coronal sections and in "flattened" maps. The representation of the central visual field is magnified relative to that of the periphery in V1. The evaluation of the cortical magnification factors measured along isoeccentric and isopolar dimensions in the partially unfolded model of V1 revealed anisotropies in the representation of the visual field with larger magnification along isopolar lines than along isoeccentric lines. Receptive field size increases with increasing eccentricity, whereas point image size decreases with increasing eccentricity.     

Area map of mouse visual cortex

It is controversial whether mouse extrastriate cortex has a "simple" organization in which lateral primary visual cortex (V1) is adjoined by a single area V2 or has a "complex" organization, in which lateral V1 is adjoined by multiple distinct areas, all of which share the vertical meridian with V1. Resolving this issue is important for understanding the evolution and development of cortical arealization. We have used triple pathway tracing combined with receptive field recordings to map azimuth and elevation in the same brain and have referenced these maps against callosal landmarks. We found that V1 projects to 15 cortical fields. At least nine of these contain maps with complete and orderly representations of the entire visual hemifield and therefore represent distinct areas. One of these, PM, adjoins V1 at the medial border. Five areas, P, LM, AL, RL, and A, adjoin V1 on the lateral border, but only LM shares the vertical meridian representation with V1. This suggests that LM is homologous to V2 and that the lateral extrastriate areas do not represent modules within a single area V2. Thus, mouse visual cortex is "simple" in the sense that lateral V1 is adjoined by a single V2-like area, LM, and "complex" in having a string of areas in lateral extrastriate cortex, which receive direct V1 input. The results suggest that large numbers of areas with topologically equivalent maps of the visual field emerge early in evolution and that homologous areas are inherited in different mammalian lineages.     

Identification and viuotopic organization of areas PO and POd in Cebus monkey

Two visual areas of the anterior bank of the parietooccipital sulcus, areas PO and POd, were identified and their visual field representations were studied in six anesthetized and paralyzed Cebus monkeys. The definition of these areas was based on electrophysiological mapping and myeloarchitecture. PO is located in the ventral aspect of the anterior bank of the parietooccipital sulcus and ventral precuneate gyrus. It borders area V2 posteriorly and ventrally in the depth of the parietooccipital sulcus, area V3d laterally, and another undescribed visual area medially. POd was located dorsal to area PO and ventral to architectonic area PE. The representations of the visual field in areas PO and POd are complex. In each hemisphere, these areas have a virtually complete representation of the contralateral visual hemifield. Different from the previously described visual areas, in PO and POd there is a precise organization of isopolar lines and a complex organization of the isoeccentric ones. In PO, as well as in POd, the representation of the horizontal meridian runs dorsoventrally along the parietooccipital sulcus. The upper visual quadrant is represented medially and the lower visual quadrant laterally. A large and complex representation of the periphery, from 20° to 60° eccentricity is present at the lateral and medial portions of these areas. By contrast, the representation of the central 20° is very small in both PO and POd. The central visual field is represented ventrally in PO and dorsally in area POd. Area POd shows a more stratified myeloarchitectonic pattern than PO and both areas can be distinguished from other surrounding areas by their heavier myelinated pattern. © Wiley-Liss, Inc.     

The locus of the posterior subdivision of the inferotemporal visual learning area in the monkey

In an attempt to define the posterior subdivision of the inferotemporal visual learning area more precisely than before, pattern discrimination retention, serial object discrimination learning and concurrent object discrimination learning were tested in 16 monkeys with lesions in one of four different cytoarchitectural areas; TEO, OA, OB and OC, and in five unoperated monkeys. Marked impairment was found only in pattern discrimination retention and only in the monkeys with lesions of area TEO. It was concluded that the posterior limit of the inferotemporal visual learning area is at the ascending limb of the inferior occipital sulcus, and that the posterior subdivision thus comprises the single anatomical area TEO and does not extend into areas OA and OB.     

Somatosensory cortex in macaque monkeys: laminar differences in receptive field size in areas 3b and 1

We have examined receptive field sizes of neurons in granular, supragranular and infragranular layers within somatosensory cortical areas 3b and 1 in macaque monkeys. Receptive fields of neurons in layer 4 are smaller than receptive fields of neurons above or below layer 4. In addition, neurons in area 1 have larger receptive fields than neurons in corresponding layers of area 3b.     

Anesthetic state does not affect the map of the hand representation within area 3b somatosensory cortex in owl monkey

Receptive fields defined for small clusters of neurons within the middle cortical layers of area 3b differed little in size or in the skin locations represented among: alert, nitrous oxide-anesthetized, deep sodium pentobarbital-anesthetized, and ketamine-anesthetized owl monkeys. Repeated recordings from nearly identical cortical loci yielded nearly identical multi-unit receptive fields under different conditions of anesthesia.     

Cytoarchitecture and visual field representation in area 17 of the tammar wallaby (Macropus eugenii).

Tritiated proline was injected into one eye in the tammar wallaby and transported label was studied in the cortex after transneuronal passage through the lateral geniculate nucleus. The autoradiographic label and cytoarchitecture were used to anatomically demarcate the borders of area 17. Electrophysiological recordings from single units were done to obtain a retinotopic map of area 17. Single units in area 17 were found to have orientation sensitivity comparable to those seen in placental mammals such as cat and monkey. They could also be classified as simple, complex, and hypercomplex cells. Changes in the cortical areal magnification factor with eccentricity were found to match the drop off in retinal ganglion cell density only along the vertical meridian representation. Along the horizontal meridian, the cortical magnification falls off significantly with eccentricity, whereas the ganglion cell density shows only a mild reduction. Thus central vision, especially the binocular segment, is heavily represented at the cost of the periphery.     

Three streams of visual information processing in V2 of Cebus monkey

Gattass and collaborators (Gattass R, Rosa MGP, Souza APB, Piñon MCG, Neuenschwander S [1990a] Braz J Med Biol Res 23:375–393) proposed that the dorsal stream of visual processing, as defined by Ungerleider and Mishkin (Ungerleider LG, Mishkin M [1982] In: Ingle DJ, Goodale MA, Mansfield RJW, editors. Analysis of visual behavior. Cambridge: Massachusetts Institute of Technology. p 549–586), can be subdivided into dorsolateral and dorsomedial streams, and suggested that they may be involved in different aspects of the processing of motion and spatial perception, respectively. The goal of the present study was to provide additional evidence for this hypothesis by using cytochrome oxidase immunohistochemistry combined with retrograde tracing techniques. In Old World monkeys, the locations of visual area 4 (V4; ventral stream) and middle temporal area (MT; dorsal stream) projecting neurons in V2 supports the hypothesis that the cytochrome oxidase (CytOx)–rich thin stripes and the CytOx‐poor interstripes are associated with the ventral stream, and that the CytOx‐rich thick stripes belong to the dorsal stream. In this study we describe, in the New World monkey Cebus, the distribution of retrogradely labeled cells in V2 relative to the CytOx compartments after fluorescent tracers were placed in areas V4, MT, and the parietooccipital area (PO). We found PO‐projecting neurons in CytOx‐rich thick stripes and CytOx‐poor interstripes in V2, whereas MT‐projecting neurons appeared almost exclusively in thick stripes. In contrast, V4‐projecting neurons were located mostly in CytOx‐poor interstripes and CytOx‐rich thin stripes. In addition, V4‐ and MT‐projecting neurons were located mainly in supragranular layers, whereas PO‐projecting neurons were located in supragranular and infragranular layers. These results support the hypothesis for the existence of three distinct streams of visual processing: ventral (including V4), dorsolateral (including MT), and dorsomedial (including PO). J. Comp. Neurol. 466:104–118, 2003. © 2003 Wiley‐Liss, Inc.     

Retinotopic mapping of the peripheral visual field to human visual cortex by functional magnetic resonance imaging

Retinotopic mapping is a key property of organization in the human occipital cortex. The retinotopic organization of the central visual field of visual areas V1, V2, and V3 has been well established. We used fMRI to measure the retinotopic map of the peripheral visual field (eccentricity up to 60°). We estimated the sizes of the visual areas between 0° and 60° and obtained results consistent with anatomical studies. We also estimated the cortical distances and magnification factors for reconstruction of the retinotopic map using the peripheral wedge dipole model. By comparing the retinotopic map with the flattened surface, we analyzed the datasets used to reconstruct the map. We found that: (1) the percentage of the striate cortex devoted to peripheral vision in humans is significantly larger than that in the macaque, (2) the estimate of the scaling factor in linear magnification is larger than that found in previous studies focusing on central vision, and (3) the estimate of the peripheral factor in the dipolar model is too large to make the curve direction of the dipolar map in the periphery equivalent to that in the center. On the basis of our results, we revised the dipolar map to fit our conditions. The revised map in humans has a similar elliptical shape to that of macaques, and the central parts of the two species are the same. The different parts of the map are the peripheral regions, for which the peripheral wedge dipole model in humans is reversed compared to that of macaques.     

Brain location and visual topography of cortical area V6A in the macaque monkey

The brain location, extent and functional organization of the cortical visual area V6A was investigated in macaque monkeys by using single cell recording techniques in awake, behaving animals. Six hemispheres of four animals were studied. Area V6A occupies a horseshoe-like region of cortex in the caudalmost part of the superior parietal lobule. It extends from the medial surface of the brain, through the anterior bank of the parieto-occipital sulcus, up to the most lateral part of the fundus of the same sulcus. Area V6A borders on areas V6 ventrally, PEc dorsally, PGm medially and MIP laterally. Of 1348 neurons recorded in V6A, 61% were visual and 39% non-visual in nature. The visual neurons were particularly sensitive to orientation and direction of movement of visual stimuli. The inferior contralateral quadrant was the most represented one. Visual receptive fields were also found in the inferior ipsilateral quadrant and in the upper visual field. Receptive fields were on average smaller in the lower visual field than in the upper one. Both central and peripheral parts of the visual field were represented. Large parts of the visual field were represented in small regions of area V6A, and the same regions of the visual field were re-represented many times in different parts of this area, without any apparent topographical order. The only reliable sign of retinotopic organization was the predominance of central representation dorsally and far periphery ventrally. The functional organization of area V6A is discussed in the view that this area could be involved in the control of reaching out and grasping objects.     

Monoclonal antibody to neurofilament protein (SMI-32) labels a subpopulation of pyramidal neurons in the human and monkey neocortex

A monoclonal antibody that recognizes a nonphosphorylated epitope on the 168 kDa and 200 kDa subunits of neurofilament proteins has been used in an immunohistochemical study of cynomolgus monkey (Macaca fascicularis) and human neocortex. This antibody, SMI-32, primarily labels the cell body and dendrites of a subset of pyramidal neurons in both species. A greater proportion of neocortical pyramidal neurons were SMI-32 immunoreactive (ir) in the human than in the monkey. SMI-32-ir neurons exhibited consistent differences in the intensity of their immunoreactivity that correlated with cell size. The cellular specificity of SMI-32 immunoreactivity suggests that a subpopulation of neurons can be distinguished on the basis of differences in the molecular characteristics of basic cytoskeletal elements such as neurofilament proteins. The size, density, and laminar distribution of SMI-32-ir neurons differed substantially across neocortical areas within each species and between species. Differences across cortical areas were particularly striking in the monkey. For example, the anterior parainsular cortex had a substantial population of large SMI-32-ir neurons in layer V and a near absence of any immunoreactive neurons in the supragranular layers. This contrasted with the cortical area located more laterally on the superior temporal gyrus, where layers III and V contained substantial populations of large SMI-32-ir neurons. Both areas differed significantly from the posterior inferior temporal gyrus, which was distinguished by a bimodal distribution of large SMI-32-ir neurons in layer III. Differences across human areas were less obvious because of the increase in the number of SMI-32-ir neurons. Perhaps the most notable differences across human areas resulted from shifts in the density of the larger SMI-32-ir neurons in deep layer III. A comparison between the species revealed that isocortical areas exhibited greater differences in their representation of SMI-32-ir neurons than primary sensory or transitional cortical areas. A comparison of distribution patterns of SMI-32-ir neurons across monkey cortical areas and data available on the laminar organization of cortical efferent neurons suggests that a common anatomic characteristic of this chemically identified subpopulation of neurons is that they have a distant axonal projection. Such correlations of cell biological characteristics with specific elements of cortical circuitry will further our understanding of the molecular and cellular properties that are critically linked to a given neuron's role in cortical structure and function.     

Activity of neurons in monkey posterior temporal cortex during multidimensional visual discrimination tasks

Unit activity was recorded from the posterior temporal cortex (PTE) of awake, behaving rhesus monkeys while they performed a series of visual discrimination tasks involving colored checkerboard patterns. The activity of 130 (91%) of 143 PTE units was altered by the presentation of a visual discriminandum; 112 of these cells (86%) exhibited a significant increase in firing after presentation of the stimulus while the remainder gave an inhibitory response. Over half (64%) the PTE units exhibited differential activity between discriminanda, i.e. they were selective for color and/or form. Six of 10 neurons, recorded when the monkey was required to shift attention from one stimulus feature to another, exhibited a difference in poststimulus neural activity even though the discriminandum remained the same. Three neurons were recorded from when the stimuli were altered by changing the check size although the relevant (i.e. rewarded) dimension (color) was left the same; two showed an invariant response to the altered stimuli and one gave the same response to one of the altered stimuli but a different response to the other. These data support the role of posterior temporal cortex in visual discrimination learning and visual attention.     

Anatomical and physiological definition of the motor cortex of the marmoset monkey

We used a combination of anatomical and physiological techniques to define the primary motor cortex (M1) of the marmoset monkey and its relationship to adjacent cortical fields. Area M1, defined as a region containing a representation of the entire body and showing the highest excitability to intracortical microstimulation, is architecturally heterogeneous: it encompasses both the caudal part of the densely myelinated "gigantopyramidal" cortex (field 4) and a lateral region, corresponding to the face representation, which is less myelinated and has smaller layer 5 pyramidal cells (field 4c). Rostral to M1 is a field that is strongly reminiscent of field 4 in terms of cyto- and myeloarchitecture but that in the marmoset is poorly responsive to microstimulation. Anatomical tracing experiments revealed that this rostral field is interconnected with visual areas of the posterior parietal cortex, whereas M1 itself has no such connections. For these reasons, we considered this field to be best described as part of the dorsal premotor cortex and adopted the designation 6Dc. Histological criteria were used to define other fields adjacent to M1, including medial and ventral subdivisions of the premotor cortex (fields 6M and 6V) and the rostral somatosensory field (area 3a), as well as a rostral subdivision of the dorsal premotor area (field 6Dr). These results suggest a basic plan underlying the histological organization of the caudal frontal cortex in different simian species, which has been elaborated during the evolution of larger species of primate by creation of further morphological and functional subdivisions.     

The second visual area in the marmoset monkey: Visuotopic organisation,magnification factors,architectonical boundaries,and modularity

The organisation of the second visual area (V2) in marmoset monkeys was studied by means of extracellular recordings of responses to visual stimulation and examination of myelin- and cytochrome oxidase-stained sections. Area V2 forms a continuous cortical belt of variable width (1–2 mm adjacent to the foveal representation of V1, and 3–3.5 mm near the midline and on the tentorial surface) bordering V1 on the lateral, dorsal, medial, and tentorial surfaces of the occipital lobe. The total surface area of V2 is approximately 100 mm², or about 50% of the surface area of V1 in the same individuals. In each hemisphere, the receptive fields of V2 neurones cover the entire contralateral visual hemifield, forming an ordered visuotopic representation. As in other simians, the dorsal and ventral halves of V2 represent the lower and upper contralateral quadrants, respectively, with little invasion of the ipsilateral hemifield. The representation of the vertical meridian forms the caudal border of V2, with V1, whereas a field discontinuity approximately coincident with the horizontal meridian forms the rostral border of V2, with other visually responsive areas. The bridge of cortex connecting dorsal and ventral V2 contains neurones with receptive fields centred within 1° of the centre of the fovea. The visuotopy, size, shape and location of V2 show little variation among individuals. Analysis of cortical magnification factor (CMF) revealed that the V2 map of the visual field is highly anisotropic: for any given eccentricity, the CMF is approximately twice as large in the dimension parallel to the V1/V2 border as it is perpendicular to this border. Moreover, comparison of V2 and V1 in the same individuals demonstrated that the representation of the central visual field is emphasised in V2, relative to V1. Approximately half of the surface area of V2 is dedicated to the representation of the central 5° of the visual field. Calculations based on the CMF, receptive field scatter, and receptive field size revealed that the point-image size measured parallel to the V1/V2 border (2–3 mm) equals the width of a full cycle of cytochrome oxidase stripes in V2, suggesting a close correspondence between physiological and anatomical estimates of the dimensions of modular components in this area. J. Comp. Neurol. 387:547–567, 1997. © 1997 Wiley-Liss, Inc.     

Representation of the visual field in the primary visual area of the marmoset monkey: Magnification factors,point‐image size,and proportionality to retinal ganglion cell density

The primary visual area (V1) forms a systematic map of the visual field, in which adjacent cell clusters represent adjacent points of visual space. A precise quantification of this map is key to understanding the anatomical relationships between neurons located in different stations of the visual pathway, as well as the neural bases of visual performance in different regions of the visual field. We used computational methods to quantify the visual topography of V1 in the marmoset (Callithrix jacchus), a small diurnal monkey. The receptive fields of neurons throughout V1 were mapped in two anesthetized animals using electrophysiological recordings. Following histological reconstruction, precise 3D reconstructions of the V1 surface and recording sites were generated. We found that the areal magnification factor (M_A) decreases with eccentricity following a function that has the same slope as that observed in larger diurnal primates, including macaque, squirrel, and capuchin monkeys, and humans. However, there was no systematic relationship between M_A and polar angle. Despite individual variation in the shape of V1, the relationship between M_A and eccentricity was preserved across cases. Comparison between V1 and the retinal ganglion cell density demonstrated preferential magnification of central space in the cortex. The size of the cortical compartment activated by a punctiform stimulus decreased from the foveal representation towards the peripheral representation. Nonetheless, the relationship between the receptive field sizes of V1 cells and the density of ganglion cells suggested that each V1 cell receives information from a similar number of retinal neurons, throughout the visual field. J. Comp. Neurol. 521:1001–1019, 2013. © 2012 Wiley Periodicals, Inc.     

Cortical connections of visual area MT in the macaque

We have identified the cortical connections of area MT and determined their topographic organization and relationship to myeloarchitectural fields. Efferents of MT were examined in seven macaques that had received injections of tritiated amino acids, and afferents were examined in one macaque that had received injections of two fluorescent dyes. The injection sites formed an orderly sequence from the representation of central to that of peripheral vision in the upper and lower visual fields. In addition to connections with the striate cortex (V1), connections were found between MT and a variety of extrastriate areas, including V2, V3, V3A, V4, V4t, VIP, MST, FST, possibly PO, and, finally, the frontal eye field. The connections of MT with V1, V2, and the dorsal and ventral portions of V3 were topographically organized and consistent with the visuotopic arrangement reported previously in these areas. V2 could be distinguished from V3 by the distinctive myeloarchitectural appearance of the former. Connections with areas V4 and V4t also displayed at least a coarse visuotopic organization, in that the central representation of MT projected laterally in these areas and the peripheral representation projected medially. The lower visual field representation of V4 was located dorsally, on the prelunate convexity, while the upper field representation was located primarily on the ventral aspect of the hemisphere. V4t had a distinctively light myeloarchitecture and received projections from only the lower field representation of MT. The remaining connections of MT were with areas located entirely in the dorsal half of the hemisphere. There were widespread connections with areas MST and FST in the superior temporal sulcus, with some evidence for a crude visuotopic organization in MST. Connections were also found with area VIP in the intraparietal sulcus, with area V3A on the annectent gyrus, possibly with area PO in the dorsomedial prestriate cortex, and, finally, with the frontal eye field on the anterior bank of the lower limb of the arcuate sulcus. Area FST and parts of both MST and VIP had a distinctive myeloarchitecture. The pattern of laminar connections with V1, V2, and V3 indicated that MT projects "back" to these areas and they project "forward" to MT. That is, the projections to these areas from MT terminated in both the supragranular and infragranular layers and the projections to MT from these areas originated predominantly from cells located above granular layer IV (above layer IVC in V1).(ABSTRACT TRUNCATED AT 400 WORDS)     

The termination of geniculocortical fibres in area 17 of the visual cortex in the macaque monkey

The termination of geniculocortical fibres within the different subdivisions of lamina IV in area 17 of the visual cortex of the monkey has been studied quantitatively with the electron microscope. In lamina IVCα the axon terminals of fibres coming from the magnocellular layers of the lateral geniculate nucleus (LGN) make significantly more synapses per bouton than those of fibres arising from the parvocellular layers and terminating in laminae IVA and IVCβ. In all parts of area 17 examined there was a clear difference in the relative proportions of multisynaptic geniculo-cortical boutons between the α and β divisions of lamina IVC. Calculations have shown that a single cell in the magnocellular laminae of the LGN may make about 6 times as many synaptic contacts within lamina IV of the visual cortex than one in the parvocellular laminae. It has also been estimated that there are at least 500 million geniculocortical boutons, or 1200 million synapses, in lamina IVCα and 1000 million boutons, 1200 million synapses, in lamina IVCβ for one hemisphere, giving an approximate total number of 1500 million boutons and 2400 million synapses.     

Organization of visual cortex in the northern quoll, Dasyurus hallucatus: evidence for a homologue of the second visual area in marsupials

Two visual areas, V1 and V2 (first and second visual areas), appear to be present in the posterior neocortex of all eutherian mammals investigated so far. However, previous studies have not established whether an area homologous to V2 also exists in metatherian mammals (marsupials). Using electrophysiological techniques, we mapped the visual receptive fields of neurons in the striate and peristriate cortices of the northern quoll, an Australian marsupial. We found that neurons in a 2-mm-wide strip of cortex rostrolateral to V1 form a single, relatively simple representation of the complete contralateral hemifield. This area resembles V2 of eutherians in several respects: (i) neurons in the medial half of the peristriate area represent the lower visual quadrant, whereas those in the lateral half represent the upper visual quadrant; (ii) the vertical meridian of the visual field is represented adjacent to V1, while the visual field periphery is represented along the lateral and rostrolateral borders of the peristriate area; (iii) there is a marked anisotropy in the representation, with a larger magnification factor parallel to the V1 border than perpendicular to this border; and (iv) receptive fields of multiunit clusters in the peristriate cortex are much larger than those of cells in V1 at comparable eccentricities. The cortex immediately rostral and lateral to V2 did not respond to visual stimulation under our recording conditions. These results suggest that V1 and V2 together form a 'core' of homologous visual areas, likely to exist in all therian mammals.     

Distribution of NADPH-diaphorase-positive neurons in the prefrontal cortex of the Cebus monkey

We studied the distribution of NADPH-diaphorase (NADPH-d) activity in the prefrontal cortex of normal adult Cebus apella monkeys using NADPH-d histochemical protocols. The following regions were studied: granular areas 46 and 12, dysgranular areas 9 and 13, and agranular areas 32 and Oap. NADPH-d-positive neurons were divided into two distinct types, both non-pyramidal. Type I neurons had a large soma diameter (17.24 +/- 1.73 microm) and were densely stained. More than 90% of these neurons were located in the subcortical white matter and infragranular layers. The remaining type I neurons were distributed in the supragranular layers. Type II neurons had a small, round or oval soma (9.83 +/- 1.03 microm), and their staining pattern varied markedly. Type II neurons were distributed throughout the cortex, with their greatest numerical density being observed in layers II and III. In granular areas, the number of type II neurons was up to 20 times that of type I neurons, but this proportion was smaller in agranular areas. Areal density of type II neurons was maximum in the supragranular layers of granular areas and minimum in agranular areas. Statistical analysis revealed that these areal differences were significant when comparing some specific areas. In conclusion, our results indicate a predominance of NADPH-d-positive cells in supragranular layers of granular areas in the Cebus prefrontal cortex. These findings support previous observations on the role of type II neurons as a new cortical nitric oxide source in supragranular cortical layers in primates, and their potential contribution to cortical neuronal activation in advanced mammals.