Anatomical evidence for classical and extra-classical receptive field completion across the discontinuous horizontal meridian representation of primate area V2

In primates, a split of the horizontal meridian (HM) representation at the V2 rostral border divides this area into dorsal (V2d) and ventral (V2v) halves (representing lower and upper visual quadrants, respectively), causing retinotopically neighboring loci across the HM to be distant within V2. How is perceptual continuity maintained across this discontinuous HM representation? Injections of neuroanatomical tracers in marmoset V2d demonstrated that cells near the V2d rostral border can maintain retinotopic continuity within their classical and extra-classical receptive field (RF), by making both local and long-range intra- and interareal connections with ventral cortex representing the upper visual quadrant. V2d neurons located <0.9-1.3 mm from the V2d rostral border, whose RFs presumably do not cross the HM, make nonretinotopic horizontal connections with V2v neurons in the supra- and infragranular layers. V2d neurons located <0.6-0.9 mm from the border, whose RFs presumably cross the HM, in addition make retinotopic local connections with V2v neurons in layer 4. V2d neurons also make interareal connections with upper visual field regions of extrastriate cortex, but not of MT or MTc outside the foveal representation. Labeled connections in ventral cortex appear to represent the "missing" portion of the connectional fields in V2d across the HM. We conclude that connections between dorsal and ventral cortex can create visual field continuity within a second-order discontinuous visual topography.     

Localization of area prostriata and its projection to the cingulate motor cortex in the rhesus monkey

Area prostriata is a poorly understood cortical area located in the anterior portion of the calcarine sulcus. It has attracted interest as a separate visual area and progenitor for the cortex of this modality. In this report we describe a direct projection from area prostriata to the rostral cingulate motor cortex (M3) that forms the fundus and lower bank of the anterior part of the cingulate sulcus. Injections of retrograde tracers in M3 resulted in labeled neurons in layers III, V and VI of prostriate cortex. However, injections of anterograde tracers in M3 did not demonstrate axon terminals in area prostriata. This connection was organized topographically such that the rostral part of M3 received input from the dorsal region of prostriate cortex, whereas middle and caudal levels of M3 received input from more ventral locations. Injections of retrograde and anterograde tracers in the caudal cingulate motor cortex (M4) did not produce labeling in prostriate cortex. Cytoarchitectural analysis confirmed the identity of area prostriata and further clarified its extent and borders with the parasubiculum of the hippocampal formation rostrally, and V1 of the visual cortex caudally. This linkage between cortex bordering V1 and cortex giving rise to a component of the corticofacial and corticospinal pathways demonstrates a more direct visuomotor route than visual association projections coursing laterally.     

Cortical connections of area V4 in the macaque

To determine the locus, full extent, and topographic organization of cortical connections of area V4 (visual area 4), we injected anterograde and retrograde tracers under electrophysiological guidance into 21 sites in 9 macaques. Injection sites included representations ranging from central to far peripheral eccentricities in the upper and lower fields. Our results indicated that all parts of V4 are connected with occipital areas V2 (visual area 2), V3 (visual area 3), and V3A (visual complex V3, part A), superior temporal areas V4t (V4 transition zone), MT (medial temporal area), and FST (fundus of the superior temporal sulcus [STS] area), inferior temporal areas TEO (cytoarchitectonic area TEO in posterior inferior temporal cortex) and TE (cytoarchitectonic area TE in anterior temporal cortex), and the frontal eye field (FEF). By contrast, mainly peripheral field representations of V4 are connected with occipitoparietal areas DP (dorsal prelunate area), VIP (ventral intraparietal area), LIP (lateral intraparietal area), PIP (posterior intraparietal area), parieto-occipital area, and MST (medial STS area), and parahippocampal area TF (cytoarchitectonic area TF on the parahippocampal gyrus). Based on the distribution of labeled cells and terminals, projections from V4 to V2 and V3 are feedback, those to V3A, V4t, MT, DP, VIP, PIP, and FEF are the intermediate type, and those to FST, MST, LIP, TEO, TE, and TF are feedforward. Peripheral field projections from V4 to parietal areas could provide a direct route for rapid activation of circuits serving spatial vision and spatial attention. By contrast, the predominance of central field projections from V4 to inferior temporal areas is consistent with the need for detailed form analysis for object vision.     

Differential architecture of multisynaptic geniculo-cortical pathways to V4 and MT

Parallel visual pathways in the primate brain known as the dorsal and ventral streams receive retinal inputs mainly through the magnocellular (M) and parvocellular (P) layers of the lateral geniculate nucleus. Inputs from these layers terminate within distinct parts of layer 4C of V1 (visual area 1). Due to the complexity of M- and P-derived neural connectivity in V1 and higher visual areas, the contributions of M and P inputs to the dorsal and ventral streams remain unclear. Employing retrograde transsynaptic transport of rabies virus, we analyzed the architecture of bottom-up pathways toward ventral stream area V4 (visual area 4) and dorsal stream area MT (middle temporal area). We found that V4 receives both M and P inputs "trisynaptically" from layer 4C via layer 2/3 of V1, whereas MT receives M-dominant input "disynaptically" from layer 4C via layer 4B of V1. V4 also receives disynaptic input from the dorsal stream portion of V2 (visual area 2) (i.e., cytochrome oxidase-stained thick stripes). Moreover, both M and P inputs reach V4 trisynaptically and MT disynaptically through "short-cut" pathways that bypass layer 4C of V1. The differential patterns of multisynaptic geniculo-cortical pathways to V4 and MT imply distinct modes of information processing in the dorsal and ventral streams.     

Long-range clustered connections within extrastriate visual area V5/MT of the rhesus macaque

Visual area V5/MT in the rhesus macaque has a distinct functional organization, where neurons with specific preferences for direction of motion and binocular disparity are co-organized in columns or clusters. Here, we analyze the pattern of intrinsic connectivity within cortical area V5/MT in both parasagittal sections of the intact brain and tangential sections from flatmounted cortex using small injections of the retrograde tracer cholera toxin subunit b. Labeled cells were predominantly found in cortical layers 2, 3, and 6. Going along the cortical layers, labeled cells were concentrated in regularly spaced clusters. The clusters nearest to the injection site were approximately 2 mm from its center. In flatmounted cortex, along the dorsoventral axis of V5/MT, we identified further clusters of labeled cells up to 10 mm from the injection site. Quantitative analysis of parasagittal sections estimated average cluster spacing at 2.2 mm; in cortical flatmounts, spacing was 2.3 mm measured radially from the injection site. The results suggest a regular pattern of intrinsic connectivity within V5/MT, which is consistent with connectivity between sites with a common preference for both direction of motion and binocular depth. The long-range connections can potentially account for the large suppressive surrounds of V5/MT neurons.     

Quantitative analysis of the corticocortical projections to the middle temporal area in the marmoset monkey: evolutionary and functional implications

The connections of the middle temporal area (MT) were investigated in the marmoset, one of the smallest primates. Reflecting the predictions of studies that modeled cortical allometric growth and development, we found that in adult marmosets MT is connected to a more extensive network of cortical areas than in larger primates, including consistent connections with retrosplenial, cingulate, and parahippocampal areas and more widespread connections with temporal, frontal, and parietal areas. Quantitative analyses reveal that MT receives the majority of its afferents from other motion-sensitive areas in the temporal lobe and from the occipitoparietal transition areas, each of these regions containing approximately 30% of the projecting cells. Projections from the primary visual area (V1) and the second visual area (V2) account for approximately 20% of projecting neurons, whereas "ventral stream" and higher-order association areas form quantitatively minor projections. A relationship exists between the percentage of supragranular layer neurons forming the projections from different areas and their putative hierarchical rank. However, this relationship is clearer for projections from ventral stream areas than it is for projections from dorsal stream or frontal areas. These results provide the first quantitative data on the connections of MT and extend current understanding of the relationship between cortical anatomy and function in evolution.     

Unilateral deficits in visual perception and learning after unilateral inferotemporal cortex lesions in macaques

This study adapted the method of partial lesions, combined with controlled fixation, to study the perceptual role of macaque inferotemporal (IT) cortex. Unilateral lesions were made in IT cortex of three monkeys, without section of the corpus callosum, and visual function was tested ipsilateral and contralateral to the lesion. The observed changes were compared to the effects of bilateral lesions of IT cortex in one monkey, the approach used in most previous studies. Unilateral lesions produced far less profound, although more selective, loss on the tested visual abilities than did bilateral lesions. All three monkeys with unilateral lesions showed decreased chromatic sensitivity, but sparing of achromatic sensitivity, and severely disrupted learning and performance of visual matching to sample, and in all cases, the visual loss was contralateral to the site of the lesion. Unexpectedly, the magnitude of the contralateral loss was not increased by later section of the corpus callosum and anterior commissure in one of the monkeys, a lesion that removes interhemispheric input to contralateral from ipsilateral temporal cortex neurons. These results support physiological findings that show that the response of IT cortex neurons is dominated by the contralateral visual field, despite the bilateral activation many IT neurons receive. Comparison to earlier studies of lesions of area V4, which provides input to IT cortex, shows that V4 and IT lesions produce qualitatively different effects.     

Functional architecture of retinotopy in visual association cortex of behaving monkey

While the receptive field properties of single neurons in the inferior parietal cortex have been quantitatively described from numerous electrical measurements, the visual topography of area 7a and the adjacent dorsal prelunate area (DP) remains unknown. This lacuna may be a technical byproduct of the difficulty of reconstructing tens to hundreds of penetrations, or may be the result of varying functional retinotopic architectures. Intrinsic optical imaging, performed in behaving monkey for extended periods of time, was used to evaluate retinotopy simultaneously at multiple positions across the cortical surface. As electrical recordings through an implanted artificial dura are difficult, the measurement and quantification of retinotopy with long-term recordings was validated by imaging early visual cortex (areas V1 and V2). Retinotopic topography was found in each of the three other areas studied within a single day's experiment. However, the ventral portion of DP (DPv) had a retinotopic topography that varied from day to day, while the more dorsal aspects (DPd) exhibited consistent retinotopy. This suggests that the dorsal prelunate gyrus may consist of more than one visual area. The retinotopy of area 7a also varied from day to day. Possible mechanisms for this variability across days are discussed as well as its impact upon our understanding of the representation of extrapersonal space in the inferior parietal cortex.     

Functional anatomy and interaction of fast and slow visual pathways in macaque monkeys

We measured the timing, areal distribution, and laminar profile of fast, wavelength-insensitive and slower, wavelength-sensitive responses in V1 and extrastriate areas, using laminar current-source density analysis in awake macaque monkeys. There were 3 main findings. 1) We confirmed previously reported significant ventral-dorsal stream latency lags at the level of V4 (V4 mean = 38.7 ms vs. middle temporal mean = 26.9 ms) and inferotemporal cortex (IT mean = 43.4 ms vs. dorsal bank of the superior temporal sulcus mean = 33.9 ms). 2) We found that wavelength-sensitive inputs in areas V1, V4, and IT lagged the wavelength-insensitive responses by significant margins; this lag increased over successive levels of the system. 3) We found that laminar activation profiles in V4 and IT were inconsistent with "feedforward" input through the ascending ventral cortical pathway; the likely alternative input routes include both lateral inputs from the dorsal stream and direct inputs from nonspecific thalamic neurons. These findings support a "Framing" Model of ventral stream visual processing in which rapidly conducted inputs, mediated by one or more accessory pathways, modulate the processing of more slowly conducted feedforward inputs.     

雄性大鼠盆底横纹肌运动神经元的定位分布

目的：探讨研究正常雄性大鼠盆底横纹肌运动神经元的定位分布。方法：分别用辣根过氧化物酶(HRP)、荧光金(FG)注射于正常雄性SD大鼠的尿道外括约肌、坐骨海绵体肌、球海绵体肌、肌门外括约肌，行逆行神经追踪。结果：尿道外括约肌或坐骨海绵体肌注射HRP或FG后，标记神经元位于脊髓L5～S1段前角背外侧核；球海绵体肌或肛门外括约肌注射HRP或FG后，标记神经元位于脊髓L5～S1段前角背内侧核。结论：雄性大鼠支配盆底横纹肌的运动神经元位于两个区域：支配坐骨海绵体肌运动神经元聚集分布于脊髓L5～S1(以L6多见)段前角背外侧核；支配球海绵体肌和肛门外括约肌的运动神经元主要聚集分布于脊髓L5～S1(以L6多见)段前角背内侧核。     

Organization of horizontal axons in the inferior temporal cortex and primary visual cortex of the macaque monkey

We investigated the organization of horizontal connections at two distinct hierarchical levels in the ventral visual cortical pathway of the monkey, the inferior temporal (TE) and primary visual (V1) cortices. After injections of anterograde tracers into layers 2 and 3, clusters of terminals ('patches') of labeled horizontal collaterals in TE appeared at various distances up to 8 mm from the injection site, while in V1 clear patches were distributed only within 2 mm. The size and spacing of these patches in TE were larger and more irregular than those observed in V1. The labeling intensity of patches in V1 declined sharply with distance from the injection site. This tendency was less obvious in TE; a number of densely labeled patches existed at distant sites beyond weakly labeled patches. While injections into both areas resulted in an elongated pattern of patches, the anisotropy was greater in TE than in V1 for injections of a similar size. Dual tracer injections and larger-sized injections further revealed that the adjacent sites in TE had spatially distinct horizontal projections, compared to those in V1. These area-specific characteristics of the horizontal connections may contribute to the differences in visual information processing of TE and V1.     

Reciprocal connectivity of identified color-processing modules in the monkey inferior temporal cortex

The inferior temporal (IT) cortex is the last unimodal visual area in the ventral visual pathway and is essential for color discrimination. Recent imaging and electrophysiological studies have revealed the presence of several distinct patches of color-selective cells in the anterior IT cortex (AIT) and posterior IT cortex (PIT). To understand the neural machinery for color processing in the IT cortex, in the present study, we combined anatomical tracing methods with electrophysiological unit recordings to investigate the anatomical connections of identified clusters of color-selective cells in monkey IT cortex. We found that a color cluster in AIT received projections from a color cluster in PIT as well as from discrete clusters of cells in other occipitotemporal areas, in the superior temporal sulcus, and in prefrontal and parietal cortices. The distribution of the labeled cells in PIT closely corresponded with that of the physiologically identified color-selective cells in this region. Furthermore, retrograde tracer injections in the posterior color cluster resulted in labeled cells in the anterior cluster. Thus, temporal lobe color-processing modules form a reciprocally interconnected loop within a distributed network.     

Color discrimination involves ventral and dorsal stream visual areas

We used positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) in human subjects to investigate whether the ventral and dorsal visual stream cooperate when active judgements about color have to be made. Color was used as the attribute, because it is processed primarily in the ventral stream. The centrally positioned stimuli were equiluminant shades of brown. The successive color discrimination task was contrasted to a dimming detection task, in which retinal input was identical but with double the number of motor responses. The stimulus presentation rate was parametrically varied and a constant performance level was obtained for all conditions. The visual activation sites were identified by retinotopic mapping and cortical flattening. In addition, one psychophysical and two fMRI experiments were performed to control for differences in visuospatial attention and motor output. Successive color discrimination involved early visual areas, including V1 and VP and the ventral color-responsive region, as well as anterior and middle dorsal intraparietal sulcus, dorsal premotor cortex and pre-SMA. Cortical regions involved in dimming detection and motor output included area V3A, hMT/V5+, lateral occipital sulcus, posterior dorsal intraparietal sulcus, primary motor cortex and SMA. These experiments demonstrated that even with color as the attribute, successive discrimination, in which a decision process has to link visual signals to motor responses, involves both ventral and dorsal visual stream areas.     

The global pattern of cytochrome oxidase stripes in visual area V2 of the macaque monkey

In primate visual area V2, histochemical staining for cytochrome oxidase (CO) reveals a tripartite pattern of densely labeled thick and thin stripes separated by pale interstripes. This modularity is believed to be related to functionally distinct processing streams that course through the hierarchy of visual areas. Here, we studied the overall pattern of CO stripes in V2 of the macaque monkey, using tissue that had been physically unfolded and flattened prior to histological sectioning. CO stripes were identified on the basis of their physical dimensions and on their differential immunoreactivity for the monoclonal antibody Cat-301. We observed several distinctive features of compartmental organization in V2. The most prominent was a dorso- ventral asymmetry in the stripe pattern, occurring in the majority of cases studied. In dorsal V2, most stripes measure approximately 10 mm in length and run roughly orthogonal to both the posterior and anterior borders of V2. In contrast, many stripes in ventral V2 have a curved or oblique trajectory, and some extend up to 20 mm in length. Stripes following a curved trajectory often become nearly parallel to the anterior border of V2. These differences imply an asymmetry in how the visual field maps onto dorsal versus ventral stripes. Occasionally, thin stripes fail to alternate with thick stripes but instead occur next to one other, separated only by interstripes. In three most complete reconstructions, we found that unfolded V2 is approximately 110 mm in length, approximately 900 mm2 in surface area, and that it contains approximately 28 complete sets of stripes (one thick, one thin and two interstripes), yielding an average of approximately 4 mm per set of stripes. The maximum width of ventral V2 (13-14 mm) exceeds that of dorsal V2 (10 mm), and there is a consistent narrowing of V2 in the region of foveal representation (3-5 mm).      

On the emergence of primary visual perception

We propose a concise novel conceptual and biological framework for the analysis of primary visual perception (PVP) that refers to the most basic levels of our awake subjective visual experiences. Neural representations for image content elaborated within V1/V2 and the early occipitotemporal (ventral) loop remain only latent with respect to PVP until spatially localized with respect to an attending observer. This process requires more than the downstream deployment of attentional resources onto targeted neurons. Additionally, the source neurons for such processes must be linked to a neural representation subserving a first-person perspective. We hypothesize that the simultaneous emergence of both the perceptual experience of image content and the personal inference of its ownership requires the resolution of any conflicting neuronal signaling between afferent and recurrent projections within and between both the ventral and dorsal streams. The V1/V2 complex and ventral cortical areas V3 and the V4 complex together with dorsal cortical areas LIP, VIP, and 7a with additional contributions from the motion areas V5/MT (middle temporal area), FST (fundus of superior temporal area), and MST (medial superior temporal area) together with their subcortical dependencies have the physiological properties required to constitute a "posterior perceptual core" that encodes the normal primary perceptual experience of image content, space, and sense of minimal self.     

Organization of Rodent Auditory Cortex: Anterograde Transport of PHA-L from MGv to Temporal Neocortex

In the present study we analyzed the organization of the thalamocorticalprojections of the specific auditory relay nucleus of the thalamus,the ventral division of the medial geniculate body (MGv), usingthe anterograde axonal tracer Phaseolus vulgaris leucoagglutinin.All injections of MGv produced dense labeling of axonal fibersin temporal cortex. In all cases, labeled axons were predominantlyconcentrated in cortical layers III and IV and, to a lesserextent, at the junction of layers V and VI. Injections confinedto the medical regions of MGv, and specifically to the avoidnucleus of MGv (OV, parsovoidea), resulted in anterograde labelingof TE1, with minor labeling of the ventral quarter of TE1, designatedsubarea TE1v. Injections placed in lateral regions of MGv andoccupying the lateral ventral subnucleus (LV), or injectionsin the mediolateral center of MGv and occupying parts of LVand OV, also resulted in labeling of area TE1 and minor labelingof TE1v. However, these injections also produced labeling inareas TE2 and TE3. Thus, area TE1 (excluding subarea TE1v) receivesheavy projections from all aspects of MGv and appears to bethe core target of MGv. While regions of MGv also project tosurrounding cortical belt areas, these projections tend to belighter and to vary depending on the region of MGv examined.These results, together with other connectional findings, andcytoarchitectonic and physiological studies, suggest that TE1(possibly excluding subarea TE1v) is the primary auditory cortexin the rat.     

Anatomical Evidence for MT and Additional Cortical Visual Areas in Humans

We stained human visual cortex for myelin. cytochrome oxidase,and the monoclonal antibody CAT-301 in an attempt to demonstrateand map MT (V5) and other visual cortical areas in humans. Bothflattened and unflattened cortical tissue was examined. A likelycandidate for area MT (V5), which we refer to as MT was demonstratedusing all three stains. Myelin and CAT-301 labels for MT weredemonstrated to be coincident by comparing results from thetwo stains in adjacent sections. In all three stains, MT wasan oval area approximately 1.2 x 2.0 cm. located 5–6 cmanterior and dorsal to the foveal V1–V2 border. The positionand size of MT as defined by the present anatomy are consistentwith MT (V5) as defined by functional measures in humans. Inaddition, flattened cortical tissue stained for cytochrome oxidaserevealed a distinctive staining topography in several corticalareas, including areas V1, V2, MT, PX, and VX. Similar studiesin flattened cortex of macaque and green monkeys demonstrateddistinctive dark cytochrome oxidase staining in MT, PX, MTc,and V3.     

Laminar specificity of intrinsic connections in Broca's area

Broca's area and its right hemisphere homologue comprise 2 cytoarchitectonic subdivisions, FDgamma and FCBm of von Economo C and Koskinas GN (1925, Die Cytoarchitektonik der Hirnrinde des erwachsenen Menschen. Vienna/Berlin [Germany]: Springer). We report here on intrinsic connections within these areas, as revealed with biotinylated dextran amine and 1,1'-dioctadecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate tracing in postmortem human brains. Injections limited to supragranular layers revealed a complex intrinsic network of horizontal connections within layers II and III spreading over several millimeters and to a lesser extent within layers IV, V, and VI. Ninety percent of the retrogradely labeled neurons (n = 734) were in supragranular layers, 4% in layer IV, and 6% in infragranular layers; most were pyramids and tended to be grouped into clusters of approximately 500 microm in diameter. Injections involving layer IV revealed extended horizontal connections within layers I-IV (up to 3.7 mm) and to a lesser extent in layers V and VI. Injections limited to the infragranular layers revealed horizontal connections mainly limited to these layers. Thus, intrinsic connections within Broca's area display a strong laminar specificity. This pattern is very similar in areas FDgamma and FCBm and in the 2 hemispheres.     

Functional response properties of neurons in the dorsomedial visual area of New World monkeys (Callithrix jacchus)

The dorsomedial visual area (DM), a subdivision of extrastriatecortex located near the dorsal midline, is characterized byheavy myelination and a relative emphasis on peripheral vision.To date, DM remains the least understood of the three primarytargets of projections from the striate cortex (V1) in New Worldmonkeys. Here, we characterize the responses of DM neurons inanaesthetized marmosets to drifting sine wave gratings. Most(82.4%) cells showed bidirectional sensitivity, with only 6.9%being strongly direction selective. The distribution of orientationsensitivity was bimodal, with a distinct population (correspondingto over half of the sample) formed by neurons with very narrowselectivity. When compared with a sample of V1 units representinga comparable range of eccentricities, DM cells revealed a preferencefor much lower spatial frequencies, and higher speeds. End inhibitionwas extremely rare, and the responses of many cells summatedover distances as large as 30°. Our results suggest cleardifferences between DM and the two other main targets of V1projections, the second (V2) and middle temporal (MT) areas,with cells in DM emphasizing aspects of visual information thatare likely to be relevant for motor control.     

Properties of Simulated Neurons from a Model of Primate Inferior Temporal Cortex

The physiological properties of neurons in inferior temporal(IT) cortex of the macaque monkey suggest that this corticalarea plays a major role in visual pattern recognition. Basedon the properties of IT, and one of its major sources of inputV4, a model is proposed that can account for some of the shaperecognition properties of IT neurons including selectivity forcomplex visual stimuli and tolerance to the size and locationof the stimuli. The model is composed of three components. Firststimulus location tolerance is modeled after the complex-cell-likeproperties observed in some V4 neurons. The second componentof the model is an attentionally controlled scaling mechanismthat facilitates size-invariant shape recognition. The transitionfrom edge orientation-selective neurons in V4 to neurons withmore complicated stimulus preference in IT is explained by thethird component of the model, a competitive learning mechanism.Single-unit analysis of receptive field properties, stimulusselectivity, and stimulus size and position tolerance was performedon "neurons" from the simulation. Comparison of results fromthe simulation and a study of actual IT neurons shows that theset of mechanisms incorporated into the simulation is sufficientto emulate the physiological data.