Retinotopic organization of striate and extrastriate visual cortex in the hooded rat

The visuotopic organization of the primary visual cortex (area 17) and the extrastriate visual regions surrounding it (areas 18a and 18) has been studied in gray rats using standard microelectrode mapping techniques. The results confirm and extend previous observations in the rat. Apart from the representation of the contralateral visual field (VF) in area 17, in which the upper VF is represented caudally and the nasal VF laterally, there are additional representations of the VF in the extrastriate cortex. In lateral extrastriate cortex (area 18a) there are at least 4 such representations, namely lateromedial (LM), anterolateral (AL), laterointermediate (LI) and laterolateral (LL). In LM (second visual area) the upper VF is represented caudally and the nasal VF medially, being thus a mirror image of V1. In AL (third visual area) the upper VF is represented rostrally and the nasal VF, medially, being thus a mirror image of LM. In LI, the upper VF is medial and the nasal VF, lateral, being thus a mirror image of LM, or a reduced copy of V1. In medial extrastriate cortex (area 18) there are two representations of the temporal VF, labeled anteromedial (AM) and posteromedial (PM). In AM, the upper temporal VF is medial and the lower temporal VF, lateral, the extreme temporal field being rostral. The 30° azimuth provides the boundary between AM and PM. Thus, AM is organized as a counter-clockwise rotation by 90° of the V1 representation. In PM, the upper lower VF topography is like in AM, but the extreme temporal VF is caudal, being thus a mirror image of AM.     

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 of early visual evoked potential generators by retinotopic and topographic analyses

This study investigated the cortical sources of the early (50-250 ms) components of the pattern-onset visual evoked potential (VEP). VEPs were recorded in response to a small circular checkerboard stimulus that was flashed over a range of visual field positions. Temporally and spatially overlapping VEP components were distinguished by differences in retinotopic sensitivity and scalp topography, and by inverse dipole modeling. The C1 component (50-80 ms) was found to change its polarity and topography systematically as a function of stimulus position in a manner consistent with the retinotopic organization of the striate cortex. The P1 component (comprised of the P75 and P100 subcomponents) had a time course that overlapped the C1 but could be distinguished from the C1 by its differing topography and reduced sensitivity to stimulus position. The P1 generators were localized to the lateral extrastriate cortex. Inverse dipole models were consistent with these striate and extrastriate source locations for the C1 and P1, respectively. The N1 component (120-180 ms) was found to originate from several spatially distinct generators that differed in their retinotopic organization. © 1995 Wiley-Liss, Inc.     

Negative BOLD in the visual cortex: evidence against blood stealing

The positive BOLD (blood oxygen level-dependent) response elicited in human visual cortex by a localized visual stimulus is accompanied by a reduction in the BOLD response in regions of the visual cortex that represent unstimulated locations in the visual field. We have suggested previously that this negative BOLD reflects attention-related suppression of neural activity, but it might also be explained in terms of "blood stealing," i.e., hemodynamic changes that have no neural correlate. We distinguish two possible hemodynamic effects of this type: (1). blood flow reduction caused by locally reduced pressure in vessels that share their blood supply with nearby dilated vessels; and (2). blood flow reduction caused by active constriction of vessels under neural control. The first is ruled out as an explanation of negative BOLD by showing that a visual stimulus that stimulates primary visual cortex in one hemisphere can cause extensive suppression in the other hemisphere i.e., it is not a local phenomenon. Negative BOLD most likely reflects suppression of neural activity, but could also reflect an active blood flow control system.     

Effects of lateral suprasylvian visual cortex lesions on visual localization, discrimination, and attention in cats

Experiments were carried out to begin to define the behavioral functions of the lateral suprasylvian (LS) visual area of the cat's cortex. Behavioral tasks were chosen for analysis on the basis of previous suggestions in the literature concerning possible functions of LS cortex and its afferent pathways. These tasks included the ability of cats to orient the head and eyes to a stimulus presented in particular locations in the visual field, the ability to learn successive reversals of a two-choice visual pattern discrimination, and the ability to maintain or shift attention between relevant or irrelevant visual form and brightness cues. Eight cats were trained on each of these tasks. Four of the cats then received bilateral lesions of LS cortex, including the AMLS and PMLS regions, and the remaining 4 cats were used to assess normal retention. The LS cortex lesions had no significant effect upon performance of any of the behaviors tested. Thus, this region of cortex appears to play no essential role in simple brightness, form, and pattern discrimination performance, visual reversal learning, maintaining and shifting visual attention, or orienting the head and eyes to stimuli in the visual field. These results are discussed in relation to previous lesion studies involving large regions of the cat's extrastriate cortex and studies in other species. Possible functions of LS cortex, based upon recent electrophysiological studies, are suggested.     

Dark rearing prolongs physiological but not anatomical plasticity of the cat visual cortex

Recent studies (Cynader and Mitchell, '80; Mower et al., '81) have shown that total dark rearing prolongs susceptibility to the physiological effects of monocular deprivation (MD) in visual cortex beyond the normal age limits. The present study addressed whether this delayed physiological plasticity is accompanied by delayed anatomical plasticity in the geniculocortical pathway. Ocular dominance (OD) columns as defined by transsynaptic autoradiography following injection of 3H proline into one eye were studied both qualitatively and quantitatively in 17 cats. Compared to normal rearing (N-3), both binocular eyelid suture (N-2) and total dark rearing (N-3) resulted in incomplete segregation of OD columns in area 17. This apparent immaturity after binocular deprivation, however, did not reflect a delayed capacity for development and plasticity. Visual experience after dark rearing produced no marked changes. In cats who experienced MD after dark rearing, injection of either the nondeprived (N-2) or deprived eye (N-3) resulted in a nearly uniform distribution of label throughout layer IV of area 17. The same result occurred with binocular vision after dark rearing (N-1). MD from birth, however, produced expansion of columns from the nondeprived eye (N-1) and contraction of columns from the deprived eye (N-1). MD imposed after 4 months of normal vision resulted in normal OD columns (N-1). Electrophysiological studies revealed a high proportion of binocular cells within layer IV in cats who experienced monocular or binocular vision after dark rearing. Outside of layer IV there were clear environmental effects on OD of single cells in these cats. Measurements of cell sizes in the clateral geniculate nucleus showed shrinkage of cells innervated by the deprived eye when MD was initiated at birth (N-3). MD after dark rearing (N-4) produced no differences in cell sizes. It is concluded that visual input is necessary for the formation of normal OD columns, the critical period for formation and environmental modification of OD columns is limited to early life, and the physiological effects of visual experience after dark rearing reflect changes occurring beyond the geniculocortical pathway.     

Organization of association projections from area 17 to areas 18 and 19 and to suprasylvian areas in the cat's visual cortex.

Cells in area 17 that are labelled by single, discrete injections of retrogradely transported tracers into extrastriate visual areas are discontinuously distributed in dense patches. In this study we made multiple, closely spaced injections of fluorescent dyes into extrastriate areas, to generate large deposits that would reveal whether the distributions of corticocortical cell bodies in area 17 are truly patchy or appear clustered only after small injections. By injecting a different tracer into each extrastriate area, or group of areas, we examined the spatial relationships between the populations of association cells. All deposits of tracers in areas 18, 19, or suprasylvian cortex, irrespective of size, label cells in a series of clusters in topographically related parts of area 17. We conclude that the complete populations of cells in area 17 that project to areas 18, 19, and the lateral suprasylvian cortex are all genuinely distributed in a patchy fashion. There appears to be a complex relationship between the sets of association cells projecting to different extrastriate regions: they do not completely overlap, only partially, and share some cortical zones but not others. In these experiments, only tiny percentages (2-5%) of labelled cells in the overlapping regions were filled with both tracers, suggesting that very few association cells in area 17 project to more than one of the extrastriate areas we studied. By comparing the dimensions of each injection site and of the labelled region in area 17, we estimated the extent of the convergence from area 17 to areas 18, 19, and posteromedial suprasylvian areas in retinotopic terms. The functional convergence was very similar in these pathways.     

Human primary visual cortex topography imaged via positron tomography

The visuotopic structure of primary visual cortex was studied in a group of 7 human volunteers using positron emission transaxial tomography (PETT) and¹⁸F-labeled 2-deoxy-2-fluoro-d-glucose ([¹⁸F]DG). A computer animation was constructed with a spatial structure which was matched to estimates of human cortical magnification factor and to striate cortex stimulus preferences. A lateralized cortical ‘checker-board’ pattern of [¹⁸F]DG was stimulated in primary visual cortex by having subjects view this computer animation following i.v. injection of [¹⁸F]DG. The spatial structure of the stimulus was designed to produce an easily recognizable ‘signature’ in a series of 9 serial PETT scans obtained from each of a group of 7 volunteers. The predicted lateralized topographic ‘signature’ was observed in 6 of 7 subjects. Applications of this method for further PETT studies of human visual cortex are discussed.     

Ventral posterior visual area of the macaque: visual topography and areal boundaries

Using both physiological and anatomical techniques, we have studied the topographic organization of extrastriate visual cortex on the ventral surface of the occipital lobe in macaque monkeys. Our results show that a topographically organized representation of the superior contralateral quadrant of the visual field lies immediately anterior to the ventral half of V2. This area is organized in a mirror symmetric fashion to ventral V2: it shares a horizontal meridian representation with V2 and a representation of the superior vertical meridian forms its anterior border. A well-defined strip of callosal inputs runs along the vertical meridian representation, thereby providing a reliable anatomical marker for areal boundaries in ventral extrastriate cortex. We refer to this area as the ventral posterior area (VP) because it is, in all these respects, notably similar to VP in the owl monkey. Ventral V2 has strong reciprocal connections with VP, and the topography of the V2 projection agrees closely with the topography revealed in our physiological mapping experiments. The visual field representation in VP is strikingly anisotropic, with linear magnification factor being much larger along contours of constant polar angle than along contours of constant eccentricity.     

Intersubject variability of functional areas in the human visual cortex

Intersubject variability of striate and extrastriate areas was mapped by conjoined use of positron emission tomography (PET) and magnetic resonance imaging (MRI). We used two dynamic bowtie-shaped random-dot patterns centered symmetrically around the vertical- and horizontal-meridian, respectively, presented during sequential PET scans in 11 subjects. Control condition was simple fixation on a central dot in absence of a surrounding random dot pattern. V1, V2, VP, V3, V3a, V4, V5, and “wordform” areas were identified. After spatial normalization to Talairach atlas space, mean locations and standard deviations about these mean locations for x-, y-, and z-axes were calculated for each area in both hemispheres and compared for differences. The mean standard deviation for all axes across all areas tested was found to be small (4.9 mm). No significant differences were found in the mean standard deviations for the x-, y-, and z-axes in the left hemisphere vs. their counterparts in the right hemisphere. However, when mean standard deviations in both hemispheres were polled together by axis, the mean standard deviation for the y-axis (5.3 mm) was found to be significantly different from the mean standard deviation for the x-axis (4.3 mm). Furthermore, in the left hemisphere, the mean standard deviation for the z-axis (5.7 mm) was significantly greater than the mean standard deviation for the x-axis (3.9 mm). The values reported in this study for mean location and standard deviation of visual areas can be used to establish confidence intervals for distinguishing normal variations from pathology and consequent brain reorganization. Hum. Brain Mapping 6:301–315, 1998. © 1998 Wiley-Liss, Inc.     

Effects of dark-rearing on the vascularization of the developmental rat visual cortex

Cerebral vascular density corresponds to metabolic demand, which increases in highly active areas. External inputs play an important role in the modeling and development of the visual cortex. Experience-mediated development is very active during the first postnatal month, when accurate simultaneous blood supply is needed to satisfy increased demand. We studied the development of visual cortex vascularization in relation to experience, comparing rats raised in darkness with rats raised in standard conditions. The parameters measured were cortical thickness, vascular density and number of perpendicular vessels, constituting the first stage of cortical vascular development. Vessels were stained using butyryl cholinesterase histochemistry, which labels some neurons and microvascularization (vessels from 5 to 50 μm). Animals from both groups were sampled at 0, 7, 14, 21 and 60 days postnatal. Vascularization of the brain starts with vertically oriented intracortical vascular trunks whose density decreases notably after birth in rats reared in standard laboratory conditions. The most striking finding of our work is the significantly lower decrease in the number of these vessels in dark-reared rats. Our results also show that cortex thickness and vessel density are significantly lower in dark-reared rats. These results suggest that the absence of visual stimuli retards the maturation of the visual cortex including its vascular bed.     

Interocular temporal delay sensitivity in the visual cortex of the awake monkey.

Due to the separation of the eyes, temporal retinal disparities are created during binocular stimulation and they have been proposed to be the basis of several stereo-visual effects. This paper studies the sensitivity of cortical neurons from area V1 to interocular temporal delay in the awake monkey (Macaca mulatta). Forty-four cells were included in this study. Temporal delay sensitivity was observed in 59% of them. About half of these temporal-delay-sensitive cells were also sensitive to the stimulation sequence of the eyes. The cells that preferred one eye to be stimulated first were termed asymmetrical (46%); those which were not sensitive to the eye sequence of stimulation were termed symmetrical (54%). No clear differences were observed in the distribution of delay-sensitive cells according to their eye dominance. Fifty-six percent of balanced cells and 65% of unbalanced cells were sensitive to interocular delay. These data underline the importance of temporal cues for depth perception.     

Rearrangement of synaptic connections with inhibitory neurons in developing mouse visual cortex

Cortical inhibition is determined in part by the organization of synaptic inputs to gamma-aminobutyric acidergic (GABAergic) neurons. In adult rat visual cortex, feedforward (FF) and feedback (FB) connections that link lower with higher areas provide approximately 10% of inputs to parvalbumin (PV)-expressing GABAergic neurons and approximately 90% to non-GABAergic cells (Gonchar and Burkhalter [1999] J. Comp. Neurol. 406:346-360). Although the proportions of these targets are similar in both pathways, FF synapses prefer larger PV dendrites than FB synapses, which may result in stronger inhibition in the FF than in the FB pathway (Gonchar and Burkhalter [1999] J. Comp. Neurol. 406:346-360). To determine when during postnatal (P) development FF and FB inputs to PV and non-PV neurons acquire mature proportions, and whether the pathway-specific distributions of FF and FB inputs to PV dendrites develop from a similar pattern, we studied FF and FB connections between area 17 and the higher order lateromedial area (LM) in visual cortex of P15-42 mice. We found that the innervation ratio of PV and non-PV neurons is mature at P15. Furthermore, the size distributions of PV dendrites contacted by FF and FB synapses were similar at P15 but changed during the third to sixth postnatal weeks so that, by P36-42, FF inputs preferred thick dendrites and FB synapses favored thin PV dendrites. These results suggest that distinct FF and FB circuits develop after eye opening by rearranging the distribution of excitatory synaptic inputs on the dendritic tree of PV neurons. The purpose of this transformation may be to adjust differentially the strengths of inhibition in FF and FB circuits.     

Projections to the superior temporal sulcus from the central and peripheral field representations of V1 and V2

In a series of three studies, we have begun to explore the sequence of visual information processing along the pathway from striate cortex (V1), through MT, into the parietal lobe. In this first study, we sought to establish the relationships among MT, the heavily myelinated zone of the superior temporal sulcus (STS), and the V1 and V2 projection fields in the STS. Autoradiographic material from seven hemispheres of six macaques injected with tritiated amino acids into either V1 or V2 was analyzed in detail, and the results were plotted onto two-dimensional reconstructions of the STS. Autoradiographic material from eight additional macaques with V2 injections was also examined. The results indicate that the central visual field representations of both V1 and V2 project into the heavily myelinated zone in the lower bank and floor of the STS, confirming prior studies, whereas the far peripheral representations of both V1 and V2 project into the cortex medial to this zone on the upper bank of the sulcus. There is no evidence that this medial cortex is a separate area that receives projections from V1 and V2 in parallel with the projections these areas send to the heavily myelinated zone. Rather, there seems to be a single projection field of V1 and V2 whose central representation lies within the heavily myelinated zone and whose most peripheral representation lies medial to it. Because of the difference in myelination between the central and peripheral field representations as well as visuotopic anomalies between them, we retain the term "MT" for the heavily myelinated zone and apply the term "MTp" to the far peripheral projection zone. Both MT and MTp are required to process the complete outputs of V1 and V2 within the STS and thus should probably be regarded as two distinctive parts of a single visual area. The difference in myelination between MT and MTp suggests that there is a difference in visual processing between the central and peripheral visual fields. The average size of MT is estimated to be 62 mm2, and the average size of MT and MTp combined to be 76 mm2, which is consistent with estimates derived from several other studies.     

Topography of the afferent connectivity of area 17 in the macaque monkey: a double-labelling study

The various structures afferent to area 17 (or V1) of the macaque monkey have widely differing retinotopic organizations. It is likely that these differences are reflected in the topographic organizations of the projections from these structures to area V1. We have investigated this issue by placing side-by-side injections of two retrograde fluorescent tracers, fast blue and diamidino yellow, in V1. By examining the extent of mixing of the two populations of singly labelled cells and the presence of doubly labelled cells, in different structures, we have characterized the topography of each projection in terms of the size of its axonal arborization and the amount of convergence and divergence. The afferents from the lateral geniculate nucleus (LGN) and from the pulvinar are organized in a point-to-point fashion. The maximum extent of axonal arborization of these afferents is 0.5 mm and these projections demonstrate little scatter (i.e., neighboring LGN neurons project to adjacent regions of V1). The other two subcortical structures examined, the claustrum and the intralaminar nuclei, demonstrate a much larger scatter and wider axonal arborizations in their projections to V1 than do the LGN and pulvinar. Two-dimensional reconstructions were made of the distribution of labelled neurons in extrastriate cortical areas. Using the separation between patches of labelled cells and transitions in myelin-stained sections, we have identified seven separate cortical regions containing labelled cells. Two of these can be identified as area V2 and the middle temporal visual area (MT). Three other regions correspond to areas V3, V3A and V4t. Finally, two more regions of labelling have been distinguished that belong to area V4. These results demonstrate that, at least within the central 6 degrees of visual field, all the presently known extrastriate visual cortical areas project to V1. This result is interesting in view of the fact that only a few extrastriate cortical areas are reported to receive afferents from V1. Three groups of cortical areas can be distinguished on the basis of the characteristics of their cortical connections to V1. The first group contains area V2, V3, and the posterior region of V4. These areas project to V1 with infra- as well as supragranular layer neurons and show limited axonal arborization and scatter in the projection. The second group consists of two regions of labelling in the superior temporal sulcus corresponding to V4t and MT and another on the annectant gyrus (V3A). These regions contain almost exclusively infragranular labelling and show wide axonal arborization and scatter in their projections to V1.(ABSTRACT TRUNCATED AT 400 WORDS)     

Modulatory actions of the gigantocellular reticular nucleus on baroreceptor reflexes in the cat

Threshold visual acuity for three cats which were reared from birth to 4–12 months of age with bilateral lid closure was measured and compared to visual acuity in three cats which had the use of a non-deprived eye. The results indicate that binocular deprivation (BD) results in significant deficits in visual acuity which are proportional to the duration of deprivation. Threshold visual acuities were 3.7 cycles/deg. following 4 months of BD, 3.25 cycles/deg. following 7 months of BD and 2.55 cycles/deg. following 12 months of BD compared to acuities of 6.0, 6.5 and 6.8 cyclesdeg. for cats using a non-deprived eye. All BD cats had recovered from the initial visuomotor deficits, seen in these cats and reported in the literature, following lid-parting. The implication of such deficits in visual acuity on visual discrimination learning in BD cats is discussed.     

Evidence from V1 connections for both dorsal and ventral subdivisions of V3 in three species of New World monkeys

We used patterns of connections of primary visual cortex (V1) to reevaluate differing proposals on the organization of extrastriate cortex in three species of New World monkeys. Several fluorescent tracers and the bidirectional tracer cholera toxin B subunit (CTB) were injected into dorsal V1 (representing the lower visual quadrant) and ventral V1 (representing the upper visual quadrant) of titi, squirrel, and owl monkeys. Labeled cells and terminals were plotted on brain sections cut parallel to the surface of flattened cortex and were related to architectonic boundaries. The results provided compelling evidence for both dorsal V3 with dorsal V1 connections and ventral V3 with ventral V1 connections. The connection pattern indicated that V3 represents the visual hemifield as a mirror image of V2. In addition, V3 could be recognized by a weak banding pattern in brain sections processed for cytochrome oxidase. V1 has connections with at least 12 subdivisions of visual cortex, with half of the connections involving V2 and 20% V3. Comparable results were obtained from all three species, suggesting that visual cortex is similarly organized.     

Enucleation demonstrates ocular dominance columns in Old World macaque but not in New World squirrel monkey visual cortex

The effect of monocular enucleation on basophilic and metabolic staining in primary (striate) visual cortex has been compared in Old and New World monkeys. Both species show a 30-40% shrinkage of neurons in the layers of dorsal lateral geniculate nucleus receiving axons from the enucleated eye. In striate cortex Old World macaque monkeys show alternating bands of increased and diminished staining in layers 3, 4 and 6, corresponding to ocular dominance columns. New World squirrel monkeys show staining patterns in all layers which are unchanged from normal cortex, suggesting that New World monkeys lack obvious ocular dominance columns.