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Biomedical subjects

H Sherk

Publications and source records attributed to H Sherk.

At least 19 recordsLinked to original sources
PubMed2001

Prolonged survival of axons terminating within lesions of cat visual cortex.

It is well known that brain lesions made by the injection of ibotenic acid destroy neuronal cell bodies but do not kill passing axons. We have found that axons terminating within such lesions in visual cortex also survive for at least 2 months, despite the absence of available synaptic sites. We made tracer injections in area 17, and observed dense patches of anterograde label within lesions in other visual cortical areas. Furthermore, because the retinotopic site of the tracer injection was known, we could conclude that the retinotopic site was encompassed within the lesion.

Animals↗
PubMed2001

Neural analysis of visual information during locomotion.

For most vertebrate species, vision is critical during locomotion. In this review, we survey what is known about neural mechanisms that might be involved in this visual analysis. Problems that such mechanisms are likely to solve include: (1) determination of heading (the discrepancy between an observer's direction of motion and direction of gaze); (2) detection of course changes; (3) setting of courses using goals and landmarks; (4) obstacle avoidance; (5) accurate foot placement. To understand how the visual system handles these tasks, we must first consider what neurons 'see' during locomotion: this is determined both by an observer's motion and by his gaze. We then review response properties of neurons in the cortical 'motion pathways' of the monkey and cat as they relate to problems encountered during locomotion. The most studied areas are the medial superior temporal area in the monkey, which has been linked to heading determination, and the lateral suprasylvian area in the cat, where many cells are sensitive to motion in depth, and some are selective for optic flow patterns generated during locomotion. A few subcortical populations have also been linked to visual analysis during locomotion. Most notable are cells in the pigeon's nucleus rotundus: these respond selectively to looming stimuli, some firing at a specific time before the stimulus collides with the bird. Another intriguing population is in the cat's visual pontine nucleus, where cells respond to large displays suggestive of optic flow during locomotion.

Animals↗
PubMed2001

Visual analysis and image motion in locomoting cats.

During locomotion, observers see a characteristic pattern of motion referred to as an optic flow field. To investigate how they make use of this pattern, we have developed a paradigm for testing visual function during locomotion. Foot placement was recorded while cats walked down an alley cluttered with a high density of small objects; the task was to avoid stepping on any object. In the experiments reported here, motion cues were eliminated by the use of low-frequency strobe lighting. In bright continuous light cats performed with great accuracy, and likewise at scotopic light levels. However, in strobe lighting their error rates increased more than threefold. This deterioration could not be attributed to lower acuity, since the cats' performance remained excellent when the light level was reduced well below that afforded by the strobe light. When very dim continuous light was combined with low-frequency strobe lighting, performance was substantially better than under strobe light alone. We conclude that motion-sensitive neurons make a major contribution to visual guidance of foot placement during locomotion. When strobe lighting is combined with very dim continuous light, even the minimal motion information available in the intervals between bright strobe flashes improves performance significantly. Cats were also trained to discriminate between complex patterns, and this discrimination was not affected by strobe lighting, suggesting that motion-sensitive neurons are not critical for this analysis.

Animals↗
PubMed1997

Simulated optic flow and extrastriate cortex. I. Optic flow versus texture.

A locomoting observer sees a very different visual scene than an observer at rest: images throughout the visual field accelerate and expand, and they follow approximately radial outward paths from a single origin. This so-called optic flow field is presumably used for visual guidance, and it has been suggested that particular areas of visual cortex are specialized for the analysis of optic flow. In the cat, the lateral suprasylvian visual area (LS) is a likely candidate. To test the hypothesis that LS is specialized for analysis of optic flow fields, we recorded cell responses to optic flow displays. Stimulus movies simulated the experience of a cat trotting slowly across an endless plain covered with small balls. In different simulations we varied the size of balls, their organization (randomly or regularly dispersed), and their color (all one gray level, or multiple shades of gray). For each optic flow movie, a "texture" movie composed of the same elements but lacking optic flow cues was tested. In anesthetized cats, > 500 neurons in LS were studied with a variety of movies. Most (70%) of 454 visually responsive cells responded to optic flow movies. Visually responsive cells generally preferred optic flow to texture movies (69% of those responsive to any movie). The direction in which a movie was shown (forward or reverse) was also an important factor. Most cells (68%) strongly preferred forward motion, which corresponded to visual experience during locomotion.

Animals↗
PubMed1997

Simulated optic flow and extrastriate cortex. II. Responses to bar versus large-field stimuli.

In the preceding paper we described the responses of cells in the cat's lateral suprasylvian visual area (LS) to large-field optic flow and texture movies. To assess response properties such as direction selectivity, cells were also tested with moving bar stimuli. We expected that there would be good agreement between response properties elicited with optic flow movies and those revealed with bar stimuli. We first asked how well bar response properties predicted responsiveness to optic flow movies. There was no correlation between responsiveness to movies and the degree of end-stopping, length summation, or preference for bars that accelerated and expanded. We then considered only the 322 cells that responded to both bars and optic flow or texture movies and asked how well the strength of their response to movies could be predicted from the direction-tuning curves generated with bar stimuli. One-third of these cells responded much more strongly to movies than could be predicted from their direction-tuning curves. Generally, such cells were rather well tuned for the direction of bar motion and preferred a direction substantially different from what they saw in optic flow movies. Optic flow movies shown in the forward direction were the most effective variety of movie for two-thirds of these cells. To see whether this outcome stemmed from differential direction tuning for bars and large multielement displays, in a second series of experiments we compared direction tuning for bars and large-field texture movies. Many cells showed substantially different direction tuning for the two kinds of stimulus: almost 1/3 of 409 cells had tuning curves that overlapped each other by < 50%. But only a small number of cells (< 10%) responded much better to texture movies than to bars in the predominant direction of image motion in optic flow movies. This result, like that reported in the preceding paper, suggests that cells in LS respond differently to optic flow than to texture displays lacking optic flow motion cues.

Animals↗
PubMed1995

Sources of subcortical afferents to the macaque's dorsal lateral geniculate nucleus.

BACKGROUND: The dorsal lateral geniculate nucleus (dLGN) is the thalamic region responsible for transmitting retina signals to cortex. Brainstem pathways to this nucleus have been described in several species and are believed to control the retinocortical pathway depending on the state of the animal (awake, asleep, drowsy, etc.). The purpose of this study was to determine all of the subcortical sources of afferents to the dLGN in a higher primate, the macaque monkey, whose visual system is similar to that of humans. METHODS: Injections of horseradish peroxidase (HRP), with or without conjugation to wheat germ agglutinin, were made into the dLGNs of seven macaque monkeys, followed by perfusion, brain sectioning, and analyses of neurons in the brainstem, thalamus, and hypothalamus that contained the retrogradely transported marker. RESULTS: The reticular nucleus of the thalamus, pedunculopontine nucleus, parabigeminal nucleus, pretectal nucleus of the optic tract, superior colliculus, dorsal raphe nucleus, and tuberomammillary region of the hypothalamus contained many retrogradely labeled neurons ipsilateral to the injections. In the contralateral brainstem, HRP-labeled cells were found only in the pedunculopontine nucleus, nucleus of the optic tract, and dorsal raphe nucleus. The number of labeled neurons on the contralateral side was about one-half of that in corresponding ipsilateral nuclei. The locus coeruleus contained no labeled neurons in four of the macaques that had injections limited to the dLGN. CONCLUSION: There are seven subcortical regions that send afferents to the dLGNs of macaque monkeys. Except for the locus coeruleus, these are the same as observed for other species, such as the cat and rat, and indicate the possible sources of subcortical control over the dLGNs of humans.

Animals↗
PubMed1993

A comparison of magnification functions in area 19 and the lateral suprasylvian visual area in the cat.

A retinotopic map can be described by a magnification function that relates magnification factor to visual field eccentricity. Magnification factor for primary visual cortex (V1) in both the cat and the macaque monkey is directly proportional to retinal ganglion cell density. However, among those extrastriate areas for which a magnification function has been described, this is often not the case. Deviations from the pattern established in V1 are of considerable interest because they may provide insight into an extrastriate area's role in visual processing. The present study explored the magnification function for the lateral suprasylvian area (LS) in the cat. Because of its complex retinotopic organization, magnification was calculated indirectly using the known magnification function for area 19. Small tracer injections were made in area 17, and the extent of anterograde label in LS and in area 19 was measured. Using the ratio of cortical area labeled in LS to that in area 19, and the known magnification factor for area 19 at the corresponding retinotopic location, we were able to calculate magnification factor for LS. We found that the magnification function for LS differed substantially from that for area 19: central visual field was expanded, and peripheral field compressed in LS compared with area 19. Additionally, we found that the lower vertical meridian's representation was compressed relative to that of the horizontal meridian. We also examined receptive field size in areas 17, 19, and LS and found that, for all three areas, receptive field size was inversely proportional to magnification factor.

Animals↗
PubMed1992

Retinotopic order is surprisingly good within cell columns in the cat's lateral suprasylvian cortex.

The retinotopic map in the striate-recipient region of the cat's lateral suprasylvian cortex (referred to here as the lateral suprasylvian area (LS)) has generally been described as quite disorderly. The disorder is commonly attributed to receptive field scatter within cell columns, reflecting the very large size of receptive fields. However, scatter within columns has never been investigated. In the experiments reported here, we examined the receptive field scatter of cells in columns, and also the scatter of a limited sample of their afferents arising from areas 17 and 18. To measure post-synaptic receptive field scatter, electrode penetrations were made parallel to columns in LS, with the electrode approaching from the medial side, traversing the suprasylvian gyrus and emerging into the suprasylvian sulcus. In all 13 such penetrations, receptive fields were clustered together despite their large size. Their centers were scattered over a region that occupied on average less than 20% of the largest field in the column. In contrast, in columns in areas 17 and 18 receptive field centers reportedly are dispersed over regions about equal to the largest of the fields (Hubel and Wiesel 1962, 1965, 1974). The scatter of afferents' receptive fields was assessed anatomically by measuring the overlap between patches of different anterograde tracers in LS. These patches represented terminal labeling from two adjacent or overlapping tracer injections in area 17. While a large degree of overlap would be predicted if afferents have substantial scatter, we found the overlap to be small unless the two injection sites themselves were highly overlapping. Scatter in afferents' receptive fields was measured more directly by physiological recording. In previous experiments, cells in LS were silenced by the local injection of kainic acid, and responses were recorded from axon terminals arising from areas 17 and 18 (Sherk 1989). We examined the receptive field scatter in three penetrations made approximately normal to the cortical surface. Scatter was modest, much less than predicted by the size of post-synaptic receptive fields. Because the degree of receptive field scatter for postsynaptic cells in LS was similar to that of inputs from areas 17 and 18, the scatter of these inputs might be entirely responsible for that seen postsynaptically. Postsynaptic receptive field scatter, on the other hand, was too small to explain the reported disorder in the map in LS.

Animals↗
PubMed1992

Flattening the cerebral cortex by computer.

A computer program was developed for unfolding the cerebral cortex so that it could be viewed as a 2-dimensional surface. Input to the program consisted of tissue sections cut in a standard plane of section. Each section was represented by one line, which corresponded to a contour line in the flattened map. From these data, the computer constructed a 3-dimensional surface representation, which it then flattened. Because the cerebral cortex has considerable intrinsic curvature, flattening required that some regions be expanded and others shrunken. These changes occurred as a natural consequence of local decisions made by the computer as it laid down successive contours. The user could intervene during both surfacing and flattening in order to shape the developing map. The program has been used to generate 37 flattened maps from various regions of cat cortex, and 1 from monkey cortex. The local topography of cortical features such as gyri, sulci, architectonic boundaries, and patches of transported tracer, appeared to be conserved fairly faithfully. Areal distortion was also modest, with an average change in surface area of only 12%.

Animals↗
PubMed1990

Functional organization of input from areas 17 and 18 to an extrastriate area in the cat.

In the cat, areas 17 and 18 have 2 main cortical targets: area 19 and a large region of suprasylvian cortex, referred to here as the Clare-Bishop area (Hubel and Wiesel, 1969). The functional organization of the latter area is not well understood. It seems likely that its organization reflects, in part, the organization of its inputs from areas 17 and 18, and I therefore studied the functional organization of these inputs. Responses were recorded directly from afferents terminating in the Clare-Bishop area after silencing its cells by local injections of kainic acid. Afferents were precisely ordered according to preferred orientation in a fashion resembling the columnar organization of area 17. Preferred direction was also an important organizing parameter. Within sizable regions of cortex, directional preference changed gradually or not at all, while at these regions' boundaries, preferred direction reversed by 180 degrees. Except for a loose grouping according to eye dominance, afferents were not ordered according to other response properties. It is possible that the afferent array is also compartmentalized according to the area of origin of each afferent. Inputs from areas 17 and 18, for example, might end in separate territories in the Clare-Bishop area. However, physiological recording, as well as anatomical experiments, failed to show a clear segregation. This study provides the first direct demonstration that a cortico-cortical pathway can be organized according to particular response properties. While the existence of an organization according to preferred orientation was not surprising, the high degree of order was unexpected. It is also interesting that afferents were well ordered according to 1 parameter, directional preference, that is weakly organized, at best, in area 17.

Afferent Pathways↗
PubMed1989

Visual response properties of cortical inputs to an extrastriate cortical area in the cat.

The existence of multiple areas of extrastriate visual cortex raises the question of how the response properties of each area are derived from its visual input. This question was investigated for one such area in the cat, referred to here as the Clare-Bishop area (Hubel & Wiesel, 1969); it is the region of lateral suprasylvian cortex that receives input from area 17. A novel approach was used, in which kainic acid was injected locally into the Clare-Bishop area, making it possible to record directly from afferent inputs. The response properties of the great majority of a sample of 424 presumed afferents resembled cells in areas 17 and 18. Thus, a systematic comparison was made with cells from area 17's upper layers, the source of its projection to the Clare-Bishop area (Gilbert & Kelly, 1975), to see whether these afferents had distinctive properties that might distinguish them from cells projecting to areas 18 or 19. Some differences did emerge: (1) The smallest receptive fields typical of area 17 were relatively scarce among afferents. (2) Direction-selective afferents were more abundant than were such cells in area 17. (3) End-stopped afferents were extremely rare, although end-stopped cells were common in area 17's upper layers. Despite these differences, afferents were far more similar in their properties to cells in areas 17 and 18 than to cells in the Clare-Bishop area. Compared to the latter, afferents showed major discrepancies in receptive-field size, in direction selectivity, in end-stopping, and in ocular dominance distribution. These differences seem most likely to stem from circuitry intrinsic to the Clare-Bishop area.

Action Potentials↗
PubMed1988

The retinotopic match between area 17 and its targets in visual suprasylvian cortex.

It is widely believed that cells in area 17 send axons specifically to neurons in other cortical areas whose receptive fields coincide with their own. We asked whether this was true in cats for area 17's projection to a large suprasylvian visual area, the Clare-Bishop area. Receptive fields were plotted at multiple sites in the Clare-Bishop area. Then, in area 17, anterograde tracer was injected at a retinotopically-characterized site, giving rise to patches of labeled terminals in the Clare-Bishop area. Receptive field centers recorded within these patches were located close to the visual field location at the injection site in area 17. Receptive fields recorded outside of labeled patches, on the other hand, were never in register with that plotted in area 17. However, due to their large size, even fields located outside of labeled patches often encompassed the visual field point injected in area 17. In other experiments, receptive fields for both neurons and presumed cortical afferents were recorded at the same site in the Clare-Bishop area. The centers of such pairs of receptive fields were on average less than 1 degree apart. Finally, the gaps between widely separated patches of label were investigated. Both physiological and anatomical evidence indicated that a different part of the visual field was represented in gaps than in the adjacent patches.

Afferent Pathways↗
PubMed1986

Coincidence of patchy inputs from the lateral geniculate complex and area 17 to the cat's Clare-Bishop area.

Pathways from a variety of structures to the largest of the cat's suprasylvian visual areas, the Clare-Bishop area, were found to patchy. These inputs arose from the lateral geniculate complex, from area 18, from area 19, and, as noted by Montero (Brain Behav. Evol. 18:194-218, '81), from area 17. The Clare-Bishop area was previously delineated on the basis of its uniform pattern of connections with cortex and thalamus (Sherk: J. Comp. Neurol. 247:1-31, '86) and found to incorporate pieces of several retinotopically defined areas (Tusa, Palmer, Rosenquist: Cortical Sensory Organization. Vol 2. Multiple Visual Areas. Clifton, NJ: Humana Press, pp. 1-31, '81). However, since individual patches did not correspond to particular retinotopically defined areas, other explanations of afferent patchiness were sought. An obvious question is whether the patches originating from different sources are systematically related to each other. Two hypotheses were considered. First, different inputs--for example, from the lateral geniculate nucleus (LGN) and from area 17--might terminate in intermingled but mutually exclusive zones in the Clare-Bishop area. Second, multiple patches of input might reflect duplicate representations of the corresponding visual field segment in the Clare-Bishop area. Both hypotheses were tested by injecting the lateral geniculate complex and either area 17 or area 19 with different anterograde tracers. In each case the two injections involved regions of the visual field that coincided to some degree, ranging from near-total overlap to almost complete exclusion. The first hypothesis predicted that the different labels in the Clare-Bishop area would never be found to overlap, while the second hypothesis predicted that when injections were closely matched retinotopically, there would be extensive overlap between patches. The results supported the second hypothesis: the better the retinotopic match between injections, the greater the overlap found between labeled geniculate and cortical input in the Clare-Bishop area. However, the multiplicity of patches seen in some experiments, and the close spacing between some patches, suggested that an additional, nonretinotopic mechanism also contributes to patchiness in the projections to the Clare-Bishop area.

Animals↗