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Judith A Hirsch

Publications and source records attributed to Judith A Hirsch.

7 recordsLinked to original sources

Laminar processing in the visual cortical column.

Sensory regions of neocortex are organized as arrays of vertical columns composed of cells that share similar response properties, with the orientation columns of the cat's visual cortex being the best known example. Interest in how sensitivity to different stimulus features first emerges in the columns and how this selectivity is refined by subsequent processing has fueled decades of research. A natural starting point in approaching these issues is anatomy. Each column traverses the six cortical layers and each layer has a unique pattern of inputs, intrinsic connections and outputs. Thus, it makes sense to explore the possibility of corresponding laminar differences in sensory function, that is, to examine relationships between morphology and physiology. In addition, to help identify general patterns of cortical organization, it is useful to compare results obtained from different sensory systems and diverse species. The picture that emerges from such comparisons is that each cortical layer serves a distinct role in sensory function. Furthermore, different cortices appear to share some common strategies for processing information but also have specialized mechanisms adapted for the demands of specific sensory tasks.

Animals↗

Circuits that build visual cortical receptive fields.

Neural sensitivity to basic elements of the visual scene changes dramatically as information is handed from the thalamus to the primary visual cortex in cats. Famously, thalamic neurons are insensitive to stimulus orientation whereas their cortical targets easily resolve small changes in stimulus angle. There are two main types of cells in the visual cortex, simple and complex, defined by the structure of their receptive fields. Simple cells are thought to lay the groundwork for orientation selectivity. This review focuses on approaches that combine anatomy with physiology at the intracellular level, to explore the circuits that build simple receptive fields and that help to maintain neural sensitivity to stimulus features even when luminance contrast changes.

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Receptive field structure varies with layer in the primary visual cortex.

Here we ask whether visual response pattern varies with position in the cortical microcircuit by comparing the structure of receptive fields recorded from the different layers of the cat's primary visual cortex. We used whole-cell recording in vivo to show the spatial distribution of visually evoked excitatory and inhibitory inputs and to stain individual neurons. We quantified the distribution of 'On' and 'Off' responses and the presence of spatially opponent excitation and inhibition within the receptive field. The thalamorecipient layers (4 and upper 6) were dominated by simple cells, as defined by two criteria: they had separated On and Off subregions, and they had push-pull responses (in a given subregion, stimuli of the opposite contrast evoked responses of the opposite sign). Other types of response profile correlated with laminar location as well. Thus, connections unique to each visual cortical layer are likely to serve distinct functions.

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Functionally distinct inhibitory neurons at the first stage of visual cortical processing.

Here we explore inhibitory circuits at the thalamocortical stage of processing in layer 4 of the cat's visual cortex, focusing on the anatomy and physiology of the interneurons themselves. Our immediate aim was to explore the inhibitory mechanisms that contribute to orientation selectivity, perhaps the most dramatic response property to emerge across the thalamocortical synapse. The broader goal was to understand how inhibitory circuits operate. Using whole-cell recording in cats in vivo, we found that layer 4 contains two populations of inhibitory cells defined by receptive field class--simple and complex. The simple cells were selective for stimulus orientation, whereas the complex cells were not. Our observations help to explain how neurons become sensitive to stimulus orientation and maintain that selectivity as stimulus contrast changes. Overall, the work suggests that different sources of inhibition, either selective for specific features or broadly tuned, interact to provide appropriate representations of elements within the environment.

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Synaptic physiology and receptive field structure in the early visual pathway of the cat.

How does the cortical circuitry analyze the visual scene? Here we explore the earliest levels of striate cortical processing: the first stage, where orientation sensitivity emerges, and the second stage, where stimulus selectivity is further refined. The approach is whole cell recording from cat in vivo. Neurons in the lateral geniculate nucleus of the thalamus have circular receptive fields whose subregions, center and surround are concentrically arranged and have the reverse sign, on or off. These neurons supply cortical simple cells, whose receptive fields have on and off subregions that are elongated and lie side by side. Feedforward models hold that orientation sensitivity depends on this thalamocortical change in receptive field structure and an arrangement within subregions such that stimuli of the reverse contrast evoke synaptic responses of the opposite polarity-push-pull. Our work provides support for feedforward models and emphasizes that push-pull is key in the geniculostriate pathway, preserved from retina by thalamic relay cells and reiterated, point by point, by cortical simple cells. Also, we help define the cortical push-pull circuit by identifying inhibitory simple cells. Lastly, separate experiments that compare the first and second levels of cortical processing suggest that differences in the synaptic physiology of connections at the two (thalamocortical versus intracortical) stages underlie differential selectivity for properties such as motion.

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Laminar processing of stimulus orientation in cat visual cortex.

One of the most salient features to emerge in visual cortex is sensitivity to stimulus orientation. Here we asked if orientation selectivity, once established, is altered by successive stages of cortical processing. We measured patterns of orientation selectivity at all depths of the cat's visual cortex by making whole-cell recordings with dye-filled electrodes. Our results show that the synaptic representation of orientation indeed changes with position in the microcircuit, as information passes from layer 4 to layer 2+3 to layer 5. At the earliest cortical stage, for simple cells in layer 4, orientation tuning curves for excitation (depolarization) and inhibition (hyperpolarization) had similar peaks (within 0-7 deg, n = 11) and bandwidths. Further, the sharpness of orientation selectivity covaried with receptive field geometry (r = 0.74) - the more elongated the strongest subregion, the shaper the tuning. Tuning curves for complex cells in layer 2+3 also had similar peaks (within 0-4 deg, n = 7) and bandwidths. By contrast, at a later station, layer 5, the preferred orientation for excitation and inhibition diverged such that the peaks of the tuning curves could be as far as 90 deg apart (average separation, 54 deg; n = 6). Our results support the growing consensus that orientation selectivity is generated at the earliest cortical level and structured similarly for excitation and inhibition. Moreover, our novel finding that the relative tuning of excitation and inhibition changes with laminar position helps resolve prior controversy about orientation selectivity at later phases of processing and gives a mechanistic view of how the cortical circuitry recodes orientation.

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Synaptic physiology of the flow of information in the cat's visual cortex in vivo.

Each stage of the striate cortical circuit extracts novel information about the visual environment. We asked if this analytic process reflected laminar variations in synaptic physiology by making whole-cell recording with dye-filled electrodes from the cat's visual cortex and thalamus; the stimuli were flashed spots. Thalamic afferents terminate in layer 4, which contains two types of cell, simple and complex, distinguished by the spatial structure of the receptive field. Previously, we had found that the postsynaptic and spike responses of simple cells reliably followed the time course of flash-evoked thalamic activity. Here we report that complex cells in layer 4 (or cells intermediate between simple and complex) similarly reprised thalamic activity (response/trial, 99 +/- 1.9 %; response duration 159 +/- 57 ms; latency 25 +/- 4 ms; average +/- standard deviation; n = 7). Thus, all cells in layer 4 share a common synaptic physiology that allows secure integration of thalamic input. By contrast, at the second cortical stage (layer 2+3), where layer 4 directs its output, postsynaptic responses did not track simple patterns of antecedent activity. Typical responses to the static stimulus were intermittent and brief (response/trial, 31 +/- 40 %; response duration 72 +/- 60 ms, latency 39 +/- 7 ms; n = 11). Only richer stimuli like those including motion evoked reliable responses. All told, the second level of cortical processing differs markedly from the first. At that later stage, ascending information seems strongly gated by connections between cortical neurons. Inputs must be combined in newly specified patterns to influence intracortical stages of processing.

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