Area V4—the focus of this review—is a mid-level processing stage along the ventral visual pathway of the macaque monkey. V4 is extensively interconnected with other visual cortical areas along the ventral and dorsal visual streams, with frontal cortical areas, and with several subcortical structures. Thus, it is well poised to play a broad and integrative role in visual perception and recognition—the functional domain of the ventral pathway. Neurophysiological studies in monkeys engaged in passive fixation and behavioral tasks suggest that V4 responses are dictated by tuning in a high-dimensional stimulus space defined by form, texture, color, depth, and other attributes of visual stimuli. This high-dimensional tuning may underlie the development of object-based representations in the visual cortex that are critical for tracking, recognizing, and interacting with objects. Neurophysiological and lesion studies also suggest that V4 responses are important for guiding perceptual decisions and higher-order behavior. Expected final online publication date for the Annual Review of Vision Science, Volume 6 is September 15, 2020. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
Divided attention has little effect for simple tasks, such as luminance detection, but it has large effects for complex tasks, such as semantic categorization of masked words. Here, we asked whether the semantic categorization of visual objects shows divided attention effects as large as those observed for words, or as small as those observed for simple feature judgments. Using a dual-task paradigm with nameable object stimuli, performance was compared with the predictions of serial and parallel models. At the extreme, parallel processes with unlimited capacity predict no effect of divided attention; alternatively, an all-or-none serial process makes two predictions: a large divided attention effect (lower accuracy for dual-task trials, compared to single-task trials) and a negative response correlation in dual-task trials (a given response is more likely to be incorrect when the response about the other stimulus is correct). These predictions were tested in two experiments examining object judgments. In both experiments, there was a large divided attention effect and a small negative correlation in responses. The magnitude of these effects was larger than for simple features, but smaller than for words. These effects were consistent with serial models, and rule out some but not all parallel models. More broadly, the results help establish one of the first examples of likely serial processing in perception.
Visual area V4 is an important midlevel cortical processing stage that subserves object recognition in primates. Studies investigating shape coding in V4 have largely probed neuronal responses with filled shapes, i.e., shapes defined by both a boundary and an interior fill. As a result, we do not know whether form-selective V4 responses are dictated by boundary features alone or if interior fill is also important. We studied 43 V4 neurons in two male macaque monkeys ( Macaca mulatta) with a set of 362 filled shapes and their corresponding outlines to determine how interior fill modulates neuronal responses in shape-selective neurons. Only a minority of neurons exhibited similar response strength and shape preferences for filled and outline stimuli. A majority responded preferentially to one stimulus category (either filled or outline shapes) and poorly to the other. Our findings are inconsistent with predictions of the hierarchical-max (HMax) V4 model that builds form selectivity from oriented boundary features and takes little account of attributes related to object surface, such as the phase of the boundary edge. We modified the V4 HMax model to include sensitivity to interior fill by either removing phase-pooling or introducing unoriented units at the V1 level; both modifications better explained our data without increasing the number of free parameters. Overall, our results suggest that boundary orientation and interior surface information are both maintained until at least the midlevel visual representation, consistent with the idea that object fill is important for recognition and perception in natural vision. NEW & NOTEWORTHY The shape of an object’s boundary is critical for identification; consistent with this idea, models of object recognition predict that filled and outline versions of a shape are encoded similarly. We report that many neurons in a midlevel visual cortical area respond differently to filled and outline shapes and modify a biologically plausible model to account for our data. Our results suggest that representations of boundary shape and surface fill are interrelated in visual cortex.
Dendritic spines form the postsynaptic half of the synapse but how they form during CNS development remains uncertain, as are the factors that promote their morphological and physiological maturation. One hypothesis posits that filopodia, long motile dendritic processes that are present prior to spine formation, are the precursors to spines. Another hypothesis posits that they form directly from the dendritic shaft. We used microphotolysis of caged glutamate to stimulate individual dendritic processes in young hippocampal slice cultures while recording their morphological and physiological responses. We observed that brief trains of stimuli delivered to immature processes triggered morphological changes within minutes that resulted in a more mature, spine-like appearance in about half of experiments, such as decreases in spine neck length, increases in spine head width. We also observed that glutamate-induced inward currents elicited from immature processes were mostly or entirely mediated by NMDARs, whereas responses in those processes with a more mature morphology, regardless of actual developmental age, were mediated by both AMPARs and NMDARs. Consistent with this observation, glutamate-induced morphological changes were largely, but not entirely, prevented by blocking NMDARs. Our observations thus favor a model in which filopodia in the developing nervous system sense and respond to release of glutamate from developing axons, resulting in physiological and morphological maturation.
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