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Matteo Carandini

Publications and source records attributed to Matteo Carandini.

8 recordsLinked to original sources

The contribution of spike threshold to the dichotomy of cortical simple and complex cells.

The existence of two classes of cells, simple and complex, discovered by Hubel and Wiesel in 1962, is one of the fundamental features of cat primary visual cortex. A quantitative measure used to distinguish simple and complex cells is the ratio between modulated and unmodulated components of spike responses to drifting gratings, an index that forms a bimodal distribution. We have found that the modulation ratio, when derived from the subthreshold membrane potential instead of from spike rate, is unimodally distributed, but highly skewed. The distribution of the modulation ratio as derived from spike rate can, in turn, be predicted quantitatively by the nonlinear properties of spike threshold applied to the skewed distribution of the subthreshold modulation ratio. Threshold also increases the spatial segregation of ON and OFF regions of the receptive field, a defining attribute of simple cells. The distinction between simple and complex cells is therefore enhanced by threshold, much like the selectivity for stimulus features such as orientation and direction. In this case, however, a continuous distribution in the spatial organization of synaptic inputs is transformed into two distinct classes of cells.

Action Potentials↗

Amplification of trial-to-trial response variability by neurons in visual cortex.

The visual cortex responds to repeated presentations of the same stimulus with high variability. Because the firing mechanism is remarkably noiseless, the source of this variability is thought to lie in the membrane potential fluctuations that result from summated synaptic input. Here this hypothesis is tested through measurements of membrane potential during visual stimulation. Surprisingly, trial-to-trial variability of membrane potential is found to be low. The ratio of variance to mean is much lower for membrane potential than for firing rate. The high variability of firing rate is explained by the threshold present in the function that converts inputs into firing rates. Given an input with small, constant noise, this function produces a firing rate with a large variance that grows with the mean. This model is validated on responses recorded both intracellularly and extracellularly. In neurons of visual cortex, thus, a simple deterministic mechanism amplifies the low variability of summated synaptic inputs into the large variability of firing rate. The computational advantages provided by this amplification are not known.

Action Potentials↗

Contrast invariance of functional maps in cat primary visual cortex.

Neurons in cat primary visual cortex (V1) are clustered according to their preference for stimulus position, orientation, spatial frequency, and eye of presentation, thereby giving rise to functional maps. It is not known, however, whether a similar arrangement is present for stimulus contrast. Neurons in cat V1 vary considerably in their contrast responses, and might be clustered in a systematic fashion in this respect. Additionally, stimulus contrast might affect other functional maps. For example, there has been debate over whether the contrast threshold of neurons in cytochrome oxidase blobs is lower than elsewhere. Here we have imaged intrinsic signals to measure orientation maps in cat V1 at a range of contrast levels. We determined, on a pixel-by-pixel basis, contrast-response functions and orientation tuning curves. The fit parameters describing contrast responses were more or less uniform: We found no regions where neurons have lower contrast threshold than elsewhere. If such regions do exist, their functional maps must be substantially weaker than maps of orientation preference. Moreover, we found that contrast has no impact on maps of orientation preference: The orientation selectivity of each pixel is invariant with stimulus contrast. The contrast invariance that we demonstrate at the level of maps is well known at the level of single neurons. It suggests that neurons contributing to a pixel response generally have similar orientation preferences or similar contrast responses. The latter explanation is likely to hold in pinwheel centers, where preferred orientation of nearby neurons can differ markedly. In summary, our data suggest that contrast is represented uniformly over the surface of cat V1, and changes in contrast do not affect maps of orientation preference.

Animals↗

A synaptic explanation of suppression in visual cortex.

The responses of neurons in the primary visual cortex (V1) are suppressed by mask stimuli that do not elicit responses if presented alone. This suppression is widely believed to be mediated by intracortical inhibition. As an alternative, we propose that it can be explained by thalamocortical synaptic depression. This explanation correctly predicts that suppression is monocular, immune to cortical adaptation, and occurs for mask stimuli that elicit responses in the thalamus but not in the cortex. Depression also explains other phenomena previously ascribed to intracortical inhibition. It explains why responses saturate at high stimulus contrast, whereas selectivity for orientation and spatial frequency is invariant with contrast. It explains why transient responses to flashed stimuli are nonlinear, whereas spatial summation is primarily linear. These results suggest that the very first synapses into the cortex, and not the cortical network, may account for important response properties of V1 neurons.

Action Potentials↗

Suppression without inhibition in visual cortex.

Neurons in primary visual cortex (V1) are thought to receive inhibition from other V1 neurons selective for a variety of orientations. Evidence for this inhibition is commonly found in cross-orientation suppression: responses of a V1 neuron to optimally oriented bars are suppressed by superimposed mask bars of different orientation. We show, however, that suppression is unlikely to result from intracortical inhibition. First, suppression can be obtained with masks drifting too rapidly to elicit much of a response in cortex. Second, suppression is immune to hyperpolarization (through visual adaptation) of cortical neurons responding to the mask. Signals mediating suppression might originate in thalamus, rather than in cortex. Thalamic neurons exhibit some suppression; additional suppression might arise from depression at thalamocortical synapses. The mechanisms of suppression are subcortical and possibly include the very first synapse into cortex.

Action Potentials↗

Testing the Bayesian model of perceived speed.

In a recent Bayesian model by Weiss, Simoncelli, and Adelson, motion perception is biased by a prior favoring slow speeds. This model predicts qualitatively an impressive variety of phenomena, including the dependence of perceived speed on contrast. We show that the model can also generate quantitative predictions: for a drifting grating with contrast c, perceived speed is proportional to c(q)/(k(q) + c(q)), with k, q constants. We tested this expression on measurements of perceived speed as a function of contrast. Observers indicated the slower of two drifting gratings, a test and a standard. For each test contrast we found the test speed that appeared to match the standard speed. The model fits the data, but only if q is less than 2, the value it would have if the internal representation of contrast were linear. The Bayesian model can make correct quantitative predictions, but needs to be extended to incorporate a more realistic, nonlinear representation of contrast.

Bayes Theorem↗

Masking by fast gratings.

Perception of an oriented pattern is impaired in the presence of a superimposed orthogonal mask. This masking effect most likely arises in visual cortex, where neuronal responses are suppressed by masks having a broad range of orientations. Response suppression is commonly ascribed to lateral inhibition between cortical neurons. Recent physiological results, however, have cast doubt on this view: powerful suppression has been observed with masks drifting too rapidly to elicit much of a response in cortex. We show here that the same is true for perceptual masking. From contrast discrimination thresholds, we estimated the cortical response to drifting patterns of various frequencies, and found it greatly reduced above 15-20 Hz. In the same subjects, we measured the strength of masking by the same patterns and found it equally strong for masks drifting slowly (2.7 Hz) as for masks drifting rapidly (27-38 Hz). Fast gratings thus cause strong masking while eliciting weak cortical responses. Our results might be explained by inhibition from cortical neurons that respond to unusually high frequencies, and yet do not make their signals fully available for perceptual judgments. A more parsimonious explanation, however, is that masking does not involve lateral inhibition from cortex. Masking might operate in retina or thalamus, which respond to much higher frequencies than cortex. Masking might also be due to thalamic signals to cortex, perhaps through depression at thalamocortical synapses.

Contrast Sensitivity↗