Modeling Invariances in Inferotemporal Cell Tuning

In macaque inferotemporal cortex (IT), neurons have been found to respond selectively to complex shapes while showing broad tuning ("invariance") with respect to stimulus transformations such as translation and scale changes and a limited tuning to rotation in depth. Training monkeys with novel, paperclip-like objects, Logothetis et al. could investigate whether these invariance properties are due to experience with exhaustively many transformed instances of an object or if there are mechanisms that allow the cells to show response invariance also to previously unseen instances of that object. They found object-selective cells in anterior IT which exhibited limited invariance to various transformations after training with single object views. While previous models accounted for the tuning of the cells for rotations in depth and for their selectivity to a specific object relative to a population of distractor objects, the model described here attempts to explain in a biologically plausible way the additional properties of translation and size invariance. Using the same stimuli as in the experiment, we find that model IT neurons exhibit invariance properties which closely parallel those of real neurons. Simulations show that the model is capable of unsupervised learning of view-tuned neurons. The model also allows to make experimentally testable predictions regarding novel stimulus transformations and combinations of stimuli.

A Detailed Look at Scale and Translation Invariance in a Hierarchical Neural Model of Visual Object Recognition

The HMAX model has recently been proposed by Riesenhuber & Poggio as a hierarchical model of position- and size-invariant object recognition in visual cortex. It has also turned out to model successfully a number of other properties of the ventral visual stream (the visual pathway thought to be crucial for object recognition in cortex), and particularly of (view-tuned) neurons in macaque inferotemporal cortex, the brain area at the top of the ventral stream. The original modeling study only used ``paperclip'' stimuli, as in the corresponding physiology experiment, and did not explore systematically how model units' invariance properties depended on model parameters. In this study, we aimed at a deeper understanding of the inner workings of HMAX and its performance for various parameter settings and ``natural'' stimulus classes. We examined HMAX responses for different stimulus sizes and positions systematically and found a dependence of model units' responses on stimulus position for which a quantitative description is offered. Interestingly, we find that scale invariance properties of hierarchical neural models are not independent of stimulus class, as opposed to translation invariance, even though both are affine transformations within the image plane.

Generalization over contrast and mirror reversal, but not figure-ground reversal, in an "edge-based

Baylis & Driver (Nature Neuroscience, 2001) have recently presented data on the response of neurons in macaque inferotemporal cortex (IT) to various stimulus transformations. They report that neurons can generalize over contrast and mirror reversal, but not over figure-ground reversal. This finding is taken to demonstrate that ``the selectivity of IT neurons is not determined simply by the distinctive contours in a display, contrary to simple edge-based models of shape recognition'', citing our recently presented model of object recognition in cortex (Riesenhuber & Poggio, Nature Neuroscience, 1999). In this memo, I show that the main effects of the experiment can be obtained by performing the appropriate simulations in our simple feedforward model. This suggests for IT cell tuning that the possible contributions of explicit edge assignment processes postulated in (Baylis & Driver, 2001) might be smaller than expected.

Stimulus Simplification and Object Representation: A Modeling Study

Tsunoda et al. (2001) recently studied the nature of object representation in monkey inferotemporal cortex using a combination of optical imaging and extracellular recordings. In particular, they examined IT neuron responses to complex natural objects and "simplified" versions thereof. In that study, in 42% of the cases, optical imaging revealed a decrease in the number of activation patches in IT as stimuli were "simplified". However, in 58% of the cases, "simplification" of the stimuli actually led to the appearance of additional activation patches in IT. Based on these results, the authors propose a scheme in which an object is represented by combinations of active and inactive columns coding for individual features. We examine the patterns of activation caused by the same stimuli as used by Tsunoda et al. in our model of object recognition in cortex (Riesenhuber 99). We find that object-tuned units can show a pattern of appearance and disappearance of features identical to the experiment. Thus, the data of Tsunoda et al. appear to be in quantitative agreement with a simple object-based representation in which an object's identity is coded by its similarities to reference objects. Moreover, the agreement of simulations and experiment suggests that the simplification procedure used by Tsunoda (2001) is not necessarily an accurate method to determine neuronal tuning.

Pre-Attentive Segmentation in the Primary Visual Cortex

Stimuli outside classical receptive fields have been shown to exert significant influence over the activities of neurons in primary visual cortexWe propose that contextual influences are used for pre-attentive visual segmentation, in a new framework called segmentation without classification. This means that segmentation of an image into regions occurs without classification of features within a region or comparison of features between regions. This segmentation framework is simpler than previous computational approaches, making it implementable by V1 mechanisms, though higher leve l visual mechanisms are needed to refine its output. However, it easily handles a class of segmentation problems that are tricky in conventional methods. The cortex computes global region boundaries by detecting the breakdown of homogeneity or translation invariance in the input, using local intra-cortical interactions mediated by the horizontal connections. The difference between contextual influences near and far from region boundaries makes neural activities near region boundaries higher than elsewhere, making boundaries more salient for perceptual pop-out. This proposal is implemented in a biologically based model of V1, and demonstrated using examples of texture segmentation and figure-ground segregation. The model performs segmentation in exactly the same neural circuit that solves the dual problem of the enhancement of contours, as is suggested by experimental observations. Its behavior is compared with psychophysical and physiological data on segmentation, contour enhancement, and contextual influences. We discuss the implications of segmentation without classification and the predictions of our V1 model, and relate it to other phenomena such as asymmetry in visual search.

Vector-Based Integration of Local and Long-Range Information in Visual Cortex

Integration of inputs by cortical neurons provides the basis for the complex information processing performed in the cerebral cortex. Here, we propose a new analytic framework for understanding integration within cortical neuronal receptive fields. Based on the synaptic organization of cortex, we argue that neuronal integration is a systems--level process better studied in terms of local cortical circuitry than at the level of single neurons, and we present a method for constructing self-contained modules which capture (nonlinear) local circuit interactions. In this framework, receptive field elements naturally have dual (rather than the traditional unitary influence since they drive both excitatory and inhibitory cortical neurons. This vector-based analysis, in contrast to scalarsapproaches, greatly simplifies integration by permitting linear summation of inputs from both "classical" and "extraclassical" receptive field regions. We illustrate this by explaining two complex visual cortical phenomena, which are incompatible with scalar notions of neuronal integration.

Computational Models of Object Recognition in Cortex: A Review

Understanding how biological visual systems perform object recognition is one of the ultimate goals in computational neuroscience. Among the biological models of recognition the main distinctions are between feedforward and feedback and between object-centered and view-centered. From a computational viewpoint the different recognition tasks - for instance categorization and identification - are very similar, representing different trade-offs between specificity and invariance. Thus the different tasks do not strictly require different classes of models. The focus of the review is on feedforward, view-based models that are supported by psychophysical and physiological data.

Visual Segmentation without Classification in a Model of the Primary Visual Cortex

Stimuli outside classical receptive fields significantly influence the neurons' activities in primary visual cortex. We propose that such contextual influences are used to segment regions by detecting the breakdown of homogeneity or translation invariance in the input, thus computing global region boundaries using local interactions. This is implemented in a biologically based model of V1, and demonstrated in examples of texture segmentation and figure-ground segregation. By contrast with traditional approaches, segmentation occurs without classification or comparison of features within or between regions and is performed by exactly the same neural circuit responsible for the dual problem of the grouping and enhancement of contours.

Categorization in IT and PFC: Model and Experiments

In a recent experiment, Freedman et al. recorded from inferotemporal (IT) and prefrontal cortices (PFC) of monkeys performing a "cat/dog" categorization task (Freedman 2001 and Freedman, Riesenhuber, Poggio, Miller 2001). In this paper we analyze the tuning properties of view-tuned units in our HMAX model of object recognition in cortex (Riesenhuber 1999) using the same paradigm and stimuli as in the experiment. We then compare the simulation results to the monkey inferotemporal neuron population data. We find that view-tuned model IT units that were trained without any explicit category information can show category-related tuning as observed in the experiment. This suggests that the tuning properties of experimental IT neurons might primarily be shaped by bottom-up stimulus-space statistics, with little influence of top-down task-specific information. The population of experimental PFC neurons, on the other hand, shows tuning properties that cannot be explained just by stimulus tuning. These analyses are compatible with a model of object recognition in cortex (Riesenhuber 2000) in which a population of shape-tuned neurons provides a general basis for neurons tuned to different recognition tasks.