Neurons in area V4 have relatively large receptive fields (RFs), so multiple visual features are simultaneously "seen" by these cells. Recordings from single V 4 neurons suggest that simultaneously presented stimuli compete to set the output firing rate, and that attention acts to isolate individual features by biasing the competition in favor of the attended object. We propose that both stimulus competition and attentional biasing arise from the spatial segregation of afferent synapses onto different regions of the excitable dendritic tree of V 4 neurons. The pattern of feedforward, stimulus-driven inputs follows from a Hebbian rule: excitatory afferents with similar RFs tend to group together on the dendritic tree, avoiding randomly located inhibitory inputs with similar RFs. The same principle guides the formation of inputs that mediate attentional modulation.

Neurons in area V4 have relatively large receptive fields (RFs), so multiple visualfeatures are simultaneously "seen" by these cells. Recordings from single V4 neurons suggest that simultaneously presented stimuli compete to set the output firing rate, and that attention acts to isolate individual features by biasing the competition in favor of the attended object. We propose that both stimulus competition and attentional biasing arisefrom the spatial segregation of afferent synapses onto different regions of the excitable dendritic tree of V4 neurons. The pattern of feedforward, stimulus-driveninputs follows from a Hebbian rule: excitatory afferents with similar RFs tend to group together on the dendritic tree, avoiding randomly located inhibitory inputs with similar RFs. The same principle guides the formation of inputs that mediate attentional modulation.

Previous biophysical modeling work showed that nonlinear interactions amongnearby synapses located on active dendritic trees can provide a large boost in the memory capacity of a cell (Mel, 1992a, 1992b).

Previous biophysical modeling work showed that nonlinear interactions among nearby synapses located on active dendritic trees can provide a large boost in the memory capacity of a cell (Mel, 1992a, 1992b). The aim of our present work is to quantify this boost by estimating the capacity of (1) a neuron model with passive dendritic integration where inputs are combined linearly across the entire cell followed by a single global threshold, and (2) an active dendrite model in which a threshold is applied separately to the output of each branch, and the branch subtotals are combined linearly. We focus here on the limiting case of binary-valued synaptic weights, and derive expressions which measure model capacity by estimating the number of distinct input-output functions available to both neuron types. We show that (1) the application of a fixed nonlinearity to each dendritic compartment substantially increases the model's flexibility, (2) for a neuron of realistic size, the capacity of the nonlinear cell can exceed that of the same-sized linear cell by more than an order of magnitude, and (3) the largest capacity boost occurs for cells with a relatively large number of dendritic subunits of relatively small size. We validated the analysis by empirically measuring memory capacity with randomized two-class classification problems, where a stochastic delta rule was used to train both linear and nonlinear models. We found that large capacity boosts predicted for the nonlinear dendritic model were readily achieved in practice.

To understand human object recognition, it is essential to understand how objects are represented in human visual memory. A central component in object recognition is the matching of the stored object representation with that derived from the image input.But the nature of the object representation has to be inferred from recognition performance, by taking into account the contribution from the image information. When evaluating human performance, how can one separate the con- 830 ZLiu and D. Kersten tributions to performance of the image information from the representation? Ideal observer analysis provides a precise computational tool to answer this question. An ideal observer's recognition performance is restricted only by the available image information and is otherwise optimal, in the sense of statistical decision theory, irrespective of how the model is implemented.

Further, the greater the similarity between objects, the stronger is the dependence on object appearance, and the more important twodimensional (2D) image information becomes. These findings, however, do not rule out the use of 3D structural information in recognition, and the degree to which 3D information is used in visual memory is an important issue. Liu, Knill, & Kersten (1995) showed that any model that is restricted to rotations in the image plane of independent 2D templates could not account for human performance in discriminating novel object views. We now present results from models of generalized radial basis functions (GRBF), 2D nearest neighbor matching that allows 2D affine transformations, and a Bayesian statistical estimator that integrates over all possible 2D affine transformations. The performance of the human observers relative to each of the models is better for the novel views than for the familiar template views, suggesting that humans generalize better to novel views from template views. The Bayesian estimator yields the optimal performance with 2D affine transformations and independent 2D templates. Therefore, models of 2D affine matching operations with independent 2D templates are unlikely to account for human recognition performance.

Biophysical modeling studies have previously shown that cortical pyramidal cells driven by strong NMDA-type synaptic currents and/or containing dendritic voltage-dependent Ca or Na channels, respond more strongly when synapses are activated in several spatially clustered groups of optimal size-in comparison to the same number of synapses activated diffusely about the dendritic arbor [8]- The nonlinear intradendritic interactions giving rise to this "cluster sensitivity" property are akin to a layer of virtual nonlinear "hidden units" in the dendrites, with implications for the cellular basis of learning and memory [7, 6], and for certain classes of nonlinear sensory processing [8]- In the present study, we show that a single neuron, with access only to excitatory inputs from unoriented ONand OFFcenter cells in the LGN, exhibits the principal nonlinear response properties of a "complex" cell in primary visual cortex, namely orientation tuning coupled with translation invariance and contrast insensitivity_ We conjecture that this type of intradendritic processing could explain how complex cell responses can persist in the absence of oriented simple cell input [13]- 84 B. W. Mel, D. L. Ruderman and K. A. Archie

Biophysical modeling studies have previously shown that cortical pyramidal cells driven by strong NMDA-type synaptic currents and/or containing dendritic voltage-dependent Ca or Na channels, respondmore strongly when synapses are activated in several spatially clustered groups of optimal size-in comparison to the same number of synapses activated diffusely about the dendritic arbor [8]- The nonlinear intradendritic interactions giving rise to this "cluster sensitivity" property are akin to a layer of virtual nonlinear "hiddenunits" in the dendrites, with implications for the cellular basis of learning and memory [7, 6], and for certain classes of nonlinear sensory processing [8]- In the present study, we show that a single neuron, with access only to excitatory inputs from unoriented ONand OFFcenter cells in the LGN, exhibits the principal nonlinear response properties of a "complex" cell in primary visual cortex, namely orientation tuning coupled with translation invariance andcontrast insensitivity_ We conjecture that this type of intradendritic processing could explain how complex cell responses can persist in the absence of oriented simple cell input [13]- 84 B. W. Mel, D. L. Ruderman and K. A. Archie

Biophysical modeling studies have previously shown that cortical pyramidal cells driven by strong NMDA-type synaptic currents and/or containing dendritic voltage-dependent Ca or Na channels, respond more strongly when synapses are activated in several spatially clustered groups of optimal size-in comparison to the same number of synapses activated diffusely about the dendritic arbor [8]- The nonlinear intradendritic interactions giving rise to this "cluster sensitivity" property are akin to a layer of virtual nonlinear "hidden units" in the dendrites, with implications for the cellular basis of learning and memory [7, 6], and for certain classes of nonlinear sensory processing [8]- In the present study, we show that a single neuron, with access only to excitatory inputs from unoriented ONand OFFcenter cells in the LGN, exhibits the principal nonlinear response properties of a "complex" cell in primary visual cortex, namely orientation tuning coupled with translation invariance and contrast insensitivity_ We conjecture that this type of intradendritic processing could explain how complex cell responses can persist in the absence of oriented simple cell input [13]- 84 B. W. Mel, D. L. Ruderman and K. A. Archie

In natural contexts, visual object recognition in humans is remarkably fast, reliable, and viewpoint invariant. The present approach to object recognition is "view-based" (e.g.