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Perceptual Coherence : Hearing and Seeing

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Characteristics of Auditory <strong>and</strong> Visual Scenes 135<br />

nervous system is organized. These outputs, when combined, would optimally<br />

reproduce the original image.<br />

Hyvärinen <strong>and</strong> Hoyer (2001) followed this strategy to derive the receptive<br />

spatial fields <strong>and</strong> spatial arrangement of the complex cells found in the<br />

primary visual cortex V1. Simple cells will increase their firing rate above<br />

baseline if a white b<strong>and</strong> falls on the excitation region <strong>and</strong> a black b<strong>and</strong> falls<br />

on the inhibition region but reduce their firing rate below baseline for the reverse<br />

arrangement. Complex cells, in contrast, will increase their firing rate<br />

for either arrangement; they are phase insensitive. Intuitively, the simple<br />

cells feed forward to the complex cells such that either an increase or decrease<br />

in firing rate of simple cells increases the firing rate of the complex<br />

cell. Hyvärinen <strong>and</strong> Hoyer (2001) modeled this process by assuming that (1)<br />

the outputs of the simple cells (the difference between the firing rate <strong>and</strong> the<br />

baseline-firing rate) are squared so that all changes in the light pattern due to<br />

movements of edges increase the firing rate, <strong>and</strong> (2) the outputs from nearby<br />

simple cells (a 5 × 5 grid of cells) converge on every complex cell. They<br />

then use independent component analysis to maximize the sparseness or independence<br />

of the complex cell responses. To maximize the sparseness of<br />

the complex cells means that the simple cells that converge on a complex<br />

cell should have as similar a filter response as possible. The authors argued<br />

that if the responses of the convergent simple cells were independent, then<br />

the response of the complex cell would be more normal <strong>and</strong> less peaky (an<br />

outcome predicted from the central limit theorem).<br />

The results can be considered in terms of the simple cells <strong>and</strong> in terms<br />

of how the activations of the simple cells create the complex cells. Each<br />

retinal region is represented by a set of adjacent overlapping simple cells.<br />

The receptive fields of the simple cells in figure 3.11 differ in retinal location<br />

<strong>and</strong> orientation of the brightness contrast, <strong>and</strong> respond to a narrow<br />

range of spatial frequencies. The topographic map of each set of simple<br />

cells shows that the cells are smoothly arranged spatially in terms of orientation<br />

<strong>and</strong> position. The majority of the receptive fields are simply scaled<br />

versions of another field. Thus the receptive fields possess differing spatial<br />

<strong>and</strong> temporal scales that create the multiresolution characteristics of perceptual<br />

systems described in chapter 1. The simple cells that are pooled to<br />

create complex cells have similar receptive fields.<br />

The results of Hyvärinen <strong>and</strong> Hoyer (2001) provide a model to explain<br />

the retinal topographic organization of the visual cortex. Starting with a<br />

simple feed-forward model in which simple cells converge on complex<br />

cells <strong>and</strong> invoking a criterion of maximal independence creates the spatial<br />

arrangement of receptive fields found in all species.<br />

Körding, Käyser, Einhouser, <strong>and</strong> König (2004) followed a different route<br />

to derive the properties of complex cells. They argued that nearly all visual<br />

objects <strong>and</strong> higher-level variables change slowly or not at all over time, even

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