Elastic net model of ocular dominance - overall stripe pattern and monocular deprivation

The elastic net (Durbin and Willshaw 1987) can account for the development of both topography and ocular dominance in the mapping from the lateral geniculate nucleus to primary visual cortex (Goodhill and Willshaw 1990). Here it is further shown for this model that (1) the overall pattern of stripes produced is strongly influenced by the shape of the cortex: in particular, stripes with a global order similar to that seen biologically can be produced under appropriate conditions, and (2) the observed changes in stripe width associated with monocular deprivation are reproduced in the model

Familiarization: A theory of repetition suppression predicts interference between overlapping cortical representations

Repetition suppression refers to a reduction in the cortical response to a novel stimulus that results from repeated presentation of the stimulus. We demonstrate repetition suppression in a well established computational model of cortical plasticity, according to which the relative strengths of lateral inhibitory interactions are modified by Hebbian learning. We present the model as an extension to the traditional account of repetition suppression offered by sharpening theory, which emphasises the contribution of afferent plasticity, by instead attributing the effect primarily to plasticity of intra-cortical circuitry. In support, repetition suppression is shown to emerge in simulations with plasticity enabled only in intra-cortical connections. We show in simulation how an extended ‘inhibitory sharpening theory’ can explain the disruption of repetition suppression reported in studies that include an intermediate phase of exposure to additional novel stimuli composed of features similar to those of the original stimulus. The model suggests a re-interpretation of repetition suppression as a manifestation of the process by which an initially distributed representation of a novel object becomes a more localist representation. Thus, inhibitory sharpening may constitute a more general process by which representation emerges from cortical re-organisation

Familiarization: A theory of repetition suppression predicts interference between overlapping cortical representations

Repetition suppression refers to a reduction in the cortical response to a novel stimulus that results from repeated presentation of the stimulus. We demonstrate repetition suppression in a well established computational model of cortical plasticity, according to which the relative strengths of lateral inhibitory interactions are modified by Hebbian learning. We present the model as an extension to the traditional account of repetition suppression offered by sharpening theory, which emphasises the contribution of afferent plasticity, by instead attributing the effect primarily to plasticity of intra-cortical circuitry. In support, repetition suppression is shown to emerge in simulations with plasticity enabled only in intra-cortical connections. We show in simulation how an extended ‘inhibitory sharpening theory’ can explain the disruption of repetition suppression reported in studies that include an intermediate phase of exposure to additional novel stimuli composed of features similar to those of the original stimulus. The model suggests a re-interpretation of repetition suppression as a manifestation of the process by which an initially distributed representation of a novel object becomes a more localist representation. Thus, inhibitory sharpening may constitute a more general process by which representation emerges from cortical re-organisation

The influence of restricted orientation rearing on map structure in primary visual cortex

Visual experience is critical to the development of the structure of the primary visual cortex and, in turn, normal functional vision. The primary visual cortex contains maps of multiple features of the visual input, and these maps are characterised by specific types of geometric relationships. Manipulations of the visual environment during development in animals such as ferrets, cats and monkeys provide an opportunity to probe the rules governing map formation via their effect on these relationships. Here we use a computational model of map formation based on dimension-reduction principles to predict the effect on map relationships of presenting only a single orientation to one eye and the orthogonal orientation to the other eye. Since orientation preference and ocular dominance are now tightly coupled one might expect orientation and ocular dominance contours to lose their normally orthogonal relationship and instead run parallel to each other. However, surprisingly, the model predicts that orthogonal intersection can sometimes be preserved in this case. The model also predicts that orientation pinwheels can migrate from the centre to the borders of ocular dominance columns, and that the wavelengths of the ocular dominance and orientation maps can become coupled. These predictions provide a way to further test the adequacy of dimension reduction principles for explaining map structure under perturbed as well as normal rearing conditions, and thus allow us to deepen our understanding of the effect of the visual environment on visual cortical development

Coverage, Continuity and Visual Cortical Architecture

The primary visual cortex of many mammals contains a continuous representation of visual space, with a roughly repetitive aperiodic map of orientation preferences superimposed. It was recently found that orientation preference maps (OPMs) obey statistical laws which are apparently invariant among species widely separated in eutherian evolution. Here, we examine whether one of the most prominent models for the optimization of cortical maps, the elastic net (EN) model, can reproduce this common design. The EN model generates representations which optimally trade of stimulus space coverage and map continuity. While this model has been used in numerous studies, no analytical results about the precise layout of the predicted OPMs have been obtained so far. We present a mathematical approach to analytically calculate the cortical representations predicted by the EN model for the joint mapping of stimulus position and orientation. We find that in all previously studied regimes, predicted OPM layouts are perfectly periodic. An unbiased search through the EN parameter space identifies a novel regime of aperiodic OPMs with pinwheel densities lower than found in experiments. In an extreme limit, aperiodic OPMs quantitatively resembling experimental observations emerge. Stabilization of these layouts results from strong nonlocal interactions rather than from a coverage-continuity-compromise. Our results demonstrate that optimization models for stimulus representations dominated by nonlocal suppressive interactions are in principle capable of correctly predicting the common OPM design. They question that visual cortical feature representations can be explained by a coverage-continuity-compromise.Comment: 100 pages, including an Appendix, 21 + 7 figure

What, if anything, are topological maps for?

What, if anything, is the functional significance of spatial patterning in cortical feature maps? We ask this question of four major theories of cortical map formation: self-organizing maps, wiring optimization, place coding, and reaction-diffusion. We argue that (i) self-organizing maps yield spatial patterning only as a by-product of efficient mechanisms for developing environmentally appropriate distributions of feature preferences, (ii) wiring optimization assumes rather than explains a map-like organization, (iii) place-coding mechanisms can at best explain only a subset of maps in functional terms, and (iv) reaction-diffusion models suggest two factors in the evolution of maps, the first based on efficient development of feature distributions, and the second based on generating feature-specific long-range recurrent cortical circuitry. None of these explanations for the existence of topological maps requires spatial patterning in maps to be useful. Thus despite these useful frameworks for understanding how maps form and how they are wired, the possibility that patterns are merely epiphenomena in the evolution of mammalian neocortex cannot be rejected. The article is intended as a nontechnical introduction to the assumptions and predictions of these four important classes of models, along with other possible functional explanations for maps

How can plasticity of lateral interactions affect cortical representation?

Lateral connectivity within cortical areas is pervasive in the mammalian neocortex. The lateral interaction between cortical minicolumns mediated by such connections has been shown to play a critical role in cortical function and cognition, and has been used to explain the emergence of large-scale patterns such as cortical maps. Further evidence suggests that aspects of cortical representation of learnt sensory stimuli may be encoded in the synaptic strengths of lateral connections. This thesis builds upon a program of existing computational neuroscience research, which has identified plasticity in lateral interactions as the key component of cortical functional organisation, to ask whether a neurobiologically plausible computational model of cortical self-organisation can be used to investigate how synaptic plasticity and adaptation in lateral cortical interactions modifies the structure of pre-existing cortical representations and how it affects their decoding. The inhibitory sharpening theory is proposed, based on computer simulations, that shows how repetition suppression is compatible with an increase in the strength of the inhibitory interactions between cortical units co-active during the presentation of the same adapter stimulus due to Hebbian learning. A key prediction of the theory is then derived, that stimuli that produce overlapping patterns of cortical activity, that is that activate a common sub-set of neurons, may produce mutual interference that should be reflected both in changes to the neural signal and in higher level cognition. The predictions of the theory are tested with two approaches, a neuroimaging experiment to measure the magnitude of repetition suppression in a protocol compatible with that used in the simulations, and a behavioural experiment

Arbitrary elastic topologies and ocular dominance

The elastic net, which has been used to produce accounts of the formation of topology preserving maps and ocular dominance columns (OD), embodies a nearest neighbour topology. A Hebbian account of OD is not so restricted – and indeed makes the prediction that the width of the columns depends on the nature of the (more general) neighbourhood relations. Elastic and Hebbian accounts have recently been unified – raising a question mark about their different determiners of column widths. This paper considers this issue, and demonstrates theoretically that it is possible to use more general topologies in the elastic net, including those effectively adopted in the Hebbian model.