Biomedical subjects
Publications and source records attributed to E Erwin.
Two sorts of subjectivity problems are discussed. The 1st concerns the characterization of psychotherapy as an attempt to alter clients' meanings. The 2nd concerns the seemingly subjective nature of value judgments about psychotherapy outcomes. It is argued that despite initial appearances, neither of these problems poses an insuperable difficulty for transforming the discipline of psychotherapy into a genuine science.
We explore the hypothesis that binocular simple cells in cat areas 17 and 18 show subregion correspondence, defined as follows: within the region of overlap of the two eye's receptive fields, their ON subregions lie in corresponding locations, as do their OFF subregions. This hypothesis is motivated by a developmental model (Erwin and Miller, 1998) that suggested that simple cells could develop binocularly matched preferred orientations and spatial frequencies by developing subregion correspondence. Binocular organization of simple cell receptive fields is commonly characterized by two quantities: interocular position shift, the distance in visual space between the center positions of the two eye's receptive fields; and interocular phase shift, the difference in the spatial phases of those receptive fields, each measured relative to its center position. The subregion correspondence hypothesis implies that interocular position and phase shifts are linearly related. We compare this hypothesis with the null hypothesis, assumed by most previous models of binocular organization, that the two types of shift are uncorrelated. We demonstrate that the subregion correspondence and null hypotheses are equally consistent with previous measurements of binocular response properties of individual simple cells in the cat and other species and with measurements of the distribution of interocular phase shifts versus preferred orientations or versus interocular position shifts. However, the observed tendency of binocular simple cells in the cat to have "tuned excitatory" disparity tuning curves with preferred disparities tightly clustered around zero (Fischer and Krüger, 1979; Ferster, 1981; LeVay and Voigt, 1988) follows naturally from the subregion correspondence hypothesis but is inconsistent with the null hypothesis. We describe tests that could more conclusively differentiate between the hypotheses. The most straightforward test requires simultaneous determination of the receptive fields of groups of three or more binocular simple cells.
The primary focus of this paper is the abrupt transition that occurs midway through the rhesus lateral geniculate nucleus (LGN) from six layers posteriorly (conventionally numbered 1-6, ventral to dorsal) to four layers anteriorly. At this transition, layers 4 and 6 fuse into a single layer, as do layers 3 and 5, requiring an inversion of the stacking order of the cell categories making up layers 4 and 5. To understand the topology of this transition and its relationship to geniculate retinotopy, we have created a functional atlas of a rhesus LGN that affords three-dimensional views of morphology and retinotopy at a resolution of 25 microm. The projection of the path of the transition into visual space is highly biased toward lower visual fields, intersecting the upper vertical meridian at 6.4 degrees , the horizonal meridian at 15.4 degrees, and the lower vertical meridian at 25.0 degrees. Between inclinations of -31 degrees and 55 degrees, layers 3 and 5 merge through an elongated tear in layer 4 that subsumes the optic disk gap and extends medially and laterally; elsewhere, layers 4 and 6 merge through a tear in layer 5. These tears cause substantial violations of retinotopy and laminar integrity, so the inversion of layers 4 and 5 requires that the forces establishing retinotopy and grouping by cell class be locally overcome during morphogenesis. The transition and associated tears are evaluated in the context of recent computational models of geniculate morphogenesis. We have also used the atlas to estimate the borders of the binocular (55 approximately 62 degrees) and monocular (91 approximately 97 degrees) visual fields. Files containing the atlas are made publicly available on a website.
The idiographic problem is said to arise because psychotherapy experiments study groups of subjects, whereas the clinician wants useful etiological or treatment information concerning a single, unique client. One response is to place little value on psychotherapy experiments and to argue for nonexperimental methods for obtaining clinical knowledge. It is argued here that these alternative methods are defective and that the idiographic problem can be solved without renouncing experimental methods.
Is the development of orientation selectivity in visual cortex instructed by the patterns of neural activity of input neurons? We review evidence as to the role of activity, review models of activity-instructed development, and discuss how these models can be tested. The models can explain the normal development of simple cells with binocularly matched orientation preferences, the effects of monocular deprivation and reverse suture on the orientation map, and the development of a full intracortical circuit sufficient to explain mature response properties including the contrast-invariance of orientation tuning. Existing experiments are consistent with the models, in that (a) selective blockade of ON-center ganglion cells, which will degrade or eliminate the information predicted to drive development of orientation selectivity, in fact prevents development of orientation selectivity; and (b) the spontaneous activities of inputs serving the two eyes are correlated in the lateral geniculate nucleus at appropriate developmental times, as was predicted to be required to achieve binocular matching of preferred orientations. However, definitive tests remain to be done to firmly establish the instructive rather than simply permissive role of activity and determine whether the retinotopically and center type-specific patterns of activity predicted by the models actually exist. We conclude by critically examining alternative scenarios for the development of orientation selectivity and maps, including the idea that maps are genetically prespecified.
We extend previous models for separate development of ocular dominance and orientation selectivity in cortical layer 4 by exploring conditions permitting combined organization of both properties. These conditions are expressed in terms of functions describing the degree of correlation in the firing of two inputs from the lateral geniculate nucleus (LGN), as a function of their retinotopic separation and their "type" (ON center or OFF center and left eye or right eye). The development of ocular dominance requires that the correlations of an input with other inputs of the same eye be stronger than or equal to its correlations with inputs of the opposite eye and strictly stronger at small retinotopic separations. This must be true after summing correlations with inputs of both center types. The development of orientation-selective simple cells requires that (1) an input's correlations with other inputs of the same center type be stronger than its correlations with inputs of the opposite center type at small retinotopic separation; and (2) this relationship reverse at larger retinotopic separations within an arbor radius (the radius over which LGN cells can project to a common cortical point). This must be true after summing correlations with inputs serving both eyes. For orientations to become matched in the two eyes, correlated activity within the receptive fields must be maximized by specific between-eye alignments of ON and OFF subregions. Thus the correlations between the eyes must differ depending on center type, and this difference must vary with retinotopic separation within an arbor radius. These principles are satisfied by a wide class of correlation functions. Combined development of ocularly matched orientation maps and ocular dominance maps can be achieved either simultaneously or sequentially. In the latter case, the model can produce a correlation between the locations of orientation map singularities and local ocular dominance peaks similar to that observed physiologically. The model's main prediction is that the above correlations should exist among inputs to cortical layer 4 simple cells before vision. In addition, mature simple cells are predicted to have certain relationships between the locations of the ON and OFF subregions of the left and right eyes' receptive fields.
Orientation and ocular dominance maps in the primary visual cortex of mammals are among the most thoroughly investigated of the patterns in the cerebral cortex. A considerable amount of work has been dedicated to unraveling both their detailed structure and the neural mechanisms that underlie their formation and development. Many schemes have been proposed, some of which are in competition. Some models focus on development of receptive fields while others focus on the structure of cortical maps, i.e., the arrangement of receptive field properties across the cortex. Each model used different means to determine its success at reproducing experimental map patterns, often relying principally on visual comparison. Experimental data are becoming available that allow a more careful evaluation of models. In this contribution more than 10 of the most prominent models of cortical map formation and structure are critically evaluated and compared with the most recent experimental findings from macaque striate cortex. Comparisons are based on properties of the predicted or measured cortical map patterns. We introduce several new measures for comparing experimental and model map data that reveal important differences between models. We expect that the use of these measures will improve current models by helping determine parameters to match model maps to experimental data now becoming available from a variety of species. Our study reveals that (1) despite apparent differences, many models are based on similar principles and consequently make similar predictions, (2) several models produce orientation map patterns that are not consistent with the experimental data from macaques, regardless of the plausibility of the models' suggested physiological implementations, and (3) no models have yet fully accounted for both the local and the global relationships between orientation and ocular dominance map patterns.
We investigate the effect of various types of neighborhood function on the convergence rates and the presence or absence of metastable stationary states of Kohonen's self-organizing feature map algorithm in one dimension. We demonstrate that the time necessary to form a topographic representation of the unit interval [0, 1] may vary over several orders of magnitude depending on the range and also the shape of the neighborhood function, by which the weight changes of the neurons in the neighborhood of the winning neuron are scaled. We will prove that for neighborhood functions which are convex on an interval given by the length of the Kohonen chain there exist no metastable states. For all other neighborhood functions, metastable states are present and may trap the algorithm during the learning process. For the widely-used Gaussian function there exists a threshold for the width above which metastable states cannot exist. Due to the presence or absence of metastable states, convergence time is very sensitive to slight changes in the shape of the neighborhood function. Fastest convergence is achieved using neighborhood functions which are "convex" over a large range around the winner neuron and yet have large differences in value at neighboring neurons.
We investigate the convergence properties of the self-organizing feature map algorithm for a simple, but very instructive case: the formation of a topographic representation of the unit interval [0, 1] by a linear chain of neurons. We extend the proofs of convergence of Kohonen and of Cottrell and Fort to hold in any case where the neighborhood function, which is used to scale the change in the weight values at each neuron, is a monotonically decreasing function of distance from the winner neuron. We prove that the learning dynamics cannot be described by a gradient descent on a single energy function, but may be described using a set of potential functions, one for each neuron, which are independently minimized following a stochastic gradient descent. We derive the correct potential functions for the one- and multi-dimensional case, and show that the energy functions given by Tolat (1990) are an approximation which is no longer valid in the case of highly disordered maps or steep neighborhood functions.
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