A synaptic model for spatial frequency adaptation

A model for visual adaptation to spatial grating is developed based on the assumption that inhibitory synapses within the visual system may be temporarily modified as a function of recent usage. Specifically, it is hypothesized that inhibitory synaptic weights are altered as a function of the correlation between recent presynaptic and postsynaptic activity. When such modifiable synapses are incorporated into a simple neural network model having the spatial filtering properties of the human visual system, two coupled equations are obtained which may be solved analytically. The model accounts for experimental data on adaptation to sinusoidal gratings, square wave gratings, single bars, and tilted gratings. The relationship of the model to single and multiple channel models of the human visual system is discussed.     

Spatial frequency and visual persistence: cortical reset

Psychophysical studies show that the duration of visual persistence increases with spatial frequency of gratings. Previous theories ascribe this finding to differences between the spatial and temporal properties of sustained and transient pathways. This paper proposes an alternative account that explains persistence as a side-effect of excitatory feedback in neural circuits for contour extraction. Mechanisms to break excitatory feedback include inhibitory reset signals at stimulus offset. Simulations demonstrate how gratings with lower spatial frequency generate stronger inhibitory reset signals, thereby resulting in shorter persistence for lower spatial frequencies. Additional simulations account for interactions of spatial frequency with stimulus duration, effects of adaptation, and properties of residual traces, as opposed to visual persistence.     

Contrast adaptation and contrast masking in human vision.

After a preliminary study of visual evoked potentials (VEPS) to a test grating seen in the presence of masks at different orientations, psychophysical data are presented showing the effects of adaptation and of masking on thresholds for detecting the same test grating. The test is a vertical grating of spatial frequency 2 cycles per degree; adapting and masking gratings differ from the test either in orientation or in spatial frequency. The effects of adaptation and masking are explained by a single mechanism model that assumes: (i) adaptation and masking both alter the contrast response (or transducer) function of the mechanism that detects the test; (ii) masks, but not adaptors, stimulate the mechanism that detects the test; and (iii) a test is detectable when it raises response level by a constant amount. The model incorporates two distinct tuning functions, a broad adaptive contrast function and a narrow effective contrast function. It accounts adequately for all the data, including the location and size of the facilitative dip found in some masking functions, the constant slopes of the threshold elevation segments of adaptation functions and the varying slopes of masking functions. It also predicts the sometimes surprising joint effects of adaptation followed by masking and of two masks operating simultaneously.     

Probabilistic secretion of quanta in the central nervous system: granule cell synaptic control of pattern separation and activity regulation.

The implications of probabilistic secretion of quanta for the functioning of neural networks in the central nervous system have been explored. A model of stochastic secretion at synapses in simple networks, consisting of large numbers of granule cells and a relatively small number of inhibitory interneurons, has been analysed. Such networks occur in the input to the cerebellum Purkinje cells as well as to hippocampal CA3 pyramidal cells and to pyramidal cells in the visual cortex. In this model the input axons terminate on granule cells as well as on an inhibitory interneuron that projects to the granule cells. Stochastic secretion at these synapses involves both temporal variability in secretion at single synapses in the network as well as spatial variability in the secretion at different synapses. The role of this stochastic variability in controlling the size of the granule cell output to a level independent of the size of the input and in separating overlapping inputs has been determined analytically as well as by simulation. The regulation of granule-cell output activity to a reasonably constant value for different size inputs does not occur in the absence of an inhibitory interneuron when both spatial and temporal stochastic variability occurs at the remaining synapses; it is still very poor in the presence of such an interneuron but in the absence of stochastic variability. However, quite good regulation is achieved when the inhibitory interneuron is present with spatial and temporal stochastic variability of secretion at synapses in the network. Excellent regulation is achieved if, in addition, allowance is made for the nonlinear behaviour of the input-output characteristics of inhibitory interneurons. The capacity of granule-cell networks to separate overlapping patterns of activity on their inputs is adequate, with spatial variability in the secretion at synapses, but is improved if there is also temporal variability in the stochastic secretion at individual synapses, although this is at the expense of reliability in the network. Other factors which improve pattern separation are control of the output to very low activity levels, and a restriction on the cumulative size of the excitatory input terminals of each granule cell. Application of the theory to the input neural networks of the cerebellum and the hippocampus shows the role of stochastic variability in quantal transmission in determining the capacity of these networks for pattern separation and activity regulation.     

Modeling spatial integration in the ocular following response using a probabilistic framework

The machinery behind the visual perception of motion and the subsequent sensori-motor transformation, such as in ocular following response (OFR), is confronted to uncertainties which are efficiently resolved in the primate's visual system. We may understand this response as an ideal observer in a probabilistic framework by using Bayesian theory [Weiss, Y., Simoncelli, E.P., Adelson, E.H., 2002. Motion illusions as optimal percepts. Nature Neuroscience, 5(6), 598-604, doi:10.1038/nn858] which we previously proved to be successfully adapted to model the OFR for different levels of noise with full field gratings. More recent experiments of OFR have used disk gratings and bipartite stimuli which are optimized to study the dynamics of center-surround integration. We quantified two main characteristics of the spatial integration of motion: (i) a finite optimal stimulus size for driving OFR, surrounded by an antagonistic modulation and (ii) a direction selective suppressive effect of the surround on the contrast gain control of the central stimuli [Barthélemy, F.V., Vanzetta, I., Masson, G.S., 2006. Behavioral receptive field for ocular following in humans: dynamics of spatial summation and center-surround interactions. Journal of Neurophysiology, (95), 3712-3726, doi:10.1152/jn.00112.2006]. Herein, we extended the ideal observer model to simulate the spatial integration of the different local motion cues within a probabilistic representation. We present analytical results which show that the hypothesis of independence of local measures can describe the spatial integration of the motion signal. Within this framework, we successfully accounted for the contrast gain control mechanisms observed in the behavioral data for center-surround stimuli. However, another inhibitory mechanism had to be added to account for suppressive effects of the surround.     

Adaptation to contingencies in macaque primary visual cortex.

We tested the hypothesis that neurons in the primary visual cortex (V1) adapt selectively to contingencies in the attributes of visual stimuli. We recorded from single neurons in macaque V1 and measured the effects of adaptation either to the sum of two gratings (compound stimulus) or to the individual gratings. According to our hypothesis, there would be a component of adaptation that is specific to the compound stimulus. In a first series of experiments, the two gratings differed in orientation. One grating had optimal orientation and the other was orthogonal to it, and therefore did not activate the neuron under study. These experiments provided evidence in favour of our hypothesis. In most cells adaptation to the compound stimulus reduced responses to the compound stimulus more than it reduced responses to the optimal grating, and the responses to the compound stimulus were reduced more by adaptation to the compound stimulus than by adaptation to the individual gratings. This suggests that a component of adaptation was specific to (and caused by) the simultaneous presence of the two orientations in the compound stimulus. To test whether V1 neurons could adapt to other contingencies in the stimulus attributes, we performed a second series of experiments, in which the component gratings were parallel but differed in spatial frequency, and were both effective in activating the neuron under study. These experiments failed to reveal convincing contingent effects of adaptation, suggesting that neurons cannot adapt equally well to all types of contingency.     

Stationary pattern adaptation and the early components in human visual evoked potentials

Pattern-onset visual evoked potentials were elicited from humans by sinusoidal gratings of 0.5, 1, 2 and 4 cpd (cycles/degree) following adaptation to a blank field or one of the gratings. The wave forms recorded after blank field adaptation showed an early positive component, P0, which decreased in amplitude with spatial frequency, whereas the immediately succeeding negative component, N1, increased in amplitude with spatial frequency. P0 and N1 components of comparable size were recorded at 1 cpd. Stationary pattern adaptation to a grating of the same spatial frequency as the test grating significantly reduced N1 amplitude at 4, 2 and 1 cpd. The N1 component elicited at 4 cpd was attenuated in log-linear fashion as the spatial frequency of the adaptation grating increased. P0, on the other hand, was unaffected by stationary pattern adaptation at all combinations of test and adapting spatial frequencies, although P0 amplitude is known to be attenuated by adaptation to a drifting grating. Since N1, but not P0, was significantly attenuated following adaptation and testing at 1 cpd, it was concluded that the neurons generating these components are functionally distinct. The use of a common adaptation grating discounted the possibility that N1, but not P0, was affected due to a difference in the rates of retinal image modulation caused by eye movements made while viewing adaptation gratings of different spatial frequencies. The neurons generating N1 were adapted at a lower rate of retinal image modulation than that apparently required for adaptation of the neurons generating P0, which suggests a difference between these neurons in the rate of stimulus modulation necessary for activation.     

Human vision combines oriented filters to compute edges.

The experiments examined the perceived spatial structure of plaid patterns, composed of two or three sinusoidal gratings of the same spatial frequency, superimposed at different orientations. Perceived structure corresponded well with the pattern of zero crossings in the output of a circular spatial filter applied to the image. This lends some support to Marr & Hildreth's (Proc. R. Soc. Lond. B 207, 187 (1980)) theory of edge detection as a model for human vision, but with a very different implementation. The perceived structure of two-component plaids was distorted by prior exposure to a masking or adapting grating, in a way that was perceptually equivalent to reducing the contrast of one of the plaid components. This was confirmed by finding that the plaid distortion could be nulled by increasing the contrast of the masked or adapted component. A corresponding reduction of perceived contrast for single gratings was observed after adaptation and in some masking conditions. I propose the outlines of a model for edge finding in human vision. The plaid components are processed through cortical, orientation-selective filters that are subject to attenuation by forward masking and adaptation. The outputs of these oriented filters are then linearly summed to emulate circular filtering, and zero crossings (zcs) in the combined output are used to determine edge locations. Masking or adapting to a grating attenuates some oriented filters more than others, and although this changes only the effective contrast of the components, it results in a geometric distortion at the zc level after different filters have been combined. The orientation of zcs may not correspond at all with the orientation of Fourier components, but they are correctly predicted by this two-stage model. The oriented filters are not 'orientation detectors', but are precursors to a more subtle stage that locates and represents spatial features.     

Visual detection of spatial contrast; Influence of location in the visual field,target extent and illuminance level

A model is proposed that permits the prediction of contrast detection thresholds for arbitrary spatial patterns. The influence of the inhomogeneous structure of the visual field and a form of spatial integration are incorporated in the model. A hypothetical density function for the spatial sampling units, which specifies the distribution of these units with respect to both size and location, is described. The density function is compared with anatomical and electrophysiological knowledge of the density of retinal and cortical receptive fields. This density function permits a particularly lucid interpretation in terms of pattern processing. It can be considered as a system that permits simultaneous global and focal views of the surroundings. The density function, together with a schematized adaptation behaviour of single units, and an incoherent summation rule permit us to calculate a measure of the mass response, and consequently the threshold function. Predictions of the model are compared with recently obtained psychophysical data. In particular an explanation is offered for certain invariance properties of spatial contrast detection that seems to possess promising generality.     

Interocular Transfer of Adaptation to Spatial Frequency during Retinal Ischaemia

PROLONGED inspection of a grating pattern of repetitive light and dark contours produces a transient reduction in the visibility of low-contrast gratings the spatial frequency and orientation of which are similar to that of the inspected grating. This after-effect, often referred to as adaptation, occurs even when one eye performs the inspection and its partner is tested. Certain properties of this after-effect, with other data, suggest that its locus is visual cortex^1–7, but the evidence is largely indirect because of the difficulty of identifying sites of action within the stages of the human visual system. One of the few techniques available for inferentially partitioning the visual system stages is retinal pressure blinding, a heroic but effective method for functionally uncoupling the retina from higher stages^8–11.     

Orientation-Specificity of Adaptation: Isotropic Adaptation Is Purely Monocular

Numerous studies have found that prolonged exposure to grating stimuli reduces sensitivity to subsequently presented gratings, most evidently when the orientations of the adapting and test patterns are similar. The rate of sensitivity loss varies with angular difference indicating both the presence and bandwidths of psychophysical ‘orientation channels’. Here we study the orientation dependency of contrast adaptation measured both monoptically and dichoptically. Earlier psychophysical reports show that orientation bandwidths are broader at lower spatial frequencies, and we confirm this with a simple von Mises model using 0.25 vs. 2 c.p.d. gratings. When a single isotropic (orientation invariant) parameter is added to this model, however, we find no evidence for any difference in bandwidth with spatial frequency. Consistent with cross-orientation masking effects, we find isotropic adaptation to be strongly low spatial frequency-biased. Surprisingly, unlike masking, we find that the effects of interocular adaptation are purely orientation-tuned, with no evidence of isotropic threshold elevation. This dissociation points to isotropic (or ‘cross-orientation’) adaptation being an earlier and more magnocellular-like process than that which supports orientation-tuned adaptation and suggests that isotropic masking and adaptation are likely mediated by separate mechanisms.     

Dependence of Visual Cell Properties on Intracortical Synapses Among Hypercolumns: Analysis by a Computer Model

The role of intracortical synapses in affecting the property of visual cells is investigated by means of an original mathematical model of cortical circuitry in V1. The model represents a compromise between computational simplicity and physiological reliability. The model incorporates four different inputs into a cortical cell: thalamic input from the lateral geniculate nucleus, according to an even Gabor function; short-range inhibition confined within the hypercolumn; a long-range excitation, which emphasizes the properties of the input; and a long-range inhibition. In the model we assume that all cells receive a similar thalamic input, which differs simply according to its position in the retina and orientation preference. Simulations were performed, with different parameter values, to assess the main characteristics of cell response (i.e., the width and locations of subregions in the receptive field (RF), orientation tuning curve, and response to drifting and counterphase gratings) as a function of the strength and extension of intracortical excitatory synapses. Results suggest that, if intracortical excitation is confined within the hypercolumn, the cells exhibit the same properties as simple cells, both with regards to the width and shape of the RF, orientation tuning curve, and response to drifting and counterphase gratings. By contrast, if excitatory synapses extend beyond the hypercolumn with sufficient strength, the cells exhibit the typical characteristics of complex cells. A progressive shift from complex to simple cells can be realized with a monotonic variation in parameters. Simulations are also performed with a hierarchical model, to suggest possible experiments able to discriminate the present recurrent mechanism from the classical hierarchical one. Results support the assumptions of previous simpler models (Chance et al., 1999) and may help to understand and assess the role of intracortical synapses in rigorous quantitative terms.     

A unified neural network model of spatiotemporal processing in X and Y retinal ganglion cells

This article makes use of a push-pull shunting network, which was introduced in the companion article, to model certain properties of X and Y retinal ganglion cells. Input to the push-pull network is preprocessed by a nonlinear mechanism for temporal adaptation, which is ascribed here to photoreceptor dynamics. The complete circuit is used to show that a simple change in receptive field morphology within a single model equation can change the network's response characteristics to closely resemble those of either X or Y cells. Specifically, an increase in width of the receptive field center mechanism is sufficient to account for generation of on-off (Y-like) instead of null (X-like) responses to modulated gratings. In agreement with experimental data, the Y cell on-off response is independent of spatial phase. Also, the model accurately predicts that on-off responses can be observed in X cells for particular stimulus configurations. Taken together, the results show how the retina combines individually inadequate modules to efficiently handle the tasks required for accurate spatial and temporal visual information processing. The model is also able to clarify a number of controversial experimental findings on the nature of spatiotemporal visual processing in the retina.     

Loss of sensory input causes rapid structural changes of inhibitory neurons in adult mouse visual cortex

A fundamental property of neuronal circuits is the ability to adapt to altered sensory inputs. It is well established that the functional synaptic changes underlying this adaptation are reflected by structural modifications in excitatory neurons. In contrast, the degree to which structural plasticity in inhibitory neurons accompanies functional changes is less clear. Here, we use two-photon imaging to monitor the fine structure of inhibitory neurons in mouse visual cortex after deprivation induced by retinal lesions. We find that a subset of inhibitory neurons carry dendritic spines, which form glutamatergic synapses. Removal of visual input correlates with a rapid and lasting reduction in the number of inhibitory cell spines. Similar to the effects seen for dendritic spines, the number of inhibitory neuron boutons dropped sharply after retinal lesions. Together, these data suggest that structural changes in inhibitory neurons may precede structural changes in excitatory circuitry, which ultimately result in functional adaptation following sensory deprivation.     

Human psychophysics: Functional interpretation for contrast sensitivity versus spatial frequency curve

The hypothesis that neural processing in the human visual pathways compensates for both optical degradation as well as noise contamination at the photoreceptor level is introduced and shown to be consistent with the high frequency portion of the contrast sensitivity function for threshold detection of sinusoidal gratings in addition to the suprathreshold phenomenon of matching sinusoidal gratings of different spatial frequencies. This offers a unifying interpretation for why, at threshold conditions, the high spatial frequency portion of the image is blurred as severely by the nervous system as it is by the optics (e.g. Campbell and Green, 1965) while in extreme suprathreshold conditions the nervous system effectively deblurs the image (e.g. Georgeson and sullivan, 1975; Kulikowski, 1976). These conclusions do not necessitate a highly specific form of visual processing such as Fourier channeling.This research was conducted at Yale University, Department of Ophthalmology and Visual Science, New Haven, Connecticut, USA, throughout which period A.W.S. was a John Simon Guggenheim fellow     

幼年大鼠视皮层神经元对闪光刺激的反应特性

哺乳动物视觉系统的发育延续到出生后,大鼠出生后 3~5 周是视觉系统发育的关键期 . 在关键期中,视皮层的兴奋性和抑制性突触连接逐渐成熟,形成有效的皮层内回路 . 为了观察发育关键期大鼠视皮层神经元的反应特性与成年大鼠的异同,使用胞外单细胞记录的方法对比研究了幼年和成年大鼠对闪光刺激的视觉反应特性 . 结果显示：与成年大鼠相比较,幼年大鼠视皮层神经元对持续闪光刺激显示出更强的适应性,对光刺激的诱发放电频率更低,而在没有光刺激时的自发放电频率更高,从而导致信噪比更低 . 这一结果表明,幼年大鼠视皮层对连续刺激的反应能力下降,对信号的分辨能力也更弱,其原因可能是兴奋性突触和抑制性突触发育的不同步所致 .     

Temporal adaptability and the inverse relationship to sensitivity: a parameter identification model

Following a prolonged period of visual adaptation to a temporally modulated sinusoidal luminance pattern, the threshold contrast of a similar visual pattern is elevated. The adaptive elevation in threshold contrast is selective for spatial frequency, may saturate at low adaptor contrast, and increases as a function of the spatio-temporal frequency of the adapting signal. A model for signal extraction that is capable of explaining these threshold contrast effects of adaptation is proposed. Contrast adaptation in the model is explained by the identification of the parameters of an environmental model: the autocorrelation function of the visualized signal. The proposed model predicts that the adaptability of threshold contrast is governed by unpredicted signal variations present in the visual signal, and thus represents an internal adjustment by the visual system that takes into account these unpredicted signal variations given the additional possibility for signal corruption by additive noise.