Model Of Primary Visual Cortex Research Articles

Population codes enable learning from few examples by shaping inductive bias.

Learning from a limited number of experiences requires suitable inductive biases. To identify how inductive biases are implemented in and shaped by neural codes, we analyze sample-efficient learning of arbitrary stimulus-response maps from arbitrary neural codes with biologically-plausible readouts. We develop an analytical theory that predicts the generalization error of the readout as a function of the number of observed examples. Our theory illustrates in a mathematically precise way how the structure of population codes shapes inductive bias, and how a match between the code and the task is crucial for sample-efficient learning. It elucidates a bias to explain observed data with simple stimulus-response maps. Using recordings from the mouse primary visual cortex, we demonstrate the existence of an efficiency bias towards low-frequency orientation discrimination tasks for grating stimuli and low spatial frequency reconstruction tasks for natural images. We reproduce the discrimination bias in a simple model of primary visual cortex, and further show how invariances in the code to certain stimulus variations alter learning performance. We extend our methods to time-dependent neural codes and predict the sample efficiency of readouts from recurrent networks. We observe that many different codes can support the same inductive bias. By analyzing recordings from the mouse primary visual cortex, we demonstrate that biological codes have lower total activity than other codes with identical bias. Finally, we discuss implications of our theory in the context of recent developments in neuroscience and artificial intelligence. Overall, our study provides a concrete method for elucidating inductive biases of the brain and promotes sample-efficient learning as a general normative coding principle.

Spatial and color hallucinations in a mathematical model of primary visual cortex

We study a simplified model of the representation of colors in the primate primary cortical visual area V1. The model is described by an initial value problem related to a Hammerstein equation. The solutions to this problem represent the variation of the activity of populations of neurons in V1 as a function of space and color. The two space variables describe the spatial extent of the cortex while the two color variables describe the hue and the saturation represented at every location in the cortex. We prove the well-posedness of the initial value problem. We focus on its stationary, i.e. independent of time, and periodic in space solutions. We show that the model equation is equivariant with respect to the direct product G of the group of the Euclidean transformations of the planar lattice determined by the spatial periodicity and the group of color transformations, isomorphic to O(2), and study the equivariant bifurcations of its stationary solutions when some parameters in the model vary. Their variations may be caused by the consumption of drugs and the bifurcated solutions may represent visual hallucinations in space and color. Some of the bifurcated solutions can be determined by applying the Equivariant Branching Lemma (EBL) by determining the axial subgroups of G . These define bifurcated solutions which are invariant under the action of the corresponding axial subgroup. We compute analytically these solutions and illustrate them as color images. Using advanced methods of numerical bifurcation analysis we then explore the persistence and stability of these solutions when varying some parameters in the model. We conjecture that we can rely on the EBL to predict the existence of patterns that survive in large parameter domains but not to predict their stability. On our way we discover the existence of spatially localized stable patterns through the phenomenon of "snaking".

Interrogating theoretical models of neural computation with emergent property inference.

A cornerstone of theoretical neuroscience is the circuit model: a system of equations that captures a hypothesized neural mechanism. Such models are valuable when they give rise to an experimentally observed phenomenon -- whether behavioral or a pattern of neural activity -- and thus can offer insights into neural computation. The operation of these circuits, like all models, critically depends on the choice of model parameters. A key step is then to identify the model parameters consistent with observed phenomena: to solve the inverse problem. In this work, we present a novel technique, emergent property inference (EPI), that brings the modern probabilistic modeling toolkit to theoretical neuroscience. When theorizing circuit models, theoreticians predominantly focus on reproducing computational properties rather than a particular dataset. Our method uses deep neural networks to learn parameter distributions with these computational properties. This methodology is introduced through a motivational example of parameter inference in the stomatogastric ganglion. EPI is then shown to allow precise control over the behavior of inferred parameters and to scale in parameter dimension better than alternative techniques. In the remainder of this work, we present novel theoretical findings in models of primary visual cortex and superior colliculus, which were gained through the examination of complex parametric structure captured by EPI. Beyond its scientific contribution, this work illustrates the variety of analyses possible once deep learning is harnessed towards solving theoretical inverse problems.

Dysregulation of excitatory neural firing replicates physiological and functional changes in aging visual cortex.

The mammalian visual system has been the focus of countless experimental and theoretical studies designed to elucidate principles of neural computation and sensory coding. Most theoretical work has focused on networks intended to reflect developing or mature neural circuitry, in both health and disease. Few computational studies have attempted to model changes that occur in neural circuitry as an organism ages non-pathologically. In this work we contribute to closing this gap, studying how physiological changes correlated with advanced age impact the computational performance of a spiking network model of primary visual cortex (V1). Our results demonstrate that deterioration of homeostatic regulation of excitatory firing, coupled with long-term synaptic plasticity, is a sufficient mechanism to reproduce features of observed physiological and functional changes in neural activity data, specifically declines in inhibition and in selectivity to oriented stimuli. This suggests a potential causality between dysregulation of neuron firing and age-induced changes in brain physiology and functional performance. While this does not rule out deeper underlying causes or other mechanisms that could give rise to these changes, our approach opens new avenues for exploring these underlying mechanisms in greater depth and making predictions for future experiments.

Understanding Eye Movements: Psychophysics and a Model of Primary Visual Cortex

Humans move their eyes in order to learn visual representations of the world. These eye movements depend on distinct factors, either by the scene that we perceive or by our own decisions. To select what is relevant to attend is part of our survival mechanisms and the way we build reality, as we constantly react both consciously and unconsciously to all the stimuli that is projected into our eyes. In this thesis we try to explain (1) how we move our eyes, (2) how to build machines that understand visual information and deploy eye movements, and (3) how to make these machines understand tasks in order to decide for eye movements.(1) We provided the analysis of eye movement behavior elicited by low-level feature distinctiveness with a dataset of 230 synthetically-generated image patterns. A total of 15 types of stimuli has been generated (e.g. orientation, brightness, color, size, etc.), with 7 feature contrasts for each feature category. Eye-tracking data was collected from 34 participants during the viewing of the dataset, using Free-Viewing and Visual Search task instructions. Results showed that saliency is predominantly and distinctively influenced by: 1. feature type, 2. feature contrast, 3. temporality of fixations, 4. task difficulty and 5. center bias. From such dataset (SID4VAM), we have computed a benchmark of saliency models by testing performance using psychophysical patterns. Model performance has been evaluated considering model inspiration and consistency with human psychophysics. Our study reveals that state-of-the-art Deep Learning saliency models do not perform well with synthetic pattern images, instead, models with Spectral/Fourier inspiration outperform others in saliency metrics and are more consistent with human psychophysical experimentation.(2) Computations in the primary visual cortex (area V1 or striate cortex) have long been hypothesized to be responsible, among several visual processing mechanisms, of bottom-up visual attention (also named saliency). In order to validate this hypothesis, images from eye tracking datasets have been processed with a biologically plausible model of V1 (named Neurodynamic Saliency Wavelet Model or NSWAM). Following Li's neurodynamic model, we define V1's lateral connections with a network of firing rate neurons, sensitive to visual features such as brightness, color, orientation and scale. Early subcortical processes (i.e. retinal and thalamic) are functionally simulated. The resulting saliency maps are generated from the model output, representing the neuronal activity of V1 projections towards brain areas involved in eye movement control. We want to pinpoint that our unified computational architecture is able to reproduce several visual processes (i.e. brightness, chromatic induction and visual discomfort) without applying any type of training or optimization and keeping the same parametrization. The model has been extended (NSWAM-CM) with an implementation of the cortical magnification function to define the retinotopical projections towards V1, processing neuronal activity for each distinct view during scene observation. Novel computational definitions of top-down inhibition (in terms of inhibition of return and selection mechanisms), are also proposed to predict attention in Free-Viewing and Visual Search conditions. Results show that our model outpeforms other biologically-inpired models of saliency prediction as well as to predict visual saccade sequences, specifically for nature and synthetic images. We also show how temporal and spatial characteristics of inhibition of return can improve prediction of saccades, as well as how distinct search strategies (in terms of feature-selective or category-specific inhibition) predict attention at distinct image contexts.(3) Although previous scanpath models have been able to efficiently predict saccades during Free-Viewing, it is well known that stimulus and task instructions can strongly affect eye movement patterns. In particular, task priming has been shown to be crucial to the deployment of eye movements, involving interactions between brain areas related to goal-directed behavior, working and long-term memory in combination with stimulus-driven eye movement neuronal correlates. In our latest study we proposed an extension of the Selective Tuning Attentive Reference Fixation Controller Model based on task demands (STAR-FCT), describing novel computational definitions of Long-Term Memory, Visual Task Executive and Task Working Memory. With these modules we are able to use textual instructions in order to guide the model to attend to specific categories of objects and/or places in the scene. We have designed our memory model by processing a visual hierarchy of low- and high-level features. The relationship between the executive task instructions and the memory representations has been specified using a tree of semantic similarities between the learned features and the object category labels. Results reveal that by using this model, the resulting object localization maps and predicted saccades have a higher probability to fall inside the salient regions depending on the distinct task instructions compared to saliency.

Functional asymmetry of local connections in V1 and its impact on orientation tuning

Abstract Classic models of primary visual cortex (V1) reproduce orientation selectivity with sufficiently high accuracy. However, they are unable to explain phenomena that are widely studied experimentally: a single cell carry additional information in response waveform and oscillations. Among the other effects not reproduced by the classic models are changes in the peak orientation during the response time course and shifts induced by inhibition blockage. We suppose that the main reason for such a failure is assumption that spatial distribution of short-range horizontal connections in V1 is symmetric. In the present study, we provide an experimental evidence that such symmetry is not the case in higher mammals. Based on this evidence we improved the classical model of V1 by replacing the symmetric spatial distribution to another one revealed from experiments. The improved model is able to reproduce orientation shifts and transmission of additional information. This means that the asymmetry shall take into consideration when simulating V1.

Short-Term Attractive Tilt Aftereffects Predicted by a Recurrent Network Model of Primary Visual Cortex.

Adaptation is a multi-faceted phenomenon that is of interest in terms of both its function and its potential to reveal underlying neural processing. Many behavioral studies have shown that after exposure to an oriented adapter the perceived orientation of a subsequent test is repulsed away from the orientation of the adapter. This is the well-known Tilt Aftereffect (TAE). Recently, we showed that the dynamics of recurrently connected networks may contribute substantially to the neural changes induced by adaptation, especially on short time scales. Here we extended the network model and made the novel behavioral prediction that the TAE should be attractive, not repulsive, on a time scale of a few 100 ms. Our experiments, using a novel adaptation protocol that specifically targeted adaptation on a short time scale, confirmed this prediction. These results support our hypothesis that recurrent network dynamics may contribute to short-term adaptation. More broadly, they show that understanding the neural processing of visual inputs that change on the time scale of a typical fixation requires a detailed analysis of not only the intrinsic properties of neurons, but also the slow and complex dynamics that emerge from their recurrent connectivity. We argue that this is but one example of how even simple recurrent networks can underlie surprisingly complex information processing, and are involved in rudimentary forms of memory, spatio-temporal integration, and signal amplification.

Deep convolutional models improve predictions of macaque V1 responses to natural images.

Despite great efforts over several decades, our best models of primary visual cortex (V1) still predict spiking activity quite poorly when probed with natural stimuli, highlighting our limited understanding of the nonlinear computations in V1. Recently, two approaches based on deep learning have emerged for modeling these nonlinear computations: transfer learning from artificial neural networks trained on object recognition and data-driven convolutional neural network models trained end-to-end on large populations of neurons. Here, we test the ability of both approaches to predict spiking activity in response to natural images in V1 of awake monkeys. We found that the transfer learning approach performed similarly well to the data-driven approach and both outperformed classical linear-nonlinear and wavelet-based feature representations that build on existing theories of V1. Notably, transfer learning using a pre-trained feature space required substantially less experimental time to achieve the same performance. In conclusion, multi-layer convolutional neural networks (CNNs) set the new state of the art for predicting neural responses to natural images in primate V1 and deep features learned for object recognition are better explanations for V1 computation than all previous filter bank theories. This finding strengthens the necessity of V1 models that are multiple nonlinearities away from the image domain and it supports the idea of explaining early visual cortex based on high-level functional goals.

Synaptic refinement during development and its effect on slow-wave activity: a computational study

Recent evidence suggests that synaptic refinement, the reorganization of synapses and connections without significant change in their number or strength, is important for the development of the visual system of juvenile rodents. Other evidence in rodents and humans shows that there is a marked drop in sleep slow-wave activity (SWA) during adolescence. Slow waves reflect synchronous transitions of neuronal populations between active and inactive states, and the amount of SWA is influenced by the connection strength and organization of cortical neurons. In this study, we investigated whether synaptic refinement could account for the observed developmental drop in SWA. To this end, we employed a large-scale neural model of primary visual cortex and sections of the thalamus, capable of producing realistic slow waves. In this model, we reorganized intralaminar connections according to experimental data on synaptic refinement: during prerefinement, local connections between neurons were homogenous, whereas in postrefinement, neurons connected preferentially to neurons with similar receptive fields and preferred orientations. Synaptic refinement led to a drop in SWA and to changes in slow-wave morphology, consistent with experimental data. To test whether learning can induce synaptic refinement, intralaminar connections were equipped with spike timing-dependent plasticity. Oriented stimuli were presented during a learning period, followed by homeostatic synaptic renormalization. This led to activity-dependent refinement accompanied again by a decline in SWA. Together, these modeling results show that synaptic refinement can account for developmental changes in SWA. Thus sleep SWA may be used to track noninvasively the reorganization of cortical connections during development.

Orientation selectivity in a model of primary visual cortex with and without orientation map

Since its discovery by Hubel and Wiesel in 1959, orientation selectivity has been observed in every mammal for which the neuronal response selectivity of primary visual cortex (V1) has been examined. In some animals, like cats and monkeys, anatomically close V1 neurons have similar preferred orientations, giving rise to maps of orientation preferences. However, sharp selectivity is also observed in mice, squirrels and rats, whose V1 has no orientation map. This means that neurons with different preferred orientations are intermixed. This second scenario is called salt-and-pepper organization and leads to question the structural organization of the intracortical connections. With this study, we intend to analyze the differences between both scenarios focusing on selectivity properties, and to clarify the effect of plasticity on orientation selectivity. We study a computational model of layer 2/3 and a reduced one-dimensional model of orientation selective neurons, both in the balanced state. We also analyze a plasticity mechanism that involves spike-timing dependent plasticity (STDP) [1]. This rule implies that if the postsynaptic spike comes after the presynaptic spike, the connection becomes stronger. For the reverse order it becomes weaker. Inhibitory synapses are taken as non-plastic. The selectivity is quantified using the Orientation Selectivity Index (OSI) of the activity profile. According to the theory of balanced networks [2] applied to spatially structured networks [3], we find that for a given profile of the input, selectivity of the cortical activity is determined by the ratio between the first Fourier component of the connectivity and its mean value. As a consequence, for large connectivity salt-and-pepper structures are more selective than systems with orientation maps: our simulations indicate that the mean OSI for salt-and-pepper is 0.57, while it is 0.27 for orientation map. Moreover, for systems with orientation maps selectivity could be increased by taking connection probabilities that are broader. Regarding plasticity, we find that under certain conditions STDP can indeed improve selectivity but it works in a somehow unexpected way, by decreasing the modulated part of the intracortical connectivity with respect to the non-modulated part. We find this conclusion to be valid both for systems with salt-and-pepper organization and with orientation maps. This can be understood in terms of the relative change between the background connectivity and the functionally modulated part.

Laminar Neural Field Model of Laterally Propagating Waves of Orientation Selectivity.

We construct a laminar neural-field model of primary visual cortex (V1) consisting of a superficial layer of neurons that encode the spatial location and orientation of a local visual stimulus coupled to a deep layer of neurons that only encode spatial location. The spatially-structured connections in the deep layer support the propagation of a traveling front, which then drives propagating orientation-dependent activity in the superficial layer. Using a combination of mathematical analysis and numerical simulations, we establish that the existence of a coherent orientation-selective wave relies on the presence of weak, long-range connections in the superficial layer that couple cells of similar orientation preference. Moreover, the wave persists in the presence of feedback from the superficial layer to the deep layer. Our results are consistent with recent experimental studies that indicate that deep and superficial layers work in tandem to determine the patterns of cortical activity observed in vivo.

A unified account of tilt illusions, association fields, and contour detection based on elastica

As expressed in the Gestalt law of good continuation, human perception tends to associate stimuli that form smooth continuations. Contextual modulation in primary visual cortex, in the form of association fields, is believed to play an important role in this process. Yet a unified and principled account of the good continuation law on the neural level is lacking. In this study we introduce a population model of primary visual cortex. Its contextual interactions depend on the elastica curvature energy of the smoothest contour connecting oriented bars. As expected, this model leads to association fields consistent with data. However, in addition the model displays tilt-illusions for stimulus configurations with grating and single bars that closely match psychophysics. Furthermore, the model explains not only pop-out of contours amid a variety of backgrounds, but also pop-out of single targets amid a uniform background. We thus propose that elastica is a unifying principle of the visual cortical network.

The effect of synaptic plasticity on orientation selectivity in a balanced model of primary visual cortex.

Orientation selectivity is ubiquitous in the primary visual cortex (V1) of mammals. In cats and monkeys, V1 displays spatially ordered maps of orientation preference. Instead, in mice, squirrels, and rats, orientation selective neurons in V1 are not spatially organized, giving rise to a seemingly random pattern usually referred to as a salt-and-pepper layout. The fact that such different organizations can sharpen orientation tuning leads to question the structural role of the intracortical connections; specifically the influence of plasticity and the generation of functional connectivity. In this work, we analyze the effect of plasticity processes on orientation selectivity for both scenarios. We study a computational model of layer 2/3 and a reduced one-dimensional model of orientation selective neurons, both in the balanced state. We analyze two plasticity mechanisms. The first one involves spike-timing dependent plasticity (STDP), while the second one considers the reconnection of the interactions according to the preferred orientations of the neurons. We find that under certain conditions STDP can indeed improve selectivity but it works in a somehow unexpected way, that is, effectively decreasing the modulated part of the intracortical connectivity as compared to the non-modulated part of it. For the reconnection mechanism we find that increasing functional connectivity leads, in fact, to a decrease in orientation selectivity if the network is in a stable balanced state. Both counterintuitive results are a consequence of the dynamics of the balanced state. We also find that selectivity can increase due to a reconnection process if the resulting connections give rise to an unstable balanced state. We compare these findings with recent experimental results.

Computational Model of Primary Visual Cortex Combining Visual Attention for Action Recognition.

Humans can easily understand other people’s actions through visual systems, while computers cannot. Therefore, a new bio-inspired computational model is proposed in this paper aiming for automatic action recognition. The model focuses on dynamic properties of neurons and neural networks in the primary visual cortex (V1), and simulates the procedure of information processing in V1, which consists of visual perception, visual attention and representation of human action. In our model, a family of the three-dimensional spatial-temporal correlative Gabor filters is used to model the dynamic properties of the classical receptive field of V1 simple cell tuned to different speeds and orientations in time for detection of spatiotemporal information from video sequences. Based on the inhibitory effect of stimuli outside the classical receptive field caused by lateral connections of spiking neuron networks in V1, we propose surround suppressive operator to further process spatiotemporal information. Visual attention model based on perceptual grouping is integrated into our model to filter and group different regions. Moreover, in order to represent the human action, we consider the characteristic of the neural code: mean motion map based on analysis of spike trains generated by spiking neurons. The experimental evaluation on some publicly available action datasets and comparison with the state-of-the-art approaches demonstrate the superior performance of the proposed model.

Uncomfortable images produce non-sparse responses in a model of primary visual cortex.

The processing of visual information by the nervous system requires significant metabolic resources. To minimize the energy needed, our visual system appears to be optimized to encode typical natural images as efficiently as possible. One consequence of this is that some atypical images will produce inefficient, non-optimal responses. Here, we show that images that are reported to be uncomfortable to view, and that can trigger migraine attacks and epileptic seizures, produce relatively non-sparse responses in a model of the primary visual cortex. In comparison with the responses to typical inputs, responses to aversive images were larger and less sparse. We propose that this difference in the neural population response may be one cause of visual discomfort in the general population, and can produce more extreme responses in clinical populations such as migraine and epilepsy sufferers.

A Unified Computational Model of Primary Visual Cortex: Consolidation of the Scattered Literature on Simple and Complex Cells

A Unified Computational Model of Primary Visual Cortex: Consolidation of the Scattered Literature on Simple and Complex Cells

Conductance-based refractory density model of primary visual cortex

A layered continual population model of primary visual cortex has been constructed, which reproduces a set of experimental data, including postsynaptic responses of single neurons on extracellular electric stimulation and spatially distributed activity patterns in response to visual stimulation. In the model, synaptically interacting excitatory and inhibitory neuronal populations are described by a conductance-based refractory density approach. Populations of two-compartment excitatory and inhibitory neurons in cortical layers 2/3 and 4 are distributed in the 2-d cortical space and connected by AMPA, NMDA and GABA type synapses. The external connections are pinwheel-like, according to the orientation of a stimulus. Intracortical connections are isotropic local and patchy between neurons with similar orientations. The model proposes better temporal resolution and more detailed elaboration than conventional mean-field models. In comparison to large network simulations, it excludes a posteriori statistical data manipulation and provides better computational efficiency and minimal parametrization.

Beyond inhibition: lateral modulation of plasticity of feedforward synapses in a spiking model of V1

Lateral inhibition is typically used to repel neural receptive fields. Here we introduce an additional learning mechanism that modifies the plasticity for feedforward synapses based on lateral interactions. We show a model, based on evidence from biological recordings[1], in which a heterosynaptic learning rule in conjunction with lateral inhibition introduces competition among neurons for input features. We demonstrate an STDP learning rule for feedforward connections to a spiking neuron where plasticity is modulated by activity of neighboring neurons. We apply the learning rule to a spiking model of primary visual cortex. In our model, input to V1 cortical neurons comes from the magnocellular pathway of a simulated spiking retina responding to saccades over a natural image. A group of neurons in V1 respond to a particular input feature and convey information that they have spiked to neighboring neurons by way of specialized lateral connections. The connections convey a direct signal of recent spiking activity. Alternatively, lateral inhibitory synapses and the level of inhibition to a neuron can also be used as this signal. If this recent neighbor activity signal is high, feedforward plasticity to the neuron becomes a simple flat depression as opposed to regular exponential STDP, as shown in Figure ​Figure1A.1A. Thus, late spiking neurons are prevented from developing receptive fields similar to those of earlier spiking neurons for a given input feature. This rule in conjunction with lateral inhibition and slow weight updrift ensures competition between neurons for features over a long timescale. Figure 1 A. Detail of the heterosynaptic rule. Neurons N1 and N2 fire early for the input, and their feedforward synapses undergo normal STDP. They send an activity signal to N3 and N4, whose feedforward synaptic weights are then depressed to prevent learning ... Our V1 model is built on a spatially distributed grid of single-compartment spiking neurons with parameters configured to model regular spiking cortical pyramidal cells, as detailed in [2]. The model has plastic excitatory feedforward connections and local lateral inhibition between spatially proximal neurons. In our model, we observe a better overall coverage of orientation selectivity of V1 Layer 4 neurons with our heterosynaptic rule than without it. We find a larger range of orientations and less redundancy of receptive field features between neighboring neurons, as shown in Figure ​Figure1B.1B. Such a rule could be used in a generic spiking cortical architecture to enforce independence of neural receptive fields.

Visual Perception: One World from Two Eyes

Binocular vision requires us to match up the different views of the world seen by each eye. Computational models of primary visual cortex describe how the brain begins this process. Recurrent connections help suppress the response to false matches.

Perceptual Decision Making “Through the Eyes” of a Large-Scale Neural Model of V1

Sparse coding has been posited as an efficient information processing strategy employed by sensory systems, particularly visual cortex. Substantial theoretical and experimental work has focused on the issue of sparse encoding, namely how the early visual system maps the scene into a sparse representation. In this paper we investigate the complementary issue of sparse decoding, for example given activity generated by a realistic mapping of the visual scene to neuronal spike trains, how do downstream neurons best utilize this representation to generate a “decision.” Specifically we consider both sparse (L1-regularized) and non-sparse (L2 regularized) linear decoding for mapping the neural dynamics of a large-scale spiking neuron model of primary visual cortex (V1) to a two alternative forced choice (2-AFC) perceptual decision. We show that while both sparse and non-sparse linear decoding yield discrimination results quantitatively consistent with human psychophysics, sparse linear decoding is more efficient in terms of the number of selected informative dimension.