Adaptive learning in a compartmental model of visual cortex—how feedback enables stable category learning and refinement

The categorization of real world objects is often reflected in the similarity of their visual appearances. Such categories of objects do not necessarily form disjunct sets of objects, neither semantically nor visually. The relationship between categories can often be described in terms of a hierarchical structure. For instance, tigers and leopards build two separate mammalian categories, but both belong to the category of felines. In other words, tigers and leopards are subcategories of the category Felidae. In the last decades, the unsupervised learning of categories of visual input stimuli has been addressed by numerous approaches in machine learning as well as in the computational neurosciences. However, the question of what kind of mechanisms might be involved in the process of subcategory learning, or category refinement, remains a topic of active investigation. We propose a recurrent computational network architecture for the unsupervised learning of categorial and subcategorial visual input representations. During learning, the connection strengths of bottom-up weights from input to higher-level category representations are adapted according to the input activity distribution. In a similar manner, top-down weights learn to encode the characteristics of a specific stimulus category. Feedforward and feedback learning in combination realize an associative memory mechanism, enabling the selective top-down propagation of a category's feedback weight distribution. We suggest that the difference between the expected input encoded in the projective field of a category node and the current input pattern controls the amplification of feedforward-driven representations. Large enough differences trigger the recruitment of new representational resources and the establishment of (sub-) category representations. We demonstrate the temporal evolution of such learning and show how the approach successully establishes category and subcategory representations

A Theory of Object Recognition: Computations and Circuits in the Feedforward Path of the Ventral Stream in Primate Visual Cortex

We describe a quantitative theory to account for the computations performed by the feedforward path of the ventral stream of visual cortex and the local circuits implementing them. We show that a model instantiating the theory is capable of performing recognition on datasets of complex images at the level of human observers in rapid categorization tasks. We also show that the theory is consistent with (and in some case has predicted) several properties of neurons in V1, V4, IT and PFC. The theory seems sufficiently comprehensive, detailed and satisfactory to represent an interesting challenge for physiologists and modelers: either disprove its basic features or propose alternative theories of equivalent scope. The theory suggests a number of open questions for visual physiology and psychophysics

Optogenetic interrogation of primary visual cortex and its impact on neural coding and behavior

Understanding the mechanism by which the brain transforms simple sensory inputs into rich perceptual experiences is one of the great mysteries of systems neuroscience. Undoubtedly this involves the activity of large populations of interconnected neurons, but while the responses of individual neurons to a variety of sensory stimuli have been well-characterized, how populations of such neurons organize their activity to create our sensory perceptions is almost entirely unknown. To investigate this complex circuitry requires the ability to causally manipulate the activity of neural populations and monitor the resultant effects. Here we focus on primary visual cortex (V1), which has been shown to be crucial for visual perception, and utilize optogenetic tools to render the activity of genetically- defined neural populations sensitive to light. By simultaneously recording and modulating (either driving or silencing) the activity of excitatory (glutamatergic) neurons, we are able to causally examine their role in visual perception. Here we report 3 major findings. First, we show that activating subpopulations of excitatory neurons can improve visual perception under certain conditions and that information in V1 used for perceptual decisions is integrated across spatially-limited populations of neurons. Further, we show that a key signature of this information integration is a reduction in correlated variability between neurons. Correlated variability has been implicated as a major source of behavioral choice related activity in the cortex, and theorized to be a major factor limiting information in cortical populations. However, until now, there has not been a way to manipulate correlations without altering firing rates or other task related variables. Here we demonstrate a novel method using optogenetic stimulation to causally manipulate correlated variability between cortical neurons without altering their firing rates. Lastly, with the goal of expanding the currently limited repertoire of optogenetic tools for non-human primates, we establish the viability of a novel optogenetic construct capable of dramatically silencing neural populations using a recently discovered anion conducting channelrhodopsin

Sensor fusion in distributed cortical circuits

The substantial motion of the nature is to balance, to survive, and to reach perfection. The evolution in biological systems is a key signature of this quintessence. Survival cannot be achieved without understanding the surrounding world. How can a fruit fly live without searching for food, and thereby with no form of perception that guides the behavior? The nervous system of fruit fly with hundred thousand of neurons can perform very complicated tasks that are beyond the power of an advanced supercomputer. Recently developed computing machines are made by billions of transistors and they are remarkably fast in precise calculations. But these machines are unable to perform a single task that an insect is able to do by means of thousands of neurons. The complexity of information processing and data compression in a single biological neuron and neural circuits are not comparable with that of developed today in transistors and integrated circuits. On the other hand, the style of information processing in neural systems is also very different from that of employed by microprocessors which is mostly centralized. Almost all cognitive functions are generated by a combined effort of multiple brain areas. In mammals, Cortical regions are organized hierarchically, and they are reciprocally interconnected, exchanging the information from multiple senses. This hierarchy in circuit level, also preserves the sensory world within different levels of complexity and within the scope of multiple modalities. The main behavioral advantage of that is to understand the real-world through multiple sensory systems, and thereby to provide a robust and coherent form of perception. When the quality of a sensory signal drops, the brain can alternatively employ other information pathways to handle cognitive tasks, or even to calibrate the error-prone sensory node. Mammalian brain also takes a good advantage of multimodal processing in learning and development; where one sensory system helps another sensory modality to develop. Multisensory integration is considered as one of the main factors that generates consciousness in human. Although, we still do not know where exactly the information is consolidated into a single percept, and what is the underpinning neural mechanism of this process? One straightforward hypothesis suggests that the uni-sensory signals are pooled in a ploy-sensory convergence zone, which creates a unified form of perception. But it is hard to believe that there is just one single dedicated region that realizes this functionality. Using a set of realistic neuro-computational principles, I have explored theoretically how multisensory integration can be performed within a distributed hierarchical circuit. I argued that the interaction of cortical populations can be interpreted as a specific form of relation satisfaction in which the information preserved in one neural ensemble must agree with incoming signals from connected populations according to a relation function. This relation function can be seen as a coherency function which is implicitly learnt through synaptic strength. Apart from the fact that the real world is composed of multisensory attributes, the sensory signals are subject to uncertainty. This requires a cortical mechanism to incorporate the statistical parameters of the sensory world in neural circuits and to deal with the issue of inaccuracy in perception. I argued in this thesis how the intrinsic stochasticity of neural activity enables a systematic mechanism to encode probabilistic quantities within neural circuits, e.g. reliability, prior probability. The systematic benefit of neural stochasticity is well paraphrased by the problem of Duns Scotus paradox: imagine a donkey with a deterministic brain that is exposed to two identical food rewards. This may make the animal suffer and die starving because of indecision. In this thesis, I have introduced an optimal encoding framework that can describe the probability function of a Gaussian-like random variable in a pool of Poisson neurons. Thereafter a distributed neural model is proposed that can optimally combine conditional probabilities over sensory signals, in order to compute Bayesian Multisensory Causal Inference. This process is known as a complex multisensory function in the cortex. Recently it is found that this process is performed within a distributed hierarchy in sensory cortex. Our work is amongst the first successful attempts that put a mechanistic spotlight on understanding the underlying neural mechanism of Multisensory Causal Perception in the brain, and in general the theory of decentralized multisensory integration in sensory cortex. Engineering information processing concepts in the brain and developing new computing technologies have been recently growing. Neuromorphic Engineering is a new branch that undertakes this mission. In a dedicated part of this thesis, I have proposed a Neuromorphic algorithm for event-based stereoscopic fusion. This algorithm is anchored in the idea of cooperative computing that dictates the defined epipolar and temporal constraints of the stereoscopic setup, to the neural dynamics. The performance of this algorithm is tested using a pair of silicon retinas

Sensor fusion in distributed cortical circuits

The substantial motion of the nature is to balance, to survive, and to reach perfection. The evolution in biological systems is a key signature of this quintessence. Survival cannot be achieved without understanding the surrounding world. How can a fruit fly live without searching for food, and thereby with no form of perception that guides the behavior? The nervous system of fruit fly with hundred thousand of neurons can perform very complicated tasks that are beyond the power of an advanced supercomputer. Recently developed computing machines are made by billions of transistors and they are remarkably fast in precise calculations. But these machines are unable to perform a single task that an insect is able to do by means of thousands of neurons. The complexity of information processing and data compression in a single biological neuron and neural circuits are not comparable with that of developed today in transistors and integrated circuits. On the other hand, the style of information processing in neural systems is also very different from that of employed by microprocessors which is mostly centralized. Almost all cognitive functions are generated by a combined effort of multiple brain areas. In mammals, Cortical regions are organized hierarchically, and they are reciprocally interconnected, exchanging the information from multiple senses. This hierarchy in circuit level, also preserves the sensory world within different levels of complexity and within the scope of multiple modalities. The main behavioral advantage of that is to understand the real-world through multiple sensory systems, and thereby to provide a robust and coherent form of perception. When the quality of a sensory signal drops, the brain can alternatively employ other information pathways to handle cognitive tasks, or even to calibrate the error-prone sensory node. Mammalian brain also takes a good advantage of multimodal processing in learning and development; where one sensory system helps another sensory modality to develop. Multisensory integration is considered as one of the main factors that generates consciousness in human. Although, we still do not know where exactly the information is consolidated into a single percept, and what is the underpinning neural mechanism of this process? One straightforward hypothesis suggests that the uni-sensory signals are pooled in a ploy-sensory convergence zone, which creates a unified form of perception. But it is hard to believe that there is just one single dedicated region that realizes this functionality. Using a set of realistic neuro-computational principles, I have explored theoretically how multisensory integration can be performed within a distributed hierarchical circuit. I argued that the interaction of cortical populations can be interpreted as a specific form of relation satisfaction in which the information preserved in one neural ensemble must agree with incoming signals from connected populations according to a relation function. This relation function can be seen as a coherency function which is implicitly learnt through synaptic strength. Apart from the fact that the real world is composed of multisensory attributes, the sensory signals are subject to uncertainty. This requires a cortical mechanism to incorporate the statistical parameters of the sensory world in neural circuits and to deal with the issue of inaccuracy in perception. I argued in this thesis how the intrinsic stochasticity of neural activity enables a systematic mechanism to encode probabilistic quantities within neural circuits, e.g. reliability, prior probability. The systematic benefit of neural stochasticity is well paraphrased by the problem of Duns Scotus paradox: imagine a donkey with a deterministic brain that is exposed to two identical food rewards. This may make the animal suffer and die starving because of indecision. In this thesis, I have introduced an optimal encoding framework that can describe the probability function of a Gaussian-like random variable in a pool of Poisson neurons. Thereafter a distributed neural model is proposed that can optimally combine conditional probabilities over sensory signals, in order to compute Bayesian Multisensory Causal Inference. This process is known as a complex multisensory function in the cortex. Recently it is found that this process is performed within a distributed hierarchy in sensory cortex. Our work is amongst the first successful attempts that put a mechanistic spotlight on understanding the underlying neural mechanism of Multisensory Causal Perception in the brain, and in general the theory of decentralized multisensory integration in sensory cortex. Engineering information processing concepts in the brain and developing new computing technologies have been recently growing. Neuromorphic Engineering is a new branch that undertakes this mission. In a dedicated part of this thesis, I have proposed a Neuromorphic algorithm for event-based stereoscopic fusion. This algorithm is anchored in the idea of cooperative computing that dictates the defined epipolar and temporal constraints of the stereoscopic setup, to the neural dynamics. The performance of this algorithm is tested using a pair of silicon retinas

The Role of Bottom-Up and Top-Down Cortical Interactions in Adaptation to Natural Scene Statistics

Adaptation is a mechanism by which cortical neurons adjust their responses according to recently viewed stimuli. Visual information is processed in a circuit formed by feedforward (FF) and feedback (FB) synaptic connections of neurons in different cortical layers. Here, the functional role of FF-FB streams and their synaptic dynamics in adaptation to natural stimuli is assessed in psychophysics and neural model. We propose a cortical model which predicts psychophysically observed motion adaptation aftereffects (MAE) after exposure to geometrically distorted natural image sequences. The model comprises direction selective neurons in V1 and MT connected by recurrent FF and FB dynamic synapses. Psychophysically plausible model MAEs were obtained from synaptic changes within neurons tuned to salient direction signals of the broadband natural input. It is conceived that, motion disambiguation by FF-FB interactions is critical to encode this salient information. Moreover, only FF-FB dynamic synapses operating at distinct rates predicted psychophysical MAEs at different adaptation time-scales which could not be accounted for by single rate dynamic synapses in either of the streams. Recurrent FF-FB pathways thereby play a role during adaptation in a natural environment, specifically in inducing multilevel cortical plasticity to salient information and in mediating adaptation at different time-scales