362 research outputs found
Neuromodulatory effects on early visual signal processing
Understanding how the brain processes information and generates simple to complex behavior constitutes one of the core objectives in systems neuroscience. However, when studying different neural circuits, their dynamics and interactions researchers often assume fixed connectivity, overlooking a crucial factor - the effect of neuromodulators. Neuromodulators can modulate circuit activity depending on several aspects, such as different brain states or sensory contexts. Therefore, considering the modulatory effects of neuromodulators on the functionality of neural circuits is an indispensable step towards a more complete picture of the brain’s ability to process information. Generally, this issue affects all neural systems; hence this thesis tries to address this with an experimental and computational approach to resolve neuromodulatory effects on cell type-level in a well-define system, the mouse retina. In the first study, we established and applied a machine-learning-based classification algorithm to identify individual functional retinal ganglion cell types, which enabled detailed cell type-resolved analyses. We applied the classifier to newly acquired data of light-evoked retinal ganglion cell responses and successfully identified their functional types. Here, the cell type-resolved analysis revealed that a particular principle of efficient coding applies to all types in a similar way. In a second study, we focused on the issue of inter-experimental variability that can occur during the process of pooling datasets. As a result, further downstream analyses may be complicated by the subtle variations between the individual datasets. To tackle this, we proposed a theoretical framework based on an adversarial autoencoder with the objective to remove inter-experimental variability from the pooled dataset, while preserving the underlying biological signal of interest. In the last study of this thesis, we investigated the functional effects of the neuromodulator nitric oxide on the retinal output signal. To this end, we used our previously developed retinal ganglion cell type classifier to unravel type-specific effects and established a paired recording protocol to account for type-specific time-dependent effects. We found that certain
retinal ganglion cell types showed adaptational type-specific changes and that nitric oxide had a distinct modulation of a particular group of retinal ganglion cells.
In summary, I first present several experimental and computational methods that allow to
study functional neuromodulatory effects on the retinal output signal in a cell type-resolved manner and, second, use these tools to demonstrate their feasibility to study the neuromodulator nitric oxide
Towards Neuromorphic Gradient Descent: Exact Gradients and Low-Variance Online Estimates for Spiking Neural Networks
Spiking Neural Networks (SNNs) are biologically-plausible models that can run on low-powered non-Von Neumann neuromorphic hardware, positioning them as promising alternatives to conventional Deep Neural Networks (DNNs) for energy-efficient edge computing and robotics. Over the past few years, the Gradient Descent (GD) and Error Backpropagation (BP) algorithms used in DNNs have inspired various training methods for SNNs. However, the non-local and the reverse nature of BP, combined with the inherent non-differentiability of spikes, represent fundamental obstacles to computing gradients with SNNs directly on neuromorphic hardware. Therefore, novel approaches are required to overcome the limitations of GD and BP and enable online gradient computation on neuromorphic hardware.
In this thesis, I address the limitations of GD and BP with SNNs by proposing three algorithms. First, I extend a recent method that computes exact gradients with temporally-coded SNNs by relaxing the firing constraint of temporal coding and allowing multiple spikes per neuron. My proposed method generalizes the computation of exact gradients with SNNs and enhances the tradeoffs between performance and various other aspects of spiking neurons. Next, I introduce a novel alternative to BP that computes low-variance gradient estimates in a local and online manner. Compared to other alternatives to BP, the proposed method demonstrates an improved convergence rate and increased performance with DNNs. Finally, I combine these two methods and propose an algorithm that estimates gradients with SNNs in a manner that is compatible with the constraints of neuromorphic hardware. My empirical results demonstrate the effectiveness of the resulting algorithm in training SNNs without performing BP
Exploring space situational awareness using neuromorphic event-based cameras
The orbits around earth are a limited natural resource and one that hosts a vast range of vital space-based systems that support international systems use by both commercial industries, civil organisations, and national defence. The availability of this space resource is rapidly depleting due to the ever-growing presence of space debris and rampant overcrowding, especially in the limited and highly desirable slots in geosynchronous orbit. The field of Space Situational Awareness encompasses tasks aimed at mitigating these hazards to on-orbit systems through the monitoring of satellite traffic. Essential to this task is the collection of accurate and timely observation data. This thesis explores the use of a novel sensor paradigm to optically collect and process sensor data to enhance and improve space situational awareness tasks. Solving this issue is critical to ensure that we can continue to utilise the space environment in a sustainable way. However, these tasks pose significant engineering challenges that involve the detection and characterisation of faint, highly distant, and high-speed targets. Recent advances in neuromorphic engineering have led to the availability of high-quality neuromorphic event-based cameras that provide a promising alternative to the conventional cameras used in space imaging. These cameras offer the potential to improve the capabilities of existing space tracking systems and have been shown to detect and track satellites or ‘Resident Space Objects’ at low data rates, high temporal resolutions, and in conditions typically unsuitable for conventional optical cameras. This thesis presents a thorough exploration of neuromorphic event-based cameras for space situational awareness tasks and establishes a rigorous foundation for event-based space imaging. The work conducted in this project demonstrates how to enable event-based space imaging systems that serve the goals of space situational awareness by providing accurate and timely information on the space domain. By developing and implementing event-based processing techniques, the asynchronous operation, high temporal resolution, and dynamic range of these novel sensors are leveraged to provide low latency target acquisition and rapid reaction to challenging satellite tracking scenarios. The algorithms and experiments developed in this thesis successfully study the properties and trade-offs of event-based space imaging and provide comparisons with traditional observing methods and conventional frame-based sensors. The outcomes of this thesis demonstrate the viability of event-based cameras for use in tracking and space imaging tasks and therefore contribute to the growing efforts of the international space situational awareness community and the development of the event-based technology in astronomy and space science applications
Insect neuroethology of reinforcement learning
Historically, reinforcement learning is a branch of machine learning founded on observations of how animals learn. This involved collaboration between the fields of biology and artificial intelligence that was beneficial to both fields, creating smarter artificial agents and improving the understanding of how biological systems function. The evolution of reinforcement learning during the past few years was rapid but substantially diverged from providing insights into how biological systems work, opening a gap between reinforcement learning and biology. In an attempt to close this gap, this thesis studied the insect neuroethology of reinforcement learning, that is, the neural circuits that underlie reinforcement-learning-related behaviours in insects. The goal was to extract a biologically plausible plasticity function from insect-neuronal data, use this to explain biological findings and compare it to more standard reinforcement
learning models. Consequently, a novel dopaminergic plasticity rule was developed to approximate the function of dopamine as the plasticity mechanism between neurons in the insect brain. This allowed a range of observed learning phenomena to happen in parallel, like memory depression, potentiation, recovery, and saturation. In addition, by using anatomical data of connections between neurons in the mushroom body neuropils of the insect brain, the neural incentive circuit of dopaminergic and output neurons was also explored. This, together with the dopaminergic plasticity rule, allowed for dynamic collaboration amongst parallel memory functions, such as acquisition, transfer, and forgetting. When tested on olfactory conditioning paradigms, the model reproduced the observed changes in the activity of the identified neurons in fruit flies. It also replicated the observed behaviour of the animals and it allowed for flexible behavioural control. Inspired by the visual navigation system of desert ants, the model was further challenged in the visual place recognition task. Although a relatively simple encoding of the olfactory information was sufficient to explain odour learning, a more sophisticated encoding of the visual input was required to increase the separability among the visual inputs and enable visual place recognition. Signal whitening and sparse combinatorial encoding were sufficient to boost the performance of the system in this task. The incentive circuit enabled the encoding of increasing familiarity along a known route, which dropped proportionally to the distance of the animal from that route. Finally, the proposed model was challenged in delayed reinforcement tasks, suggesting that it might take the role of an adaptive critic in the context of reinforcement learning
Unsupervised space-time learning in primary visual cortex
The mammalian visual system is an incredibly complex computation device, capable of performing the various tasks of seeing: navigation, pattern and object recognition, motor coordination, trajectory extrapolation, among others. Decades of research has shown that experience-dependent plasticity of cortical circuitry underlies the impressive ability to rapidly learn many of these tasks and to adjust as required. One particular thread of investigation has focused on unsupervised learning, wherein changes to the visual environment lead to corresponding changes in cortical circuits. The most prominent example of unsupervised learning is ocular dominance plasticity, caused by visual deprivation to one eye and leading to a dramatic re-wiring of cortex. Other examples tend to make more subtle changes to the visual environment through passive exposure to novel visual stimuli. Here, we use one such unsupervised paradigm, sequence learning, to study experience-dependent plasticity in the mouse visual system. Through a combination of theory and experiment, we argue that the mammalian visual system is an unsupervised learning device.
Beginning with a mathematical exploration of unsupervised learning in biology, engineering, and machine learning, we seek a more precise expression of our fundamental hypothesis. We draw connections between information theory, efficient coding, and common unsupervised learning algorithms such as Hebbian plasticity and principal component analysis. Efficient coding suggests a simple rule for transmitting information in the nervous system: use more spikes to encode unexpected information, and fewer spikes to encode expected information. Therefore, expectation violations ought to produce prediction errors, or brief periods of heightened firing when an unexpected event occurs. Meanwhile, modern unsupervised learning algorithms show how such expectations can be learned.
Next, we review data from decades of visual neuroscience research, highlighting the computational principles and synaptic plasticity processes that support biological learning and seeing. By tracking the flow of visual information from the retina to thalamus and primary visual cortex, we discuss how the principle of efficient coding is evident in neural activity. One common example is predictive coding in the retina, where ganglion cells with canonical center-surround receptive fields compute a prediction error, sending spikes to the central nervous system only in response to locally-unpredictable visual stimuli. This behavior can be learned through simple Hebbian plasticity mechanisms. Similar models explain much of the activity of neurons in primary visual cortex, but we also discuss ways in which the theory fails to capture the rich biological complexity.
Finally, we present novel experimental results from physiological investigations of the mouse primary visual cortex. We trained mice by passively exposing them to complex spatiotemporal patterns of light: rapidly-flashed sequences of images. We find evidence that visual cortex learns these sequences in a manner consistent with efficient coding, such that unexpected stimuli tend to elicit more firing than expected ones. Overall, we observe dramatic changes in evoked neural activity across days of passive exposure. Neural responses to the first, unexpected sequence element increase with days of training while responses at other, expected time points either decrease or stay the same. Furthermore, substituting an unexpected element for an expected one or omitting an expected element both cause brief bursts of increased firing. Our results therefore provide evidence for unsupervised learning and efficient coding in the mouse visual system, especially because unexpected events drive prediction errors. Overall, our analysis suggests novel experiments, which could be performed in the near future, and provides a useful framework to understand visual perception and learning
Artificial Dendritic Neuron: A Model of Computation and Learning Algorithm
Dendrites are root-like extensions from the neuron cell body and have long been thought to serve as the predominant input structures of neurons. Since the early twentieth century, neuroscience research has attempted to define the dendrite’s contribution to neural computation and signal integration. This body of experimental and modeling research strongly indicates that dendrites are not just input structures but are crucial to neural processing. Dendritic processing consists of both active and passive elements that utilize the spatial, electrical and connective properties of the dendritic tree.
This work presents a neuron model based around the structure and properties of dendrites. This research assesses the computational benefits and requirements of adding dendrites to a spiking artificial neuron model. A list of the computational properties of actual dendrites that have shaped this work is given. An algorithm capable of generating and training a network of dendritic neurons is created as an investigative tool through which computational challenges and attributes are explored.
This work assumes that dendrites provide a necessary and beneficial function to biological intelligence (BI) and their translation into the artificial intelligence (AI) realm would broaden the capabilities and improve the realism of artificial neural network (ANN) research. To date there have been only a few instances in which neural network-based AI research has ventured beyond the point neuron; therefore, the work presented here should be viewed as exploratory. The contribution to AI made by this work is an implementation of the artificial dendritic (AD) neuron model and an algorithm for training AD neurons with spatially distributed inputs with dendrite-like connectivity
Brain Computations and Connectivity [2nd edition]
This is an open access title available under the terms of a CC BY-NC-ND 4.0 International licence. It is free to read on the Oxford Academic platform and offered as a free PDF download from OUP and selected open access locations.
Brain Computations and Connectivity is about how the brain works. In order to understand this, it is essential to know what is computed by different brain systems; and how the computations are performed.
The aim of this book is to elucidate what is computed in different brain systems; and to describe current biologically plausible computational approaches and models of how each of these brain systems computes.
Understanding the brain in this way has enormous potential for understanding ourselves better in health and in disease. Potential applications of this understanding are to the treatment of the brain in disease; and to artificial intelligence which will benefit from knowledge of how the brain performs many of its extraordinarily impressive functions.
This book is pioneering in taking this approach to brain function: to consider what is computed by many of our brain systems; and how it is computed, and updates by much new evidence including the connectivity of the human brain the earlier book: Rolls (2021) Brain Computations: What and How, Oxford University Press.
Brain Computations and Connectivity will be of interest to all scientists interested in brain function and how the brain works, whether they are from neuroscience, or from medical sciences including neurology and psychiatry, or from the area of computational science including machine learning and artificial intelligence, or from areas such as theoretical physics
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Understanding the molecular, cellular, and circuit defects characterizing the early stages of Alzheimer’s disease
One of the most foundational and personal philosophical questions one can ask is what makes you, you? In large part, you are made up of your relationships, experiences, and memories. The hippocampus, a brain region which is critical for the formation of memories, has been the focus of neuroscience research for decades due partially to this function, which is foundational to our individuality. In Alzheimer’s disease (AD), one of the most common and well-researched neurodegenerative diseases in the world, the hippocampus is one of the earliest targets. Despite extensive work on AD, we still lack a coherent understanding of what is causing the disease, the mechanisms by which it is causing neuronal dysfunction and death within the hippocampus and other brain regions, and how it ultimately causes deficits in cognition and behavior, leading to an erosion of our selves.
In this thesis, I explore three independent but related questions: 1) what molecular mechanisms are causing early synaptic loss in AD, specifically within the hippocampus, 2) what molecular effectors are responsible for establishing and maintaining intracellular architecture in hippocampal neurons, which are exploited in early AD, and 3) how and when does the hippocampal circuit dysfunction in AD progression?
Using a variety of experimental techniques, ranging from in utero and ex utero electroporation, primary murine and human neuronal cell culture, longitudinal confocal microscopy, immunohistochemistry, biochemistry, cell and molecular biology, in vivo two-photon calcium imaging, and behavioral assays, I have found that, within CA1 of the hippocampus, synapse loss requires degradation of the dendritic mitochondrial network, activity and input specificity are driving mitochondrial compartmentalization within CA1 neurons through the same pathway that is aberrantly overactivated in AD, and the hippocampal circuit is overly rigid in encoding the environment as the disease progresses
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