There is substantial experimental evidence that learning and memory-related behaviours rely on local synaptic changes, but the search for distinct plasticity rules has been driven by human intuition, with limited success for multiple, co-active plasticity rules in biological networks. More recently, automated meta-learning approaches have been used in simplified settings, such as rate networks and small feed-forward spiking networks. Here, we develop a simulation-based inference (SBI) method for sequentially filtering plasticity rules through an increasingly fine mesh of constraints that can be modified on-the-fly. This method, _filter SBI_, allows us to infer entire families of complex and co-active plasticity rules in spiking networks. We first consider flexibly parameterized doublet (Hebbian) rules, and find that the set of inferred rules contains solutions that extend and refine -and also reject- predictions from mean-field theory. Next, we expand the search space of plasticity rules by modelling them as multi-layer perceptrons that combine several plasticity-relevant factors, such as weight, voltage, triplets and co-dependency. Out of the millions of possible rules, we identify thousands of unique rule combinations that satisfy biological constraints like plausible activity and weight dynamics. The resulting rules can be used as a starting point for further investigations into specific network computations, and already suggest refinements and predictions for classical experimental approaches on plasticity. This flexible approach for principled exploration of complex plasticity rules in large recurrent spiking networks presents the most advanced search tool to date for enabling robust predictions and deep insights into the plasticity mechanisms underlying brain function.

Neuronal networks encode information through patterns of activity that define the networks' function. The neurons' activity relies on specific connectivity structures, yet the link between structure and function is not fully understood. Here, we tackle this structure-function problem with a new conceptual approach. Instead of manipulating the connectivity directly, we focus on upper triangular matrices, which represent the network dynamics in a given orthonormal basis obtained by the Schur decomposition. This abstraction allows us to independently manipulate the eigenspectrum and feedforward structures of a connectivity matrix. Using this method, we describe a diverse repertoire of non-normal transient amplification, and to complement the analysis of the dynamical regimes, we quantify the geometry of output trajectories through the effective rank of both the eigenvector and the dynamics matrices. Counter-intuitively, we find that shrinking the eigenspectrum's imaginary distribution leads to highly amplifying regimes in linear and long-lasting dynamics in nonlinear networks. We also find a trade-off between amplification and dimensionality of neuronal dynamics, i.e., trajectories in neuronal state-space. Networks that can amplify a large number of orthogonal initial conditions produce neuronal trajectories that lie in the same subspace of the neuronal state-space. Finally, we examine networks of excitatory and inhibitory neurons. We find that the strength of global inhibition is directly linked with the amplitude of amplification, such that weakening inhibitory weights also decreases amplification, and that the eigenspectrum's imaginary distribution grows with an increase in the ratio between excitatory-to-inhibitory and excitatory-to-excitatory connectivity strengths. Consequently, the strength of global inhibition reveals itself as a strong signature for amplification and a potential control mechanism to switch dynamical regimes. Our results shed a light on how biological networks, i.e., networks constrained by Dale's law, may be optimised for specific dynamical regimes.

Neuronal computations depend on synaptic connectivity and intrinsic electrophysiological properties. Synaptic connectivity determines which inputs from presynaptic neurons are integrated, while cellular properties determine how inputs are filtered over time. Unlike their biological counterparts, most computational approaches to learning in simulated neural networks are limited to changes in synaptic connectivity. However, if intrinsic parameters change, neural computations are altered drastically. Here, we include the parameters that determine the intrinsic properties, e.g., time constants and reset potential, into the learning paradigm. Using sparse feedback signals that indicate target spike times, and gradient-based parameter updates, we show that the intrinsic parameters can be learned along with the synaptic weights to produce specific input-output functions. Specifically, we use a teacher-student paradigm in which a randomly initialised leaky integrate-and-fire or resonate-and-fire neuron must recover the parameters of a teacher neuron. We show that complex temporal functions can be learned online and without backpropagation through time, relying on event-based updates only. Our results are a step towards online learning of neural computations from ungraded and unsigned sparse feedback signals with a biologically inspired learning mechanism.

Brains process information in spiking neural networks. Their intricate connections shape the diverse functions these networks perform. Yet how network connectivity relates to function is poorly understood, and the functional capabilities of models of spiking networks are still rudimentary. The lack of both theoretical insight and practical algorithms to find the necessary connectivity poses a major impediment to both studying information processing in the brain and building efficient neuromorphic hardware systems. The training algorithms that solve this problem for artificial neural networks typically rely on gradient descent. But doing so in spiking networks has remained challenging due to the nondifferentiable nonlinearity of spikes. To avoid this issue, one can employ surrogate gradients to discover the required connectivity. However, the choice of a surrogate is not unique, raising the question of how its implementation influences the effectiveness of the method. Here, we use numerical simulations to systematically study how essential design parameters of surrogate gradients affect learning performance on a range of classification problems. We show that surrogate gradient learning is robust to different shapes of underlying surrogate derivatives, but the choice of the derivative's scale can substantially affect learning performance. When we combine surrogate gradients with suitable activity regularization techniques, spiking networks perform robust information processing at the sparse activity limit. Our study provides a systematic account of the remarkable robustness of surrogate gradient learning and serves as a practical guide to model functional spiking neural networks.

The search for biologically faithful synaptic plasticity rules has resulted in a large body of models. They are usually inspired by -- and fitted to -- experimental data, but they rarely produce neural dynamics that serve complex functions. These failures suggest that current plasticity models are still under-constrained by existing data. Here, we present an alternative approach that uses meta-learning to discover plausible synaptic plasticity rules. Instead of experimental data, the rules are constrained by the functions they implement and the structure they are meant to produce. Briefly, we parameterize synaptic plasticity rules by a Volterra expansion and then use supervised learning methods (gradient descent or evolutionary strategies) to minimize a problem-dependent loss function that quantifies how effectively a candidate plasticity rule transforms an initially random network into one with the desired function. We first validate our approach by re-discovering previously described plasticity rules, starting at the single-neuron level and ``Oja’s rule'', a simple Hebbian plasticity rule that captures the direction of most variability of inputs to a neuron (i.e., the first principal component). We expand the problem to the network level and ask the framework to find Oja’s rule together with an anti-Hebbian rule such that an initially random two-layer firing-rate network will recover several principal components of the input space after learning. Next, we move to networks of integrate-and-fire neurons with plastic inhibitory afferents. We train for rules that achieve a target firing rate by countering tuned excitation. Our algorithm discovers a specific subset of the manifold of rules that can solve this task. Our work is a proof of principle of an automated and unbiased approach to unveil synaptic plasticity rules that obey biological constraints and can solve complex functions.

Understanding the brain is one of the most challenging scientific problems faced by humanity.In addition to offering an improved understanding of who we are, deciphering how our brains operate will provide payoffs for the treatment of mental and neurological disease as well as the creation of intelligent systems.Consequently, developed countries are pouring ever-increasing research funds into brain science.Prominent examples include the BRAIN initiative in the United States and the Human Brain Project in Europe.Although Africa may not yet possess financial and technological resources on this scale, Africans represent a vast and growing human resource poised to make important contributions to neuroscience.As the third yearly International Brain Research Organisation (IBRO)-Simons Computational Neuroscience Imbizo recently wrapped up in South Africa, we call for greater efforts to empower Africans to join the global neuroscience community and argue that the subfield of computational neuroscience is the ideal vehicle to do so.Computational and theoretical neuroscience, also known as neurotheory, or just "theory" within neurobiology, is a young, rapidly expanding field within neuroscience; its goal is to determine and articulate the general principles by which nervous systems perform their computations.We think it is a good example of how theory, as a subfield, can serve as a catalyst to accelerate the development of science in Africa.Due to the diversity of methods required for understanding the brain at multiple levels [1], skills are leveraged from a range of fields, such as biology, computer science, electrical engineering, mathematics, physics, and psychology.Beyond just increasing our body of scientific knowledge, the impact of this interdisciplinary field is already significant to the clinic and industry: decoding algorithms are helping restore movement in paralyzed patients [2,3], and machine learning is continually influenced by engagement with neuroscience [4].A notable aspect of this growing field is its accessibility: computational neuroscience requires only data and individual expertise, independent of geographic location.There is growing momentum for experimentalists to make their data available for public access and for increased collaboration between groups, a movement lead by flagship projects that curate, systematize, and make freely available large-scale neuronal data [5-8].A parallel development has been the recent surge in open-access publishing and the establishment of alternative methods for accessing scientific papers outside of well-endowed institutions (e.g., ResearchGate, bioRxiv, and especially important in Africa, SciHub).These dual developments mean that aspiring African scientists in resource-limited settings have increasing access to the latest in both data and theory from across the broad field of neuroscience.From an African

Cortical circuits exhibit intricate recurrent architectures that are remarkably similar across different brain areas. Such stereotyped structure suggests the existence of common computational principles. However, such principles have remained largely elusive. Inspired by gated-memory networks, namely long short-term memory networks (LSTMs), we introduce a recurrent neural network in which information is gated through inhibitory cells that are subtractive (subLSTM). We propose a natural mapping of subLSTMs onto known canonical excitatory-inhibitory cortical microcircuits. Our empirical evaluation across sequential image classification and language modelling tasks shows that subLSTM units can achieve similar performance to LSTM units. These results suggest that cortical circuits can be optimised to solve complex contextual problems and proposes a novel view on their computational function. Overall our work provides a step towards unifying recurrent networks as used in machine learning with their biological counterparts.