Introduction: Machine learning provides fundamental tools both for scientific research and for the development of technologies with significant impact on society. It provides methods that facilitate the discovery of regularities in data and that give predictions without explicit knowledge of the rules governing a system. However, a price is paid for exploiting such flexibility: machine learning methods are typically black-boxes where it is difficult to fully understand what the machine is doing or how it is operating. This poses constraints on the applicability and explainability of such methods. Methods: Our research aims to open the black-box of recurrent neural networks, an important family of neural networks used for processing sequential data. We propose a novel methodology that provides a mechanistic interpretation of behaviour when solving a computational task. Our methodology uses mathematical constructs called excitable network attractors, which are invariant sets in phase space composed of stable attractors and excitable connections between them. Results and Discussion: As the behaviour of recurrent neural networks depends both on training and on inputs to the system, we introduce an algorithm to extract network attractors directly from the trajectory of a neural network while solving tasks. Simulations conducted on a controlled benchmark task confirm the relevance of these attractors for interpreting the behaviour of recurrent neural networks, at least for tasks that involve learning a finite number of stable states and transitions between them.as reservoir computing [32,33]. Echo state networks (ESNs) [21,30] constitute an important example of reservoir computing, where a recurrent layer (called a reservoir) is composed of a large number of neurons with randomly initialised connections that are not fine-tuned via gradient-based optimisation mechanisms. The main idea behind ESNs is to exploit the rich dynamics generated by the reservoir with an output layer, the read-out that is optimised to solve a specific task. Problem statement and research hypothesisThe high-dimensional and non-linear nature of RNNs complicates interpretability of their internal dynamics, which are characterised by complex, input-dependent spatio-temporal patterns of activity [47,55]. This poses constraints on understanding the behaviour of RNNs: they are usually viewed as black-boxes from which it is hard to extract useful knowledge about their inner workings. As highlighted by recent research efforts [10,24,40], similar interpretability issues affect many other machine learning methods. Furthermore, an increasing societal need to develop accountability and explainability of decision making by AI [17] is driving the development of methodologies for explaining the behaviour of such methods.Our aim in this paper is to develop effective models that capture the essential dynamical behaviour of RNNs on computational tasks as input-driven responses of a dynamical system, while neglecting microscopic details of the RNN dynamics in phase...

Coupling among neural rhythms is one of the most important mechanisms at the basis of cognitive processes in the brain. In this study, we consider a neural mass model, rigorously obtained from the microscopic dynamics of an inhibitory spiking network with exponential synapses, able to autonomously generate collective oscillations (COs). These oscillations emerge via a super-critical Hopf bifurcation, and their frequencies are controlled by the synaptic time scale, the synaptic coupling, and the excitability of the neural population. Furthermore, we show that two inhibitory populations in a master–slave configuration with different synaptic time scales can display various collective dynamical regimes: damped oscillations toward a stable focus, periodic and quasi-periodic oscillations, and chaos. Finally, when bidirectionally coupled, the two inhibitory populations can exhibit different types of θ–γ cross-frequency couplings (CFCs): phase-phase and phase-amplitude CFC. The coupling between θ and γ COs is enhanced in the presence of an external θ forcing, reminiscent of the type of modulation induced in hippocampal and cortex circuits via optogenetic drive.

A recurrent neural network (RNN) possesses the echo state property (ESP) if, for a given input sequence, it "forgets" any internal states of the driven (nonautonomous) system and asymptotically follows a unique, possibly complex trajectory. The lack of ESP is conventionally understood as a lack of reliable behaviour in RNNs. Here, we show that RNNs can reliably perform computations under a more general principle that accounts only for their local behaviour in phase space. To this end, we formulate a generalisation of the ESP and introduce an echo index to characterise the number of simultaneously stable responses of a driven RNN. We show that it is possible for the echo index to change with inputs, highlighting a potential source of computational errors in RNNs due to characteristics of the inputs driving the dynamics.

Coupling among neural rhythms is one of the most important mechanisms at the basis of cognitive processes in the brain. In this study we consider a neural mass model, rigorously obtained from the microscopic dynamics of an inhibitory spiking network with exponential synapses, able to autonomously generate collective oscillations (COs). These oscillations emerge via a super-critical Hopf bifurcation, and their frequencies are controlled by the synaptic time scale, the synaptic coupling and the excitability of the neural population. Furthermore, we show that two inhibitory populations in a master-slave configuration with different synaptic time scales can display various collective dynamical regimes: namely, damped oscillations towards a stable focus, periodic and quasi-periodic oscillations, and chaos. Finally, when bidirectionally coupled the two inhibitory populations can exhibit different types of θ-γ cross-frequency couplings (CFCs): namely, phase-phase and phase-amplitude CFC. The coupling between θ and γ COs is enhanced in presence of a external θ forcing, reminiscent of the type of modulation induced in Hippocampal and Cortex circuits via optogenetic drive.In healthy conditions, the brain’s activity reveals a series of intermingled oscillations, generated by large ensembles of neurons, which provide a functional substrate for information processing. How single neuron properties influence neuronal population dynamics is an unsolved question, whose solution could help in the understanding of the emergent collective behaviors arising during cognitive processes. Here we consider a neural mass model, which reproduces exactly the macroscopic activity of a network of spiking neurons. This mean-field model is employed to shade some light on an important and ubiquitous neural mechanism underlying information processing in the brain: the θ-γ cross-frequency coupling. In particular, we will explore in detail the conditions under which two coupled inhibitory neural populations can generate these functionally relevant coupled rhythms.

Since the recognition in the early nineties of the vanishing/exploding (V/E) gradient issue plaguing the training of neural networks (NNs), significant efforts have been exerted to overcome this obstacle. However, a clear solution to the V/E issue remained elusive so far. In this manuscript a new architecture of NN is proposed, designed to mathematically prevent the V/E issue to occur. The pursuit of approximate dynamical isometry, i.e. parameter configurations where the singular values of the input-output Jacobian are tightly distributed around 1, leads to the derivation of a NN's architecture that shares common traits with the popular Residual Network model. Instead of skipping connections between layers, the idea is to filter the previous activations orthogonally and add them to the nonlinear activations of the next layer, realising a convex combination between them. Remarkably, the impossibility for the gradient updates to either vanish or explode is demonstrated with analytical bounds that hold even in the infinite depth case. The effectiveness of this method is empirically proved by means of training via backpropagation an extremely deep multilayer perceptron of 50k layers, and an Elman NN to learn long-term dependencies in the input of 10k time steps in the past. Compared with other architectures specifically devised to deal with the V/E problem, e.g. LSTMs for recurrent NNs, the proposed model is way simpler yet more effective. Surprisingly, a single layer vanilla RNN can be enhanced to reach state of the art performance, while converging super fast; for instance on the psMNIST task, it is possible to get test accuracy of over 94% in the first epoch, and over 98% after just 10 epochs.

Reservoir computing (RC) is a state-of-the-art approach for efficient training in temporal domains. In this paper, we explore new RC architectures that generalise the popular leaky echo state network model (leaky-ESN) introducing an additive orthogonal term outside the nonlinear part of the ESN equation. We investigate the benefits of employing orthogonal matrices in ESNs both inside the nonlinearity and outside of it. We show empirically how to boost the memory capacity towards the theoretical maximum value while still preserving the power of nonlinear computations. Ergo, we optimise the compromise between computing with memory and computing with nonlinearity. The proposed model demonstrates to outperform both leaky-ESN and orthogonal reservoir ESN models on tasks requiring nonlinear computations with memory.