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Carina Curto

Publications and source records attributed to Carina Curto.

At least 19 recordsLinked to original sources
arXiv2026

Sequential chaotic oscillations in excitatory-inhibitory threshold-linear networks

Metastable states, a phenomenon observed in brain dynamics and many other systems, have been proposed as a key feature of healthy brain function, reflecting a balance between integration and segregation. However, it remains unclear how to capture this behavior within a dynamical-systems framework. In this paper, we propose sequential chaotic oscillations (SCOs), arising in excitatory-inhibitory threshold-linear networks (E-I TLNs), as a candidate dynamical mechanism for sequential metastability. As a simple form of chaotic itinerancy, SCOs occur under constant input and consist of a sequence of metastable states whose transition order can be predicted by the underlying graph. To identify the parameter regime for SCOs, we develop new graph rules for E-I TLNs and use them to characterize the fixed point structure of E-I TLNs on paths and cycles. Our results show that the emergence of SCOs requires unstable singleton fixed points and sufficiently strong inhibition. In addition to SCOs, we find that E-I oscillations need not be synchronized. Motivated by this, we introduce a decomposition into the z-mode and the mean mode, which capture excitatory differences and overall network activity, respectively. These modes are then used to distinguish attractors associated with the full-support fixed point of E-I TLNs on cycles.

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arXiv2025

On graphical domination for threshold-linear networks with recurrent excitation and global inhibition

Graphical domination was first introduced in [1] in the context of combinatorial threshold-linear networks (CTLNs). There it was shown that when a domination relationship exists between a pair of vertices in a graph, certain fixed points in the corresponding CTLN can be ruled out. Here we prove two new theorems about graphical domination, and show that they apply to a significantly more general class of recurrent networks called generalized CTLNs (gCTLNs). Theorem 1 establishes that if a dominated node is removed from a network, the reduced network has exactly the same fixed points. Theorem 2 tells us that by iteratively removing dominated nodes from an initial graph $G$, the final (irreducible) graph $\widetilde{G}$ is unique. We also introduce another new family of TLNs, called E-I TLNs, consisting of $n$ excitatory nodes and a single inhibitory node providing global inhibition. We provide a concrete mapping between the parameters of gCTLNs and E-I TLNs built from the same graph such that corresponding networks have the same fixed points. We also show that Theorems 1 and 2 apply equally well to E-I TLNs, and that the dynamics of gCTLNs and E-I TLNs with the same underlying graph $G$ exhibit similar behavior that is well predicted by the fixed points of the reduced graph $\widetilde{G}$.

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arXiv2023

Stable fixed points of combinatorial threshold-linear networks

Combinatorial threshold-linear networks (CTLNs) are a special class of recurrent neural networks whose dynamics are tightly controlled by an underlying directed graph. Recurrent networks have long been used as models for associative memory and pattern completion, with stable fixed points playing the role of stored memory patterns in the network. In prior work, we showed that target-free cliques of the graph correspond to stable fixed points of the dynamics, and we conjectured that these are the only stable fixed points possible. In this paper, we prove that the conjecture holds in a variety of special cases, including for networks with very strong inhibition and graphs of size $n \leq 4$. We also provide further evidence for the conjecture by showing that sparse graphs and graphs that are nearly cliques can never support stable fixed points. Finally, we translate some results from extremal combinatorics to obtain an upper bound on the number of stable fixed points of CTLNs in cases where the conjecture holds.

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arXiv2023

Diversity of emergent dynamics in competitive threshold-linear networks

Threshold-linear networks consist of simple units interacting in the presence of a threshold nonlinearity. Competitive threshold-linear networks have long been known to exhibit multistability, where the activity of the network settles into one of potentially many steady states. In this work, we find conditions that guarantee the absence of steady states, while maintaining bounded activity. These conditions lead us to define a combinatorial family of competitive threshold-linear networks, parametrized by a simple directed graph. By exploring this family, we discover that threshold-linear networks are capable of displaying a surprisingly rich variety of nonlinear dynamics, including limit cycles, quasiperiodic attractors, and chaos. In particular, several types of nonlinear behaviors can co-exist in the same network. Our mathematical results also enable us to engineer networks with multiple dynamic patterns. Taken together, these theoretical and computational findings suggest that threshold-linear networks may be a valuable tool for understanding the relationship between network connectivity and emergent dynamics. The new Matlab package CTLN Basic 2.0 can be used to reproduce the simulations in Figures 1-13. The package is available at https://github.com/nebneuron/CTLN-Basic-2.0

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arXiv2022

Sequential attractors in combinatorial threshold-linear networks

Sequences of neural activity arise in many brain areas, including cortex, hippocampus, and central pattern generator circuits that underlie rhythmic behaviors like locomotion. While network architectures supporting sequence generation vary considerably, a common feature is an abundance of inhibition. In this work, we focus on architectures that support sequential activity in recurrently connected networks with inhibition-dominated dynamics. Specifically, we study emergent sequences in a special family of threshold-linear networks, called combinatorial threshold-linear networks (CTLNs), whose connectivity matrices are defined from directed graphs. Such networks naturally give rise to an abundance of sequences whose dynamics are tightly connected to the underlying graph. We find that architectures based on generalizations of cycle graphs produce limit cycle attractors that can be activated to generate transient or persistent (repeating) sequences. Each architecture type gives rise to an infinite family of graphs that can be built from arbitrary component subgraphs. Moreover, we prove a number of graph rules for the corresponding CTLNs in each family. The graph rules allow us to strongly constrain, and in some cases fully determine, the fixed points of the network in terms of the fixed points of the component subnetworks. Finally, we also show how the structure of certain architectures gives insight into the sequential dynamics of the corresponding attractor.

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arXiv2022

Core motifs predict dynamic attractors in combinatorial threshold-linear networks

Combinatorial threshold-linear networks (CTLNs) are a special class of inhibition-dominated TLNs defined from directed graphs. Like more general TLNs, they display a wide variety of nonlinear dynamics including multistability, limit cycles, quasiperiodic attractors, and chaos. In prior work, we have developed a detailed mathematical theory relating stable and unstable fixed points of CTLNs to graph-theoretic properties of the underlying network. Here we find that a special type of fixed points, corresponding to core motifs, are predictive of both static and dynamic attractors. Moreover, the attractors can be found by choosing initial conditions that are small perturbations of these fixed points. This motivates us to hypothesize that dynamic attractors of a network correspond to unstable fixed points supported on core motifs. We tested this hypothesis on a large family of directed graphs of size $n=5$, and found remarkable agreement. Furthermore, we discovered that core motifs with similar embeddings give rise to nearly identical attractors. This allowed us to classify attractors based on structurally-defined graph families. Our results suggest that graphical properties of the connectivity can be used to predict a network's complex repertoire of nonlinear dynamics.

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arXiv2021

Nerve theorems for fixed points of neural networks

Nonlinear network dynamics are notoriously difficult to understand. Here we study a class of recurrent neural networks called combinatorial threshold-linear networks (CTLNs) whose dynamics are determined by the structure of a directed graph. They are a special case of TLNs, a popular framework for modeling neural activity in computational neuroscience. In prior work, CTLNs were found to be surprisingly tractable mathematically. For small networks, the fixed points of the network dynamics can often be completely determined via a series of graph rules that can be applied directly to the underlying graph. For larger networks, it remains a challenge to understand how the global structure of the network interacts with local properties. In this work, we propose a method of covering graphs of CTLNs with a set of smaller directional graphs that reflect the local flow of activity. While directional graphs may or may not have a feedforward architecture, their fixed point structure is indicative of feedforward dynamics. The combinatorial structure of the graph cover is captured by the nerve of the cover. The nerve is a smaller, simpler graph that is more amenable to graphical analysis. We present three nerve theorems that provide strong constraints on the fixed points of the underlying network from the structure of the nerve. We then illustrate the power of these theorems with some examples. Remarkably, we find that the nerve not only constrains the fixed points of CTLNs, but also gives insight into the transient and asymptotic dynamics. This is because the flow of activity in the network tends to follow the edges of the nerve.

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arXiv2021

Betti Curves of Rank One Symmetric Matrices

Betti curves of symmetric matrices were introduced in (Giusti et. al., 2015) as a new class of matrix invariants that depend only on the relative ordering of matrix entries. These invariants are computed using persistent homology, and can be used to detect underlying structure in biological data that may otherwise be obscured by monotone nonlinearities. Here we prove three theorems that fully characterize the Betti curves of rank 1 symmetric matrices. We then illustrate how these Betti curve signatures arise in natural data obtained from calcium imaging of neural activity in zebrafish.

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arXiv2021

Periodic Neural Codes and Sound Localization in Barn Owls

Inspired by the sound localization system of the barn owl, we define a new class of neural codes, called periodic codes, and study their basic properties. Periodic codes are binary codes with a special patterned form that reflects the periodicity of the stimulus. Because these codes can be used by the owl to localize sounds within a convex set of angles, we investigate whether they are examples of convex codes, which have previously been studied for hippocampal place cells. We find that periodic codes are typically not convex, but can be completed to convex codes in the presence of noise. We introduce the convex closure and Hamming distance completion as ways of adding codewords to make a code convex, and describe the convex closure of a periodic code. We also find that the probability of the convex closure arising stochastically is greater for sparser codes. Finally, we provide an algebraic method using the neural ideal to detect if a code is periodic. We find that properties of periodic codes help to explain several aspects of the behavior observed in the sound localization system of the barn owl, including common errors in localizing pure tones.

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arXiv2020

Combinatorial Geometry of Threshold-Linear Networks

The architecture of a neural network constrains the potential dynamics that can emerge. Some architectures may only allow for a single dynamic regime, while others display a great deal of flexibility with qualitatively different dynamics that can be reached by modulating connection strengths. In this work, we develop novel mathematical techniques to study the dynamic constraints imposed by different network architectures in the context of competitive threshold-linear networks (TLNs). Any given TLN is naturally characterized by a hyperplane arrangement in $\mathbb{R}^n$, and the combinatorial properties of this arrangement determine the pattern of fixed points of the dynamics. This observation enables us to recast the question of network flexibility in the language of oriented matroids, allowing us to employ tools and results from this theory in order to characterize the different dynamic regimes a given architecture can support. In particular, fixed points of a TLN correspond to cocircuits of an associated oriented matroid; and mutations of the matroid correspond to bifurcations in the collection of fixed points. As an application, we provide a complete characterization of all possible sets of fixed points that can arise in networks through size $n=3$, together with descriptions of how to modulate synaptic strengths of the network in order to access the different dynamic regimes. These results provide a framework for studying the possible computational roles of various motifs observed in real neural networks.

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arXiv2019

Robust motifs of threshold-linear networks

To any inhibition-dominated threshold-linear network (TLN) we can associate a directed graph that captures the pattern of strong and weak inhibition between neurons. Robust motifs are graphs for which the structure of fixed points in the network is independent of the choice of connectivity matrix $W$, and whose dynamics are thus greatly constrained. This makes them ideal building blocks for constructing larger networks whose behaviors are robust to changes in connection strengths. In contrast, flexible motifs correspond to networks with multiple dynamic regimes. In this work, we give a purely graphical characterization of both flexible and robust motifs of any size. We find that all but a few robust motifs fall into two infinite families of graphs with simple feedforward architectures.

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arXiv2019

Neural ring homomorphisms and maps between neural codes

Neural codes are binary codes that are used for information processing and representation in the brain. In previous work, we have shown how an algebraic structure, called the {\it neural ring}, can be used to efficiently encode geometric and combinatorial properties of a neural code [1]. In this work, we consider maps between neural codes and the associated homomorphisms of their neural rings. In order to ensure that these maps are meaningful and preserve relevant structure, we find that we need additional constraints on the ring homomorphisms. This motivates us to define {\it neural ring homomorphisms}. Our main results characterize all code maps corresponding to neural ring homomorphisms as compositions of 5 elementary code maps. As an application, we find that neural ring homomorphisms behave nicely with respect to convexity. In particular, if $\mathcal{C}$ and $\mathcal{D}$ are convex codes, the existence of a surjective code map $\mathcal{C}\rightarrow \mathcal{D}$ with a corresponding neural ring homomorphism implies that the minimal embedding dimensions satisfy $d(\mathcal{D}) \leq d(\mathcal{C})$.

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arXiv2018

Fixed points of competitive threshold-linear networks

Threshold-linear networks (TLNs) are models of neural networks that consist of simple, perceptron-like neurons and exhibit nonlinear dynamics that are determined by the network's connectivity. The fixed points of a TLN, including both stable and unstable equilibria, play a critical role in shaping its emergent dynamics. In this work, we provide two novel characterizations for the set of fixed points of a competitive TLN: the first is in terms of a simple sign condition, while the second relies on the concept of domination. We apply these results to a special family of TLNs, called combinatorial threshold-linear networks (CTLNs), whose connectivity matrices are defined from directed graphs. This leads us to prove a series of graph rules that enable one to determine fixed points of a CTLN by analyzing the underlying graph. Additionally, we study larger networks composed of smaller "building block" subnetworks, and prove several theorems relating the fixed points of the full network to those of its components. Our results provide the foundation for a kind of "graphical calculus" to infer features of the dynamics from a network's connectivity.

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arXiv2018

Algebraic signatures of convex and non-convex codes

A convex code is a binary code generated by the pattern of intersections of a collection of open convex sets in some Euclidean space. Convex codes are relevant to neuroscience as they arise from the activity of neurons that have convex receptive fields. In this paper, we use algebraic methods to determine if a code is convex. Specifically, we use the neural ideal of a code, which is a generalization of the Stanley-Reisner ideal. Using the neural ideal together with its standard generating set, the canonical form, we provide algebraic signatures of certain families of codes that are non-convex. We connect these signatures to the precise conditions on the arrangement of sets that prevent the codes from being convex. Finally, we also provide algebraic signatures for some families of codes that are convex, including the class of intersection-complete codes. These results allow us to detect convexity and non-convexity in a variety of situations, and point to some interesting open questions.

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arXiv2018

Predicting neural network dynamics via graphical analysis

Neural network models in neuroscience allow one to study how the connections between neurons shape the activity of neural circuits in the brain. In this chapter, we study Combinatorial Threshold-Linear Networks (CTLNs) in order to understand how the pattern of connectivity, as encoded by a directed graph, shapes the emergent nonlinear dynamics of the corresponding network. Important aspects of these dynamics are controlled by the stable and unstable fixed points of the network, and we show how these fixed points can be determined via graph-based rules. We also present an algorithm for predicting sequences of neural activation from the underlying directed graph, and examine the effect of graph symmetries on a network's set of attractors.

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arXiv2016

The Leray Dimension of a Convex Code

Convex codes were recently introduced as models for neural codes in the brain. Any convex code $\C$ has an associated minimal embedding dimension $d(\C)$, which is the minimal Euclidean space dimension such that the code can be realized by a collection of convex open sets. In this work we import tools from combinatorial commutative algebra in order to obtain better bounds on $d(\C)$ from an associated simplicial complex $Δ(\C)$. In particular, we make a connection to minimal free resolutions of Stanley-Reisner ideals, and observe that they contain topological information that provides stronger bounds on $d(\C)$. This motivates us to define the Leray dimension $d_L(\C),$ and show that it can be obtained from the Betti numbers of such a minimal free resolution. We compare $d_L(\C)$ to two previously studied dimension bounds, obtained from Helly's theorem and the simplicial homology of $Δ(\C)$. Finally, we show explicitly how $d_L(\C)$ can be computed algebraically, and illustrate this with examples.

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