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Convex Efficient Coding

William Dorrell, Peter E. Latham, James Whittington

TL;DR

This work develops a tractable, convex framework for normative neural coding by reformulating optimizations over neural representations as convex problems on the representational similarity matrix $\mathbf{Q}=\mathbf{Z}^T\mathbf{Z}$. By showing convexity across a wide suite of constraints and objectives, the authors construct a family of problems that encompass linear and certain nonlinear networks, including nonnegative affine autoencoders and nonlinear similarity matching. They derive a tight identifiability criterion for semi-nonnegative matrix factorisation, establish when population-level representational similarity uniquely determines single-neuron tunings, and use tractable nonlinear formulations to explain ON-OFF coding in retinal-like settings, with sparsity setting the transition between single-channel and dual-channel codes. The framework thus links population representations to neuron-level tuning, enabling analytic insights into neural codes and potentially guiding interpretability in AI models. Overall, the paper provides a versatile, convex analytic toolkit for addressing core questions in neural coding and representation learning, with implications for both neuroscience and machine learning interpretability.

Abstract

Why do neurons encode information the way they do? Normative answers to this question model neural activity as the solution to an optimisation problem; for example, the celebrated efficient coding hypothesis frames neural activity as the optimal encoding of information under efficiency constraints. Successful normative theories have varied dramatically in complexity, from simple linear models (Atick & Redlich '90), to complex deep neural networks (Lindsay '21). What complex models gain in flexibility, they lose in tractability and often understandability. Here, we split the difference by constructing a set of tractable but flexible normative representational theories. Instead of optimising the neural activities directly, following Sengupta et al. '18, we optimise the representational similarity, a matrix formed from the dot products of each pair of neural responses. Using this, we show that a large family of interesting optimisation problems are convex. This family includes problems corresponding to linear and some non-linear neural networks, and problems from the literature not previously recognised as convex, such as modified versions of semi-nonnegative matrix factorisation or nonnegative sparse coding. We put these findings to work in three ways. First, we provide the first necessary and sufficient identifiability result for a form of semi-nonnegative matrix factorisation. Second, we show that if neural tunings are `different enough' then they are uniquely linked to the optimal representational similarity, partially justifying the use of single neuron tuning analysis in neuroscience. Finally, we use the tractable nonlinearity of some of our problems to explain why dense retinal codes, but not sparse cortical codes, optimally split the coding of a single variable into ON & OFF channels. In sum, we identify a space of convex problems, and use them to derive neural coding results.

Convex Efficient Coding

TL;DR

This work develops a tractable, convex framework for normative neural coding by reformulating optimizations over neural representations as convex problems on the representational similarity matrix . By showing convexity across a wide suite of constraints and objectives, the authors construct a family of problems that encompass linear and certain nonlinear networks, including nonnegative affine autoencoders and nonlinear similarity matching. They derive a tight identifiability criterion for semi-nonnegative matrix factorisation, establish when population-level representational similarity uniquely determines single-neuron tunings, and use tractable nonlinear formulations to explain ON-OFF coding in retinal-like settings, with sparsity setting the transition between single-channel and dual-channel codes. The framework thus links population representations to neuron-level tuning, enabling analytic insights into neural codes and potentially guiding interpretability in AI models. Overall, the paper provides a versatile, convex analytic toolkit for addressing core questions in neural coding and representation learning, with implications for both neuroscience and machine learning interpretability.

Abstract

Why do neurons encode information the way they do? Normative answers to this question model neural activity as the solution to an optimisation problem; for example, the celebrated efficient coding hypothesis frames neural activity as the optimal encoding of information under efficiency constraints. Successful normative theories have varied dramatically in complexity, from simple linear models (Atick & Redlich '90), to complex deep neural networks (Lindsay '21). What complex models gain in flexibility, they lose in tractability and often understandability. Here, we split the difference by constructing a set of tractable but flexible normative representational theories. Instead of optimising the neural activities directly, following Sengupta et al. '18, we optimise the representational similarity, a matrix formed from the dot products of each pair of neural responses. Using this, we show that a large family of interesting optimisation problems are convex. This family includes problems corresponding to linear and some non-linear neural networks, and problems from the literature not previously recognised as convex, such as modified versions of semi-nonnegative matrix factorisation or nonnegative sparse coding. We put these findings to work in three ways. First, we provide the first necessary and sufficient identifiability result for a form of semi-nonnegative matrix factorisation. Second, we show that if neural tunings are `different enough' then they are uniquely linked to the optimal representational similarity, partially justifying the use of single neuron tuning analysis in neuroscience. Finally, we use the tractable nonlinearity of some of our problems to explain why dense retinal codes, but not sparse cortical codes, optimally split the coding of a single variable into ON & OFF channels. In sum, we identify a space of convex problems, and use them to derive neural coding results.
Paper Structure (49 sections, 5 theorems, 156 equations, 4 figures)

This paper contains 49 sections, 5 theorems, 156 equations, 4 figures.

Key Result

Theorem 1

Given a dataset ${\bm{X}} = {\bm{A}}{\bm{S}}$, if and only if the matrix $\bar{{\bm{S}}}$ is tightly scattered with respect to ${\bm{A}}$ then the optimal positive affine autoencoder recovers the sources: each neuron's activity is an affine function of one source and every source has at least one ne

Figures (4)

  • Figure 1: (A) We schematise the the identifiability conditions for two sources; the conditions specify a set (e.g. the red ellipse that depends on the linear mixing of the sources in the observed data) that the convex hull of the underlying empirical source distribution has to engulf. If this condition is satisfied the empirical source data is 'rectangular enough', there is no better linear transformation, and the sources are recovered, else the optimal representation is mixed. (B) When the data consists of linearly mixed sources via ${\bm{A}}$, then this equates to warping the identifiability conditions: either via aligning (blue) or antialigning (green). (C) Source alignment can make the optimal solution mix when it would have otherwise modularised. We show such a dataset in which the orthogonal but not an example antialigned identifiability conditions are satisfied. Matching the theory, numerical solutions are modular for the orthogonally encoded sources (rightmost column), but not for the antialigned (middle column). We plot the linear conditional mutual information hsu2023disentanglementdorrell2025range between each neuron and source scaled by the neuron's peak activity. Below we display a highlighted (purple) neuron's tuning to sources. (D) Similarly, source alignment can cause recovery of sources that would otherwise not be. We show an example dataset where aligning the sources by a specific amount causes the warped identifiability conditions to be satisfied (blue). Matching this, the aligned sources are recovered (middle column), but not the orthogonal ones (right column).
  • Figure 2: A) Illustration of identifiability condition: the condition states that the convex hull of the neural responses must engulf one of a set of ellipses. Plotting the data for two neurons, and drawing two ellipses for different choices of ${\bm{F}}$ shows us an example where the ellipse is not (purple) and is (blue) engulfed. Since there is at least one, the condition is satisfied. (B) We try this on two neurons, tuned like place cells to a single 1D latent. Plotting the neural responses and an example ellipse shows us that these tunings are identifiable, since the convex hull of the data engulfs the ellipse. However, (C) moving the place cells to overlap leads to non-identifiability. Indeed, (D) we can rotate the responses to find two different tuning curves with the same optimal dot-product structure. (E) We apply this to grid cells. Grid cells from two different modules are identifiable as long as their wavevectors are not integer multiples of one another, otherwise (F) they become non-identifiable, and we can numerically find a rotation of these neural responses that preserves nonnegativity.
  • Figure 3: A) We numerically find a solution to \ref{['eq:ON-OFF_problem']}, and plot the firing rates of each neuron in the population as a function of $I$. We find that all tuning curves correspond to either OFF or ON channels. B). We add a point mass at $I = 0$ to the distribution over $I$, thus increasing the sparsity. For low sparsity the optimal representation contains both ON and OFF neurons. As sparsity increases the coding range of the ON neurons increase while the range of the OFF neurons decreases, until at the threshold (eq. \ref{['eq:sparsity']}) the OFF neuron disappears and we are left with ON neurons only. We display only the unique tuning curves, the population is comprised of copies of these tuning curves. C) We quantify the degree of single channel coding, \ref{['app:ON_OFF_details']}, and find it slowly increases to a ceiling at the predicted sparsity threshold, corresponding to a completely single channel representation.
  • Figure 4: Similarly to \ref{['fig:Linear_Mixing']}, in (A) We schematise identifiability conditions for two sources; which are either satisfied (red) or brocken (blue) by this dataset depending on the regularisation hyperaparameter. (B) Matching the theory, numerical solutions are modular for the less regularised network (left), but not for the more regularised (right). We plot the linear conditional mutual information hsu2023disentanglementdorrell2025range between each neuron and source scaled by the neuron's peak activity. Below we display a highlighted (purple) neuron's tuning to sources.

Theorems & Definitions (10)

  • Definition 1: Tight Scattering
  • Theorem 1: Identifiability of Nonnegative Affine Autoencoders
  • Theorem 2: Tight Scattering Implies Unique Neurons
  • Definition 1: Tight Scattering
  • Theorem 1: Identifiability of Nonnegative Affine Autoencoders
  • Definition 2: Tight Scattering
  • Theorem 3: Identifiability of Imperfect Nonnegative Affine Autoencoders
  • proof
  • Theorem 2: Tight Scattering Implies Unique Neurons
  • proof