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Title: Additive function approximation in the brain

Paper content:
Many biological learning systems such as the mushroom body, hippocampus, and cerebellum are built from sparsely connected networks of neurons.
For a new understanding of such networks, we study the function spaces induced by sparse random features and characterize what functions may and may not be learned.
A network with d inputs per neuron is found to be equivalent to an additive model of order d, whereas with a degree distribution the network combines additive terms of different orders.
We identify three specific advantages of sparsity: additive function approximation is a powerful inductive bias that limits the curse of dimensionality, sparse networks are stable to outlier noise in the inputs, and sparse random features are scalable.
Thus, even simple brain architectures can be powerful function approximators.
Finally, we hope that this work helps popularize kernel theories of networks among computational neuroscientists.
Kernel function spaces are popular among machine learning researchers as a potentially tractable framework for understanding artificial neural networks trained via gradient descent [e.g. 1, 2, 3, 4, 5, 6].
Artificial neural networks are an area of intense interest due to their often surprising empirical performance on a number of challenging problems and our still incomplete theoretical understanding.
Yet computational neuroscientists have not widely applied these new theoretical tools to describe the ability of biological networks to perform function approximation.
The idea of using fixed random weights in a neural network is primordial, and was a part of Rosenblatt's perceptron model of the retina [7] .
Random features have then resurfaced under many guises: random centers in radial basis function networks [8] , functional link networks [9] , Gaussian processes (GPs) [10, 11] , and so-called extreme learning machines [12] ; see [13] for a review.
Random feature networks, where the neurons are initialized with random weights and only the readout layer is trained, were proposed by Rahimi and Recht in order to improve the performance of kernel methods [14, 15] and can perform well for many problems [13] .
In parallel to these developments in machine learning, computational neuroscientists have also studied the properties of random networks with a goal towards understanding neurons in real brains.
To a first approximation, many neuronal circuits seem to be randomly organized [16, 17, 18, 19, 20] .
However, the recent theory of random features appears to be mostly unknown to the greater computational neuroscience community.
Here, we study random feature networks with sparse connectivity: the hidden neurons each receive input from a random, sparse subset of input neurons.
This is inspired by the observation that the connectivity in a variety of predominantly feedforward brain networks is approximately random and sparse.
These brain areas include the cerebellar cortex, invertebrate mushroom body, and dentate gyrus of the hippocampus [21] .
All of these areas perform pattern separation and associative learning.
The cerebellum is important for motor control, while the mushroom body and dentate gyrus are The function shown is the sparse random feature approximation to an additive sum of sines, learned from poorly distributed samples (red crosses).
Additivity offers structure which may be leveraged for fast and efficient learning.
general learning and memory areas for invertebrates and vertebrates, respectively, and may have evolved from a similar structure in the ancient bilaterian ancestor [22] .
Recent work has argued that the sparsity observed in these areas may be optimized to balance the dimensionality of representation with wiring cost [20] .
Sparse connectivity has been used to compress artificial networks and speed up computation [23, 24, 25] , whereas convolutions are a kind of structured sparsity [26, 27] .
We show that sparse random features approximate additive kernels [28, 29, 30, 31] with arbitrary orders of interaction.
The in-degree of the hidden neurons d sets the order of interaction.
When the degrees of the neurons are drawn from a distribution, the resulting kernel contains a weighted mixture of interactions.
These sparse features offer advantages of generalization in high-dimensions, stability under perturbations of their input, and computational and biological efficiency.
Inspired by their ubiquity in biology, we have studied sparse random networks of neurons using the theory of random features, finding the advantages of additivity, stability, and scalability.
This theory shows that sparse networks such as those found in the mushroom body, cerebellum, and hippocampus can be powerful function approximators.
Kernel theories of neural circuits may be more broadly applicable in the field of computational neuroscience.
Expanding the theory of dimensionality in neuroscience Learning is easier in additive function spaces because they are low-dimensional, a possible explanation for few-shot learning in biological systems.
Our theory is complementary to existing theories of dimensionality in neural systems [16, 44, 45, 46, 47, 20, 48, 49, 50] , which defined dimensionality using a skewness measure of covariance eigenvalues.
Kernel theory extends this concept, measuring dimensionality similarly [51] in the space of nonlinear functions spanned by the kernel.
Limitations We model biological neurons as simple scalar functions, completely ignoring time and neuromodulatory context.
It seems possible that a kernel theory could be developed for timeand context-dependent features.
Our networks suppose i.i.d. weights, but weights that follow Dale's law should also be considered.
We have not studied the sparsity of activity, postulated to be relevant in cerebellum.
It remains to be demonstrated how the theory can make concrete, testable predictions, e.g. whether this theory may explain identity versus concentration encoding of odors or the discrimination/generalization tradeoff under experimental conditions.
Appendices: Additive function approximation in the brain As said in the main text, Kandasamy and Yu [1] created a theory of the generalization properties of higher-order additive models.
They supplemented this with an empirical study of a number of datasets using their Shrunk Additive Least Squares Approximation (SALSA) implementation of the additive kernel ridge regression (KRR).
Their data and code were obtained from https: //github.com/kirthevasank/salsa.
We compared the performance of SALSA to the sparse random feature approximation of the same kernel.
We employ random sparse Fourier features with Gaussian weights N (0, σ 2 I) with σ = 0.05 · √ dn 1/5 in order to match the Gaussian radial basis function used by Kandasamy and Yu [1] .
We use m = 300l features for every problem, with regular degree d selected equal to the one chosen by SALSA.
The regressor on the features is cross-validated ridge regression (RidgeCV from scikit-learn) with ridge penalty selected from 5 logarithmically spaced points between 10 −4 · n and 10 2 · n.
In Figure 2 , we compare the performance of sparse random features to SALSA.
Generally, the training and testing errors of the sparse model are slightly higher than for the kernel method, except for the forestfires dataset.

assistant: text: We advocate for random features as a theory of biological neural networks, focusing on sparsely connected networks