Sampling and Variational Inference (VI) are two large families of methods for approximate inference that have complementary strengths. Sampling methods excel at approximating arbitrary probability distributions, but can be inefficient. VI methods are efficient, but may misrepresent the true distribution. Here, we develop a general framework where approximations are stochastic mixtures of simple component distributions. Both sampling and VI can be seen as special cases: in sampling, each mixture component is a delta-function and is chosen stochastically, while in standard VI a single component is chosen to minimize divergence. We derive a practical method that interpolates between sampling and VI by analytically solving an optimization problem over a mixing distribution. Intermediate inference methods then arise by varying a single parameter. Our method provably improves on sampling (reducing variance) and on VI (reducing bias+variance despite increasing variance). We demonstrate our method’s bias/variance trade-off in practice on reference problems, and we compare outcomes to commonly used sampling and VI methods. This work takes a step towards a highly flexible yet simple family of inference methods that combines the complementary strengths of sampling and VI.

Perception is often characterized computationally as an inference process in which uncertain or ambiguous sensory inputs are combined with prior expectations. Although behavioral studies have shown that observers can change their prior expectations in the context of a task, robust neural signatures of task-specific priors have been elusive. Here, we analytically derive such signatures under the general assumption that the responses of sensory neurons encode posterior beliefs that combine sensory inputs with task-specific expectations. Specifically, we derive predictions for the task-dependence of correlated neural variability and decision-related signals in sensory neurons. The qualitative aspects of our results are parameter-free and specific to the statistics of each task. The predictions for correlated variability also differ from predictions of classic feedforward models of sensory processing and are therefore a strong test of theories of hierarchical Bayesian inference in the brain. Importantly, we find that Bayesian learning predicts an increase in so-called "differential correlations" as the observer's internal model learns the stimulus distribution, and the observer's behavioral performance improves. This stands in contrast to classic feedforward encoding/decoding models of sensory processing, since such correlations are fundamentally information-limiting. We find support for our predictions in data from existing neurophysiological studies across a variety of tasks and brain areas. Finally, we show in simulation how measurements of sensory neural responses can reveal information about a subject's internal beliefs about the task. Taken together, our results reinterpret task-dependent sources of neural covariability as signatures of Bayesian inference and provide new insights into their cause and their function.

Making good decisions requires updating beliefs according to new evidence. This is a dynamical process that is prone to biases: in some cases, beliefs become entrenched and resistant to new evidence (leading to primacy effects), while in other cases, beliefs fade over time and rely primarily on later evidence (leading to recency effects). How and why either type of bias dominates in a given context is an important open question. Here, we study this question in classic perceptual decision-making tasks, where, puzzlingly, previous empirical studies differ in the kinds of biases they observe, ranging from primacy to recency, despite seemingly equivalent tasks. We present a new model, based on hierarchical approximate inference and derived from normative principles, that not only explains both primacy and recency effects in existing studies, but also predicts how the type of bias should depend on the statistics of stimuli in a given task. We verify this prediction in a novel visual discrimination task with human observers, finding that each observer's temporal bias changed as the result of changing the key stimulus statistics identified by our model. The key dynamic that leads to a primacy bias in our model is an overweighting of new sensory information that agrees with the observer's existing belief-a type of 'confirmation bias'. By fitting an extended drift-diffusion model to our data we rule out an alternative explanation for primacy effects due to bounded integration. Taken together, our results resolve a major discrepancy among existing perceptual decision-making studies, and suggest that a key source of bias in human decision-making is approximate hierarchical inference.

Sensory processing is often characterized as implementing probabilistic inference: networks of neurons compute posterior beliefs over unobserved causes given the sensory inputs. How these beliefs are computed and represented by neural responses is much-debated (Fiser et al. 2010, Pouget et al. 2013). A central debate concerns the question of whether neural responses represent samples of latent variables (Hoyer & Hyvarinnen 2003) or parameters of their distributions (Ma et al. 2006) with efforts being made to distinguish between them (Grabska-Barwinska et al. 2013). A separate debate addresses the question of whether neural responses are proportionally related to the encoded probabilities (Barlow 1969), or proportional to the logarithm of those probabilities (Jazayeri & Movshon 2006, Ma et al. 2006, Beck et al. 2012). Here, we show that these alternatives -- contrary to common assumptions -- are not mutually exclusive and that the very same system can be compatible with all of them. As a central analytical result, we show that modeling neural responses in area V1 as samples from a posterior distribution over latents in a linear Gaussian model of the image implies that those neural responses form a linear Probabilistic Population Code (PPC, Ma et al. 2006). In particular, the posterior distribution over some experimenter-defined variable like "orientation" is part of the exponential family with sufficient statistics that are linear in the neural sampling-based firing rates.

Following earlier studies which showed that a sparse coding principle may explain the receptive field properties of complex cells in primary visual cortex, it has been concluded that the same properties may be equally derived from a slowness principle. In contrast to this claim, we here show that slowness and sparsity drive the representations towards substantially different receptive field properties. To do so, we present complete sets of basis functions learned with slow subspace analysis (SSA) in case of natural movies as well as translations, rotations, and scalings of natural images. SSA directly parallels independent subspace analysis (ISA) with the only difference that SSA maximizes slowness instead of sparsity. We find a large discrepancy between the filter shapes learned with SSA and ISA. We argue that SSA can be understood as a generalization of the Fourier transform where the power spectrum corresponds to the maximally slow subspace energies in SSA. Finally, we investigate the trade-off between slowness and sparseness when combined in one objective function.

Many studies have explored the impact of response variability on the quality of sensory codes. The source of this variability is almost always assumed to be intrinsic to the brain. However, when inferring a particular stimulus property, variability associated with other stimulus attributes also effectively act as noise. Here we study the impact of such stimulus-induced response variability for the case of binocular disparity inference. We characterize the response distribution for the binocular energy model in response to random dot stereograms and find it to be very different from the Poisson-like noise usually assumed. We then compute the Fisher information with respect to binocular disparity, present in the monocular inputs to the standard model of early binocular processing, and thereby obtain an upper bound on how much information a model could theoretically extract from them. Then we analyze the information loss incurred by the different ways of combining those inputs to produce a scalar single-neuron response. We find that in the case of depth inference, monocular stimulus variability places a greater limit on the extractable information than intrinsic neuronal noise for typical spike counts. Furthermore, the largest loss of information is incurred by the standard model for position disparity neurons (tuned-excitatory), that are the most ubiquitous in monkey primary visual cortex, while more information from the inputs is preserved in phase-disparity neurons (tuned-near or tuned-far) primarily found in higher cortical regions.

A crucial part of developing mathematical models of how the brain works is the quantification of their success. One of the most widely-used metrics yields the percentage of the variance in the data that is explained by the model. Unfortunately, this metric is biased due to the intrinsic variability in the data. This variability is in principle unexplainable by the model. We derive a simple analytical modification of the traditional formula that significantly improves its accuracy (as measured by bias) with similar or better precision (as measured by mean-square error) in estimating the true underlying Variance Explained by the model class. Our estimator advances on previous work by a) accounting for the uncertainty in the noise estimate, b) accounting for overfitting due to free model parameters mitigating the need for a separate validation data set and c) adding a conditioning term. We apply our new estimator to binocular disparity tuning curves of a set of macaque V1 neurons and find that on a population level almost all of the variance unexplained by Gabor functions is attributable to noise.