Efficient coding provides a direct link between prior and likelihood in perceptual Bayesian inference.

This paper makes two contributions. First, this paper continues a trend of recent work by Park and Pillow (2011) and Ramirez & Paninski (2012) to provide a firm grounding of the spike-triggered methods STA and STC within forward, likelihood-based modelling. STA and STC are calculated as moments from the stimulus/response ensemble, and are popular for constructing stimulus/response models due to their speed and simplicity. Paninski 2003, Sharpee et al 2004, Samengo & Gollisch 2013), the validity and utility of these moments depends on particular restrictive conditions on the stimulus set. The prior work of Park & Pillow (2011) was able to shed light on this, by relating these moments to the parameters of a Generalised Linear Model (GLM), a popular forward model for neural responses.

We describe a set of fast, tractable methods for characterizing neural responses to high-dimensional sensory stimuli using a model we refer to as the generalized quadratic model (GQM). The GQM consists of a low-rank quadratic function followed by a point nonlinearity and exponential-family noise. The quadratic function characterizes the neuron's stimulus selectivity in terms of a set linear receptive fields followed by a quadratic combination rule, and the invertible nonlinearity maps this output to the desired response range.

We describe a set of fast, tractable methods for characterizing neural responses to high-dimensional sensory stimuli using a model we refer to as the generalized quadratic model (GQM). The GQM consists of a low-rank quadratic form followed by a point nonlinearity and exponential-family noise. The quadratic form characterizes the neuron's stimulus selectivity in terms of a set linear receptive fields followed by a quadratic combination rule, and the invertible nonlinearity maps this output to the desired response range. Special cases of the GQM include the 2nd-order Volterra model (Marmarelis and Marmarelis 1978, Koh and Powers 1985) and the elliptical Linear-Nonlinear-Poisson model (Park and Pillow 2011). Here we show that for canonical form" GQMs, spectral decomposition of the first two response-weighted moments yields approximate maximum-likelihood estimators via a quantity called the expected log-likelihood. The resulting theory generalizes moment-based estimators such as the spike-triggered covariance, and, in the Gaussian noise case, provides closed-form estimators under a large class of non-Gaussian stimulus distributions. We show that these estimators are fast and provide highly accurate estimates with far lower computational cost than full maximum likelihood. Moreover, the GQM provides a natural framework for combining multi-dimensional stimulus sensitivity and spike-history dependencies within a single model. We show applications to both analog and spiking data using intracellular recordings of V1 membrane potential and extracellular recordings of retinal spike trains."

Neurons typically respond to a restricted number of stimulus features within the high-dimensional space of natural stimuli. Here we describe an explicit model-based interpretation of traditional estimators for a neuron's multi-dimensional feature space, which allows for several important generalizations and extensions. First, we show that traditional estimators based on the spike-triggered average (STA) and spike-triggered covariance (STC) can be formalized in terms of the "expected log-likelihood" of a Linear-Nonlinear-Poisson (LNP) model with Gaussian stimuli. This model-based formulation allows us to define maximum-likelihood and Bayesian estimators that are statistically consistent and efficient in a wider variety of settings, such as with naturalistic (non-Gaussian) stimuli. It also allows us to employ Bayesian methods for regularization, smoothing, sparsification, and model comparison, and provides Bayesian confidence intervals on model parameters. We describe an empirical Bayes method for selecting the number of features, and extend the model to accommodate an arbitrary elliptical nonlinear response function, which results in a more powerful and more flexible model for feature space inference. We validate these methods using neural data recorded extracellularly from macaque primary visual cortex.

Self-organizing feature maps like the Kohonen map (Kohonen, 1989, Ritter et al., 1990) not only provide a plausible explanation for the formation of maps in brains, e.g. in the visual system (Obermayer et al., 1990), but have also been applied to problems like vector quantization, or robot arm control (Martinetz et al., 1990). The underlying organizing principle is the preservation of neighborhood relations. For this principle to lead to a most useful map, the topological structure of the output space must roughly fit the structure of the input data. However, in technical 1141 1142 Bauer, Pawelzik, and Geisel applications this structure is often not a priory known. For this reason several attempts have been made to modify the Kohonen-algorithm such, that not only the weights, but also the output space topology itself is adapted during learning (Kangas et al., 1990, Martinetz et al., 1991). Our contribution is also concerned with optimal output space topologies, but we follow a different approach, which avoids a possibly complicated structure of the output space. First we describe a quantitative measure for the preservation of neighborhood relations in maps, the topographic product P. The topographic product had been invented under the name of" wavering product" in nonlinear dynamics in order to optimize the embeddings of chaotic attractors (Liebert et al., 1991).

Self-organizing feature maps like the Kohonen map (Kohonen, 1989, Ritter et al., 1990) not only provide a plausible explanation for the formation of maps in brains, e.g. in the visual system (Obermayer et al., 1990), but have also been applied to problems like vector quantization, or robot arm control (Martinetz et al., 1990). The underlying organizing principle is the preservation of neighborhood relations. For this principle to lead to a most useful map, the topological structure of the output space must roughly fit the structure of the input data. However, in technical 1141 1142 Bauer, Pawelzik, and Geisel applications this structure is often not a priory known. For this reason several attempts have been made to modify the Kohonen-algorithm such, that not only the weights, but also the output space topology itself is adapted during learning (Kangas et al., 1990, Martinetz et al., 1991). Our contribution is also concerned with optimal output space topologies, but we follow a different approach, which avoids a possibly complicated structure of the output space. First we describe a quantitative measure for the preservation of neighborhood relations in maps, the topographic product P. The topographic product had been invented under the name of" wavering product" in nonlinear dynamics in order to optimize the embeddings of chaotic attractors (Liebert et al., 1991).