An active neural mechanism for relational learning and fast knowledge reassembly.

How do we gain general insights from limited novel experiences? Humans and animals have a striking ability to learn relationships between experienced items, enabling efficient generalization and rapid assimilation of new information. One fundamental instance of such relational learning is transitive inference (learn A>B and B>C, infer A>C), which can be quickly and globally reorganized upon learning a new item (learn A>B>C and D>E>F, then C>D, and infer B>E). Despite considerable study, neural mechanisms of transitive inference and fast reassembly of existing knowledge remain elusive. Here we adopt a meta-learning ("learning-to-learn") approach. We train artificial neural networks, endowed with synaptic plasticity and neuromodulation, to be able to learn novel orderings of arbitrary stimuli from repeated presentation of stimulus pairs. We then obtain a complete mechanistic understanding of this discovered neural learning algorithm. Remarkably, this learning involves active cognition: items from previous trials are selectively reinstated in working memory, enabling delayed, self-generated learning and knowledge reassembly. These findings identify a new mechanism for relational learning and insight, suggest new interpretations of neural activity in cognitive tasks, and highlight a novel approach to discovering neural mechanisms capable of supporting cognitive behaviors.

Biologically plausible learning in recurrent neural networks reproduces neural dynamics observed during cognitive tasks.

Neural activity during cognitive tasks exhibits complex dynamics that flexibly encode task-relevant variables. Chaotic recurrent networks, which spontaneously generate rich dynamics, have been proposed as a model of cortical computation during cognitive tasks. However, existing methods for training these networks are either biologically implausible, and/or require a continuous, real-time error signal to guide learning. Here we show that a biologically plausible learning rule can train such recurrent networks, guided solely by delayed, phasic rewards at the end of each trial. Networks endowed with this learning rule can successfully learn nontrivial tasks requiring flexible (context-dependent) associations, memory maintenance, nonlinear mixed selectivities, and coordination among multiple outputs. The resulting networks replicate complex dynamics previously observed in animal cortex, such as dynamic encoding of task features and selective integration of sensory inputs. We conclude that recurrent neural networks offer a plausible model of cortical dynamics during both learning and performance of flexible behavior.

Spontaneous emergence of fast attractor dynamics in a model of developing primary visual cortex.

Recent evidence suggests that neurons in primary sensory cortex arrange into competitive groups, representing stimuli by their joint activity rather than as independent feature analysers. A possible explanation for these results is that sensory cortex implements attractor dynamics, although this proposal remains controversial. Here we report that fast attractor dynamics emerge naturally in a computational model of a patch of primary visual cortex endowed with realistic plasticity (at both feedforward and lateral synapses) and mutual inhibition. When exposed to natural images (but not random pixels), the model spontaneously arranges into competitive groups of reciprocally connected, similarly tuned neurons, while developing realistic, orientation-selective receptive fields. Importantly, the same groups are observed in both stimulus-evoked and spontaneous (stimulus-absent) activity. The resulting network is inhibition-stabilized and exhibits fast, non-persistent attractor dynamics. Our results suggest that realistic plasticity, mutual inhibition and natural stimuli are jointly necessary and sufficient to generate attractor dynamics in primary sensory cortex.

Defining and simulating open-ended novelty: requirements, guidelines, and challenges.

The open-endedness of a system is often defined as a continual production of novelty. Here we pin down this concept more fully by defining several types of novelty that a system may exhibit, classified as variation, innovation, and emergence. We then provide a meta-model for including levels of structure in a system's model. From there, we define an architecture suitable for building simulations of open-ended novelty-generating systems and discuss how previously proposed systems fit into this framework. We discuss the design principles applicable to those systems and close with some challenges for the community.

A Feedback Model of Attention Explains the Diverse Effects of Attention on Neural Firing Rates and Receptive Field Structure.

Visual attention has many effects on neural responses, producing complex changes in firing rates, as well as modifying the structure and size of receptive fields, both in topological and feature space. Several existing models of attention suggest that these effects arise from selective modulation of neural inputs. However, anatomical and physiological observations suggest that attentional modulation targets higher levels of the visual system (such as V4 or MT) rather than input areas (such as V1). Here we propose a simple mechanism that explains how a top-down attentional modulation, falling on higher visual areas, can produce the observed effects of attention on neural responses. Our model requires only the existence of modulatory feedback connections between areas, and short-range lateral inhibition within each area. Feedback connections redistribute the top-down modulation to lower areas, which in turn alters the inputs of other higher-area cells, including those that did not receive the initial modulation. This produces firing rate modulations and receptive field shifts. Simultaneously, short-range lateral inhibition between neighboring cells produce competitive effects that are automatically scaled to receptive field size in any given area. Our model reproduces the observed attentional effects on response rates (response gain, input gain, biased competition automatically scaled to receptive field size) and receptive field structure (shifts and resizing of receptive fields both spatially and in complex feature space), without modifying model parameters. Our model also makes the novel prediction that attentional effects on response curves should shift from response gain to contrast gain as the spatial focus of attention drifts away from the studied cell.

There's Waldo! A Normalization Model of Visual Search Predicts Single-Trial Human Fixations in an Object Search Task.

When searching for an object in a scene, how does the brain decide where to look next? Visual search theories suggest the existence of a global "priority map" that integrates bottom-up visual information with top-down, target-specific signals. We propose a mechanistic model of visual search that is consistent with recent neurophysiological evidence, can localize targets in cluttered images, and predicts single-trial behavior in a search task. This model posits that a high-level retinotopic area selective for shape features receives global, target-specific modulation and implements local normalization through divisive inhibition. The normalization step is critical to prevent highly salient bottom-up features from monopolizing attention. The resulting activity pattern constitues a priority map that tracks the correlation between local input and target features. The maximum of this priority map is selected as the locus of attention. The visual input is then spatially enhanced around the selected location, allowing object-selective visual areas to determine whether the target is present at this location. This model can localize objects both in array images and when objects are pasted in natural scenes. The model can also predict single-trial human fixations, including those in error and target-absent trials, in a search task involving complex objects.

The gamma slideshow: object-based perceptual cycles in a model of the visual cortex.

While recent studies have shed light on the mechanisms that generate gamma (>40 Hz) oscillations, the functional role of these oscillations is still debated. Here we suggest that the purported mechanism of gamma oscillations (feedback inhibition from local interneurons), coupled with lateral connections implementing "Gestalt" principles of object integration, naturally leads to a decomposition of the visual input into object-based "perceptual cycles," in which neuron populations representing different objects within the scene will tend to fire at successive cycles of the local gamma oscillation. We describe a simple model of V1 in which such perceptual cycles emerge automatically from the interaction between lateral excitatory connections (linking oriented cells falling along a continuous contour) and fast feedback inhibition (implementing competitive firing and gamma oscillations). Despite its extreme simplicity, the model spontaneously gives rise to perceptual cycles even when faced with natural images. The robustness of the system to parameter variation and to image complexity, together with the paucity of assumptions built in the model, support the hypothesis that perceptual cycles occur in natural vision.

Evolution and complexity: the double-edged sword.

We attempt to provide a comprehensive answer to the question of whether, and when, an arrow of complexity emerges in Darwinian evolution. We note that this expression can be interpreted in different ways, including a passive, incidental growth, or a pervasive bias towards complexification. We argue at length that an arrow of complexity does indeed occur in evolution, which can be most reasonably interpreted as the result of a passive trend rather than a driven one. What, then, is the role of evolution in the creation of this trend, and under which conditions will it emerge? In the later sections of this article we point out that when certain proper conditions (which we attempt to formulate in a concise form) are met, Darwinian evolution predictably creates a sustained trend of increase in maximum complexity (that is, an arrow of complexity) that would not be possible without it; but if they are not, evolution will not only fail to produce an arrow of complexity, but may actually prevent any increase in complexity altogether. We conclude that, with regard to the growth of complexity, evolution is very much a double-edged sword.