A scientific theory of ars memoriae : spatial view cells in a continuous attractor network with linked items

The art of memory (ars memoriae) used since classical times includes using a well-known scene to associate each view or part of the scene with a different item in a speech. This memory technique is also known as the “method of loci.” The new theory is proposed that this type of memory is implemented in the CA3 region of the hippocampus where there are spatial view cells in primates that allow a particular view to be associated with a particular object in an event or episodic memory. Given that the CA3 cells with their extensive recurrent collateral system connecting different CA3 cells, and associative synaptic modifiability, form an autoassociation or attractor network, the spatial view cells with their approximately Gaussian view fields become linked in a continuous attractor network. As the view space is traversed continuously (e.g., by self-motion or imagined self-motion across the scene), the views are therefore successively recalled in the correct order, with no view missing, and with low interference between the items to be recalled. Given that each spatial view has been associated with a different discrete item, the items are recalled in the correct order, with none missing. This is the first neuroscience theory of ars memoriae. The theory provides a foundation for understanding how a key feature of ars memoriae, the ability to use a spatial scene to encode a sequence of items to be remembered, is implemented

The mechanisms for pattern completion and pattern separation in the hippocampus

The mechanisms for pattern completion and pattern separation are described in the context of a theory of hippocampal function in which the hippocampal CA3 system operates as a single attractor or autoassociation network to enable rapid, one-trial, associations between any spatial location (place in rodents, or spatial view in primates) and an object or reward, and to provide for completion of the whole memory during recall from any part. The factors important in the pattern completion in CA3 together with a large number of independent memories stored in CA3 include a sparse distributed representation which is enhanced by the graded firing rates of CA3 neurons, representations that are independent due to the randomizing effect of the mossy fibers, heterosynaptic long-term depression as well as long-term potentiation in the recurrent collateral synapses, and diluted connectivity to minimize the number of multiple synapses between any pair of CA3 neurons which otherwise distort the basins of attraction. Recall of information from CA3 is implemented by the entorhinal cortex perforant path synapses to CA3 cells, which in acting as a pattern associator allow some pattern generalization. Pattern separation is performed in the dentate granule cells using competitive learning to convert grid-like entorhinal cortex firing to place-like fields. Pattern separation in CA3, which is important for completion of any one of the stored patterns from a fragment, is provided for by the randomizing effect of the mossy fiber synapses to which neurogenesis may contribute, by the large number of dentate granule cells each with a sparse representation, and by the sparse independent representations in CA3. Recall to the neocortex is achieved by a reverse hierarchical series of pattern association networks implemented by the hippocampo-cortical backprojections, each one of which performs some pattern generalization, to retrieve a complete pattern of cortical firing in higher-order cortical areas

Decision time, slow inhibition, and theta rhythm

In this paper, we examine decision making in a spiking neuronal network and show that longer time constants for the inhibitory neurons can decrease the reaction times and produce theta rhythm.We analyze the mechanism and find that the spontaneous firing rate before the decision cues are applied can drift, and thereby influence the speed of the reaction time when the decision cues are applied. The drift of the firing rate in the population that will win the competition is larger if the time constant of the inhibitory interneurons is increased from 10 to 33 ms, and even larger if there are two populations of inhibitory neurons with time constants of 10 and 100 ms. Of considerable interest is that the decision that will be made can be influenced by the noise-influenced drift of the spontaneous firing rate over many seconds before the decision cues are applied. The theta rhythm associated with the longer time constant networks mirrors the greater integration in the firing rate drift produced by the recurrent connections over long time periods in the networks with slow inhibition. The mechanism for the effect of slow waves in the theta and delta range on decision times is suggested to be increased neuronal spiking produced by depolarization of the membrane potential on the positive part of the slow waves when the neuron’s membrane potential is close to the firing threshold

Cortical coding

The encoding of information in the primate inferior temporal visual cortex, hippocampus, orbitofrontal cortex, and insula is described. All these areas have sparse distributed graded firing rate representations. The firing rate probability distribution is close to exponential. The information increases approximately linearly with the number of neurons. Consistent with this relative independence, there is little extra information that is available because of stimulus-dependent synchrony, and little redundancy. The code can be read very fast, in 20–50 ms, by dot-product, biologically plausible decoding. The advantages of this code include high capacity, generalisation, graceful degradation, and rapid readout of the information by biologically plausible dot-product decoding. None of these are properties of local or “grandmother-cell’ representations. Consistent evidence is becoming available for humans. These sparse distributed graded firing rate representations have major computational advantages for the brain including for language that are not met by local or ‘grandmother-cell’ representations

Neurobiological foundations of aesthetics and art

A theory of the neurobiological foundations of aesthetics and art is described. This has its roots in emotion, in which what is pleasant or unpleasant, a reward or punisher, is the result of an evolutionary process in which genes define the (pleasant or unpleasant) goals for action. To this is added the operation of the reasoning, syntactic, brain system which evolved to help solve difficult, multistep, problems, and the use of which is encouraged by pleasant feelings when elegant, simple, and hence aesthetic solutions are found that are advantageous because they are parsimonious, and follow Occam's Razor. The combination of these two systems, and the interactions between them, provide an approach to understanding aesthetics that is rooted in evolution and its effects on brain design and function

The roles of the orbitofrontal cortex via the habenula in non-reward and depression, and in the responses of serotonin and dopamine neurons

Cortical regions such as the orbitofrontal cortex involved in reward and in non-reward and which are implicated in depression, and the amygdala, are connected to the habenula via the striatum and pallidum, and via subcortical limbic structures. The habenula in turn projects to the raphe nuclei, the source of the serotonin-containing neurons that project to the forebrain. It is proposed that this provides a route for cortical signals related to reward, and to not obtaining expected rewards, to influence the serotonin-containing neuronal system that is influenced by many antidepressant treatments. This helps to provide a more circuit-based understanding of the brain mechanisms related to depression, and how some treatments influence this system. The habenula also projects via the rostromedial tegmental nucleus to the dopamine-containing neurons, and this, it is proposed, provides a route for reward prediction error signals and other reward- and punishment-related signals of cortical and striatal origin to influence the dopamine system

A non-reward attractor theory of depression

A non-reward attractor theory of depression is proposed based on the operation of the lateral orbitofrontal cortex and supracallosal cingulate cortex. The orbitofrontal cortex contains error neurons that respond to non-reward for many seconds in an attractor state that maintains a memory of the non-reward. The human lateral orbitofrontal cortex is activated by non-reward during reward reversal, and by a signal to stop a response that is now incorrect. Damage to the human orbitofrontal cortex impairs reward reversal learning. Not receiving reward can produce depression. The theory proposed is that in depression, this lateral orbitofrontal cortex non-reward system is more easily triggered, and maintains its attractor-related firing for longer. This triggers negative cognitive states, which in turn have positive feedback top-down effects on the orbitofrontal cortex non-reward system. Treatments for depression, including ketamine, may act in part by quashing this attractor. The mania of bipolar disorder is hypothesized to be associated with oversensitivity and overactivity in the reciprocally related reward system in the medial orbitofrontal cortex and pregenual cingulate cortex

Chemosensory Learning in the Cortex

Taste is a primary reinforcer. Olfactory–taste and visual–taste association learning takes place in the primate including human orbitofrontal cortex to build representations of flavor. Rapid reversal of this learning can occur using a rule-based learning system that can be reset when an expected taste or flavor reward is not obtained, that is by negative reward prediction error, to which a population of neurons in the orbitofrontal cortex responds. The representation in the orbitofrontal cortex but not the primary taste or olfactory cortex is of the reward value of the visual/olfactory/taste input as shown by devaluation experiments in which food is fed to satiety, and by correlations of the activations with subjective pleasantness ratings in humans. Sensory-specific satiety for taste, olfactory, visual, and oral somatosensory inputs produced by feeding a particular food to satiety is implemented it is proposed by medium-term synaptic adaptation in the orbitofrontal cortex. Cognitive factors, including word-level descriptions, modulate the representation of the reward value of food in the orbitofrontal cortex, and this effect is learned it is proposed by associative modification of top-down synapses onto neurons activated by bottom-up taste and olfactory inputs when both are active in the orbitofrontal cortex. A similar associative synaptic learning process is proposed to be part of the mechanism for the top-down attentional control to the reward value vs. the sensory properties such as intensity of taste and olfactory inputs in the orbitofrontal cortex, as part of a biased activation theory of selective attention

Invariant Visual Object and Face Recognition: Neural and Computational Bases, and a Model, VisNet

Neurophysiological evidence for invariant representations of objects and faces in the primate inferior temporal visual cortex is described. Then a computational approach to how invariant representations are formed in the brain is described that builds on the neurophysiology. A feature hierarchy model in which invariant representations can be built by self-organizing learning based on the temporal and spatial statistics of the visual input produced by objects as they transform in the world is described. VisNet can use temporal continuity in an associative synaptic learning rule with a short-term memory trace, and/or it can use spatial continuity in continuous spatial transformation learning which does not require a temporal trace. The model of visual processing in the ventral cortical stream can build representations of objects that are invariant with respect to translation, view, size, and also lighting. The model has been extended to provide an account of invariant representations in the dorsal visual system of the global motion produced by objects such as looming, rotation, and object-based movement. The model has been extended to incorporate top-down feedback connections to model the control of attention by biased competition in, for example, spatial and object search tasks. The approach has also been extended to account for how the visual system can select single objects in complex visual scenes, and how multiple objects can be represented in a scene. The approach has also been extended to provide, with an additional layer, for the development of representations of spatial scenes of the type found in the hippocampus

Neural computations underlying phenomenal consciousness : a higher order syntactic thought theory

Problems are raised with the global workspace hypothesis of consciousness, for example about exactly how global the workspace needs to be for consciousness to suddenly be present. Problems are also raised with Carruthers's (2019) version that excludes conceptual (categorical or discrete) representations, and in which phenomenal consciousness can be reduced to physical processes, with instead a different levels of explanation approach to the relation between the brain and the mind advocated. A different theory of phenomenal consciousness is described, in which there is a particular computational system involved in which Higher Order Syntactic Thoughts are used to perform credit assignment on first order thoughts of multiple step plans to correct them by manipulating symbols in a syntactic type of working memory. This provides a good evolutionary reason for the evolution of this kind of computational module, with which, it is proposed, phenomenal consciousness is associated. Some advantages of this HOST approach to phenomenal consciousness are then described with reference not only to the global workspace approach, but also to Higher Order Thought (HOT) theories. It is hypothesized that the HOST system which requires the ability to manipulate first order symbols in working memory might utilize parts of the prefrontal cortex implicated in working memory, and especially the left inferior frontal gyrus, which is involved in language and probably syntactical processing. Overall, the approach advocated is to identify the computations that are linked to consciousness, and to analyze the neural bases of those computations. [Abstract copyright: Copyright © 2020 Rolls.