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W B Levy

Publications and source records attributed to W B Levy.

At least 37 records · Page 2Linked to original sources

Energy efficient neural codes.

In 1969 Barlow introduced the phrase "economy of impulses" to express the tendency for successive neural systems to use lower and lower levels of cell firings to produce equivalent encodings. From this viewpoint, the ultimate economy of impulses is a neural code of minimal redundancy. The hypothesis motivating our research is that energy expenditures, e.g., the metabolic cost of recovering from an action potential relative to the cost of inactivity, should also be factored into the economy of impulses. In fact, coding schemes with the largest representational capacity are not, in general, optimal when energy expenditures are taken into account. We show that for both binary and analog neurons, increased energy expenditure per neuron implies a decrease in average firing rate if energy efficient information transmission is to be maintained.

Action Potentials↗

A sequence predicting CA3 is a flexible associator that learns and uses context to solve hippocampal-like tasks.

The model discussed in this paper is, by hypothesis, a minimal, biologically plausible model of hippocampal region CA3. Because cognitive mapping can be viewed as a sequence prediction problem, we qualify this model as a successful sequence predictor. Since the model solves problems which require the use of context, the model is also able to learn and use context. The model also solves configural learning problems of which, at least one, requires a hippocampus. Thus, by solving sequence problems, by solving configural learning problems, and by creating codes for context, this model provides a computational unification of hippocampal functions which are often viewed as disparate.

Cognition↗

Context codes and the effect of noisy learning on a simplified hippocampal CA3 model.

This paper investigates how noise affects a minimal computational model of the hippocampus and, in particular, region CA3. The architecture and physiology employed are consistent with the known anatomy and physiology of this region. Here, we use computer simulations to demonstrate and quantify the ability of this model to create context codes in sequential learning problems. These context codes are mediated by local context neurons which are analogous to hippocampal place-coding cells. These local context neurons endow the network with many of its problem-solving abilities. Our results show that the network encodes context on its own and then uses context to solve sequence prediction under ambiguous conditions. Noise during learning affects performance, and it also affects the development of context codes. The relationship between noise and performance in a sequence prediction is simple and corresponds to a disruption of local context neuron firing. As noise exceeds the signal, sequence completion and local context neuron firing are both lost. For the parameters investigated, extra learning trials and slower learning rates do not overcome either of the effects of noise. The results are consistent with the important role played, in this hippocampal model, by local context neurons in sequence prediction and for disambiguation across time.

Artifacts↗

The relationship of local context codes to sequence length memory capacity.

This paper pursues part of a theory to understand sequence prediction networks that use local context neuron encodings. In particular, a previously described neural network model of hippocampal CA3 is studied. General expressions relating CA3 interpattern distances (which reflect local context codes) to sequence length memory capacity are created and verified by computer simulations. As a result, we confirm a very simple relationship between the sequence length memory capacity and a combination of average activity level and the average local context lifetimes. Sequence length memory capacity is also bounded for networks using such local context neuronal codes. Thus, this simple theory quantifies an important limitation on a fully adaptive (i.e. self-supervising) neural model capable of creating context-dependent learning and memory.

Journal Article↗

A simple spike train decoder inspired by the sampling theorem.

Reconstructing a time-varying stimulus estimate from a spike train (Bialek's "decoding" of a spike train) has become an important way to study neural information processing. In this paper, we describe a simple method for reconstructing a time-varying current injection signal from the simulated spike train it produces. This technique extracts most of the information from the spike train, provided that the input signal is appropriately matched to the spike generator. To conceptualize this matching, we consider spikes as instantaneous "samples" of the somatic current. The Sampling Theorem is then applicable, and it suggests that the bandwidth of the injected signal not exceed half the spike generator's average firing rate. The average firing rate, in turn, depends on the amplitude range and DC bias of the injected signal. We hypothesize that nature faces similar problems and constraints when transmitting a time-varying waveform from the soma of one neuron to the dendrite of the postsynaptic cell.

Action Potentials↗

Astrocytic volume fluctuates in the hippocampal CA1 region across the estrous cycle.

The number of dendritic spine synapses in the hippocampal CA1 stratum radiatum fluctuates across the rat estrous cycle, being high on proestrus and low on estrus [20]. We hypothesized that the volume occupied by astrocytic processes changes in a complementary manner. The volume fraction of astrocytic processes was determined stereologically in CA1 s. radiatum and s. lacunosum-moleculare of cycling female rats. Consistent with our hypothesis, the volume fraction was significantly lower on the afternoon of proestrus than on the afternoon of estrus in both laminae.

Animals↗

Rapid development of somatic spines in stratum granulosum of the adult hippocampus in vitro.

Somatic spines on granule cells have been observed occasionally in the adult rat dentate gyrus in vivo. Here we evaluate the appearance and formation of somatic spines in s. granulosum from adult rat hippocampal slices immediately after slice preparation, 45 min, 3 h, and 5 h later. Initially somatic spines are extremely rare but, after 3 h in vitro, they are readily apparent. Some of these somatic spines form asymmetric synapses that have a spherical vesicle-containing presynaptic bouton and a postsynaptic density. Other somatic spines lack a postsynaptic density and may also lack an apposed presynaptic bouton (free somatic spines) as observed in single thin sections. Both the number of these free somatic spines and the number of somatic spine synapses increase in a time-dependent manner. No somatic spines were observed on CA1 pyramidal cells in the same hippocampal slices. Electrophysiological observations indicate that the formation of somatic spine synapses on granule cells in the hippocampal slice occurs without any apparent granule cell activation. The trigger event for this very rapid synaptogenesis in the adult dentate gyrus remains to be determined.

Animals↗

Ultrastructural identification of entorhinal cortical synapses in CA1 stratum lacunosum-moleculare of the rat.

The goal of the present study was to identify and characterize, at the electron microscopic level, the synapses formed by entorhinal cortical (EC) axons in the hippocampal CA1 stratum lacunosum-moleculare of the adult rat. Phaseolus vulgaris leucoagglutinin was ionotophoresed at various loci throughout the mediolateral and dorsoventral extent of the EC to label EC-CA1 synapses. Virtually all labeled synapses were asymmetric and axospinous. EC axons did not preferentially synapse with any particular type of dendritic spine; rather, EC axons formed synapses with the range of dendritic spine morphologies observed in CA1 s. lacunosum-moleculare. Spines with either perforated or nonperforated postsynaptic densities were contacted by EC axons. Occasionally both a labeled and an unlabeled axon synapsed on a single dendritic spine head. The data are discussed in relation to the morphology of other afferent systems synapsing in s. lacunosum-moleculare and to the physiology of the EC-CA1 system.

Animals↗

The influence of limited presynaptic growth and synapse removal on adaptive synaptogenesis.

This report continues our research into the effectiveness of adaptive synaptogenesis in constructing feed-forward networks which perform good transformations on their inputs. Good transformations are characterized by the maintenance of input information and the removal of statistical dependence. Adaptive synaptogenesis stochastically builds and sculpts a synaptic connectivity in initially unconnected networks using two mechanisms. The first, synaptogenesis, creates new, excitatory, feed-forward connections. The second, associative modification, adjusts the strength of existing synapses. Our previous implementations of synaptogenesis only incorporated a postsynaptic regulatory process, receptivity to new innervation (Adelsberger-Mangan and Levy 1993a, b). In the present study, a presynaptic regulatory process, presynaptic avidity, which regulates the tendency of a presynaptic neuron to participate in a new synaptic connection as a function of its total synaptic weight, is incorporated into the synaptogenesis process. In addition, we investigate a third mechanism, selective synapse removal. This process removes synapses between neurons whose firing is poorly correlated. Networks that are constructed with the presynaptic regulatory process maintain more information and remove more statistical dependence than networks constructed with postsynaptic receptivity and associative modification alone. Selective synapse removal also improves network performance, but only when implemented in conjunction with the presynaptic regulatory process.

Computer Simulation↗

Somatic ribosomal changes induced by long-term potentiation of the perforant path-hippocampal CA1 synapses.

The present study quantified ribosomes, as an ultrastructural marker of neuronal protein synthesis, following long-term potentiation (LTP) in the hippocampal CA1 region in vitro. Sixty min after LTP-inducing, high-frequency stimulation of the perforant path, the total number of ribosomes, the number of polysomes, and the number of membrane-bound ribosomes increased significantly. These increases are a postsynaptic morphological correlate consistent with enhanced protein synthesis following the induction of LTP in the perforant path-CA1 system.

Animals↗

Bicuculline permits the induction of long-term depression by heterosynaptic, translaminar conditioning in the hippocampal dentate gyrus.

Using intact, urethane-anesthetized albino rats, we examine long-term depression (LTD) of the ipsilateral medial entorhinal cortical (MEC) response in the dentate gyrus (DG) that is induced by conditioning of the ipsilateral lateral entorhinal cortex (LEC). Although the LEC-DG response potentiates, conditioning the LEC induces little or no LTD of the MEC-DG response. However, bicuculline methiodide, and the correlated presence of population spikes, enable the LEC input to induce LTD of the MEC-DG response. This result suggests that inhibition can control the selectivity of translaminar interactions.

Animals↗

Long-term potentiation of perforant path synapses in hippocampal CA1 in vitro.

This paper reports a study of long-term potentiation (LTP) of perforant path synapses in CA1. Using rat hippocampal slices with CA3 and the dentate gyrus removed, stimulation of the perforant path evoked a population excitatory postsynaptic potential (pEPSP) that was negative-going in s. lacunosum-moleculare of CA1. High-frequency conditioning stimulation of the perforant pathway induced LTP of the perforant path pEPSP in slices disinhibited by the GABAA receptor antagonist bicuculline methiodide (20 microM). Conditioning of the perforant pathway in normal medium, however, failed to induce LTP. Potentiation of the perforant path pEPSP in the presence of bicuculline lasted at least 1 h, was specific to the tetanized pathway, and based on a threshold property, appeared associative in nature.

Animals↗

The dynamics of sparse random networks.

Recurrent neural networks with full symmetric connectivity have been extensively studied as associative memories and pattern recognition devices. However, there is considerable evidence that sparse, asymmetrically connected, mainly excitatory networks with broadly directed inhibition are more consistent with biological reality. In this paper, we use the technique of return maps to study the dynamics of random networks with sparse, asymmetric connectivity and nonspecific inhibition. These networks show three qualitatively different kinds of behavior: fixed points, cycles of low period, and extremely long cycles verging on aperiodicity. Using statistical arguments, we relate these behaviors to network parameters and present empirical evidence for the accuracy of this statistical model. The model, in turn, leads to methods for controlling the level of activity in networks. Studying random, untrained networks provides an understanding of the intrinsic dynamics of these systems. Such dynamics could provide a substrate for the much more complex behavior shown when synaptic modification is allowed.

Animals↗

Adaptive synaptogenesis constructs networks that maintain information and reduce statistical dependence.

This report demonstrates the effectiveness of two processes in constructing simple feedforward networks which perform good transformations on their inputs. Good transformations are characterized by the minimization of two information measures: the information loss incurred with the transformation and the statistical dependency of the output. The two processes build appropriate synaptic connections in initially unconnected networks. The first process, synaptogenesis, creates new synaptic connections; the second process, associative synaptic modification, adjusts the connection strength of existing synapses. Synaptogenesis produces additional innervation for each output neuron until each output neuron achieves a firing rate of approximately 0.50. Associative modification of existing synaptic connections lends robustness to network construction by adjusting suboptimal choices of initial synaptic weights. Networks constructed using synaptogenesis and synaptic modification successfully preserve the information content of a variety of inputs. By recording a high-dimensional input into an output of much smaller dimension, these networks drastically reduce the statistical dependence of neuronal representations. Networks constructed with synaptogenesis and associative modification perform good transformations over a wide range of neuron firing thresholds.

Adaptation, Physiological↗

Ketamine blocks the induction of LTP at the lateral entorhinal cortex-dentate gyrus synapses.

In many regions NMDA receptor activation is required for the synaptic induction of long-term potentiation (LTP). This role for NMDA receptors is controversial at the synapses formed between the cells of the lateral entorhinal cortex (LEC) and the dentate gyrus (DG). Using anesthetized rats, the present study shows that ketamine reversibly blocks the induction of LTP at the LEC-DG synapses, thus favoring a role for NMDA receptors in the induction of LTP there. Ketamine also reversibly blocks the induction of the small translaminar depression of the medial EC response or of the LEC response by conditioning the other system while the test system is inactive.

Animals↗

Longevity of synaptic depression in the hippocampal dentate gyrus.

This study used urethane-anesthetized rats to investigate the longevity of heterosynaptically evoked depression of the monosynaptic response generated by synapses between entorhinal cortical (EC) afferents and the cells of the dentate gyrus (DG). Brief, high-frequency activation of the converging ipsilateral EC-DG input depressed the synaptic response of the contralateral EC-DG synapses without prior experimentally induced potentiation. This depression lasted for hours. Such observations are consistent with a role for heterosynaptically induced long-term depression in the encoding functions of synapses.

Animals↗

Information maintenance and statistical dependence reduction in simple neural networks.

This study compares the ability of excitatory, feed-forward neural networks to construct good transformations on their inputs. The quality of such a transformation is judged by the minimization of two information measures: the information loss of the transformation and the statistical dependency of the output. The networks that are compared differ from each other in the parametric properties of their neurons and in their connectivity. The particular network parameters studied are output firing threshold, synaptic connectivity, and associative modification of connection weights. The network parameters that most directly affect firing levels are threshold and connectivity. Networks incorporating neurons with dynamic threshold adjustment produce better transformations. When firing threshold is optimized, sparser synaptic connectivity produces a better transformation than denser connectivity. Associative modification of synaptic weights confers only a slight advantage in the construction of optimal transformations. Additionally, our research shows that some environments are better suited than others for recording. Specifically, input environments high in statistical dependence, i.e. those environments most in need of recoding, are more likely to undergo successful transformations.

Action Potentials↗