Toward a dynamical systems analysis of neuromodulation

This work presents some first steps toward a more thorough understanding of the control systems employed in evolutionary robotics. In order to choose an appropriate architecture or to construct an effective novel control system we need insights into what makes control systems successful, robust, evolvable, etc. Here we present analysis intended to shed light on this type of question as it applies to a novel class of artificial neural networks that include a neuromodulatory mechanism: GasNets. We begin by instantiating a particular GasNet subcircuit responsible for tuneable pattern generation and thought to underpin the attractive property of “temporal adaptivity”. Rather than work within the GasNet formalism, we develop an extension of the well-known FitzHugh-Nagumo equations. The continuous nature of our model allows us to conduct a thorough dynamical systems analysis and to draw parallels between this subcircuit and beating/bursting phenomena reported in the neuroscience literature. We then proceed to explore the effects of different types of parameter modulation on the system dynamics. We conclude that while there are key differences between the gain modulation used in the GasNet and alternative schemes (including threshold modulation of more traditional synaptic input), both approaches are able to produce tuneable pattern generation. While it appears, at least in this study, that the GasNet’s gain modulation may not be crucial to pattern generation , we go on to suggest some possible advantages it could confer

Direct evidence for local oscillatory current sources and intracortical phase gradients in turtle visual cortex

Visual stimuli induce oscillations in the membrane potential of neurons in cortices of several species. In turtle, these oscillations take the form of linear and circular traveling waves. Such waves may be a consequence of a pacemaker that emits periodic pulses of excitation that propagate across a network of excitable neuro-nal tissue or may result from continuous and possibly reconfigu-rable phase shifts along a network with multiple weakly coupled neuronal oscillators. As a means to resolve the origin of wave propagation in turtle visual cortex, we performed simultaneous measurements of the local field potential at a series of depths throughout this cortex. Measurements along a single radial pen-etration revealed the presence of broadband current sources, with a center frequency near 20 Hz ( g band), that were activated by visual stimulation. The spectral coherence between sources at two well-separated loci along a rostral– caudal axis revealed the pres-ence of systematic timing differences between localized cortical oscillators. These multiple oscillating current sources and their timing differences in a tangential plane are interpreted as the neuronal activity that underlies the wave motion revealed in previous imaging studies. The present data provide direct evidence for the inference from imaging of bidirectional wave motion that the stimulus-induced electrical waves in turtle visual cortex corre-spond to phase shifts in a network of coupled neuronal oscillators

Metabotropic Glutamate Receptor Activation in Cerebelar Purkinje Cells as Substrate for Adaptive Timing of the Classicaly Conditioned Eye Blink Response

To understand how the cerebellum adaptively times the classically conditioned nictitating membrane response (NMR), a model of the metabotropic glutamate receptor (mGluR) second messenger system in cerebellar Purkinje cells is constructed. In the model slow responses, generated postsynaptically by mGluR-mediated phosphoinositide hydrolysis, and calcium release from intracellular stores, bridge the interstimulus interval (ISI) between the onset of parallel fiber activity associated with the conditioned stimulus (CS) and climbing fiber activity associated with unconditioned stimulus (US) onset. Temporal correlation of metabotropic responses and climbing fiber signals produces persistent phosphorylation of both AMPA receptors and Ca2+-dependent K+ channels. This is responsible for long-term depression (LTD) of AMPA receptors. The phosphorylation of Ca2+-dependent K+ channels leads to a reduction in baseline membrane potential and a reduction of Purkinje cell population firing during the CS-US interval. The Purkinje cell firing decrease disinhibits cerebellar nuclear cells which then produce an excitatory response corresponding to the learned movement. Purkinje cell learning times the response, while nuclear cell learning can calibrate it. The model reproduces key features of the conditioned rabbit NMR: Purkinje cell population response is properly timed, delay conditioning occurs for ISIs of up to four seconds while trace conditioning occurs only at shorter ISIs, mixed training at two different ISis produces a double-peaked response, and ISIs of 200-400ms produce maximal responding. Biochemical similarities between timed cerebellar learning and photoreceptor transduction, and circuit similarities between the timed cerebellar circuit and a timed dentate-CA3 hippocampal circuit, are noted.Office of Naval Research (N00014- 92-J-4015, N00014-92-J-1309, N00014-95-1-0409); Air Force Office of Scientific Research (F49620-92-J-0225);National Science Foundation (IRI-90-24877

A Neural Model of Timed Response Learning in the Cerebellum

A spectral timing model is developed to explain how the cerebellum learns adaptively timed responses during the rabbit's conditioned nictitating membrane response (NMR). The model posits two learning sites that respectively enable conditioned excitation and timed disinhibition of the response. Long-term potentiation of mossy fiber pathways projecting to interpositus nucleus cells allows conditioned excitation of the response's adaptive gain. Long-term depression of parallel fiber- Purkinje cell synapses in the cerebellar cortex allows learning of an adaptively timed reduction in Purkinje cell inhibition of the same nuclear cells. A spectrum of partially timed responses summate to generate an accurately timed population response. In agreement with physiological data, the model Purkinje cell activity decreases in the interval following the onset of the conditioned stimulus, and nuclear cell responses match conditioned response (CR) topography. The model reproduces key behavioral features of the NMR, including the properties that CR peak amplitude occurs at the unconditioned stimulus (US) onset, a discrete CR peak shift occurs with a change in interstimulus interval (ISI) between conditioned stim- ulus (CS) and US, mixed training at two different ISis produces a double-peaked CR, CR acquisition and rate of responding depend unimodally on the lSI, CR onset latency decreases during training, and maladaptively-timed, small-amplitude CRs result from ablation of cerebellar cortex.National Science Foundation (IRI-90-24877); Office of Naval Research (N00014-92-J-1309); Air Force Office of Scientific Research (F49620-92-J-0225

A Neural Circuit Model for Prospective Control of Interceptive Reaching

Two prospective controllers of hand movements in catching -- both based on required velocity control -- were simulated. Under certain conditions, this required velocity controlled to overshoots of the future interception point. These overshoots were absent in pertinent experiments. To remedy this shortcoming, the required velocity model was reformulated in terms of a neural network, the Vector Integration To Endpoint model, to create a Required Velocity Integration To Endpoint modeL Addition of a parallel relative velocity channel, resulting in the Relative and Required Velocity Integration To Endpoint model, provided a better account for the experimentally observed kinematics than the existing, purely behavioral models. Simulations of reaching to intercept decelerating and accelerating objects in the presence of background motion were performed to make distinct predictions for future experiments.Vrije Universiteit (Gerrit-Jan van Jngen-Schenau stipend of the Faculty of Human Movement Sciences); Royal Netherlands Academy of Arts and Sciences; Defense Advanced Research Projects Agency and Office of Naval Research (N00014-95-1-0409

Can the mid-Holocene provide suitable models for rewilding the landscape in Britain?

Palaeoecologists have been encouraging us to think about the relevance of the Holocene fossil record for nature conservation for many years (e.g. Buckland 1993) but this information seems slow to filter through to the conservation community. Indeed, Willis et al. (2005) report that recently published biodiversity reports and policy documents rarely look back more than 50 years and may ignore the historical context entirely. This has been a lost opportunity for understanding ecological systems. Many natural processes occur over timescales that confound our attempts to understand them, so the vast temporal perspective provided by palaeoecological studies can provide important guidance for nature conservation (Willis & Birks 2006). However, accurate vegetation mapping is difficult enough in modern landscapes (Cherrill & McLean 1999), so the challenge of describing prehistoric environments is immeasurably greater. Nevertheless, pioneering work in the mid 20th century showed that pollen and spores extracted from peat bogs were so perfectly preserved thatthey could be used to demonstrate sequences of vegetation change since the last glaciation (Godwin 1956). Since then, the science has burgeoned: ancient deposits of beetles, snails, fungal spores and plant macrofossils add to the picture, as does the chemistry of ancient lake sediments (Bell & Walker 2004). Many questions still remain to be answered by this fascinating research and one aspect has received considerable attention in the last decade. This concerns the nature of the ‘primeval’ landscapes, in other words our understanding of natural systems prior to significant human impact. The debate was kindled by a thesis by the Dutch forest ecologist Frans Vera in 2000 (see also Vera & Buissink 2007). Vera effectively challenged established views about the primeval landscapes and argued that the refutation, and the resulting alternative landscape models, had critical importance for modern conservation practice. Vera’s thesis is focused on the pre-Neolithic (ca 8000-5000bp) landscape in the lowlands of central and western Europe, with the assumption that this period represents an almost pristine or ‘natural’ state which should provide a suitable conservation benchmark. Vera contends (i) that this landscape was not closed woodland but a relatively open park-like mosaic of wood and grassland,and (ii) that large wild herbivores were an essential driving force behind woodland-grassland vegetation cycles. The advocacy in his argument and the timing of the publication, when grazingwas seen as increasingly important in conservation in Europe, have combined to raise the profile of this issue. If Vera is correct, the open park-like landscapes were inherited rather than created by people; this may have implications for conservation practice in Europe. The adoption of Vera’s ideas into conservation management plans in the UK (see Box 1) gives an indication of the influence that this work has had. Indeed, Vera’s ideas have been described as a ‘challenge to orthodox thinking’ (Miller 2002) and considerable debate has been stimulated centering on the ecological validity of Vera’s hypothesis and its relevance for modern conservation. In this article, we attempt to address these issues on the basis of results from a literature review, web-debate and discussions with Dutch and British ecologists, prepared for English Nature with a view to informing conservation strategies (Hodder & Bullock 2005a)