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Models of neural computation are attempts to elucidate, in an abstract and mathematical fashion, the core principles that underlie information processing in biological nervous systems, or functional components thereof. This article aims to provide an overview of the most definitive models of neuro-biological computation as well as the tools commonly used to construct and analyze them.

Introduction

Due to the complexity of nervous system behavior, the associated experimental error bounds are ill-defined, but the relative merit of the different models of a particular subsystem can be compared according to how closely they reproduce real-world behaviors or respond to specific input signals. In the closely related field of computational neuroethology, the practice is to include the environment in the model in such a way that the loop is closed. In the cases where competing models are unavailable, or where only gross responses have been measured or quantified, a clearly formulated model can guide the scientist in designing experiments to probe biochemical mechanisms or network connectivity.

In all but the simplest cases, the mathematical equations that form the basis of a model cannot be solved exactly. Nevertheless, computer technology, sometimes in the form of specialized software or hardware architectures, allow scientists to perform iterative calculations and search for plausible solutions. A computer chip or a robot that can interact with the natural environment in ways akin to the original organism is one embodiment of a useful model. The ultimate measure of success is however the ability to make testable predictions.

General criteria for evaluating models

Speed of information processing

The rate of information processing in biological neural systems are constrained by the speed at which an action potential can propagate down a nerve fibre. This conduction velocity ranges from 1 m/s to over 100 m/s, and generally increases with the diameter of the neuronal process. Slow in the timescales of biologically-relevant events dictated by the speed of sound or the force of gravity, the nervous system overwhelmingly prefers parallel computations over serial ones in time-critical applications.

Robustness

A model is robust if it continues to produce the same computational results under small variations in inputs or operating parameters. For example, the direction of motion as computed by a robust motion detector would not change under small changes of luminance, contrast or velocity jitter.

Gain control

This refers to the principle that the response of a nervous system should stay within certain bounds even as the inputs from the environment change drastically. For example, when adjusting between a sunny day and a moonless night, the retina changes the relationship between light level and neuronal output by a factor of more than so that the signals sent to later stages of the visual system always remain within a much narrower range of amplitudes.[1] [2] [3]

Linearity versus nonlinearity

A linear system is one whose response in a specified unit of measure, to a set of inputs considered at once, is the sum of its responses due to the inputs considered individually.

Linear systems are easier to analyze mathematically. Linearity may occur in the basic elements of a neural circuit such as the response of a postsynaptic neuron, or as an emergent property of a combination of nonlinear subcircuits.[4]

Examples

A computational neural model may be constrained to the level of biochemical signalling in individual neurons or it may describe an entire organism in its environment. The examples here are grouped according to their scope.

Models of information transfer in neurons

Mining Engineer (Excluding Oil ) Truman from Alma, loves to spend time knotting, largest property developers in singapore developers in singapore and stamp collecting. Recently had a family visit to Urnes Stave Church. The most widely used models of information transfer in biological neurons are based on analogies with electrical circuits. The equations to be solved are time-dependent differential equations with electro-dynamical variables such as current, conductance or resistance, capacitance and voltage.

Hodgkin–Huxley model and its derivatives

Mining Engineer (Excluding Oil ) Truman from Alma, loves to spend time knotting, largest property developers in singapore developers in singapore and stamp collecting. Recently had a family visit to Urnes Stave Church. The Hodgkin–Huxley model, widely regarded as one of the great achievements of 20th-century biophysics, describes how action potentials in neurons are initiated and propagated in axons via voltage-gated ion channels. It is a set of nonlinear ordinary differential equations that were introduced by Alan Lloyd Hodgkin and Andrew Huxley in 1952 to explain the results of voltage clamp experiments on the squid giant axon. Analytic solutions do not exist, but the Levenberg–Marquardt algorithm, a modified Gauss–Newton algorithm, is often used to fit these equations to voltage-clamp data.

The FitzHugh–Nagumo model is a simplication of the Hodgkin–Huxley model. The Hindmarsh–Rose model is an extension which describes neuronal spike bursts. The Morris–Lecar model is a modification which does not generate spikes, but describes slow-wave propagation, which is implicated in the inhibitory synaptic mechanisms of central pattern generators.

Solitons

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Transfer functions and linear filters

This approach, influenced by control theory and signal processing, treats neurons and synapses as time-invariant entities that produce outputs that are linear combinations of input signals, often depicted as sine waves with a well-defined temporal or spatial frequencies.

The entire behavior of a neuron or synapse are encoded in a transfer function, lack of knowledge concerning the exact underlying mechanism notwithstanding. This brings a highly developed mathematics to bear on the problem of information transfer.

The accompanying taxonomy of linear filters turns out to be useful in characterizing neural circuitry. Both low- and high-pass filters are postulated to exist in some form in sensory systems, as they act to prevent information loss in high and low contrast environments, respectively.

Indeed, measurements of the transfer functions of neurons in the horseshoe crab retina according to linear systems analysis show that they remove short-term fluctuations in input signals leaving only the long-term trends, in the manner of low-pass filters. These animals are unable to see low-contrast objects without the help of optical distortions caused by underwater currents.[4][5]

Models of computations in sensory systems

Lateral inhibition in the retina: Hartline–Ratliff equations

In the retina, an excited neural receptor can suppress the activity of surrounding neurons within an area called the inhibitory field. This effect, known as lateral inhibition, increases the contrast and sharpness in visual response, but leads to the epiphenomenon of Mach bands. This is often illustrated by the optical illusion of light or dark stripes next to a sharp boundary between two regions in an image of different luminance.

The Hartline-Ratliff model describes interactions within a group of p photoreceptor cells.[6] Assuming these interactions to be linear, they proposed the following relationship for the steady-state response rate of the given p-th photoreceptor in terms of the steady-state response rates of the j surrounding receptors:

.

Here,

is the excitation of the target p-th receptor from sensory transduction

is the associated threshold of the firing cell, and

is the coefficient of inhibitory interaction between the p-th and the jth receptor. The inhibitory interaction decreases with distance from the target p-th receptor.

Cross-correlation in sound localization: Jeffress model

According to Jeffress,[7] in order to compute the location of a sound source in space from interaural time differences, an auditory system relies on delay lines: the induced signal from an ipsilateral auditory receptor to a particular neuron is delayed for the same time as it takes for the original sound to go in space from that ear to the other. Each postsynaptic cell is differently delayed and thus specific for a particular inter-aural time difference. This theory is equivalent to the mathematical procedure of cross-correlation.

Following Fischer and Anderson,[8] the response of the postsynaptic neuron to the signals from the left and right ears is given by

where

and

represents the delay function. This is not entirely correct and a clear eye is needed to put the symbols in order.

Structures have been located in the barn owl which are consistent with Jeffress-type mechanisms.[9]

Cross-correlation for motion detection: Hassenstein–Reichardt model

A motion detector needs to satisfy three general requirements: pair-inputs, asymmetry and nonlinearity.[10] The cross-correlation operation implemented asymmetrically on the responses from a pair of photoreceptors satisfies these minimal criteria, and furthermore, predicts features which have been observed in the response of neurons of the lobula plate in bi-wing insects.[11]

The master equation for response is

The HR model predicts a peaking of the response at a particular input temporal frequency. The conceptually similar Barlow–Levick model is deficient in the sense that a stimulus presented to only one receptor of the pair is sufficient to generate a response. This is unlike the HR model, which requires two correlated signals delivered in a time ordered fashion. However the HR model does not show a saturation of response at high contrasts, which is observed in experiment. Extensions of the Barlow-Levick model can provide for this discrepancy.[12]

Watson–Ahumada model for motion estimation in humans

This uses a cross-correlation in both the spatial and temporal directions, and is related to the concept of optical flow. [13]

Neurophysiological metronomes: neural circuits for pattern generation

Mutually inhibitory processes are a unifying motif of all central pattern generators. This has been demonstrated in the stomatogastric (STG) nervous system of crayfish and lobsters.[14] Two and three-cell oscillating networks based on the STG have been constructed which are amenable to mathematical analysis, and which depend in a simple way on on synaptic strengths and overall activity, presumably the knobs on these things.[15] The mathematics involved is the theory of dynamical systems.

Anti-Hebbian adaptation: spike-timing dependent plasticity

[16]

[17]

Feedback and control: models of flight control in the fly

Flight control in the fly is believed to be mediated by inputs from the visual system and also the halteres, a pair of knob-like organs which measure angular velocity. Integrated computer models of Drosophila, short on neuronal circuitry but based on the general guidelines given by control theory and data from the tethered flights of flies, have been constructed to investigate the details of flight control [1][2].

Software modelling approaches and tools

Neural networks

Mining Engineer (Excluding Oil ) Truman from Alma, loves to spend time knotting, largest property developers in singapore developers in singapore and stamp collecting. Recently had a family visit to Urnes Stave Church. In this approach the strength and type, excitatory or inhibitory, of synaptic connections are represented by the magnitude and sign of weights, that is, numerical coefficients in front of the inputs x to a particular neuron. The response of the j-th neuron is given by a sum of nonlinear, usually "sigmoidal" functions of the inputs as:

.

This response is then fed as input into other neurons and so on. The goal is to solve for the weights given a desired response at the output layer to a set given inputs at the input layer.

Genetic algorithms

Genetic algorithms are used to evolve neural (and sometimes body) properties in a model brain-body-environment system so as to exhibit some desired behavioral performance. The evolved agents can then be subjected to a detailed analysis to uncover their principles of operation. Evolutionary approaches are particularly useful for exploring spaces of possible solutions to a given behavioral task because these approaches minimize a priori assumptions about how a given behavior ought to be instantiated. They can also be useful for exploring different ways to complete a computational neuroethology model when only partial neural circuitry is available for a biological system of interest [3].

MATLAB

MATLAB is a programming environment that is used globally in virtually all neuroscience and cognitive psychology laboratories [4]. MATLAB integrates the modelling and experimental processes by bringing together, under the aegis of an intuitive scripting language, powerful data analysis and mathematical modelling tools.

NEURON

NEURON, developed at Duke University, is a simulation environment for modeling individual neurons and networks of neurons [5]. With the NEURON interface, it is possible to generate publication-quality results without having to write any program code at all. The NEURON simulation engine is based on a Hodgkin–Huxley type model with a Borg–Graham formulation.

Embodiment in electronic hardware

Conductance-based silicon neurons

Nervous systems differ from the majority of silicon-based computing devices in that they resemble analog computers (not digital data processors) and massively parallel processors, not sequential processors. To model nervous systems accurately, in real-time, alternative hardware is required.

The most realistic circuits to date make use of analog properties of existing digital electronics (operated under non-standard conditions) to realize Hodgkin–Huxley-type models in silico[18][6].

Retinomorphic chips

[19]

See also

References

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External links

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de:Neuroethologie es:Neuroetología it:Neuroetologia pl:Neuroetologia sv:Neuroetologi tr:Nöroetoloji ur:عصبی سلوکیات

  1. Template:Cite news
  2. Klein, S. A., Carney, T., Barghout-Stein, L., & Tyler, C. W. (1997, June). Seven models of masking. In Electronic Imaging'97 (pp. 13-24). International Society for Optics and Photonics.
  3. Barghout-Stein, Lauren. On differences between peripheral and foveal pattern masking. Diss. University of California, Berkeley, 1999.
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    Extending the tax exemption would help. The exemption, which may be as a lot as $2 million per family, covers individuals who negotiate a principal reduction on their existing mortgage, sell their house short (i.e., for lower than the excellent loans), or take part in a foreclosure course of. An extension of theexemption would seem like a common-sense means to assist stabilize the housing market, but the political turmoil around the fiscal-cliff negotiations means widespread sense could not win out. Home Minority Chief Nancy Pelosi (D-Calif.) believes that the mortgage relief provision will be on the table during the grand-cut price talks, in response to communications director Nadeam Elshami. Buying or promoting of blue mild bulbs is unlawful.

    A vendor's stamp duty has been launched on industrial property for the primary time, at rates ranging from 5 per cent to 15 per cent. The Authorities might be trying to reassure the market that they aren't in opposition to foreigners and PRs investing in Singapore's property market. They imposed these measures because of extenuating components available in the market." The sale of new dual-key EC models will even be restricted to multi-generational households only. The models have two separate entrances, permitting grandparents, for example, to dwell separately. The vendor's stamp obligation takes effect right this moment and applies to industrial property and plots which might be offered inside three years of the date of buy. JLL named Best Performing Property Brand for second year running

    The data offered is for normal info purposes only and isn't supposed to be personalised investment or monetary advice. Motley Fool Singapore contributor Stanley Lim would not personal shares in any corporations talked about. Singapore private home costs increased by 1.eight% within the fourth quarter of 2012, up from 0.6% within the earlier quarter. Resale prices of government-built HDB residences which are usually bought by Singaporeans, elevated by 2.5%, quarter on quarter, the quickest acquire in five quarters. And industrial property, prices are actually double the levels of three years ago. No withholding tax in the event you sell your property. All your local information regarding vital HDB policies, condominium launches, land growth, commercial property and more

    There are various methods to go about discovering the precise property. Some local newspapers (together with the Straits Instances ) have categorised property sections and many local property brokers have websites. Now there are some specifics to consider when buying a 'new launch' rental. Intended use of the unit Every sale begins with 10 p.c low cost for finish of season sale; changes to 20 % discount storewide; follows by additional reduction of fiftyand ends with last discount of 70 % or extra. Typically there is even a warehouse sale or transferring out sale with huge mark-down of costs for stock clearance. Deborah Regulation from Expat Realtor shares her property market update, plus prime rental residences and houses at the moment available to lease Esparina EC @ Sengkang
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    In its statement, the singapore property listing - website link, government claimed that the majority citizens buying their first residence won't be hurt by the new measures. Some concessions can even be prolonged to chose teams of consumers, similar to married couples with a minimum of one Singaporean partner who are purchasing their second property so long as they intend to promote their first residential property. Lower the LTV limit on housing loans granted by monetary establishments regulated by MAS from 70% to 60% for property purchasers who are individuals with a number of outstanding housing loans on the time of the brand new housing purchase. Singapore Property Measures - 30 August 2010 The most popular seek for the number of bedrooms in Singapore is 4, followed by 2 and three. Lush Acres EC @ Sengkang

    Discover out more about real estate funding in the area, together with info on international funding incentives and property possession. Many Singaporeans have been investing in property across the causeway in recent years, attracted by comparatively low prices. However, those who need to exit their investments quickly are likely to face significant challenges when trying to sell their property – and could finally be stuck with a property they can't sell. Career improvement programmes, in-house valuation, auctions and administrative help, venture advertising and marketing, skilled talks and traisning are continuously planned for the sales associates to help them obtain better outcomes for his or her shoppers while at Knight Frank Singapore. No change Present Rules

    Extending the tax exemption would help. The exemption, which may be as a lot as $2 million per family, covers individuals who negotiate a principal reduction on their existing mortgage, sell their house short (i.e., for lower than the excellent loans), or take part in a foreclosure course of. An extension of theexemption would seem like a common-sense means to assist stabilize the housing market, but the political turmoil around the fiscal-cliff negotiations means widespread sense could not win out. Home Minority Chief Nancy Pelosi (D-Calif.) believes that the mortgage relief provision will be on the table during the grand-cut price talks, in response to communications director Nadeam Elshami. Buying or promoting of blue mild bulbs is unlawful.

    A vendor's stamp duty has been launched on industrial property for the primary time, at rates ranging from 5 per cent to 15 per cent. The Authorities might be trying to reassure the market that they aren't in opposition to foreigners and PRs investing in Singapore's property market. They imposed these measures because of extenuating components available in the market." The sale of new dual-key EC models will even be restricted to multi-generational households only. The models have two separate entrances, permitting grandparents, for example, to dwell separately. The vendor's stamp obligation takes effect right this moment and applies to industrial property and plots which might be offered inside three years of the date of buy. JLL named Best Performing Property Brand for second year running

    The data offered is for normal info purposes only and isn't supposed to be personalised investment or monetary advice. Motley Fool Singapore contributor Stanley Lim would not personal shares in any corporations talked about. Singapore private home costs increased by 1.eight% within the fourth quarter of 2012, up from 0.6% within the earlier quarter. Resale prices of government-built HDB residences which are usually bought by Singaporeans, elevated by 2.5%, quarter on quarter, the quickest acquire in five quarters. And industrial property, prices are actually double the levels of three years ago. No withholding tax in the event you sell your property. All your local information regarding vital HDB policies, condominium launches, land growth, commercial property and more

    There are various methods to go about discovering the precise property. Some local newspapers (together with the Straits Instances ) have categorised property sections and many local property brokers have websites. Now there are some specifics to consider when buying a 'new launch' rental. Intended use of the unit Every sale begins with 10 p.c low cost for finish of season sale; changes to 20 % discount storewide; follows by additional reduction of fiftyand ends with last discount of 70 % or extra. Typically there is even a warehouse sale or transferring out sale with huge mark-down of costs for stock clearance. Deborah Regulation from Expat Realtor shares her property market update, plus prime rental residences and houses at the moment available to lease Esparina EC @ Sengkang
  7. Jeffress, L.A., 1948. A place theory of sound localization. Journal of Comparative and Physiological Psychology 41, 35–39.
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  16. Tzounopoulos T, Kim Y, Oertel D, Trussell LO (2004) Cell-specific, spike timing-dependent plasticities in the dorsal cochlear nucleus. Nat Neurosci 7:719–725
  17. Patrick D. Roberts, Christine V. Portfors (2008) Cell-specific, spike timing-dependent plasticities in the dorsal cochlear nucleus. Biological Cybernetics 98:491–507
  18. L. Alvadoa, J. Tomasa, S. Saghia, S. Renauda, T. Balb, A. Destexheb, G. Le Masson, 2004. Hardware computation of conductance-based neuron models. Neurocomputing 58–60 (2004) 109 – 115
  19. Kwabena Boahen, "A Retinomorphic Chip with Parallel Pathways: Encoding INCREASING, ON, DECREASING, and OFF Visual Signals", Analog Integrated Circuits and Signal Processing, 30, 121–135, 2002