WCCI 2012 IEEE World Congress on Computational Intelligence June, 10-15, 2012 - Brisbane, Australia

IJCNN

Learning Transmission Delays in Spiking Neural Networks: A Novel Approach to Sequence Learning Based on Spike Delay Variance
Paul W Wright, Janet Wiles
School of Information Technology and Electrical Engineering University of Queensland Brisbane, Queensland 4072, Australia paul.wright1@uqconnect.edu.au, wiles@itee.uq.edu.au
Abstract Transmission delays are an inherent component of spiking neural networks (SNNs) but relatively little is known about how delays are adapted in biological systems and studies on computational learning mechanisms have focused on spiketiming-dependent plasticity (STDP) which adjusts synaptic weights rather than synaptic delays. We propose a novel algorithm for learning temporal delays in SNNs with Gaussian synapses, which we call spike-delay-variance learning (SDVL). A key feature of the algorithm is adaptation of the shape (mean and variance) of the postsynaptic release profiles only, rather than the conventional STDP approach of adapting the networks synaptic weights. The algorithms ability to learn temporal input sequences was tested in three studies using supervised and unsupervised learning within feed-forward networks. SDVL was able to successfully classify forty spatiotemporal patterns without supervision by providing robust, effective adaption of the postsynaptic release profiles. The results demonstrate how delay learning can contribute to the stability of spiking sequences, and that there is a potential role for adaption of variance as well as mean values in learning algorithms for spiking neural networks. spiking neural networks; transmission delays; delay learning; sequence learning; spike-delay-variance learning; STDP

types creates variable axonal delay. The synaptic process, where a presynaptic potential gets converted to a postsynaptic potential, takes time. Electrical synapses and chemical synapses have different transmission properties, however in general a postsynaptic potential will have a slower rise time than its presynaptic impulse, causing a synaptic delay in the order of milliseconds [6]. The synapses location on the dendritic tree will also create variable electronic transmission delay [7]. All of these delays combine to create an overall transmission delay, which is highly variable between transmission lines but extremely repeatable on the same transmission line. Some observed mammalian transmission delays are 1 to 17 ms in Sprague-Dawley rats [8], 1 to 32 ms in rabbits [9] and 1 to 30 ms in cats [10]. The ability of a network to produce repeatable temporal spike chains, involving multiple neurons, has been likened to the ability to store and reference memories [11]. Repeatable spike trains require neurons to be capable of consistently responding to a familiar temporal sequence of input spikes. Spike-timing-dependent plasticity (STDP) is a phenomenological learning rule [12] observed in many real neural networks. STDP represents a family of Hebbian learning kernels that adjusts the synaptic transmission strength based on the relative timing of the presynaptic and postsynaptic neuron [13]. [ID_1.2] Placed in a neural network containing randomised fixed delays, STDP aids the formation of small scale sequence recognition by fine-tuning weights to reward correlated activity. However, the restriction of fixed delays potentially limits the capabilities of STDP in performing this task. Although uncommon, there exists some evidence of dynamic delay learning within biological networks. Variable transmission delays, assumed to be facilitating learning, have been observed in vivo in Mongolian gerbils [14]. Senn, Schneider and Ruf suggest that fast delay adaption is unnecessary in the presence of temporal synaptic strength learning rules. They argue that beneficial delays are naturally selected through asymmetric weight adjustment, although at much slower rate [15]. The role of delays and the extent and mechanisms of delay learning in mammalian brains is remains unclear. Computational approaches used to model delay learning vary considerably in their level of biological fidelity. Top-

I.

INTRODUCTION

Delays are ubiquitous in the brain. Analog information can be coded within temporal spikes, allowing both static and learnable delays to benefit neural computation in spiking networks [1]. Real world stimulus analysis is fundamentally a temporal process. Delays play a key role in the dynamics of neuron communication and computation, regularly demonstrated in natural biological systems. For example, barn owls precisely localise the position of fast moving rodents in absolute darkness using only their auditory input streams [2]. Delays created by the magnocellular afferents make it possible for the laminaris neurons to relate phase differences to spatial position, acting as the required mechanisms for Jeffress style interaural coincidence detection [3] [4]. The mechanisms that allow an action potential to affect the membrane potential of another neuron are far from instant. The structural properties of an axon greatly influence the transmission speed such as axon diameter and myelin thickness [5]. The inconsistency of such properties across regions and neuron

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down approaches abstract from the biology details and directly adjust the timing: Supervised learning has been used to directly optimise the delays between internal neurons and output neurons in order to achieve the best possible classification performance [16]. Another method increases the delay if a postsynaptic pulse arrives before a defined temporal reference point, and decreases the delay if it arrives after, synchronising multiple postsynaptic pulses [17]. Bottom up approaches use more biologically-motivated mechanisms, such as adjusting the concentration of glutamate receptors in order to vary the perceived transmission delay [18]. An alternative to learning mechanisms which directly adapt delays is to use a network with intrinsic unstable dynamics that are deterministic but vary over a wide range. In such a network, a specific time delayed output can be selected rather than learned. This principle underpins a new approach to temporal networks called liquid state machines [19]. Many computational models have been developed to simulate spiking neurons such as the Hodgkin-Huxley model [20], the leaky integrate and fire model [21] and the Izhikevich model [22]. They range from high biological fidelity to mathematically abstract, generally with an associated cost in terms of computational speed [23]. Simulation of membrane potentials alone is insufficient to accurately replicate brain activity due to the huge diversity and complexity of synaptic plasticity [24]. Synapses and transmission delays have been modelled in a multitude of ways [25]. We focus in this paper on two important features of the synaptic input to a spiking neuron, the temporal variance and the average value. Both interact with the baseline parameters of a neuron to produce a wide range of non-linear behaviour [26]. Analysis of these two features, when a single neuron receives input from a population, reveals intriguing results. Information can be encoded using both features, by either changing the average synaptic input to neuron (corresponding to changing average firing rates of the presynaptic neurons) or by changing the temporal variance of synaptic input (corresponding to the temporal irregularity of the summed postsynaptic potentials). The former cannot produce noticeable network responses at realistic speeds, whereas the latter can [27]. This result suggests that temporal synaptic variance is a better medium for quick data transfer and processing than the actual magnitude of input current. Synaptic variance has thus been utilised as a mechanism for effective spatiotemporal sequence classification, a common task in many biological systems. Barak and Tsodyks adjusted synaptic weights in order to maximise the postsynaptic current variance observed by the integrating neuron, yielding successful classification [28]. Two problems with Barak and Tsodyks approach to sequence recognition are the insensitivity to temporal reversal of the input pattern and the inability for a neural system to respond more quickly with increasing familiarities. Brains are well known for their ability to not only improve accuracy with learning, but also improve the processing speed of known patterns. Such optimisation occurs without reduced robustness. In the research described in this paper, we aimed to design and test a delay-learning algorithm that could take advantage of synaptic variance as an integral part of adjusting delays. We

designed the algorithm to achieve effective spatiotemporal recognition and classification in small noiseless networks. The following sections describe our novel transmission delay learning algorithm and an associated synaptic transmission model, and demonstrate its performance on sequence tasks in which the order of elements in a sequence changes over time. The learning algorithm is tested in two forms, initially for supervised learning and then applied in unsupervised learning tasks. The results demonstrate robust learning of temporal order which can adapt over time. II. METHODS

The model is comprised of key components, including a model neuron (see section II-A), synapse model and learning algorithm (II-B), network and sequence structures (II-C), difference measure and classification success rate (II-D), and learning tasks (II-E). A. Neuron Model The model of neuron used in this study was proposed by Izhikevich as a computationally tractable model of spiking neurons that balances both simplicity in parameters and complexity in the range of possible neuron behaviours [22]. It is a part of a set of hybrid dynamical systems that makes use of dual ordinary differential equations to replicate the complex neuronal firing patterns such as bursting, threshold variability and bistabilty [29]. In this paper, regular spiking Izhikevich neurons were used to balance complexity and efficiency, although other neuron models such as the integrate-and-fire model are capable of performing similar computation. The first order ordinary differential equations (see [22] for details) are solved with an improved Euler method, using a time step of 1ms. Subsequently, all delays and synaptic profiles are quantised to a resolution of 1ms. Smaller time steps were also tested and had insignificant impact on the membrane and synaptic dynamics. B. Synapse Model In real neural networks, delays are a combination of the temporal nature of dendritic, axonal and synaptic conduction. For this study, for the sake of model simplicity and efficiency, transmission delays were simulated as a single variable acting in the synapse. 1) Governing Gaussian Function Each synapse releases postsynaptic current into its target neuron over time which creates conduction delays. Spike delay-variance learning requires a postsynaptic release modelling equation that has variable delay and temporal delivery width, for example, the Gaussian function or the sum of rising and decaying exponentials. A sampled Gaussian function was used in this paper for computational efficiency. The release of post synaptic current, I, from a synapse is modelled with a Gaussian function. (1)

where p is the peak postsynaptic current, t0 is the time since a presynaptic spike in ms (t0 0), is the mean and v is the variance. The parameters p, v and are variable for each synapse, and their selection determines the release profile. The synaptic mean represents the peak-timing of postsynaptic current, such that the transmission delay of the synapse is equal to . Some current will be delivered before and after the peak timing, depending on the variance. A large variance indicates a wide release profile, whereas a small variance results in a narrow one. The mean must be non-negative as postsynaptic current cannot arrive before the presynaptic spike. 2) Fixed Integral The integral of the profile determines the amount of postsynaptic current delivered. As a result, the strength (or weight) of a Gaussian synapse is not dependent on the peak, but by the integral of the curve. The peak is proportional to the integral, hence for a given integral value there will only be one corresponding peak value. Globally fixing the integral of all synapses in a network removes the peak as a synaptic variable. The required peak value for a given variance, mean and global integral can be computationally determined using an optimisation algorithm such as gradient descent. 3) The Algorithm The mean and variance of a synapse are only adjusted when a postsynaptic spike occurs. The temporal error of the synapse is defined as the absolute value of the temporal difference between the synapses mean and t0, where t0 is the time difference between the presynaptic and postsynaptic spike. The mean is adjusted to reduce any significant error, helping to replicate the

relative spike timing of the presynaptic and postsynaptic neurons. If the error is insignificant but the magnitude of t0 is significant, the mean is reduced to improve the response time of the postsynaptic neuron. The change of mean, , is determined by (2) where: sgn(.) is the signum function is the mean of the synapse (ms) k(v) is the learning accelerator v is the variance of the synapse is the mean learning rate 1, 2 are constants

The variance is increased if the synapses temporal error is significant, searching by broadening the postsynaptic release profile. The variance is decreased if the error is sufficiently small, locking on to a familiar temporal spike pattern by narrowing the release profile. The change of variance, v, is determined by (3) where: v is the variance learning rate 1, 2 are constants

In this paper, is restricted to the range [0, 15], v is restricted to the range [0.1, 10] and, for simplicity, k(v) = (v + 0.9)2 so that k = 1 at minimum v as the choice of the magnitude of k is arbitrary. C. Network Structure and Sequence Definition The networks used for algorithm evaluation comprise of two layers of spiking neurons, an input layer connected to an output layer with a feed-forward structure. Each input neuron i {1...N} is connected to each output neuron j {1...M} with a single dynamic Gaussian synapse. The effects of non-uniform network weights and weight adaption are eliminated by enforcing that all Gaussian synapses have the same global fixed integral. Each output neuron is connected to all the other output neurons with strong static instant inhibitory synapses, acting as a winner takes all mechanism. During a presentation of a spatiotemporal sequence/pattern, Q input neurons fire exactly once within a given time window, Tm, from a global temporal reference point (the beginning of the presentation). Hence an input sequence is a vector of input spike times seq = {t1, t2, ... ti} where ti {0,1...Tm, }, where an infinite spike time denotes silence. If Q is less than the number of input neurons then there is a multivariable component, which in this paper is referred to as spatial. Supervised learning enforces that the correct output neuron fires at Ts after the presentation of the sequence, unless it has already fired naturally. Thus supervised learning only affects the network if the correct output neuron fails to fire within Ts from the start of a presentation. The transmission delay learning algorithm is

Figure 1. Post synaptic release profiles for variable variance and fixed mean (top), variable mean and fixed variance (bottom), all restricted by a fixed integral. When the variance is low (0.1), all the current is delivered as a single burst at a delay determined by the mean. At high variances (>5), current is slowly released from the synapse over a large period of time, peaking at the delay determined by the mean.

always active whereas supervised learning can be switched on or off during a trial. D. Sequence Difference Measure and Classification Success Rate The task of classifying temporal sequences has variable difficultly, inversely proportional to the input sequences absolute temporal difference. So, in order to quantify algorithm performance in context of task difficulty, a method of quantifying the temporal difference between two input sequences needed to be defined. The difference measure, D, between two normalised sequences is determined by (4) where a = (a1, a2, ... aN) and b = (b1, b2, ... bN) are the two normalised input sequences, Tm is the maximum spike phase offset. Pattern normalisation is achieved by subtracting the minimum spike phase offset from all vector elements of a sequence, so that at least one term is zero, forcing at least one input neuron to fire at the beginning of the presentation. Normalisation enforces that two patterns cannot have a large D if one is a time-shifted version of the other (i.e. temporally identical). During a classification task, the classification success rate (CSR) is the percentage of patterns that are represented by a single unique output neuron, i.e. one particular output neuron always solely spikes when its corresponding pattern is presented to the network, but remains silent when all other patterns are presented. E. Network Tasks The algorithm was evaluated using three learning tasks, labelled as Condition 1, 2 and 3. 1) Condition 1. Single Output Neuron Under Condition 1, the circuit had 3 inputs (N = 3), 1 output (M = 1) and 2 patterns, seq a = {0, 3, 7} and seq b = {7, 3, 0} (P = 2, Q = 3). Condition 1 comprised of a single 60 second trial. Seq a was solely presented during 0-25 and 55-60 seconds, seq b was solely presented during 25-55 seconds. Note that seq b is a time-reversed version of seq a. Sequences were presented to the network at a frequency of 2 Hz to allow membrane potentials to return to baseline before each presentation. Supervised learning was only switched on during 0-15 and 30-45 seconds with Ts = 12ms. The Gaussian synapses were initialised with v = 2 and = 5. The global integral was fixed at 13 so that at all three postsynaptic pulses are required to arrive at similar times to induce a spike. 2) Condition 2. Classification of Two Patterns Under Condition 2, the circuit had 10 inputs (N = 10), 2 outputs (M = 2) and 2 patterns (P = 2, Q = 10). Two random input sequences, s1 and s2, are generated at the beginning of each trial with a particular D. During each trial, the network was trained over 100 epochs (an epoch comprises of all patterns being presented once to the network at a frequency of 5 Hz) and then the classification success rate (CSR) was calcu-

lated. A trial was only considered successful if the CSR was 100%, otherwise it was considered unsuccessful. The Gaussian synapses were initialised with v ~ U (1, 3) and ~ U (2, 10) at the beginning of each trial, where U is the uniform distribution. Each integer sequence element is an independent random variable uniformly distributed on [0, Tm], where Tm = 12ms. The average D between two randomly generated patterns is 0.3563, with a standard deviation of 0.0823. This intuitively low value is a consequence of pattern normalisation. Thus the algorithms classification performance is evaluated uniformly on the range D = [0, 0.5], representing the range of sequence pairs from identical to significantly different. There are 61 possible values of D is this range, as D is a quantised value. Every value of D is trialled 30 times and the percentage of successful trials was calculated. Hence there were a total of 1830 uniquely generated experiments, each trial having different initial Gaussian parameters and input patterns. The high number of independent experiments is required to offset any dependence on initial conditions. a) Condition 2a. Supervised Learning Supervised classification activates supervised learning (Ts = 14ms) during the entire 100 training epochs. b) Condition 2b. Unsupervised Learning Unsupervised classification represents a different learning paradigm. There is no teacher to force the output neurons to fire, which is required by SDVL to change synaptic parameters. Initially, a large integral is required, to allow the output neurons to spike before the Gaussian synapses are correctly tuned. The integral needs to be reduced over time, so that the synapses can converge to a solution. Integral reduction is a form of simulated annealing, and many annealing functions could have been used. Linear annealing was chosen for parameter simplicity. The three linear annealing parameters are: I0 (initial integral), If (final integral) and Tf (time to reach the final integral). The integral starts at I0, linearly descends and reaches If after Tf seconds. The integral is then held constant at If for the remaining time. There are six algorithm parameters to be tuned, , v, 1, 2, 1, 2, in addition to the three simulated annealing parameters. This large parameter space is difficult to manually explore, thus a genetic algorithm (GA) was used to find solutions. The GA uses a hybrid of mutation and crossover, with a cost of population diversity. Details of the GAs implementation and performance are available in [30]. c) Condition 3. Classification of Forty Patterns Under Condition 3, the circuit had 40 inputs (N = 40), 40 outputs (M = 40) and 40 patterns (P = 40, Q = 10). Patterns and Gaussian parameters were initialised as per Condition 2. SDVL and simulated annealing parameters were evolved as per Condition 2b. The unsupervised network was trained for 80 epochs. The CSR was calculated after every epoch and averaged over 20 independent randomly generated trials.

III.

RESULTS

A. Condition 1 Results Under Condition 1, SDVL adapts the synaptic parameters over time to produce effective delay learning (see Fig. 2). Supervised learning forced the output neuron to fire with a 12ms phase offset at the beginning of the trial, allowing SDVL to change the synaptic parameters from their initial values. During the first 3 seconds, the synapse connected to input 1 searched for the correct delay by spreading out its postsynaptic release profile via an increasing variance. The three means deviate to

find the optimal relative delay configuration such that all postsynaptic peaks arrive at the output neuron simultaneously. The high mean learning rate allowed the means to converge after only 6 seconds. At this point, the delays perfectly matched the input pattern, enabling the variances to reduce to the smallest allowed value (observed as the narrowing of the profiles in Fig. 2 from 10 to 30 seconds). The delays are relatively correct, however are not optimal in terms of response time. SDVL improved the response time by reducing the means in a parallel manner, so that their relative timing is conserved. The delays reached the optimal configuration by 80 seconds, indicated by the accelerated output neuron spike phase offset (decreasing

Figure 2. Synaptic parameter traces and neuron spike times during a sequence recognition task. From top to bottom: A) Task design: periods where supervised learning is active. B) The input and output spike times, shown as a phase offset from the beginning of each sequence presentation. C) Mean traces for all synapses in the network. D) Variance traces for all synapses in the network. E) Gaussian postsynaptic release profiles at various stages throughout the trial.

from 12ms to 9ms). The input sequence was reversed after 120 seconds, desynchronising the arrival times of the postsynaptic pulses. The output membrane potential was unable to reach threshold due to leak currents in the soma. The unresponsiveness of the output neuron indicates that the original input pattern had been learnt: the output neuron spikes when presented with a known sequence, but remains silent when presented with a different unknown sequence. Supervised learning was turned back on after 150 seconds (see Fig. 2A), forcing the output neuron to fire, thus changing synaptic parameters. The synaptic means adjusted to match the new input pattern. The variance of the synapse connected to input 3 experienced a huge increase, broadening its profile as the temporal error between its mean and t0 was now large due to the reversed input pattern. Similar to the beginning of the trial, the means converged, and then descended in parallel, which reduces the response time to the optimal value. After 270 seconds, the input sequence was reversed back to the original pattern, rendering the output neuron unable to fire. The unresponsiveness of the output neuron, once again, confirms that the new pattern had been learnt, and the old pattern had been forgotten. Similar results were observed when replicating the experiment with a reduced global integral of 11. However, the key difference was the inability of the output neuron to speed up its response time. Output spikes always occurred with a 12ms phase offset, unable to decrease to 9ms as observed under Condition 1. Increasing the integral to 19 allowed only two input spikes to induce an output spike. The output spike reduced its response time to 5ms, only requiring the first two spikes in the sequence to fire. The unused synapse diverged to maximum values. SDVL performs effectively as well when in the input sequences are distorted with 2ms of uniform temporal noise (see [30]).

B. Classification Performance 1) Supervised Classification of Two Patterns Under Condition 2a, the average trial success rate was tested the discrete range of D = [0, 0.5] (see sup in Fig. 3). For identical sequences (D = 0) classification was always unsuccessful, as it is impossible to uniquely identify two identical input patterns. As the patterns became more diverse, the success rate rapidly increased to over 90% at D = 0.075. The success rate reached 100% soon after, when D = 0.16. The average distance between two random sequences was 0.35630.0823 (see Fig. 3). The classification success rate was perfect within this range, demonstrating that SDVL can accurately classify two arbitrary input sequences when aided by supervised learning. 2) Unsupervised Classification of Two Patterns The GA consistently found well-performing solutions in the parameter space. The average trial success rate of a typical solution (see Table I for parameters) across the entire range of D under Condition 2b was compared to the supervised case (see Fig. 3). The success rate was slightly worse than the supervised case, however still could successful classify two arbitrary patterns at least 90% of the time, as long as the patterns had a minimum D of 0.1. Unsupervised classification has a large dependence on the initial random configurations of the Gaussian synapses making a 100% trial success rate difficult to reach for the average range of D. The lack of supervision caused a slight decrease in performance as suspected, however effective unsupervised classification is clearly obtainable.

Figure 3. Averaged two pattern classification trial success rate over a wide range of sequence differences for both supervised and unsupervised learning paradigms.

TABLE I.

SIMULATED ANNEALING (IF APPLICABLE) AND SDVL PARAMETERS FOR EACH EXPERIMENTAL CONDITION.

Simulated Annealing Parameters Condition 1 2a 2b 3 I0 13 3 6.752 6.754 If 4.026 5.969 Tf 31.4 514.6 0.03 0.025 0.0338 0.0315 v 0.01 0.025 0.0357 0.0218

SDVL Parameters 1 3 3 1 4 2 2 2 3 2 1 5 5 8 1 2 5 3 5 1

is sensitive to the temporal direction of the patterns and it can speed up the response speed of a known sequence. A key finding of the results is the ability to achieve robust spatiotemporal processing in a network where the weights remain globally uniform. Individual synapse variance and delays are rapidly adjusted to explore the temporal landscape, attempting to achieve synchronised and optimised postsynaptic pulse timing. The results are evidence that varying the timing and shape of a synaptic release profile is computationally advantageous, independent of strength adaption. SDVL proved to be a robust classifier of two arbitrary input sequences. The algorithm performs extremely well as a spatiotemporal pattern classifier using both supervised and, the more realistic, unsupervised learning. Without supervision, patterns are required to be slightly more diverse in time, but can still achieve high accuracy and reliability when their temporal dissimilarity remains within a realistic range. The number of patterns was increased to 40 without disturbing the effectiveness of SDVL as a classifier. Exploring the algorithms parameter space can be a lengthy and tedious process. The usefulness of GAs to tune the algorithm parameters automatically is an additional result. Sequences were only presented in feed-forward networks once membrane potentials had stabilised and returned to baseline. Disturbing influences present in interconnected networks, such as high network activity and background noise, will force neurons to process postsynaptic potentials when they are initially not at baseline. The consistency of algorithm performance potentially could suffer from such influences. Synapses based on a Gaussian postsynaptic release profile proved to have interesting features. They can widen their response in order to search for the correct temporal relationship with a presynaptic neuron. Once a beneficial delay is found, they can narrow their response to increase temporal accuracy. If the presynaptic neuron repeatedly fires unpredictably, the release profile gets wider and wider, to a point where the synapse has little computation use. Destroying a synapse when it reaches maximum variance and mean would allow integrating neurons to filter out useless spikes, thus only allowing meaningful data to reach its soma. The biological significance of SDVL is yet to be established. However, there are many examples of biological systems optimising their processing time for frequently encountered input patterns. The algorithm provides a potential method to achieve temporal optimisation of recurrent computation routines via decreased delays. The algorithm could also be applied to nested sequences, providing robust classification over a much longer time scale. To establish a biological significance, empirical studies of dynamic transmission delays are needed.

Figure 4. Unsupervised CSR learning curves for forty noiseless and noisy spike patterns and forty output neurons.

3) Forty Pattern Unsupervised Performance A well-performing solution was found by the GA (see Table I for parameters). The solutions unsupervised classification performance was averaged over 20 trials (see Fig. 4). SDVL consistently associated each of the 40 output neurons with a unique pattern, completely characterising the input space after approximately 20-60 training epochs. SDVL still performed well even when all input spikes were randomly time shifted U[0,4] ms at each presentation. IV. DISCUSSION A novel transmission delay learning rule, spike-delayvariance learning (SDVL) was proposed and evaluated, contributing to our understanding of the range of computations possible with spiking neurons. The algorithm is different from other delay learning rules as it achieves postsynaptic pulse synchronisation and response time optimisation in unison. Its stability, reliability and effectiveness for spike sequence recognition and classification were demonstrated in a variety of contexts. The experiments clearly show that dynamic learnable delays have computational advantages in terms of spatiotemporal pattern recognition and classification. Deterministic spatiotemporal activity patterns appear in real biological systems [31]. Many previous approaches to spatiotemporal spike processing rely on the adaption of weights, either through combining spike-timing based learning kernels like STDP with variable readout delays [16], or to maximise the combined postsynaptic current variance [28]. SDVL resolves two issues experienced by Barak and Tsodyks [28] approach to variance-based spatiotemporal pattern learning, as it

Modern in vivo imaging techniques have increased in accuracy, resolution and duration [32], making such studies achievable. In a recent observation of dynamic transmission delays in vivo in Mongolian gerbils, a phenomenological model was used to estimated that the change of delay per spike was between 5-30 s [14]. Under Condition 2b and Condition 3, the genetic algorithm produced delay learning rates of 31.5 and 33.8 s per spike respectively (see Table I), similar to the biological estimate. However, it must be noted in SDVL the actual change of delay is amplified whenever the synapses variance is higher than minimum, according to the learning accelerator in (5). Static delays are clearly important in temporal processing [1], for example, Reichardt detectors [33], hypothesised as a neural mechanism for motion detection in many biological systems, especially insects [34]. It is possible that delays are initially learnt, converging once an optimal configuration is reached. Other learning rules could then perform learning tasks on a shorter time scale, complimented by the delay arrangement. In conclusion, SDVLs successful performance suggests that combining dynamic delays with synaptic variance adaption enhances the spatiotemporal processing capabilities of spiking neural networks. V. FUTURE WORK

Stable transmission delay learning promotes the sustainable repeatability of network activity. Izhikevich has proposed that polychronous neuron groups play a fundamental role in memory and consciousness [11]. Future work could include investigating the possible non-random formation of polychronous neuron groups, through robust delay learning. Transmission delay learning could also be linked with a reward signal. Neuromodulated delay learning rates would achieve effective reinforcement learning. Changing the circuit structure to include memory capacity would allow for serial input of patterns, eliminating the current limitation of one input neuron per sequence member and enabling more interesting datasets to be evaluated. The Gaussian integral is considered to be the weight of a Gaussian synapse. Applying spike-time dependent plasticity and weight normalisation to the value of the Gaussian integral would dramatically increase the processing potential of the algorithm. REFERENCES
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