JBiSE20080100007 78803519
JBiSE20080100007 78803519
JBiSE20080100007 78803519
indicate that sensory neurons of the nervous Behavioral tasks with temporal information process-
system do not show selective response to ing falling within this scale include speech discrimi-
temporal properties of external stimuli. On nation in the auditory system, motion information
the other hand, temporal selective neurons in processing in the visual systems, and movement
the cortex have been reported in many coordination in the motor system [1-3].
species. Thus, processes which realize the Information processing in neural systems normally
temporal-to-spatial transformation of consists of a number of successive stages. Neural
neuronal activities might be required for activities in a certain stage are mostly determined by
temporal information processing. In the neural activities of the preceding stages and our
present study, we propose a computational perception of the world in the brain is based on the
model to explore possible roles of electrical spatio-temporal patterns of neuronal activities
synapses in processing the duration of produced at sensory stages [4-5]. Physiological
external stimuli. Firstly, we construct a observations indicate that neurons in the sensory
small-scale network with neurons intercon- levels do not respond selectively to the temporal
nected by electrical synapses in addition to properties of external stimuli. Temporal information
chemical synapses. Basic properties of this is thus suggested to be contained in the temporal
small-scale neural network in processing patterns of neuronal activities in the sensory layer.
duration information are analyzed. Secondly, On the other hand, neurons which show selective
a large-scale neural network which is more response to specific temporal properties, especially
biologically realistic is further explored. Our the duration content, have been reported in the cortex
results suggest that neural networks with of many species [6-10]. Temporal information is
electrical synapses functioning together therefore suggested to be transformed into the
spatially distributed neuronal activities in the cortex
with chemical synapses can effectively work
and neural mechanisms which contribute to the
for the temporal-to-spatial transformation of
temporalto-spatial transformation of neuronal
neuronal activities, and the spatially distrib-
activities are required.
uted sequential neural activities can poten-
Electrical synapse is another type of widely
tially represent temporal information.
distributed neuronal connection in the neural systems
in addition to chemical synapse [11-12]. Functional
Keywords: Model; Neural network; Electrical role of electrical synapse has been identified in fine
synapse; Temporal information processing motor coordination which requires temporal infor-
mation processing in milliseconds scale [13]. In the
1. INTRODUCTION present work, we try to explore possible neural
Biological neural systems are endowed with the mechanisms of electrical synapse in processing the
ability to process temporal information given the duration content of external stimuli via computa-
inherent temporal nature of sensory environments tional approach. Briefly, we construct neural net-
Figure 1. A. Schematic structure of the small-scale neural network model. The input neuron (S) is connected to 4 of the 10
the excitatory neurons (E). All excitatory neurons are connected to each other in a recurrent way and each excitatory neuron
is coupled with an inhibitory neuron (I). Excitatory and inhibitory synapses are represented by open and solid circles,
respectively. Neurons in grey shadow are electrically coupled together recurrently.B. Schematic structure of the large-scale
neural network model. Input neuron is connected to excitatory (E) and inhibitory (I) neurons in the network on a random basis.
All excitatory neurons are further connected with each other probabilistically in a recurrent way. Electrical synapses are
formed between some of the excitatory neurons randomly.
(1)
in which syn =15 ms , E thr = - 40 mV, and (u)
follows a step function:
(2)
Figure 2. Spike activities of the input neuron (S) in response to constant injected currents with various intensities and
magnitudes.
3.1 Stimulus duration is represented by spike neuronal groups are electrically coupled together
trains of input neuron which contain 2, 3 and 4 neurons, respectively.
The injected current is first transformed into a spike Raster plots of the firing performances of the model
train of the input neuron. Spiking properties of the neurons in absence and presence of electrical
input neuron (S) are shown in Figure 2. Injected synapses are compared with stimulus duration being
currents with different magnitudes and durations are 50 ms ( Figure 3A&B) and 100 ms ( Figure 3C&D),
applied to the input neuron to test its performance. A respectively.
sustained current elicits periodic spikes from the Results given in Figure 3B&D suggest that
input neuron and the duration of the spike train is electrical synapses in a neural network can effec-
determined by the stimulus duration. Input neuron tively transform the temporal domain spike train of
can therefore mimic the function of sensory neuron in the input neuron into the spatial-temporal firing
neural system. pattern of a group of neurons. Each activated neuron
in the group fires within a specific time window,
3.2 Performance of the small-scale neural which is determined by the configuration of the
network model synaptic connection of the neural network. Fur therm ore,
3.2.1 Temporal information can be represented by stimulus with longer duration can evoke spikes from
the spatially distributed activities of a group of more neurons and therefore the stimulus durations
neurons can be represented by the spatial and temporal
Representative firing patterns of the simple model structure of the sequential neuronal activities.
are given in Figure 3. Parameters used for Figure 3
are listed in Table 2 and the synaptic connection 3.2.2 The output pattern is closely related to the
follows that illustrated in Figure 1A. Input neuron is electrical coupling configuration
connected to four of the ten excitatory neurons. Three Electrical synapses between excitatory neurons and
Figure 3. Raster plots for neuronal activities of the small-scale model elicited by 50 and 100 ms stimulus durations. Stimuli
are indicated by grey shadows. A, 50 ms duration, without electrical synapses; B, 50 ms duration, with electrical synapses; C,
100 ms duration, without electrical synapses; D, 100 ms, with electrical synapses.
Figure 4. Raster plots for spike activities of threeneuron group with different synaptic configurations. Neurons receive
synaptic input from input neuron are represented by solid circle. Electrical synapses are represented by solid lines. The
stimulus duration is 100 ms with the current intensity to input neuron being 2.0 pA.
Figure 5. A and B are representative raster plots of the neuronal activities of the large-scale model in absence and presence
of electrical synapses, respectively. The stimulus duration is 100 ms. Inset graphs represent the processes of spike activity
recruitment. C and D show the recruitment processes in absence and presence of electrical synapses, respectively. Data are
averaged based on 10 independent trails (Mean S.D.).
Figure 6. Raster plots of the large-scale neural network in response to stimuli with different durations. The configuration of
the model is identical for Figure A to F.
Figure 7. Recruitment of neuronal activities (activated numbers) for the large-scale model in response to stimuli with
durations ranging from 50 to 100 ms (A, step 10 ms) and 100 to 200 ms (B, step 20 ms). The mean values of synaptic strength
from input to excitatory neurons are 0.055 and 0.038 for results in Figure A and B, respectively. Data are analyzed from 10
independent trials in the form of (Mean S.D.).
act ivi tie s are l imi ted w ith in a na rro w tem por al temporal firing pattern of neuronal ensembles while
window in absence of electrical coupling ( A & C ). ea ch ne ur on wi th in th e en se mb le fi re s wi th in
The firing patterns of the large-scale model in different time windows, and the spatio-temporal
response to stimuli with various durations are further pat ter n of the neu ron al act ivi tie s is cap abl e of
tested. Stimuli with durations varying from 50 ms t o representing stimulus duration in the form of
100 ms are applied to the network, with steps being 10 sequential firing activities of the spatially distributed
ms. Raster plots of typical spike activities of the neurons.
network are given in Figure 6, A to F . It is revealed The contribution of electrical synapses in the
that the model neurons fire in a sequential pattern, fo rm at io n o f s pa ti o- te mp or al fi ri ng pa tt er n i s
with more neurons being sequentially recruited in particularly examined in the present study. However,
response to longer duration. Such recruitment i t i s n ec es sa ry t o m en ti on t ha t o th er f ac to rs c an a ls o
process in response to durations ranging from 50 ms contribute to this process. For example, membrane
to 100 ms is averaged based on ten independent trials capacitance of specific neurons can be variable
and the result is shown in Figure 7A. because of variation in surface area as well as the
Stimuli with durations varying from 50 ms to 100 membrane capacitance value per unit area [25-28].
ms are app lied an d relev ant res ults ar e given in These changes can function in parallel to electrical
Figure 6 and Figure 7A. However, models with this synapses in influencing the sequential firing patterns
structure can effectively represent durations in other of neuronal ensembles.
ranges while relevant parameters are changed. These Special role of electrical synapse is proposed in
parameters include the capacitance value of the I-F our mod els and there a re also experimen tal clu es
neu ron al mod el, th e time c ons tan t for ch emi cal which indicated possible roles of electrical synapse
synaptic strength, the synaptic strengths from input in te mporal informa tion process ing. Data demon-
neuron to the network et al. Stimuli with durations strated that gap junction coupling within inferior
ranging from 100 ms to 200 ms are applied to the olive mediated by connexin 36 could add 10-20 of
ne tw or k, i n wh ic h th e me an v al ue o f sy na pt ic precision to the fine temporal coordination of muscle
strength from input neuron to the neural network firing during movement [13].
(Cm se ) are changed (from 0.055 S to 0.038 S ). Neurons in the present work are modeled follow-
The performance of the model (averaged across ten ing the classic I-F neuron fashion without any
independent trials) is plotted in Figure 7B. specific properties for temporal information process-
ing. These neurons can be tuned to response to any
4. DISCUSSION non-temporal properties of natural stimulus and
Temporal information processing in neural system is thereby function for the corresponding behavioral
critical for animal behavior. Neuroscientists have tasks. For example, these neurons could be tone
tried a lot in understanding the neural basis of selective neuron which function for auditory behav-
relevant processes via both experimental [6-10] and ior, or mechanosensory neurons which function for
computational approaches [19-24]. mechanosensation. While both electrical and
In the present study, the computational results chemical synapses are universal in the central
demonstrate that electrical synapses could effec- nervous system, the model results suggest that both
tively contribute to the formation of a spatio- the spatial and temporal neuronal activities produced
at the sensory layer of neural system could be dependent plasticity. Phys Rev E Stat Nonlin Soft Matter Phys
processed together by sharing the same neural circuit. 2003, 68:011908.
[22]Buonomano DV & Merzenich MM. Temporal information
Temporal content of external stimulus could be read transformed into a spatial code by a neural network with
out from spike patterns of neuronal ensembles in the realistic properties. Science 1995, 267:1028-1030.
brain. [23]Mauk MD & Donegan NH. A model of Pavlovian eyelid
conditioning based on the synaptic organization of the
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[24]Medina JF, Garcia KS, Nores WL, Taylor NM & Mauk MD.
Timing mechanisms in the cerebellum: testing predictions of a
ACKNOWLEDGEMENTS large-scale computer simulation. J Neurosci 2000, 20:5516-
The authors would like to thank Dr. Shi-Yong Huang for helpful 5525.
discussion. This work was supported by grants from the Hi-Tech [25]Chitwood RA, Hubbard A. & Jaffe DB. Passive electrotonic
Research and Development Program of China (No. 2006AA01Z125). properties of rat hippocampal CA3 interneurones. J Physiol
1999, 515 ( Pt 3):743-756.
[26]Gentet LJ, Stuart GJ & Clements JD. Direct measurement of
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