† Corresponding author. E-mail:
Project supported by the National Natural Science Foundation of China (Grant No. 11675096), the Fundamental Research Funds for the Central Universities of China (Grant No. GK201702001), and the Fund for the Academic Leaders and Academic Backbones, Shaanxi Normal University of China (Grant No. 16QNGG007).
Effects of refractory period on the dynamical range in excitable networks are studied by computer simulations and theoretical analysis. The first effect is that the maximum or peak of the dynamical range appears when the largest eigenvalue of adjacent matrix is larger than one. We present a modification of the theory of the critical point by considering the correlation between excited nodes and their neighbors, which is brought by the refractory period. Our analysis provides the interpretation for the shift of the peak of the dynamical range. The effect is negligible when the average degree of the network is large. The second effect is that the dynamical range increases as the length of refractory period increases, and it is independent of the average degree. We present the mechanism of the second effect. As the refractory period increases, the saturated response decreases. This makes the bottom boundary of the dynamical range smaller and the dynamical range extend.
Critical avalanche is the characteristic feature of the self-organized criticality and is observed in a variety of natural phenomena.[1–3] The principle of neural network functions has been widely studied using dynamic models,[4–8] and critical avalanches have been found in neural networks.[9] The studied systems in the critical states exhibit several advantages.[9–12] One advantage is that the dynamical range is maximal in the critical state.[11,13,14] The critical states have optimal information processing by responding stimulus spanning several orders of magnitude.[11] The property was proposed in modeling study on networks of coupled excitable elements,[11] which are used widely as the simplified model of neural networks.[10,11,15–19] The property that neural network exhibits critical avalanche and largest dynamical range has been observed in experiments.[13,14] In research of excitable networks, the theory of predicating the critical state and the dynamical range has been proposed. It has been shown that criticality and the maximal dynamical range occur in networks in which the largest eigenvalue of adjacency matrix is one.[17]
The properties of networked systems depend on both the network structure and the dynamics of network elements. The biological realistic dynamic properties of network elements may play an important role in the criticality of the network. The way of enhancing the dynamical range was studied by including biological realistic features in the model.[20,21] For example, the inhibitory element was considered in the study of enhancing the dynamical range in excitable networks.[20]
The refractory period of neurons is an essential dynamical feature in excitable networks. In the refractory period, neurons do not response to signals. The dynamical features have important role in the collective behavior of the network. It was shown that refractory period leads to a wide range of scaling as well as periodic behavior.[22] The refractory period was considered in the theory predicting the criticality.[18] However, the effect of the refractory period on the dynamical range has not been studied.
In the present work, we study how the dynamical range changes with the refractory period. We show that the largest eigenvalue of the network corresponding to the critical point and the maximal dynamic range depends on the averaged degree of the network. In the critical state, the maximal dynamical range is enhanced by the refractory period.
The critical branching model[10,11] is used. The network consists of N nodes and each node has r + 2 states: si = 0 is the resting state, si = 1 is the excited state, and si = 2,3,…,r + 1 are refractory states. Time is discrete t = 0,1,… The node state is updated at each step. After excitation, the evolution of node state is deterministic. The state si is increased by 1 until si = r + 1, then the state changes to si = 0. The length of refractory state is r steps. If node i is at the resting state
We create networks in three steps: First, we create an ER random network with NK/2 links. The links are assigned to randomly chosen pairs of nodes. The average connection degree of each node in the network is K. Second, we assign a weight to each link. If node i and node j are connected, the entries of adjacent matrix Aij and Aji will be random numbers drawn from a uniform distribution between 0 and 1. Third, we calculate the largest eigenvalue λ of adjacency matrix, and multiply the adjacency matrix
In addition, we calculate the largest eigenvalue of adjacency matrix by
The response F is defined as[11]
The dynamical range was proposed to measure the ability of a network to robustly code the input stimuli.[11] As a function of the stimulus intensity η, networks have a minimum response F0 and a maximum response Fmax. Discarding stimuli that are too weak to be distinguished from F0 or close to saturation, the dynamical range is defined as
We simulate the response of the network to external stimulus and compute the dynamical range of networks with or without the refractory period. Figure
It has been shown that criticality exhibits the optimal dynamical range.[11] Here the maximal dynamical range occurs at λ = 1.0 for networks without refractory period. It is the same as the predicated critical point that the largest eigenvalue of the network adjacent matrix equals one (λ = 1). However, the maximal dynamical range occurs at λ > 1.0 for the network with refractory period.
To understand the shift of the maximal dynamical range in the relation between the dynamical range and the largest eigenvalue, we study the the transition of the response of the network. In the absence of external stimulus, sub-critical networks and critical networks do not exhibit self-sustained activity, whereas super-critical networks do. The critical point can be obtained from the transition of network response from zero to nonzero with η → 0.[11] In our simulation, the stimulus strength η = 10−5 is used to study the transition of response. Figure
We next analyze the relation between the critical point and the refractory period. In Ref. [17] the network without refractory period was considered, and it was assumed that no correlation exists between nodes in the excited states and nodes in the resting states. In networks with refractory period, the neighborhood of the excited nodes becomes different from the whole network. We first show this difference by numerical simulations.
The probability that the ith node is in the resting state (si = 0) at time t is denoted by
In networks without refractory period r = 0, the probability z satisfies z = 1 − F0, where F0 represents the fraction of nodes in the excited state under η → 0. When the external stimulus is absent, in the network in critical states, the F0 can be ignored. For deriving z′, we assume that the node i is excited. Besides the node i, the number of excited nodes is NF0 −1. Except for the node i, nodes are in the excited state with the probability (NF0 − 1)/(N − 1). Assuming that all excited nodes are independent, the probability that a neighbor of node i is in resting state is
In the network with refractory period (r > 0), nodes in excited states are not independent of their neighbor’s states. If node i is excited by the signals from nodes in the networks, the predecessor node must be in the refractory state (s = 2). Therefore, in the neighborhood of the node i, a node is in the state s = 2 with the probability 1/K. One can randomly select a node from N − 2 nodes except the node i, and its predecessor is in the excited state with the probability (NF0 − 1)/(N − 2) or one of r refractory states with the probability (NF0 r − 1)/(N − 2). In the neighborhood of node i, a node is in the resting state with the probability
The shift of the critical point with the refractory period can be understood through the approach proposed in Refs. [11] and [17]. The probability that the ith node at t is in the excited state is denoted by
Using the largest eigenvalue of the adjacency matrix
To show that the analytic result is reasonable, we plot the simulation result of the relation between F0 and β in Fig.
Furthermore, the analysis shows that when the average degree of the network is large, the shift of critical point can be ignored. We show the simulation results of the relation between F0 and λ for the network with K = 200 in Fig.
The effect of the refractory period on the dynamical range is independent of the average degree. We show the relation between the dynamical range and λ for networks with K = 200 in Fig.
We show the response F versus the input signal intensity η in Fig.
We have studied the dynamical range of networks consisting of excitable elements with the refractory period, and obtained that in networks with small average degree, the maximal dynamical range appears when the value of the largest eigenvalue of adjacent matrix is larger than 1, and the dynamical range is enhanced by the refractory period.
We present the mechanism of the effect of refractory period. The refractory period brings the correlation between the excited nodes and its neighbors. As a result, the fraction of resting state node in the neighborhood of excited nodes is less than that in the whole network. Considering the correlation, we obtain that the critical point of the network with refractory period appears when the largest eigenvalue of adjacency matrix is larger than one. The critical point is also the condition that the dynamical range reaches the maximal value. In networks with refractory period, the critical value of the largest eigenvalue decreases monotonically with the average degree. As the average degree is large, the critical condition becomes such that the largest eigenvalue of adjacency matrix is one, which was proposed in Ref. [17].
The effect that refractory period enhances the dynamical range is independent of the average degree. We present the mechanism of the effect. The saturated response decreases as the refractory period becomes longer. As a result, the low boundary of stimulus intensity becomes smaller. The decrease of low boundary induces the increase of the dynamical range.
We obtain the effects of the refractory period on enhancing the dynamical range and the shift of the critical point. The results provide a new way of enhancing dynamical range. The mechanism of these effects is helpful for understanding the critical behaviors of neural networks.
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