† Corresponding author. E-mail:
Project supported by the National Natural Science Foundation of China (Grant No. 11672233), the Fundamental Research Funds for the Central Universities, China (Grant No. 3102017AX008), and the Seed Foundation of Innovation and Creation for Graduate Student in Northwestern Polytechnical University, China (Grant No. ZZ2018173).
With the help of a magnetic flux variable, the effects of stochastic electromagnetic disturbances on autapse Hodgkin–Huxley neuronal systems are studied systematically. Firstly, owing to the autaptic function, the inter-spike interval series of an autapse neuron not only bifurcates, but also presents a quasi-periodic characteristic. Secondly, an irregular mixed-mode oscillation induced by a specific electromagnetic disturbance is analyzed using the coefficient of variation of inter-spike intervals. It is shown that the neuronal discharge activity has certain selectivity to the noise intensity, and the appropriate noise intensity can induce the significant mixed-mode oscillations. Finally, the modulation effects of electromagnetic disturbances on a ring field-coupled neuronal network with autaptic structures are explored quantitatively using the average spiking frequency and the average coefficient of variation. The electromagnetic disturbances can not only destroy the continuous and synchronous discharge state, but also induce the resting neurons to generate the intermittent discharge mode and realize the transmission of neural signals in the neuronal network. The studies can provide some theoretical guidance for applying electromagnetic disturbances to effectively control the propagation of neural signals and treat mental illness.
In 1952, Hodgkin and Huxley proposed a mathematical model of biological neurons by studying the membrane current of squid nerve fibers, which is the so-called Hodgkin–Huxley neuron model.[1] This biological neuron model marked an important breakthrough in the study of biological nervous systems, and resulted in some typical neuron models, such as FitzHugh–Nagumo model,[2] Morris–Lecar model,[3] and Hindmarsh–Rose model.[4,5] References [6] and [7] summarized twenty typical characteristics of biological neuronal discharge activities and compared seventeen representative neuron models, respectively. For computational neurosciences, it is necessary not only to study the mechanisms of various factors such as channels, synapses and electromagnetic fields on the neuronal dynamic behavior, but also to study the relationship between neuronal dynamic behavior and neural information coding, which will help to reveal the neural mechanisms of learning and memory.[8–12]
Anatomical and electrophysiological experiments showed that neurons interact through synapses, which are the key functional structures for achieving signal transmissions and information exchanges between neurons. Autapse is a special synapse structure which enables a neuron to connect with itself through a closed loop and is mathematically depicted by a time-delayed feedback term.[13,14] Autapse divided into electric and chemical autapse is usually studied to explore the effects of autaptic conductance and delay time on neuronal firing patterns.[15–17] The self-adaption of autapse driving can induce different modes of electrical activities in a simple network of neuron-coupled astrocyte.[18] Also, autaptic structures can be considered as a pacemaker which can regulate the neuronal collective behaviors in a forward feedback Hindmarsh–Rose neuronal network.[19] In addition, appropriate autaptic parameters can enhance the propagation of the discharge rhythm and induce more pronounced stochastic resonance in a scale-free Hodgkin–Huxley neuronal network.[20] Physiological anatomic experiments have confirmed the existence of autaptic structures in some neurons of human brain, especially in most cortical pyramidal neurons.[21] In a nearestneighbor synaptic coupled neuronal network, the time delay can also enhance the neuronal firing rate.[22]
Because of the ubiquity of electromagnetic radiations, it is necessary to study its effects on electrical activities of neurons and neuronal networks. On the other hand, it is necessary to explore how to use electromagnetic radiations to treat mental illness. At present, there are mainly two different mechanisms of electromagnetic radiations on nervous systems. Firstly, the electromagnetic radiation absorbed by a neuron can be converted into the power of its own electrical energy, and the neuron model resulting from the electromagnetic radiation can be obtained according to the neuronal energy theory.[23] Secondly, based on the phenomenon of electromagnetic induction induced by ion transmembrane motions, the relationship between magnetic flux and induced current can be established according to the working principle of memristors. Thus, the effects of electromagnetic radiations or disturbances on the discharge behaviors of neurons or neuronal networks can be detected by introducing a magnetic induction current.[24–28] Reference [29] commented on the dynamic behaviors of biological neuron models, summarized the effects of autaptic structures and ion channel noises on neuronal firing activities, and proposed a new coupled mode between neurons.
With the help of a magnetic flux variable, this paper systematically studies the modulations of electromagnetic disturbances to the discharge behaviors of improved Hodgkin–Huxley biological neuronal systems. Firstly, the bifurcation dynamic behavior of an autapse neuron is analyzed to investigate the effect of the self-feedback memory ability induced by its autaptic structure on neuronal electrical activities. Secondly, the stochastic discharge behaviors induced by a stochastic electromagnetic disturbance are studied by using the coefficient of variation of inter-spiking intervals, and the selectivity of the neuronal discharge mode to the noise intensity is detected. Finally, the mechanism of electromagnetic disturbances on a Hodgkin–Huxley neuronal network with autaptic structures is explored by using the average spiking frequency and the average coefficient of variation, and the feasibility of applying electromagnetic disturbances to regulate its dynamic behavior is discussed.
Considering the magnetic induction current caused by ion exchanges or ion concentration fluctuations, a typical 4-dimensional Hodgkin–Huxley neuron model can be modified by introducing the magnetic flux variable. At the same time, considering the autaptic function, an improved 5-dimensional Hodgkin–Huxley neuron model is obtained as follows:
A ring field-coupled neuronal network with autaptic structures is given as follows:
In this study, the second-order stochastic Runge–Kutta algorithm[30] is used and the time step is set as Δ t=0.01 ms for all numerical computations. In particular, some parameters are also set as the fixed constants, such as gNa =120 mS/cm2, gK =36 mS/cm2, gL =0.3 mS/cm2,
During the numerical experiments, the autaptic function is activated at t = 120 ms, and the inter-spike interval (ISI) series are only recorded from t = 3500 ms to t = 5500 ms. As can be seen from Fig.
To detect the effect of the delay time on the neuronal dynamic behavior, Figure
However, it suddenly decreases when the delay time increases to a certain value, and then restarts the next cycle-like change process. This change should be a reflection of the intrinsic periodicity of the neuron. Finally, for the autaptic conductance g=0.1, the neuronal dynamic behavior has a two-cycle bifurcation phenomenon when the delay time
To explore the mechanism of the autaptic function, we further provide the spiking frequency diagram for different autaptic parameters. As can be seen from Fig.
Thus, the autaptic function can have a great influence on the neuronal dynamic behavior. On the one hand, it can change the neuronal excitability and the neuronal spiking frequency. On the other hand, a bifurcation phenomenon usually occurs when the autaptic function inhibits the neuronal discharge activity. In addition, the effect of the autaptic function on the neuronal dynamic behavior is diverse. This conclusion includes two meanings here, one is that the neuronal discharge activity has certain selectivity to the autaptic parameters, and the different autaptic parameters can induce the different discharge modes. The two is that the influence of the autaptic parameters on the neuronal dynamic behavior is local discontinuous and indicates the local sensitivity of the neuronal firing activity to the autaptic parameters. At last, the bifurcation behavior in the inter-spike interval series is not only affected by the autaptic parameters, but also by the neuronal intrinsic periodicity, which together determine the self-feedback memory ability induced by the autaptic structure. The above analysis could help to apply the autaptic structure in neuronal networks to obtain certain neurophysiological phenomena, such as spiral waves and target waves.
The coefficient of variation of the inter-spike interval series is introduced to measure the effect of electromagnetic disturbances on the neuronal discharge activity, which is defined as follows:
To further investigate the above extreme coherent resonance phenomenon, the neuronal membrane potentials are simulated by selecting two different noise intensities, as shown in Fig.
Figure
In summary, the electromagnetic disturbance in the form of Gaussian white noise can cause the transition of neuronal discharge modes. Particularly under the condition of the appropriate noise intensity, the autapse neuron can produce a significant intermittent discharge phenomenon, which can be measured by the coefficient of variation of the inter-spike interval series. According to the characteristics of neuronal firing activities induced by the autaptic function, the autaptic structure could be the cause of some mental states, and the different autaptic conditions could correspond to the different mental states. However, the electromagnetic disturbance can effectively modulate the discharge behavior induced by the autaptic structure and will be helpful for the treatment of mental illness.
There are mainly three types of connections among neurons in the central nervous system, i.e., convergence, divergence, and ring, and the ring structure chosen here is easily realized in computational neuroscience.[10] In order to study the modulation effect of electromagnetic disturbances on a ring field-coupled neuronal network consisting of 40 neurons, the average coefficient of variation
To explore the sensitivity of the discharge pattern to the noise intensity, Figure
To further study the effects of electromagnetic disturbances on neuronal discharge behavior and neural signal propagation in the network, only four neurons in the network are given the autaptic structures, that is, only if i = 19, 20, 21, and 22, then
In summary, when some neurons in the neuronal network have autaptic structures, the electromagnetic disturbances can act as a bridge or a catalyst to some extent, not only affecting the ability of electromagnetic field coupling between neurons in the network, but also changing the self-feedback effect of neurons with autaptic structures. Furthermore, the stochastic disturbances can help to release the neural electrical signals carried by these neurons, and then excite the resting neurons in the network to produce the intermittent firing activities. Thus, the electromagnetic disturbances can help to achieve the purpose of controlling the signal transmission in the neuronal network.
The modulation effects of electromagnetic disturbances on the dynamic behaviors of autapse Hodgkin–Huxley neurons and neuronal networks are studied systematically, which can be converted into induced current by introducing a magnetic flux variable.
First, due to the self-feedback memory ability induced by an autaptic structure and the neuronal intrinsic periodic property, the neuronal excitability can change and different autaptic parameters can induce different discharge modes. In addition, the neuronal discharge activity changes discontinuously, and its dynamic behavior not only presents bifurcation phenomena but also has a periodic characteristic. Second, the discharge activity of the autapse neuron has some selectivity to the noise intensity, and the appropriate noise intensity can induce MMOs. Numerical experiments show that the neuronal discharge modes can be transited after the electromagnetic disturbance is triggered. In particular, under the condition of the appropriate noise intensity, the neuron can produce a more significant intermittent firing mode. Finally, for the ring field-coupled neuronal network with autaptic structures, the electromagnetic disturbances can change the continuous and synchronous discharge state of neurons in the network, and result in a desynchronization phenomenon. With the increase of the noise intensity, the average spiking frequency and the average coefficient of variation of all neurons show the opposite change trends and have the significant extremum points, which indicate that the dynamic behavior of the network has certain dependence on the noise intensity. Especially, when only some neurons in the network have the autaptic structures, the electromagnetic disturbances can have some impact on the ability of the electromagnetic field coupling between neurons and the self-feedback memory function induced by the special structures, which can excite the resting neurons in the network to generate an intermittent firing mode and ultimately ensure the transmission of neural signals between neurons.
The above studies can deepen our understanding of the mechanism of electromagnetic disturbances on the dynamic behaviors of neuronal systems. Furthermore, the studies can also provide a valuable reference for effectively controlling the propagation of neural signals in the network and applying electromagnetic disturbances to treat mental illness.
[1] | |
[2] | |
[3] | |
[4] | |
[5] | |
[6] | |
[7] | |
[8] | |
[9] | |
[10] | |
[11] | |
[12] | |
[13] | |
[14] | |
[15] | |
[16] | |
[17] | |
[18] | |
[19] | |
[20] | |
[21] | |
[22] | |
[23] | |
[24] | |
[25] | |
[26] | |
[27] | |
[28] | |
[29] | |
[30] | |
[31] | |
[32] | |
[33] | |
[34] | |
[35] | |
[36] | |
[37] | |
[38] | |
[39] | |
[40] |