Snippe H P, van Hateren J H
Department of Neurobiophysics, University of Groningen, Groningen, The Netherlands.
Neural Comput. 2007 May;19(5):1179-214. doi: 10.1162/neco.2007.19.5.1179.
Feedback control in neural systems is ubiquitous. Here we study the mathematics of nonlinear feedback control. We compare models in which the input is multiplied by a dynamic gain (multiplicative control) with models in which the input is divided by a dynamic attenuation (divisive control). The gain signal (resp. the attenuation signal) is obtained through a concatenation of an instantaneous nonlinearity and a linear low-pass filter operating on the output of the feedback loop. For input steps, the dynamics of gain and attenuation can be very different, depending on the mathematical form of the nonlinearity and the ordering of the nonlinearity and the filtering in the feedback loop. Further, the dynamics of feedback control can be strongly asymmetrical for increment versus decrement steps of the input. Nevertheless, for each of the models studied, the nonlinearity in the feedback loop can be chosen such that immediately after an input step, the dynamics of feedback control is symmetric with respect to increments versus decrements. Finally, we study the dynamics of the output of the control loops and find conditions under which overshoots and undershoots of the output relative to the steady-state output occur when the models are stimulated with low-pass filtered steps. For small steps at the input, overshoots and undershoots of the output do not occur when the filtering in the control path is faster than the low-pass filtering at the input. For large steps at the input, however, results depend on the model, and for some of the models, multiple overshoots and undershoots can occur even with a fast control path.
神经系统中的反馈控制无处不在。在此,我们研究非线性反馈控制的数学原理。我们将输入乘以动态增益的模型(乘法控制)与输入除以动态衰减的模型(除法控制)进行比较。增益信号(相应地,衰减信号)是通过对反馈回路输出进行瞬时非线性变换和线性低通滤波的级联得到的。对于输入阶跃,增益和衰减的动态特性可能非常不同,这取决于非线性的数学形式以及反馈回路中非线性和滤波的顺序。此外,对于输入的增量和减量阶跃,反馈控制的动态特性可能具有强烈的不对称性。然而,对于所研究的每个模型,可以选择反馈回路中的非线性,使得在输入阶跃之后,反馈控制的动态特性相对于增量和减量是对称的。最后,我们研究控制回路输出的动态特性,并找出当模型受到低通滤波阶跃激励时,输出相对于稳态输出出现超调量和欠调量的条件。对于输入处的小阶跃,当控制路径中的滤波比输入处的低通滤波更快时,输出不会出现超调量和欠调量。然而,对于输入处的大阶跃,结果取决于模型,并且对于某些模型,即使控制路径很快,也可能会出现多次超调量和欠调量。