Nick Teresa A
Department of Neuroscience, Graduate Program in Neuroscience, Center for Neurobehavioral Development, Center for Neuroengineering, The University of Minnesota, Twin Cities, Minneapolis, Minnesota, 55455.
Dev Neurobiol. 2015 Oct;75(10):1091-113. doi: 10.1002/dneu.22189. Epub 2014 Jun 4.
Birdsong is a form of sensorimotor learning that involves a mirror-like system that activates with both song hearing and production. Early models of song learning, based on behavioral measures, identified key features of vocal plasticity, such as the requirements for memorization of a tutor song and auditory feedback during song practice. The concept of a comparator, which compares the memory of the tutor song to auditory feedback, featured prominently. Later models focused on linking anatomically-defined neural modules to behavioral concepts, such as the comparator. Exploiting the anatomical modularity of the songbird brain, localized lesions illuminated mechanisms of the neural song system. More recent models have integrated neuronal mechanisms identified in other systems with observations in songbirds. While these models explain multiple aspects of song learning, they must incorporate computational elements based on unknown biological mechanisms to bridge the motor-to-sensory delay and/or transform motor signals into the sensory domain. Here, I introduce the stabilizing critic hypothesis, which enables sensorimotor learning by (1) placing a purely sensory comparator afferent of the song system and (2) endowing song system disinhibitory interneuron networks with the capacity both to bridge the motor-sensory delay through prolonged bursting and to stabilize song segments selectively based on the comparator signal. These proposed networks stabilize an otherwise variable signal generated by both putative mirror neurons and a cortical-basal ganglia-thalamic loop. This stabilized signal then temporally converges with a matched premotor signal in the efferent song motor cortex, promoting spike-timing-dependent plasticity in the premotor circuitry and behavioral song learning.
鸟鸣是一种感觉运动学习形式,涉及一个类似镜像的系统,该系统在听到歌声和发出歌声时都会被激活。早期基于行为测量的歌曲学习模型确定了发声可塑性的关键特征,例如在歌曲练习过程中对记忆示范鸟歌声和听觉反馈的要求。比较器的概念,即将示范鸟歌声的记忆与听觉反馈进行比较,占据了突出地位。后来的模型专注于将解剖学定义的神经模块与行为概念(如比较器)联系起来。利用鸣禽大脑的解剖模块化,局部损伤揭示了神经歌曲系统的机制。最近的模型将在其他系统中确定的神经元机制与在鸣禽中的观察结果相结合。虽然这些模型解释了歌曲学习的多个方面,但它们必须纳入基于未知生物学机制的计算元素,以弥合运动到感觉的延迟和/或将运动信号转换到感觉领域。在这里,我介绍稳定批评家假说,该假说通过(1)在歌曲系统的纯感觉比较器传入神经中放置一个,以及(2)赋予歌曲系统去抑制性中间神经元网络通过长时间爆发来弥合运动-感觉延迟并基于比较器信号选择性稳定歌曲片段的能力,从而实现感觉运动学习。这些提出的网络稳定了由假定的镜像神经元和皮质-基底神经节-丘脑环路产生的原本可变的信号。然后,这个稳定的信号在传出歌曲运动皮层中与匹配的运动前信号在时间上汇聚,促进运动前电路中的尖峰时间依赖性可塑性和行为歌曲学习。