Department of Neurobiology, Yale University School of Medicine, New Haven, CT 06510, United States.
Department of Neurobiology, Yale University School of Medicine, New Haven, CT 06510, United States; Department of Ophthalmology and Visual Science, Yale University School of Medicine, New Haven, CT 06510, United States; Kavli Institute for Neuroscience, Yale University School of Medicine, New Haven, CT 06510, United States.
Curr Opin Neurobiol. 2014 Feb;24(1):166-75. doi: 10.1016/j.conb.2013.11.011. Epub 2013 Dec 22.
The initial structural and functional development of visual circuits in reptiles, birds, and mammals happens independent of sensory experience. After eye opening, visual experience further refines and elaborates circuits that are critical for normal visual function. Innate genetic programs that code for gradients of molecules provide gross positional information for developing nerve cells, yet much of the cytoarchitectural complexity and synaptogenesis of neurons depends on calcium influx, neurotransmitter release, and neural activity before the onset of vision. In fact, specific spatiotemporal patterns of neural activity, or 'retinal waves', emerge amidst the development of the earliest connections made between excitable cells in the developing eye. These patterns of spontaneous activity, which have been observed in all amniote retinae examined to date, may be an evolved adaptation for species with long gestational periods before the onset of functional vision, imparting an informational robustness and redundancy to guide development of visual maps across the nervous system. Recent experiments indicate that retinal waves play a crucial role in the development of interconnections between different parts of the visual system, suggesting that these spontaneous patterns serve as a template-matching mechanism to prepare higher-order visually associative circuits for the onset of visuomotor learning and behavior. Key questions for future studies include determining the exact sources and nature of spontaneous activity during development, characterizing the interactions between neural activity and transcriptional gene regulation, and understanding the extent of circuit connectivity governed by retinal waves within and between sensory-motor systems.
在爬虫类、鸟类和哺乳动物中,视觉回路的初始结构和功能发育是独立于感觉体验的。在睁眼后,视觉体验进一步细化和阐述了对正常视觉功能至关重要的回路。编码分子梯度的先天遗传程序为发育中的神经细胞提供了大致的位置信息,但神经元的大部分细胞结构复杂性和突触发生都依赖于钙内流、神经递质释放和视觉出现前的神经活动。事实上,在发育中的眼睛中兴奋性细胞之间最早形成的连接中,出现了特定的时空模式的神经活动,或“视网膜波”。这些自发活动模式在迄今为止检查的所有羊膜类动物视网膜中都有观察到,可能是为具有较长妊娠期的物种进化适应的结果,为引导整个神经系统中的视觉图发育提供了信息鲁棒性和冗余性。最近的实验表明,视网膜波在视觉系统不同部分之间的连接发育中起着至关重要的作用,表明这些自发模式充当模板匹配机制,为视觉运动学习和行为的出现准备更高阶的视觉联想回路。未来研究的关键问题包括确定发育过程中自发活动的确切来源和性质,描述神经活动和转录基因调控之间的相互作用,以及了解视网膜波在感觉运动系统内和之间控制的回路连接的程度。