Department of Biomedical Sciences, Program in Neuronal Growth and Development, and Program in Cell and Molecular Biology, Campus Delivery 1617, Colorado State University, Fort Collins, CO 80523, USA.
Neuroscience. 2010 Jul 28;168(4):982-1008. doi: 10.1016/j.neuroscience.2009.10.028. Epub 2009 Oct 20.
The panglial syncytium maintains ionic conditions required for normal neuronal electrical activity in the central nervous system (CNS). Vital among these homeostatic functions is "potassium siphoning," a process originally proposed to explain astrocytic sequestration and long-distance disposal of K(+) released from unmyelinated axons during each action potential. Fundamentally different, more efficient processes are required in myelinated axons, where axonal K(+) efflux occurs exclusively beneath and enclosed within the myelin sheath, precluding direct sequestration of K(+) by nearby astrocytes. Molecular mechanisms for entry of excess K(+) and obligatorily-associated osmotic water from axons into innermost myelin are not well characterized, whereas at the output end, axonally-derived K(+) and associated osmotic water are known to be expelled by Kir4.1 and aquaporin-4 channels concentrated in astrocyte endfeet that surround capillaries and that form the glia limitans. Between myelin (input end) and astrocyte endfeet (output end) is a vast network of astrocyte "intermediaries" that are strongly inter-linked, including with myelin, by abundant gap junctions that disperse excess K(+) and water throughout the panglial syncytium, thereby greatly reducing K(+)-induced osmotic swelling of myelin. Here, I review original reports that established the concept of potassium siphoning in unmyelinated CNS axons, summarize recent revolutions in our understanding of K(+) efflux during axonal saltatory conduction, then describe additional components required by myelinated axons for a newly-described process of voltage-augmented "dynamic" potassium siphoning. If any of several molecular components of the panglial syncytium are compromised, K(+) siphoning is blocked, myelin is destroyed, and axonal saltatory conduction ceases. Thus, a common thread linking several CNS demyelinating diseases is the disruption of potassium siphoning/water transport within the panglial syncytium. Continued progress in molecular identification and subcellular mapping of glial ion and water channels will lead to a better understanding of demyelinating diseases of the CNS and to development of improved treatment regimens.
神经胶质合胞体能维持中枢神经系统(CNS)中神经元电活动所需的离子条件。在这些动态平衡功能中,至关重要的是“钾虹吸”,这一过程最初被提出用于解释少突胶质细胞对无髓轴突在每个动作电位期间释放的 K+的隔离和长距离处理。在有髓鞘的轴突中,需要完全不同且更有效的过程,因为轴突 K+的外排仅发生在髓鞘下方并被其包围,从而排除了附近少突胶质细胞对 K+的直接隔离。进入最内层髓鞘的过量 K+和必需伴随的渗透水的分子机制尚未很好地描述,而在输出端,已知由 Kir4.1 和水通道蛋白-4 通道将轴突来源的 K+和相关的渗透水排出,这些通道集中在包围毛细血管的星形胶质细胞足突中,并形成胶质界膜。在髓鞘(输入端)和星形胶质细胞足突(输出端)之间是一个巨大的星形胶质细胞“中间物”网络,它们通过丰富的缝隙连接强烈地相互连接,包括与髓鞘的连接,这些连接将过量的 K+和水分散到整个神经胶质合胞体中,从而大大降低了 K+诱导的髓鞘渗透肿胀。在这里,我回顾了最初确立无髓鞘 CNS 轴突中钾虹吸概念的报告,总结了我们最近对轴突跳跃传导期间 K+外排的理解的革命,然后描述了有髓鞘轴突新描述的电压增强“动态”钾虹吸过程所需的其他成分。如果神经胶质合胞体的几个分子成分受损,钾虹吸就会受阻,髓鞘被破坏,轴突跳跃传导停止。因此,几种 CNS 脱髓鞘疾病的一个共同线索是破坏神经胶质合胞体中的钾虹吸/水转运。对神经胶质离子和水通道的分子鉴定和亚细胞定位的持续进展将导致更好地理解 CNS 的脱髓鞘疾病,并开发出改进的治疗方案。