Wong-Riley Margaret T T
Department of Cell Biology, Neurobiology and Anatomy, Medical College of Wisconsin, Milwaukee, WI 53226, USA.
Eye Brain. 2010;2:99-116. doi: 10.2147/EB.S9078. Epub 2010 Jul 22.
The visual system is one of the most energetically demanding systems in the brain. The currency of energy is ATP, which is generated most efficiently from oxidative metabolism in the mitochondria. ATP supports multiple neuronal functions. Foremost is repolarization of the membrane potential after depolarization. Neuronal activity, ATP generation, blood flow, oxygen consumption, glucose utilization, and mitochondrial oxidative metabolism are all interrelated. In the retina, phototransduction, neurotransmitter utilization, and protein/organelle transport are energy-dependent, yet repolarization-after-depolarization consumes the bulk of the energy. Repolarization in photoreceptor inner segments maintains the dark current. Repolarization by all neurons along the visual pathway following depolarizing excitatory glutamatergic neurotransmission preserves cellular integrity and permits reactivation. The higher metabolic activity in the magno- versus the parvo-cellular pathway, the ON- versus the OFF-pathway in some (and the reverse in other) species, and in specialized functional representations in the visual cortex all reflect a greater emphasis on the processing of specific visual attributes. Neuronal activity and energy metabolism are tightly coupled processes at the cellular and even at the molecular levels. Deficiencies in energy metabolism, such as in diabetes, mitochondrial DNA mutation, mitochondrial protein malfunction, and oxidative stress can lead to retinopathy, visual deficits, neuronal degeneration, and eventual blindness.
视觉系统是大脑中能量需求最大的系统之一。能量货币是三磷酸腺苷(ATP),它在线粒体中通过氧化代谢最有效地产生。ATP支持多种神经元功能。最重要的是在去极化后使膜电位复极化。神经元活动、ATP生成、血流、氧消耗、葡萄糖利用和线粒体氧化代谢都是相互关联的。在视网膜中,光转导、神经递质利用以及蛋白质/细胞器运输都依赖能量,但去极化后的复极化消耗了大部分能量。光感受器内段的复极化维持暗电流。在去极化兴奋性谷氨酸能神经传递后,视觉通路上所有神经元的复极化维持细胞完整性并允许再次激活。在某些物种中,大细胞通路与小细胞通路、ON通路与OFF通路(在其他物种中则相反)以及视觉皮层中的特殊功能表征中较高的代谢活性,都反映出对特定视觉属性处理的更大重视。在细胞甚至分子水平上,神经元活动和能量代谢是紧密耦合的过程。能量代谢缺陷,如糖尿病、线粒体DNA突变、线粒体蛋白质功能障碍和氧化应激,可导致视网膜病变、视觉缺陷、神经元变性,最终导致失明。