Moffett John R, Ross Brian, Arun Peethambaran, Madhavarao Chikkathur N, Namboodiri Aryan M A
Department of Anatomy, Physiology and Genetics, Uniformed Services University of the Health Sciences, Building C, 4301 Jones Bridge Rd., Bethesda, MD 20814, USA.
Prog Neurobiol. 2007 Feb;81(2):89-131. doi: 10.1016/j.pneurobio.2006.12.003. Epub 2007 Jan 5.
The brain is unique among organs in many respects, including its mechanisms of lipid synthesis and energy production. The nervous system-specific metabolite N-acetylaspartate (NAA), which is synthesized from aspartate and acetyl-coenzyme A in neurons, appears to be a key link in these distinct biochemical features of CNS metabolism. During early postnatal central nervous system (CNS) development, the expression of lipogenic enzymes in oligodendrocytes, including the NAA-degrading enzyme aspartoacylase (ASPA), is increased along with increased NAA production in neurons. NAA is transported from neurons to the cytoplasm of oligodendrocytes, where ASPA cleaves the acetate moiety for use in fatty acid and steroid synthesis. The fatty acids and steroids produced then go on to be used as building blocks for myelin lipid synthesis. Mutations in the gene for ASPA result in the fatal leukodystrophy Canavan disease, for which there is currently no effective treatment. Once postnatal myelination is completed, NAA may continue to be involved in myelin lipid turnover in adults, but it also appears to adopt other roles, including a bioenergetic role in neuronal mitochondria. NAA and ATP metabolism appear to be linked indirectly, whereby acetylation of aspartate may facilitate its removal from neuronal mitochondria, thus favoring conversion of glutamate to alpha ketoglutarate which can enter the tricarboxylic acid cycle for energy production. In its role as a mechanism for enhancing mitochondrial energy production from glutamate, NAA is in a key position to act as a magnetic resonance spectroscopy marker for neuronal health, viability and number. Evidence suggests that NAA is a direct precursor for the enzymatic synthesis of the neuron specific dipeptide N-acetylaspartylglutamate, the most concentrated neuropeptide in the human brain. Other proposed roles for NAA include neuronal osmoregulation and axon-glial signaling. We propose that NAA may also be involved in brain nitrogen balance. Further research will be required to more fully understand the biochemical functions served by NAA in CNS development and activity, and additional functions are likely to be discovered.
大脑在许多方面在器官中独树一帜,包括其脂质合成和能量产生机制。神经系统特异性代谢物N-乙酰天门冬氨酸(NAA)由神经元中的天门冬氨酸和乙酰辅酶A合成,它似乎是中枢神经系统新陈代谢这些独特生化特征的关键环节。在出生后早期中枢神经系统(CNS)发育过程中,少突胶质细胞中脂肪生成酶的表达,包括降解NAA的酶天门冬氨酸酰化酶(ASPA),随着神经元中NAA产量的增加而增加。NAA从神经元转运到少突胶质细胞的细胞质中,在那里ASPA裂解乙酸部分用于脂肪酸和类固醇合成。然后产生的脂肪酸和类固醇继续用作髓磷脂脂质合成的组成部分。ASPA基因的突变会导致致命的脑白质营养不良——卡纳万病,目前尚无有效治疗方法。一旦出生后髓鞘形成完成,NAA可能会继续参与成人的髓磷脂脂质周转,但它似乎也承担其他作用,包括在神经元线粒体中的生物能量作用。NAA和ATP代谢似乎间接相关,即天门冬氨酸的乙酰化可能有助于其从神经元线粒体中去除,从而有利于谷氨酸转化为α-酮戊二酸,后者可进入三羧酸循环进行能量产生。作为增强谷氨酸线粒体能量产生的一种机制,NAA处于关键位置,可作为神经元健康、活力和数量的磁共振波谱标记物。有证据表明,NAA是神经元特异性二肽N-乙酰天门冬氨酰谷氨酸酶促合成的直接前体,N-乙酰天门冬氨酰谷氨酸是人类大脑中浓度最高的神经肽。NAA的其他假定作用包括神经元渗透调节和轴突-胶质细胞信号传导。我们认为NAA可能还参与大脑的氮平衡。需要进一步研究以更全面地了解NAA在中枢神经系统发育和活动中所起的生化功能,并且可能会发现更多功能。