Ledeen Robert W, Wu Gusheng
Department of Neurology and Neurosciences, New Jersey Medical School, UMDNJ, 185 So Orange Ave, MSB-H506, Newark, NJ 07103, USA.
Neurochem Res. 2010 Dec;35(12):1867-74. doi: 10.1007/s11064-010-0286-0. Epub 2010 Nov 13.
Among the many glycoconjugates contributing to the sugar code, gangliosides have drawn special attention owing to their predominance as the major sialoglycoconjugate category within the nervous system. However, their occurrence, albeit at lower levels, appears ubiquitous in vertebrate cells and even some invertebrate tissues. Now that over 100 gangliosides have been structurally characterized, their diverse physiological functions constitute a remaining enigma. This has been especially true of GM1, for which a surprising array of functions has already been revealed. Our current research has focused on two areas of GM1 function: (a) signaling induced in neural and immune cells by cross-linking of GM1 in the plasma membrane that leads to activation of TRPC5 (transient receptor potiential, canonical form 5) channels, a process important in neuritogenesis and autoimmune suppression; (b) activation by GM1 of a sodium-calcium exchanger (NCX) in the inner membrane of the nuclear envelope (NE) with resulting modulation of nuclear and cellular calcium. The latter has a role in maintaining neuronal viability, loss of which renders neurons vulnerable to Ca(2+) overload. Pathological manifestations in mutant mice and their cultured neurons lacking GM1 have shown dramatic rescue with a membrane permeable derivative of GM1 that enters the nucleus and restores NCX activity. Nuclear function of GM1 is related to the presence of neuraminidase in the NE, an enzyme that generates GM1 through hydrolysis of GD1a. A different isoform of this enzyme was found in each of the two membranes of the NE.
在众多构成糖代码的糖缀合物中,神经节苷脂因其在神经系统中作为主要唾液酸糖缀合物类别的优势地位而备受关注。然而,它们的存在虽然水平较低,但在脊椎动物细胞甚至一些无脊椎动物组织中似乎普遍存在。既然已经有超过100种神经节苷脂的结构得到了表征,它们多样的生理功能仍是一个未解之谜。对于GM1来说尤其如此,它已经被揭示出一系列惊人的功能。我们目前的研究集中在GM1功能的两个方面:(a) 质膜上GM1交联在神经和免疫细胞中诱导的信号传导,这会导致TRPC5(瞬时受体电位,典型形式5)通道的激活,这一过程在神经突生成和自身免疫抑制中很重要;(b) GM1激活核膜(NE)内膜中的钠钙交换器(NCX),从而调节细胞核和细胞内的钙。后者在维持神经元活力方面发挥作用,其丧失会使神经元易受Ca(2+)过载的影响。缺乏GM1的突变小鼠及其培养的神经元中的病理表现已通过一种可穿透膜的GM1衍生物得到显著挽救,该衍生物进入细胞核并恢复NCX活性。GM1的核功能与NE中神经氨酸酶的存在有关,这种酶通过水解GD1a产生GM1。在NE的两层膜中分别发现了这种酶的不同同工型。