The Jungers Center for Neurosciences Research, Department of Neurology, Oregon Health & Science University, Portland, OR 97239, USA.
Hum Mol Genet. 2013 Apr 15;22(8):1493-506. doi: 10.1093/hmg/dds562. Epub 2013 Jan 7.
The demyelinating peripheral neuropathy Charcot-Marie-Tooth type 4B (CMT4B) is characterized by axonal degeneration and myelin outfoldings. CMT4B results from mutations in either myotubularin-related protein 2 (MTMR2; CMT4B1) or MTMR13 (CMT4B2), phosphoinositide (PI) 3-phosphatases that dephosphorylate phosphatidylinositol 3-phosphate (PtdIns3P) and PtdIns(3,5)P2, lipids which regulate endo-lysosomal membrane traffic. The catalytically active MTMR2 and catalytically inactive MTMR13 physically associate, although the significance of this association is not well understood. Here we show that Mtmr13 loss leads to axonal degeneration in sciatic nerves of older mice. In addition, CMT4B2-like myelin outfoldings are present in Mtmr13(-/-) nerves at postnatal day 3. Thus, Mtmr13(-/-) mice show both the initial dysmyelination and later degenerative pathology of CMT4B2. Given the key role of PI 3-kinase-Akt signaling in myelination, we investigated the state of the pathway in nerves of CMT4B models. We found that Akt activation is unaltered in Mtmr13(-/-) and Mtmr2(-/-) mice. Mtmr2 and Mtmr13 are found within the Schwann cell cytoplasm, where the proteins are partially localized to punctate compartments, suggesting that Mtmr2-Mtmr13 may dephosphorylate their substrates on specific intracellular compartments. Mtmr2-Mtmr13 substrates play essential roles in endo-lysosomal membrane traffic. However, endosomes and lysosomes of Mtmr13(-/-) and Mtmr2(-/-) Schwann cells are morphologically indistinguishable from those of controls, indicating that loss of these proteins does not cause wholesale dysregulation of the endo-lysosomal system. Notably, Mtmr2 and Mtmr13 depend upon each other to achieve wild-type levels of protein expression. Mtmr2 stabilizes Mtmr13 on membranes, indicating that the Mtmr13 pseudophosphatase is regulated by its catalytically active binding partner.
脱髓鞘周围神经病型 4B(CMT4B)的特征是轴突变性和髓鞘折叠。CMT4B 是由肌管相关蛋白 2(MTMR2;CMT4B1)或 MTMR13(CMT4B2)突变引起的,这两种蛋白都是磷酸肌醇 3-磷酸(PtdIns3P)和 PtdIns(3,5)P2的磷酸酶,这些脂质调节内溶酶体膜运输。催化活性的 MTMR2 和无催化活性的 MTMR13 物理结合,尽管这种结合的意义尚不清楚。在这里,我们表明 Mtmr13 的缺失会导致老年小鼠坐骨神经中的轴突变性。此外,在出生后第 3 天的 Mtmr13(-/-)神经中也存在 CMT4B2 样的髓鞘折叠。因此,Mtmr13(-/-)小鼠既有 CMT4B2 的早期脱髓鞘和后期退行性病变。鉴于 PI 3-激酶-Akt 信号通路在髓鞘形成中的关键作用,我们研究了 CMT4B 模型中神经的通路状态。我们发现 Akt 的激活在 Mtmr13(-/-)和 Mtmr2(-/-)小鼠中没有改变。Mtmr2 和 Mtmr13 存在于施万细胞的细胞质中,在那里蛋白部分定位于点状隔室中,这表明 Mtmr2-Mtmr13 可能在特定的细胞内隔室中对其底物进行去磷酸化。Mtmr2-Mtmr13 的底物在内溶酶体膜运输中起着至关重要的作用。然而,Mtmr13(-/-)和 Mtmr2(-/-)施万细胞的内体和溶酶体在形态上与对照无区别,这表明这些蛋白的缺失不会导致内溶酶体系统的全面失调。值得注意的是,Mtmr2 和 Mtmr13 相互依赖以达到野生型蛋白表达水平。Mtmr2 稳定 Mtmr13 膜结合,表明 Mtmr13 假磷酸酶受其催化活性结合伙伴的调节。