Walker Linda C, Overstreet Mayra A, Yeowell Heather N
Division of Dermatology, Duke University Medical Center, Durham, Box 3135, NC 27710, USA.
Matrix Biol. 2005 Jan;23(8):515-23. doi: 10.1016/j.matbio.2004.11.002. Epub 2004 Dec 10.
Lysyl hydroxylases 1, 2, and 3 catalyse the hydroxylation of specific lysines in collagen. A small percentage of these hydroxylysine residues are precursors for the cross-link formation essential for the tensile strength of collagen. Lysyl hydroxylase 2 (LH2) exists as two alternatively-spliced forms; the long transcript (the major ubiquitously-expressed form) includes a 63 bp exon (13A) that is spliced out in the short form (expressed, together with the long form, in human kidney, spleen, liver, and placenta). This study shows that this alternative splicing event can be regulated by both cell density and cycloheximide (CHX). Although only the long form of LH2 is detected in untreated confluent human skin fibroblasts, after 24 h treatment with CHX the short LH2 transcript is also expressed. In kidney cells, in which both LH2 transcripts are equally expressed, the long LH2 transcript is significantly decreased after 24 h CHX treatment, whereas expression of the short transcript is slightly increased. This suggests that, in kidney cells, the splicing mechanism for the inclusion of exon 13A in LH2 requires a newly-synthesized protein factor that is suppressed by CHX, whereas, in skin fibroblasts in which levels of LH2 (long) are unaffected, CHX appears to suppress a factor that inhibits exclusion of exon 13A, thereby promoting expression of LH2 (short). As these alternate transcripts of LH2 may have specificity for hydroxylation of lysines in either telopeptide or helical collagen domains, their relative expression determines the type of cross-links formed, thereby affecting collagen strength. Therefore, any perturbation of the regulation of LH2 splicing could influence the stability of the extracellular matrix and contribute to specific connective tissue disorders.
赖氨酰羟化酶1、2和3催化胶原蛋白中特定赖氨酸的羟化反应。这些羟赖氨酸残基中有一小部分是形成交联所必需的前体,而交联对于胶原蛋白的抗张强度至关重要。赖氨酰羟化酶2(LH2)以两种可变剪接形式存在;长转录本(主要在全身广泛表达的形式)包含一个63 bp的外显子(13A),该外显子在短形式中被剪接掉(短形式与长形式一起在人肾、脾、肝和胎盘中表达)。本研究表明,这种可变剪接事件可受细胞密度和环己酰亚胺(CHX)的调控。虽然在未处理的汇合人皮肤成纤维细胞中仅检测到LH2的长形式,但用CHX处理24小时后,短的LH2转录本也会表达。在两种LH2转录本均等量表达的肾细胞中,CHX处理24小时后,长的LH2转录本显著减少,而短转录本的表达略有增加。这表明,在肾细胞中,LH2中包含外显子13A的剪接机制需要一种新合成的蛋白质因子,而该因子会被CHX抑制;而在LH2(长形式)水平不受影响的皮肤成纤维细胞中,CHX似乎抑制了一种抑制外显子13A排除的因子,从而促进了LH2(短形式)的表达。由于LH2的这些可变转录本可能对端肽或螺旋胶原蛋白结构域中的赖氨酸羟化具有特异性,它们的相对表达决定了形成的交联类型,从而影响胶原蛋白强度。因此,LH2剪接调控的任何扰动都可能影响细胞外基质的稳定性,并导致特定的结缔组织疾病。