Imai Y, Busby W H, Smith C E, Clarke J B, Garmong A J, Horwitz G D, Rees C, Clemmons D R
Department of Medicine, University of North Carolina School of Medicine, Chapel Hill 27599-7170, USA.
J Clin Invest. 1997 Nov 15;100(10):2596-605. doi: 10.1172/JCI119803.
IGFs are pleiotrophic mitogens for porcine smooth muscle cells (pSMC) in culture. The effects of IGFs on cells are modulated by various insulin-like growth factor-binding proteins (IGFBP). IGFBP-5 is synthesized by pSMC and binds to the extracellular matrix. However, IGFBP-5 is also secreted into conditioned medium of cultured cells and is cleaved into fragments by a concomitantly produced protease. These fragments have reduced affinity for the IGFs and cleavage makes it difficult to assess the role of intact IGFBP-5. To study the consequence of accumulation of intact IGFBP-5 in medium, we determined the cleavage site in IGFBP-5 and prepared a protease resistant mutant. Amino acid sequencing of purified IGFBP-5 fragments suggested Arg138-Arg139 as the primary cleavage site. Arg138-Arg139-->Asn138-Asn139 mutations were introduced to create protease-resistant IGFBP-5, which has the same affinity for IGF-I as the native protein. This mutant IGFBP-5 remained intact even after 24 h of incubation and it inhibited several IGF-I actions when added to pSMC culture medium. The mutant IGFBP-5 (500 ng/ml) decreased IGF-I stimulated cellular DNA synthesis by 84%, protein synthesis by 77%, and it inhibited IGF-I stimulated migration of pSMC by 77%. It also inhibited IGF-I stimulation of IRS-1 phosphorylation. In contrast, the same amount of native IGFBP-5 did not inhibit IGF-I actions. The significance of inhibitory effects of the protease resistant IGFBP-5 was further demonstrated in pSMC transfected with mutant or native IGFBP-5 cDNAs. The mutant IGFBP-5 accumulated in culture medium of transfected cells, while native IGFBP-5 was degraded into fragments, PSMC overexpressing the mutant IGFBP-5 also responded poorly to IGF-I compared with mock transfected cells. IGF-I (5 ng/ml) increased [35S]methionine incorporation into control cells by 36% above the basal level, but it did not significantly change (4%) in pSMC cultures that were producing the mutant IGFBP-5. In conclusion, the accumulation of protease-resistant IGFBP-5 in the medium was inhibitory to IGF-I actions on pSMC. This suggests that proteolysis can prevent IGFBP-5 from acting as an inhibitor of IGF-I-stimulated effects and that it serves as an important mechanism for regulating cellular responsiveness to IGF-I.
胰岛素样生长因子(IGFs)是培养的猪平滑肌细胞(pSMC)的多效性促有丝分裂原。IGFs对细胞的作用受到多种胰岛素样生长因子结合蛋白(IGFBP)的调节。IGFBP - 5由pSMC合成并与细胞外基质结合。然而,IGFBP - 5也分泌到培养细胞的条件培养基中,并被同时产生的蛋白酶切割成片段。这些片段对IGFs的亲和力降低,切割使得评估完整IGFBP - 5的作用变得困难。为了研究完整IGFBP - 5在培养基中积累的后果,我们确定了IGFBP - 5中的切割位点并制备了一种抗蛋白酶突变体。纯化的IGFBP - 5片段的氨基酸测序表明Arg138 - Arg139是主要切割位点。引入Arg138 - Arg139→Asn138 - Asn139突变以产生抗蛋白酶的IGFBP - 5,其对IGF - I的亲和力与天然蛋白相同。这种突变的IGFBP - 5即使在孵育24小时后仍保持完整,并且当添加到pSMC培养基中时,它抑制了几种IGF - I的作用。突变的IGFBP - 5(500 ng/ml)使IGF - I刺激的细胞DNA合成减少84%,蛋白质合成减少77%,并且它抑制IGF - I刺激的pSMC迁移77%。它还抑制IGF - I对胰岛素受体底物1(IRS - 1)磷酸化的刺激作用。相比之下,相同量的天然IGFBP - 5不抑制IGF - I的作用。抗蛋白酶IGFBP - 5的抑制作用的重要性在转染了突变或天然IGFBP - 5 cDNA的pSMC中得到进一步证明。突变的IGFBP - 5在转染细胞的培养基中积累,而天然IGFBP - 5被降解成片段,与mock转染细胞相比,过表达突变IGFBP - 5的PSMC对IGF - I的反应也较差。IGF - I(5 ng/ml)使对照细胞中[35S]甲硫氨酸掺入量比基础水平增加36%,但在产生突变IGFBP - 5的pSMC培养物中没有显著变化(4%)。总之,培养基中抗蛋白酶IGFBP - 5的积累抑制了IGF - I对pSMC的作用。这表明蛋白水解可以防止IGFBP - 5作为IGF - I刺激作用的抑制剂起作用,并且它是调节细胞对IGF - I反应性的重要机制。