Department of Genome Sciences, University of Washington, Seattle, Washington, USA.
PLoS Genet. 2013;9(2):e1003287. doi: 10.1371/journal.pgen.1003287. Epub 2013 Feb 7.
Sperm and egg proteins constitute a remarkable paradigm in evolutionary biology: despite their fundamental role in mediating fertilization (suggesting stasis), some of these molecules are among the most rapidly evolving ones known, and their divergence can lead to reproductive isolation. Because of strong selection to maintain function among interbreeding individuals, interacting fertilization proteins should also exhibit a strong signal of correlated divergence among closely related species. We use evidence of such molecular co-evolution to target biochemical studies of fertilization in North Pacific abalone (Haliotis spp.), a model system of reproductive protein evolution. We test the evolutionary rates (d(N)/d(S)) of abalone sperm lysin and two duplicated egg coat proteins (VERL and VEZP14), and find a signal of co-evolution specific to ZP-N, a putative sperm binding motif previously identified by homology modeling. Positively selected residues in VERL and VEZP14 occur on the same face of the structural model, suggesting a common mode of interaction with sperm lysin. We test this computational prediction biochemically, confirming that the ZP-N motif is sufficient to bind lysin and that the affinities of VERL and VEZP14 are comparable. However, we also find that on phylogenetic lineages where lysin and VERL evolve rapidly, VEZP14 evolves slowly, and vice versa. We describe a model of sexual conflict that can recreate this pattern of anti-correlated evolution by assuming that VEZP14 acts as a VERL mimic, reducing the intensity of sexual conflict and slowing the co-evolution of lysin and VERL.
尽管它们在介导受精(暗示稳定)方面起着根本作用,但这些分子中的一些是已知进化最快的分子之一,它们的分化可能导致生殖隔离。由于在杂交个体中维持功能的强烈选择,相互作用的受精蛋白也应该在密切相关的物种中表现出强烈的相关分化信号。我们利用这种分子协同进化的证据来针对北太平洋鲍鱼(Haliotis spp.)的受精生化研究,这是生殖蛋白进化的一个模型系统。我们测试了鲍鱼精子溶菌酶和两个复制的卵壳蛋白(VERL 和 VEZP14)的进化速率(d(N)/ d(S)),并发现了一个与 ZP-N 相关的协同进化信号,ZP-N 是以前通过同源建模确定的一个假定的精子结合基序。VERL 和 VEZP14 中的正选择残基出现在结构模型的同一面上,这表明它们与精子溶菌酶的相互作用模式相同。我们从生化角度验证了这个计算预测,证实了 ZP-N 基序足以结合溶菌酶,并且 VERL 和 VEZP14 的亲和力相当。然而,我们还发现,在溶菌酶和 VERL 快速进化的进化谱系中,VEZP14 进化缓慢,反之亦然。我们描述了一个性冲突模型,通过假设 VEZP14 作为 VERL 的模拟物,可以重现这种反相关进化的模式,从而降低性冲突的强度并减缓溶菌酶和 VERL 的协同进化。