Bachtrog Doris
Department of Integrative Biology, University of California Berkeley, Berkeley, CA 94720, USA.
BMC Evol Biol. 2008 Dec 18;8:334. doi: 10.1186/1471-2148-8-334.
Adaptive protein evolution is common in several Drosophila species investigated. Some studies point to very weak selection operating on amino-acid mutations, with average selection intensities on the order of Nes approximately in D. melanogaster and D. simulans. Species with lower effective population sizes should undergo less adaptation since they generate fewer mutations and selection is ineffective on a greater proportion of beneficial mutations.
Here I study patterns of polymorphism and divergence at 91 X-linked loci in D. miranda, a species with a roughly 5-fold smaller effective population size than D. melanogaster. Surprisingly, I find a similar fraction of amino-acid mutations being driven to fixation by positive selection in D. miranda and D. melanogaster. Genes with higher rates of amino-acid evolution show lower levels of neutral diversity, a pattern predicted by recurrent adaptive protein evolution. I fit a hitchhiking model to patterns of polymorphism in D. miranda and D. melanogaster and estimate an order of magnitude higher selection coefficients for beneficial mutations in D. miranda.
This analysis suggests that effective population size may not be a major determinant in rates of protein adaptation. Instead, adaptation may not be mutation-limited, or the distribution of fitness effects for beneficial mutations might differ vastly between different species or populations. Alternative explanation such as biases in estimating the fraction of beneficial mutations or slightly deleterious mutation models are also discussed.
适应性蛋白质进化在已研究的几种果蝇物种中很常见。一些研究指出,作用于氨基酸突变的选择非常微弱,在黑腹果蝇和拟暗果蝇中,平均选择强度约为Nes量级。有效种群规模较小的物种应该经历较少的适应性进化,因为它们产生的突变较少,而且选择对更大比例的有益突变无效。
在这里,我研究了米兰达果蝇中91个X连锁基因座的多态性和分化模式,该物种的有效种群规模比黑腹果蝇小约5倍。令人惊讶的是,我发现在米兰达果蝇和黑腹果蝇中,由正选择驱动固定的氨基酸突变比例相似。氨基酸进化速率较高的基因显示出较低水平的中性多样性,这是反复适应性蛋白质进化所预测的模式。我将一个搭便车模型应用于米兰达果蝇和黑腹果蝇的多态性模式,并估计米兰达果蝇中有益突变的选择系数要高一个数量级。
该分析表明,有效种群规模可能不是蛋白质适应性进化速率的主要决定因素。相反,适应性进化可能不受突变限制,或者有益突变的适合度效应分布在不同物种或种群之间可能有很大差异。还讨论了其他解释,如估计有益突变比例的偏差或轻微有害突变模型。