Department of Biology, Indiana University, USA.
Mol Biol Evol. 2010 Jun;27(6):1404-14. doi: 10.1093/molbev/msq020. Epub 2010 Jan 29.
A central problem in evolutionary theory concerns the mechanisms by which adaptations requiring multiple mutations emerge in natural populations. We develop a series of expressions that clarify the scaling of the time to establishment of complex adaptations with population size, mutation rate, magnitude of the selective disadvantage of intermediate-state alleles, and the complexity of the adaptation. In general, even in the face of deleterious intermediate steps, the time to establishment is minimized in populations with very large size. Under a broad range of conditions, the time to establishment also scales by no more than the square of the mutation rate, regardless of the number of sites contributing to the adaptive change, demonstrating that the emergence of complex adaptations is only weakly constrained by the independent acquisition of mutations at the underlying sites. Mutator alleles with deleterious side effects have only moderate effects on the rate of adaptation in large populations but can cause a quantum decrease in the time to establishment of some adaptive alleles in small populations, although probably not at a high enough rate to offset the increased deleterious mutation load. Transient hypermutability, whereby a subset of gamete-producing cells mutate at an elevated rate in a nonheritable manner, may also elevate the rate of adaptation, although the effect is modest and appears to result from a simple increase in the rate of transitions between intermediate states rather than from the saltational production of doublet mutations. Taken together, these results illustrate the plausibility of the relatively rapid emergence of specific complex adaptations by conventional population genetic mechanisms and provide insight into the relative incidences of various paths of allelic adaptation in organisms with different population genetic features.
进化理论中的一个核心问题涉及到在自然种群中,需要多个突变才能出现的适应性机制。我们提出了一系列表达式,可以清晰地阐明复杂适应性在种群规模、突变率、中间状态等位基因选择劣势的幅度以及适应性的复杂性等因素下建立时间的比例关系。一般来说,即使面对有害的中间步骤,具有非常大种群规模的群体也能将建立时间最小化。在广泛的条件下,建立时间的比例关系也不会超过突变率的平方,无论对适应性变化有贡献的位点数量如何,这表明复杂适应性的出现仅受到独立获得基础位点突变的微弱限制。具有有害副作用的突变体等位基因在大群体中的适应率仅有中等影响,但在小群体中可能会导致某些适应性等位基因建立时间的量子下降,尽管可能不足以抵消增加的有害突变负荷。瞬时超突变性,即一部分产生配子的细胞以非遗传性的方式以较高的速率突变,也可能会提高适应率,尽管这种影响是适度的,并且似乎是由于中间状态之间的跃迁率简单增加而不是来自双重突变的突发产生。总的来说,这些结果说明了特定复杂适应性可以通过传统的群体遗传机制快速出现的合理性,并为不同群体遗传特征的生物体中各种等位基因适应途径的相对发生率提供了深入的见解。