生殖隔离与物种形成：基因和分子的作用

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"这篇文章是关于物种形成的，特别是通过后期合子隔离的方式，由H. Allen Orr和Daven C. Presgraves两位专家撰写的综述论文。他们探讨了新物种如何在不同分化群体间的生殖隔离演化中产生，重点关注后期合子隔离的作用，包括杂种不育和不可存活性。文章涵盖了近年来在后期合子隔离遗传学方面的研究进展，特别是对推动这种隔离演化的种群遗传力以及导致杂种问题的具体基因和分子特征的了解。" 正文: 物种形成是生物学中的核心概念，它描述了原本属于同一物种的群体在经过一段时间的地理隔离或生态隔离后，逐渐发展出生殖隔离，最终形成新的物种。这篇论文“通过后期合子隔离的物种形成：力、基因和分子”深入剖析了这一过程。后期合子隔离是物种隔离的一种类型，不同于前期合子隔离，如求偶行为差异等。后期合子隔离主要涉及到杂种的不育或不可存活，这是阻止两个物种间基因交流的关键机制。近年来的研究开始揭示驱动后期合子隔离演化的种群遗传力，例如性选择和减数分裂驱动等冲突过程可能对杂种不育的进化有贡献。性选择，作为达尔文理论的一部分，可能导致物种间的适应性差异，这些差异可能加剧了生殖隔离。另一方面，减数分裂驱动是一种自私遗传元素，可以提高其自身的遗传频率，但也可能影响其他基因，从而促进后期合子隔离的形成。此外，论文还关注到导致杂种问题的具体基因和分子特征。尽管目前的分子遗传数据有限，但初步发现表明，“物种形成基因”可能对应于在物种内具有正常功能的基因座，这些基因座有时会因基因调控的进化而发生分歧。这表明，物种间的分化可能并不总是由于新基因的出现，而是现有基因在表达调控上的改变。论文引用了2000年发表在《BioEssays》上的研究，强调了2000年以来在这个领域的进步，展示了John Wiley & Sons, Inc.出版的这一研究成果对理解物种形成机制的重要性。通过对后期合子隔离的遗传学和分子层面的探索，科学家们正在逐步揭示生物多样性产生的复杂过程。

Speciation by postzygotic isolation:

forces, genes and molecules

H. Allen Orr* and Daven C. Presgraves

Summary

New species arise as reproductive isolation evolves

between diverging populations. Here we review recent

work in the genetics of postzygotic reproductive isola-

tionÐthe sterility and inviability of species hybrids. Over

the last few years, research has taken two new directions.

First, we have begun to learn a good deal about the

population genetic forces driving the evolution of post-

zygotic isolation. It has, for instance, become increas-

ingly clear that conflict-driven processes, like sexual

selection and meiotic drive, may contribute to the

evolution of hybrid sterility. Second, we have begun to

learn something about the identity and molecular char-

acteristics of the actual genes causing hybrid problems.

Although molecular genetic data are limited, early find-

ings suggest that ``speciation genes'' correspond to loci

having normal functions within species and that these

loci sometimes diverge as a consequence of evolution in

gene regulation. BioEssays 22:1085±1094, 2000.

ß 2000 John Wiley & Sons, Inc.

Introduction

SpeciationÐthe splitting of one species into twoÐoccupies a

unique place in the theory of evolution. Although a micro-

evolutionary process, speciation ultimately gives rise to the

macroevolutionary relationships we see reflected in phylo-

geny. Despite this special position, the history of speciation

research has been curiously episodic: periods of intense work

have been separated by many years of neglect.

In the first period of sustained work, Darwin, Wallace,

Jordan, Wagner and others came to realize that species are

mutable and wrestled with the roles of adaptation and geo-

graphic isolation in their origin. In the second period, the foun-

ders of the Modern Synthesis profoundly transformed our

understanding of just what species are and, in turn, of what it

means for species to split. Dobzhansky and Mayr, in particular,

championed the Biological Species Concept, the view that

species are characterized by their reproductive isolation from

each other, not by differences in morphology. This reproduc-

tive isolation, they argued, takes two forms: prezygotic and

postzygotic. In the former, barriers such as courtship

differences prevent the actual formation of hybrid zygotes

while, in the latter, barriers such as hybrid sterility or inviability

bar the flow of genes between species through hybrids. In both

cases, reproductive isolation ensures that species remain

genetically distinct and, consequently, that they can undergo

independent evolutionary fates.

In the third periodÐwhich began in the early 1980's and

which followed forty years of relative neglectÐattention

shifted to the genetics of speciation. Although much has been

learned during this period, progress has centered on a fairly

narrow class of problems. We now know a good deal, for

instance, about the number of genes that cause reproductive

isolation as well as their locations in the genome. We also

understand, at least roughly, the causes of several patterns

that characterize speciation. The best known of these is

Haldane's rule, the preferential sterility and inviability of

hybrids of the heterogametic [XY] sex.

(1±3)

And last, we now

possess a reasonably rich population genetic theory of

speciation, a theory that did not exist fifteen years ago and

that has yielded a number of novel predictions about the

evolution of reproductive isolation.

(4±6)

While these develop-

ments represent real accomplishments, they share a certain

focus. They all concern what might be called the classical or

``black box'' genetics of speciation. Only now are we beginning

to open this box, getting our first glimpse of the detailed forces,

genes and molecules that underlie reproductive isolation. We

believe that these new studies may represent a nascent fourth

period in the study of speciation.

Our goal here is to summarize some of these recent

developments. Given, however, that the speciation literature

has been heavily reviewed, it might be best if we first make

clear what we will

not do. We will not be concerned here with

the geography or ecology of speciation. Nor will we consider

prezygotic isolation in any detail. Instead, we focus on intrinsic

postzygotic isolation, i.e., on hybrid sterility and inviability,

particularly on the genetics and molecular bases of hybrid

problems, rather than on comparative patterns like Haldane's

rule. We make these restrictions for several reasons. For one,

we know much more about the genetics of hybrid sterility and

inviability than about any other form of isolation and there is

good reason to believe that there will be major advances in this

field soon. For another, while broad patterns like Haldane's

rule have received a great deal of attention, progress on the

BioEssays 22:1085±1094, ß 2000 John Wiley & Sons, Inc. BioEssays 22.12 1085

Department of Biology, University of Rochester, Rochester, New York.

Funding agencies: NIH; Grant Number: GM51932; David and Lucile

Packard Foundation.

*Correspondence to: H. Allen Orr, Department of Biology, University of

Rochester, Rochester, NY 14627.

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