Sander Klaus, Schmidt-Ott Urs
Institut für Biologie I (Zoologie), D-79104 Freiburg, Germany.
J Exp Zool B Mol Dev Evol. 2004 Jan 15;302(1):69-91. doi: 10.1002/jez.b.20003.
We discuss the interplay between evolution and development as reflected in data and concepts since about 1800. Darwin and his "continental apostle" Haeckel put the striking similarity between early vertebrate embryos in an evolutionary context. Haeckel's partly illicit generalizations discredited evolutionary thinking among early experimental embryologists who moreover noted riddles incompatible with contemporary concepts of homology and evolution. Relevant solutions were suggested by the more recent concept of ontogenetic networks that embody complex regulatory properties and genes with partly overlapping functions. Molecular data on development increasingly reveal evolutionary opportunism, for instance when a widespread signaling chain involved in primitive immune defense was apparently recruited later on for dorso-ventral axis determination in some evolutionarily advanced insect groups. Recently, Rickettsia-related bacteria colonizing many arthropod species were found to "manipulate" the first steps of host development to the advantage of their own propagation, but by ways that could also promote host speciation. Molecular genetics can now document evolutionary steps in ontogenetic networks. In the fruit fly for instance, the novel bicoid gene has superseded a crucial patterning function within a pre-existing network--a case of "molecular caenogenesis." The expression patterns of conserved genes that antagonistically determine dorso-ventral polarity support a literal revolution envisioned almost 200 years ago. This is the dorso-ventral inversion of the body plan in some metazoans--ascribed then to the Articulata, now to the Chordata. The final section posits that the opportunistic character of evolutionary innovations is detrimental to parsimony in development.
我们探讨自1800年左右以来,数据和概念中所反映的进化与发育之间的相互作用。达尔文及其“大陆使徒”海克尔将早期脊椎动物胚胎之间惊人的相似性置于进化背景之中。海克尔部分非法的归纳法使早期实验胚胎学家对进化思维产生了怀疑,此外,他们还注意到一些与当代同源性和进化概念不相容的谜题。个体发育网络的最新概念提出了相关解决方案,该概念体现了复杂的调控特性以及功能部分重叠的基因。关于发育的分子数据越来越多地揭示了进化机会主义,例如,参与原始免疫防御的广泛信号链后来显然在一些进化上更高级的昆虫群体中被用于背腹轴的确定。最近发现,许多节肢动物物种中定殖的立克次氏体相关细菌会“操纵”宿主发育的最初步骤,以利于自身繁殖,但其方式也可能促进宿主物种形成。分子遗传学现在可以记录个体发育网络中的进化步骤。例如,在果蝇中,新的双尾基因取代了一个预先存在的网络中的关键模式形成功能——这是一个“分子新发生”的例子。拮抗地决定背腹极性的保守基因的表达模式支持了近200年前设想的一场真正的革命。这就是一些后生动物身体结构的背腹反转——当时归因于节肢动物,现在归因于脊索动物。最后一部分认为,进化创新的机会主义特征不利于发育中的简约性。