Theissen Guenter, Melzer Rainer
Friedrich-Schiller-Universitaet Jena, Lehrstuhl fuer Genetik, Philosophenweg 12, D-07743 Jena, Germany.
Ann Bot. 2007 Sep;100(3):603-19. doi: 10.1093/aob/mcm143. Epub 2007 Jul 31.
Understanding the mode and mechanisms of the evolution of the angiosperm flower is a long-standing and central problem of evolutionary biology and botany. It has essentially remained unsolved, however. In contrast, considerable progress has recently been made in our understanding of the genetic basis of flower development in some extant model species. The knowledge that accumulated this way has been pulled together in two major hypotheses, termed the 'ABC model' and the 'floral quartet model'. These models explain how the identity of the different types of floral organs is specified during flower development by homeotic selector genes encoding transcription factors.
We intend to explain how the 'ABC model' and the 'floral quartet model' are now guiding investigations that help to understand the origin and diversification of the angiosperm flower.
Investigation of orthologues of class B and class C floral homeotic genes in gymnosperms suggest that bisexuality was one of the first innovations during the origin of the flower. The transition from dimer to tetramer formation of floral homeotic proteins after establishment of class E proteins may have increased cooperativity of DNA binding of the transcription factors controlling reproductive growth. That way, we hypothesize, better 'developmental switches' originated that facilitated the early evolution of the flower. Expression studies of ABC genes in basally diverging angiosperm lineages, monocots and basal eudicots suggest that the 'classical' ABC system known from core eudicots originated from a more fuzzy system with fading borders of gene expression and gradual transitions in organ identity, by sharpening of ABC gene expression domains and organ borders. Shifting boundaries of ABC gene expression may have contributed to the diversification of the angiosperm flower many times independently, as may have changes in interactions between ABC genes and their target genes.
了解被子植物花的进化模式和机制是进化生物学和植物学中长期存在的核心问题。然而,该问题基本上仍未得到解决。相比之下,最近我们在了解一些现存模式物种花发育的遗传基础方面取得了相当大的进展。通过这种方式积累的知识被整合到两个主要假说中,即“ABC模型”和“花四重奏模型”。这些模型解释了在花发育过程中,编码转录因子的同源异型选择基因是如何决定不同类型花器官的特征的。
我们旨在解释“ABC模型”和“花四重奏模型”如今如何指导有助于理解被子植物花的起源和多样化的研究。
对裸子植物中B类和C类花同源异型基因直系同源物的研究表明,两性花是花起源过程中的首批创新之一。在E类蛋白建立后,花同源异型蛋白从二聚体形成向四聚体形成的转变可能增强了控制生殖生长的转录因子与DNA结合的协同性。我们据此推测,由此产生了更好的“发育开关”,促进了花的早期进化。对基部发散的被子植物谱系、单子叶植物和基部真双子叶植物中ABC基因的表达研究表明,核心真双子叶植物中已知的“经典”ABC系统起源于一个基因表达边界更模糊、器官特征逐渐过渡的更模糊系统,通过ABC基因表达域和器官边界的锐化而来。ABC基因表达边界的移动可能多次独立地促成了被子植物花的多样化,ABC基因与其靶基因之间相互作用的变化也可能如此。