Janssen Ralf, Damen Wim G M, Budd Graham E
Department of Earth Sciences, Uppsala University, Villavägen 16, 752 36, Uppsala, Sweden.
BMC Dev Biol. 2012 May 17;12:15. doi: 10.1186/1471-213X-12-15.
A hallmark of Drosophila segmentation is the stepwise subdivision of the body into smaller and smaller units, and finally into the segments. This is achieved by the function of the well-understood segmentation gene cascade. The first molecular sign of a segmented body appears with the action of the pair rule genes, which are expressed as transversal stripes in alternating segments. Drosophila development, however, is derived, and in most other arthropods only the anterior body is patterned (almost) simultaneously from a pre-existing field of cells; posterior segments are added sequentially from a posterior segment addition zone. A long-standing question is to what extent segmentation mechanisms known from Drosophila may be conserved in short-germ arthropods. Despite the derived developmental modes, it appears more likely that conserved mechanisms can be found in anterior patterning.
Expression analysis of pair rule gene orthologs in the blastoderm of the pill millipede Glomeris marginata (Myriapoda: Diplopoda) suggests that these genes are generally involved in segmenting the anterior embryo. We find that the Glomeris pairberry-1 ( pby-1) gene is expressed in a pair rule pattern that is also found in insects and a chelicerate, the mite Tetraynchus urticae. Other Glomeris pair rule gene orthologs are expressed in double segment wide domains in the blastoderm, which at subsequent stages split into two stripes in adjacent segments.
The expression patterns of the millipede pair rule gene orthologs resemble pair rule patterning in Drosophila and other insects, and thus represent evidence for the presence of an ancestral pair rule-like mechanism in myriapods. We discuss the possibilities that blastoderm patterning may be conserved in long-germ and short-germ arthropods, and that a posterior double segmental mechanism may be present in short-germ arthropods.
果蝇体节形成的一个标志是身体逐步细分为越来越小的单位,最终形成体节。这是通过已被充分理解的体节基因级联反应实现的。体节化身体的第一个分子迹象出现在成对规则基因的作用下,这些基因在交替的体节中以横向条纹的形式表达。然而,果蝇的发育是衍生而来的,在大多数其他节肢动物中,只有身体前部(几乎)同时从预先存在的细胞区域形成模式;后部体节从后体节添加区依次添加。一个长期存在的问题是,果蝇中已知的体节形成机制在短胚节肢动物中在多大程度上是保守的。尽管发育模式有所不同,但在前部模式形成中似乎更有可能找到保守机制。
在马陆球马陆(多足纲:倍足纲)胚盘中成对规则基因直系同源物的表达分析表明,这些基因通常参与前部胚胎的体节形成。我们发现球马陆的成对莓-1(pby-1)基因以成对规则模式表达,这种模式也在昆虫和一种螯肢动物叶螨中发现。其他球马陆成对规则基因直系同源物在胚盘中以双节宽域表达,在随后的阶段在相邻体节中分裂成两条条纹。
马陆成对规则基因直系同源物的表达模式类似于果蝇和其他昆虫中的成对规则模式,因此代表了多足纲中存在祖先成对规则样机制的证据。我们讨论了胚盘模式形成在长胚节和短胚节肢动物中可能保守的可能性,以及短胚节肢动物中可能存在后部双节机制的可能性。