Grimmel Jan, Dorresteijn Adriaan W C, Fröbius Andreas C
Institut für Allgemeine und Spezielle Zoologie, Abteilung Entwicklungsbiologie, Justus-Liebig-Universität Gießen, Stephanstraße 24, 35390 Gießen, Germany.
Evodevo. 2016 Apr 12;7:10. doi: 10.1186/s13227-016-0046-6. eCollection 2016.
Platynereis and other polychaete annelids with homonomous segmentation are regarded to closely resemble ancestral forms of bilateria. The head region comprises the prostomium, the peristomium, a variable number of cephalized body segments and several appendages, like cirri, antennae and palps. The trunk of such polychaetes shows numerous, nearly identical segments. Each segment bears a parapodium with species-specific morphology on either side. The posterior end of the trunk features a segment proliferation zone and a terminal pygidium with the anus and anal cirri. The removal of a substantial part of the posterior trunk is by no means lethal. Cells at the site of injury dedifferentiate and proliferate forming a blastema to regenerate both the pygidium and the proliferation zone. The pygidium forms new anal cirri, and the proliferation zone generates new segments at a rapid pace. The formation of body appendages like the cirri and the segmental parapodia can thus be studied in the caudal regenerate of Platynereis within only a few days.
The development of body appendages in Platynereis is regulated by a network of genes common to polychaetes but also shared by distant taxa. We isolated DNA sequences from P. dumerilii of five genes known to be involved in appendage formation within other groups: Meis/homothorax, Pbx1/extradenticle, Dlx/Distal-less, decapentaplegic and specific protein 1/buttonhead. Analyses of expression patterns during caudal regeneration by in situ hybridization reveal striking similarities related to expression in arthropods and vertebrates. All genes exhibit transient expression during differentiation and growth of segments. As was shown previously in other phyla Pdu-Meis/hth and Pdu-Pbx1/exd are co-expressed, although the expression is not limited to the proximal part of the parapodia. Pdu-Dll is prominent in parapodia but upregulated in the anal cirri. No direct dependence concerning Pdu-Dll and Pdu-sp/btd expression is observed in Platynereis. Pdu-dpp shows an expression pattern not comparable to its expression in other taxa.
The expression patterns observed suggest conserved roles of these genes during appendage formation across different clades, but the underlying mechanisms utilizing this toolset might not be identical. Some genes show broad expression along the proximodistal axis indicating a possible role in proximodistal patterning of body appendages. Other genes exhibit expression patterns limited to specific parts and tissues of the growing parapodia, thus presumably being involved in formation of taxon-specific morphological differences.
多毛纲动物沙蚕及其他具有同律分节的多毛类环节动物被认为与两侧对称动物的祖先形态极为相似。头部区域包括口前叶、围口节、数量可变的头部化身体节段以及一些附肢,如触须、触角和触须。此类多毛类动物的躯干有许多几乎相同的节段。每个节段两侧都有一对具有物种特异性形态的疣足。躯干后端有一个节段增殖区和一个带有肛门及肛触须的末端尾节。切除躯干后部的大部分并非致命。损伤部位的细胞去分化并增殖形成芽基,以再生尾节和增殖区。尾节形成新的肛触须,增殖区快速产生新的节段。因此,在短短几天内就可以在沙蚕的尾部再生体中研究触须和节段性疣足等身体附肢的形成。
沙蚕身体附肢的发育受多毛类动物共有的基因网络调控,这些基因在远缘类群中也有共享。我们从杜氏沙蚕中分离出了五个已知参与其他类群附肢形成的基因的DNA序列:Meis/同胸节、Pbx1/额外齿、Dlx/无远端、果蝇decapentaplegic基因和特定蛋白1/钮头蛋白。通过原位杂交分析尾部再生过程中的表达模式,发现与节肢动物和脊椎动物中的表达有显著相似之处。所有基因在节段分化和生长过程中均表现出瞬时表达。如先前在其他门中所示,Pdu-Meis/hth和Pdu-Pbx1/exd共同表达,尽管表达不限于疣足的近端部分。Pdu-Dll在疣足中突出,但在肛触须中上调。在沙蚕中未观察到Pdu-Dll和Pdu-sp/btd表达之间的直接依赖性。Pdu-dpp显示出与在其他类群中的表达不同的表达模式。
观察到的表达模式表明这些基因在不同进化枝的附肢形成过程中具有保守作用,但利用这一工具集的潜在机制可能并不相同。一些基因沿近端 - 远端轴广泛表达,表明可能在身体附肢的近端 - 远端模式形成中发挥作用。其他基因的表达模式仅限于生长中的疣足的特定部位和组织,因此可能参与形成类群特异性形态差异。