Kuratani Evolutionary Morphology Laboratory, RIKEN Center for Developmental Biology, 2-2-3 Minatojima-Minamimachi, Chuo-ku Kobe, 650-0047 Japan.
Pattern Formation Group, Graduate School of Frontier Biosciences, Osaka University, 1-3 Yamadaoka, Suita, Osaka 565-0871 Japan.
Zoological Lett. 2015 Nov 26;1:33. doi: 10.1186/s40851-015-0033-0. eCollection 2015.
Somites, blocks of mesoderm tissue located on either side of the neural tube in the developing vertebrate embryo, are derived from mesenchymal cells in the presomitic mesoderm (PSM) and are a defining characteristic of vertebrates. In vertebrates, the somite segmental boundary is determined by Notch signalling and the antagonistic relationship of the downstream targets of Notch, Lfng, and Delta1 in the anterior PSM. The presence of somites in the basal chordate amphioxus (Branchiostoma floridae) indicates that the last common ancestor of chordates also had somites. However, it remains unclear how the genetic mechanisms underlying somitogenesis in vertebrates evolved from those in ancestral chordates.
We demonstrate that during the gastrula stages of amphioxus embryos, BfFringe expression in the endoderm of the archenteron is detected ventrally to the ventral limit of BfDelta expression in the presumptive rostral somites along the dorsal/ventral (D/V) body axis. Suppression of Notch signalling by DAPT (a γ-secretase inhibitor that indirectly inhibits Notch) treatment from the late blastula stage reduced late gastrula stage expression of BfFringe in the endodermal archenteron and somite markers BfDelta and BfHairy-b in the mesodermal archenteron. Later in development, somites in the DAPT-treated embryo did not separate completely from the dorsal roof of the archenteron. In addition, clear segmental boundaries between somites were not detected in DAPT-treated amphioxus embryos at the larva stage. Similarly, in vertebrates, DAPT treatment from the late blastula stage in Xenopus (Xenopus laevis) embryos resulted in disruption of somite XlDelta-2 expression at the late gastrula stage. At the tail bud stage, the segmental expression of XlMyoD in myotomes was diminished.
We propose that Notch signalling and the Fringe/Delta cassette for dorso-ventral boundary formation in the archenteron that separates somites from the gut in an amphioxus-like ancestral chordate were co-opted for anteroposterior segmental boundary formation in the vertebrate anterior PSM during evolution.
体节是位于发育中脊椎动物胚胎神经管两侧的中胚层组织块,由前体节中胚层(PSM)中的间充质细胞衍生而来,是脊椎动物的一个定义特征。在脊椎动物中,体节的节段边界由 Notch 信号确定,而 Notch 的下游靶标 Lfng 和 Delta1 在 PSM 的前侧的拮抗关系决定。基础脊索动物文昌鱼(Branchiostoma floridae)中存在体节表明,脊索动物的最后共同祖先也有体节。然而,脊椎动物中体节发生的遗传机制如何从前脊索动物中进化而来,仍不清楚。
我们证明,在文昌鱼胚胎的原肠胚阶段,BfFringe 在原肠胚的内胚层中的表达位于假定的头侧体节的 BfDelta 表达的腹侧限的腹侧,沿着背/腹(D/V)体轴。从晚期囊胚期开始用 DAPT(一种间接抑制 Notch 的γ-分泌酶抑制剂)处理抑制 Notch 信号,减少了晚期原肠胚阶段内胚层原肠胚中的 BfFringe 和中胚层原肠胚中的 BfDelta 和 BfHairy-b 的表达。在发育后期,DAPT 处理的胚胎中的体节不能完全与原肠胚的背顶分离。此外,在 DAPT 处理的文昌鱼胚胎中,幼虫阶段没有检测到体节之间明显的节段边界。同样,在脊椎动物中,从晚期囊胚期开始在非洲爪蟾(Xenopus laevis)胚胎中用 DAPT 处理,导致晚期原肠胚阶段体节 XlDelta-2 表达中断。在尾芽阶段,肌节中的 XlMyoD 的节段性表达减少。
我们提出,在文昌鱼等祖先脊索动物中,Notch 信号和 Fringe/Delta 盒用于形成原肠胚中的背腹边界,将体节与肠道分开,在进化过程中被脊椎动物前 PSM 中的前后节段边界形成所采用。