Atreya Krishan B, Fernandes Joyce J
Department of Zoology, Miami University, Oxford, OH 45056, USA.
Dev Biol. 2008 Sep 1;321(1):123-40. doi: 10.1016/j.ydbio.2008.06.023. Epub 2008 Jun 24.
During insect myogenesis, myoblasts are organized into a pre-pattern by specialized organizer cells. In the Drosophila embryo, these cells have been termed founder cells and play important roles in specifying muscle identity and in serving as targets for myoblast fusion. A group of adult muscles, the dorsal longitudinal (flight) muscles, DLMs, is patterned by persistent larval scaffolds; the second set, the dorso-ventral muscles, DVMs is patterned by mono-nucleate founder cells (FCs) that are much larger than the surrounding myoblasts. Both types of organizer cells express Dumbfounded, which is known to regulate fusion during embryonic myogenesis. The role of DVM founder cells as well as the DLM scaffolds was tested in genetic ablation studies using the UAS/Gal4 system of targeted transgene expression. In both cases, removal of organizer cells prior to fusion, causes formation of supernumerary fibers, suggesting that cells in the myoblast pool have the capacity to initiate fiber formation, which is normally inhibited by the organizers. In addition to the large DVM FCs, some (smaller) cells in the myoblast pool also express Dumbfounded. We propose that these cells are responsible for seeding supernumerary fibers, when DVM FCs are eliminated prior to fusion. When these cells are also eliminated, myogenesis fails to occur. In the second set of studies, targeted expression of constitutively active Ras(V12) also resulted in the appearance of supernumerary fibers. In this case, the original DVM FCs are present, suggesting alterations in cell fate. Taken together, these data suggest that DVM myoblasts are able to respond to cues other than the original founder cell, to initiate fusion and fiber formation. Thus, the role of the large DVM founder cells is to generate the correct number of fibers, but they are not required for fiber formation itself. We also present evidence that the DVM FCs may arise from the leg imaginal disc.
在昆虫肌生成过程中,成肌细胞由特化的组织者细胞组织成一种前模式。在果蝇胚胎中,这些细胞被称为起始细胞,在确定肌肉特性以及作为成肌细胞融合的靶点方面发挥着重要作用。一组成年肌肉,即背纵(飞行)肌(DLMs),由持续存在的幼虫支架形成模式;第二组,即背腹肌(DVMs),由比周围成肌细胞大得多的单核起始细胞(FCs)形成模式。两种类型的组织者细胞都表达“呆立蛋白”,已知该蛋白在胚胎肌生成过程中调节融合。使用靶向转基因表达的UAS/Gal4系统,在基因消融研究中测试了DVM起始细胞以及DLM支架的作用。在这两种情况下,在融合前去除组织者细胞会导致形成多余的纤维,这表明成肌细胞池中的细胞具有启动纤维形成的能力,而这种能力通常受到组织者的抑制。除了大的DVM FCs外,成肌细胞池中的一些(较小的)细胞也表达“呆立蛋白”。我们提出,当DVM FCs在融合前被消除时,这些细胞负责产生多余的纤维。当这些细胞也被消除时,肌生成就无法发生。在第二组研究中,组成型活性Ras(V12)的靶向表达也导致了多余纤维的出现。在这种情况下,原始的DVM FCs存在,这表明细胞命运发生了改变。综上所述,这些数据表明DVM成肌细胞能够对除原始起始细胞之外的其他信号作出反应,以启动融合和纤维形成。因此,大的DVM起始细胞的作用是产生正确数量的纤维,但纤维形成本身并不需要它们。我们还提供证据表明,DVM FCs可能起源于腿部成虫盘。