Yamamoto Maki, Nishikawa Toshikazu, Kajitani Hiroyuki, Kawano Shigeyuki
Institute of Natural Sciences, Senshu University, 2-1-1, Tama, Kawasaki, Kanagawa 214-8580, Japan.
Planta. 2007 Sep;226(4):917-27. doi: 10.1007/s00425-007-0538-7. Epub 2007 May 23.
Non-flagellated vegetative green algae of the Trebouxiophyceae propagate mainly by autosporulation. In this manner, the mother cell wall is shed following division of the protoplast in each round of cell division. Binary fission type Nannochloris and budding type Marvania are also included in the Trebouxiophyceae. Phylogenetic trees based on the actin sequences of Trebouxiophyceae members revealed that the binary fission type Nannochloris bacillaris and the budding type Marvania geminata are closely related in a distal monophyletic group. Our results suggest that autosporulation is the ancestral mode of cell division in Trebouxiophyceae. To elucidate how non-autosporulative mechanisms such as binary fission and budding evolved, we focused on the cleavage of the mother cell wall. Cell wall development was analyzed using a cell wall-specific fluorescent dye, Fluostain I. Exfoliation of the mother cell wall was not observed in either N. bacillaris or M. geminata. We then compared the two algae by transmission electron microscopy with rapid freeze fixation and freeze substitution; in both algae, the mother cell wall was cleaved at the site of cell division, but remained adhered to the daughter cell wall. In N. bacillaris, the cleaved mother cell wall gradually degenerated and was not observed in the next cell cycle. In contrast, M. geminata daughter cells entered the growth phase of the next cell cycle bearing the mother and grandmother cell walls, causing the uncovered portion of the plane of division to bulge outward. Such a delay in the degeneration and shedding of the mother cell wall probably led to the development of binary fission and budding.
绿球藻纲的无鞭毛营养绿藻主要通过自体孢子形成进行繁殖。通过这种方式,在每一轮细胞分裂中,原生质体分裂后母细胞壁会脱落。绿球藻纲还包括二分分裂型的微绿球藻属和出芽型的马尔瓦藻属。基于绿球藻纲成员肌动蛋白序列构建的系统发育树表明,二分分裂型的纤细微绿球藻和出芽型的双生马尔瓦藻在一个远缘单系群中密切相关。我们的结果表明,自体孢子形成是绿球藻纲细胞分裂的原始模式。为了阐明二分分裂和出芽等非自体孢子形成机制是如何进化的,我们重点研究了母细胞壁的裂解。使用细胞壁特异性荧光染料荧光增白剂I分析细胞壁的发育。在纤细微绿球藻或双生马尔瓦藻中均未观察到母细胞壁的剥落。然后,我们通过快速冷冻固定和冷冻置换的透射电子显微镜对这两种藻类进行了比较;在这两种藻类中,母细胞壁在细胞分裂部位裂解,但仍附着在子细胞壁上。在纤细微绿球藻中,裂解的母细胞壁逐渐退化,在下一个细胞周期中未观察到。相比之下,双生马尔瓦藻的子细胞进入下一个细胞周期的生长阶段时,仍带有母细胞壁和祖母细胞壁,导致分裂平面未覆盖的部分向外凸出。母细胞壁退化和脱落的这种延迟可能导致了二分分裂和出芽的发展。