Carothers Z B, Kreitner G L
J Cell Biol. 1967 Apr;33(1):43-51. doi: 10.1083/jcb.33.1.43.
Spermatids of the liverwort Marchantia polymorpha were shown by analysis of thin-sections to contain a four-layered organelle, the Vierergruppe, lying between the flagellar bases above and a subjacent mitochondrial body. The uppermost layer consists of parallel tubules structurally similar to those of the axoneme. Some of the tubules extend beyond the four-layered region to form a long ribbon-like appendage. The second layer consists of low, vertical, membranous fins connected by numerous short links to the tubules above. The third layer, limited above and below by two horizontal membranes, consists of vertical partitions, apparently continuous with the fins. The fourth and lowermost layer consists of narrow parallel tubules aligned beneath alternate partitions. The long axes of the lower three layers are parallel to each other and lie at an angle of 48 degrees to the tubule axis of the uppermost stratum. A graphic representation of a three-dimensional scale model interpreting organelle structure is given. The Vierergruppe supersedes Dreiergruppe interpretation in most if not all of the bryophytes studied to date.
通过对切片的分析表明，地钱（Marchantia polymorpha）的精细胞含有一种四层细胞器，即四联体（Vierergruppe），位于上方的鞭毛基部和下方相邻的线粒体之间。最上层由结构上类似于轴丝的平行微管组成。一些微管延伸到四层区域之外，形成一条长长的带状附属物。第二层由低矮的垂直膜状鳍片组成，通过许多短连接与上方的微管相连。第三层由垂直隔板组成，上下由两个水平膜界定，显然与鳍片相连。第四层也是最底层由狭窄的平行微管组成，排列在交替隔板下方。下面三层的长轴相互平行，与最上层微管轴成48度角。给出了一个解释细胞器结构的三维比例模型的图形表示。在迄今为止研究的大多数（如果不是全部）苔藓植物中，四联体取代了三联体的解释。
