Wolf Rainer, Krause Gerhard
Heiligenberg-Institut für experimentelle Biologie, Heiligenberg, Baden.
Zoologisches Institut I der Universität Würzburg, Deutschland.
Wilhelm Roux Arch Entwickl Mech Org. 1971 Sep;167(3):266-287. doi: 10.1007/BF00584253.
In the eggs ofPimpla turionellae, which are characterized by a long germ anlage ("long-germ egg" type), the cleavage nuclei primarily populate the anterior part and only later appear in the posterior of the egg lumen during the intravitelline cleavage. Gastrulation and segmentation also start within this anterior region. Time-lapse motion pictures served to observe and to check quantitatively even slow movements during cleavage and blastogenesis. In motion diagrams made by means of microkymographic technics the flow within the ooplasm along the longer axis of the egg has been timed.Shortly before the first cleavage in thestrictly unfertilized male eggs a short-time"unipolar flow" sets in from a primary initial region at 90% of their length. Thus a pillar of "central plasm" between both of the poles becomes shifted towards the posterior, while its outer coating layer of "marginal-plasm" is displaced forwards by the same distance. In eggs from fertilized females two successive flows of the same "unipolar" type have been observed.At the end of the third cleavage the energids, heretofore loosely grouped together, become distributed within the central plasm to form a "nuclear column". At the same time a fluently pulsatory "bipolar flow" sets in, within asecondary initial region at 80% of the egg length. Comparable to two mirror-image fountains, parts of the central plasm are carried towards the front pole and to the rear pole of the egg, respectively, while the marginal plasm, together with the oolemma, flows in opposite directions at times. With each pulsation the moving areas of the bipolar flow are shifted more and more towards the egg poles. The occurrence of bipolar flow pulsations, amounting to five, is correlated with the nuclear divisions in a still unknown way. In the rhythm of the bipolar flow, the energids become dispersed within the central plasm with a certain spatial lagging.After the bipolar flow has come to a halt, four further cleavages are indicated by faint local pulsations of the ooplasm. The cleavage nuclei move to the egg surface and pole cells become separatedtied off During blastoderm formation another four faint pulsations are observed, especially within the central ooplasm, all of them clearly synchronized with superficial cleavages. Occurring in mitotic waves, these cleavages indicate a third initial region, with the individual position varying between 10 and 28% of the egg length.Furthermore the technics of time-lapse motion pictures permit a local and temporal determination of extravitelline pole space formation, of a ring-shaped contracted region of slightly thickening periplasm within the secondary initial region, and the dislocation of the oosome towards the egg surface, which results from the activity of the posterior fountain during the phase of bipolar flow. Invagination and segmentation of the embryo become distinct within the secondary initial region, thus identifying this region as a differentiation centre.The correlation of plasm flow and nuclear divisions is discussed as well as the correlation of the initial regions to the different patterns of egg architecture in the longgerm egg type. The correlation between bipolar pulsations and the development of the metameric pattern including the function of the oosomal region is also discussed. The ooplasmic movements as known from egg types other thanPimpla are compared to the above observations.
在以长胚盘(“长胚卵”类型)为特征的图氏肿腿蜂卵中,卵裂核主要分布在卵的前部,在卵黄内卵裂期间,只是后来才出现在卵腔的后部。原肠胚形成和分节也在这个前部区域开始。延时动态影片用于观察和定量检查卵裂和胚泡形成过程中即使是缓慢的运动。借助显微记波技术制作的动态图中,已对卵质沿卵长轴的流动进行了计时。在严格未受精的雄卵第一次卵裂前不久,从其长度90%处的一个初级起始区域开始出现短时间的“单极流动”。于是,两极之间的“中央质”柱向后移动,而其“边缘质”的外层则向前移动相同的距离。在受精雌蜂的卵中,观察到了两次连续的相同“单极”类型的流动。在第三次卵裂结束时,此前松散聚集在一起的卵裂球,在中央质内分布形成一个“核柱”。与此同时,在卵长80%处的一个次级起始区域内开始出现平稳脉动的“双极流动”。类似于两个镜像喷泉,中央质的部分分别被带向卵的前极和后极,而边缘质与卵膜有时则以相反方向流动。随着每次脉动,双极流动的移动区域越来越向卵极移动。双极流动脉动出现了五次,其与核分裂以一种尚不清楚的方式相关。在双极流动的节律中,卵裂球在中央质内以一定的空间滞后分散开来。双极流动停止后,卵质微弱的局部脉动表明又进行了四次卵裂。卵裂核移向卵表面,极细胞分离出来。在胚盘形成过程中,又观察到四次微弱的脉动,特别是在中央卵质内,所有这些脉动都与表面卵裂明显同步。这些卵裂以有丝分裂波的形式出现,表明存在第三个起始区域,其个体位置在卵长的10%至28%之间变化。此外,延时动态影片技术可以局部和定时地确定卵黄外极空间的形成、次级起始区域内稍厚的周质环形收缩区域以及卵核向卵表面的移位,后者是双极流动阶段后喷泉活动的结果。胚胎的内陷和分节在次级起始区域内变得明显,从而将该区域确定为一个分化中心。讨论了质流与核分裂的相关性,以及起始区域与长胚卵类型中不同卵结构模式的相关性。还讨论了双极脉动与包括卵核区域功能在内的体节模式发育的相关性。将图氏肿腿蜂以外的卵类型中已知的卵质运动与上述观察结果进行了比较。