Wolf Rainer
Zoologisches Institut I der Universität Würzburg, Deutschland.
Wilhelm Roux Arch Entwickl Mech Org. 1969 Jun;162(2):121-160. doi: 10.1007/BF00573537.
The experimental results ofGEYER-DUSZYNSKA (1959), speaking in favour of three ooplasmic factors localized in the pole plasm, in the basophilic oosome material contained therein, as well as in the periplasm of the posterior egg pole ofWachtliella persicariae, suggested to investigate for further factor regions with other technical means. Since ooplasmic factor regions may be indicated by initial regions of morphogenetic development, kinematics were used for in vivo analysis of early embryonic development by means of time-lapse motion pictures. Electron microscopic investigations added to the micro-morphological aspects of plasmic systems within the egg for a better understanding of nature and effectivity of ooplasmic factors.Cleavage nuclei do not move exclusively by means of their spindle activity during anaphase movement. The nuclear envelope of cleavage energides consists of either two unit membranes with pores or of many tube-like as well as membranous elements. The appearence of a complex multi-layered nuclear envelope coincides with the moving phase of energides, an observation which is discussed in relation to the possibility of active nuclear movement. During late preblastoderm the entoplasm contains horse-shoe-shaped and multilobed vitellophagues with dense karyoplasm. With the blastoderm formed, the nuclei may become pycnotic, their membranes fragmenting at the same time. These fragments probably are piled up to form annulated membranes.The pole plasm does not show specific structures apart from the oosome material, contained therein. It is free of yolk material and nearly exclusively consists of ground plasm. The basophilic oosome material within the pole plasm is not surrounded by any membranes. It consists of numerous ribosome-like units and is restricted to the plasm of the future pole cells. The micro structure of the oosome material is preserved at least till the germ band has reached its maximal length.The cell membranes develop by invagination of the oolemma which penetrates into the egg interior. While pole cells and blastoderm cells become tied off, the ground plasm possibly participates in the growing-in process of the cell membranes by developing fibrous differentiations at the terminal extensions of oolemma folds.There is no clear cut limitation between periplasm and entoplasm. The periplasm which is without yolk material, appears rich in ground plasm and does not contain specific ultra structures. During the process of cleavage external ooplasmic regions of the egg are shifted in rhythmical pulsation parallel to its longer axis by a maximum of about 6 % of the entire egg length. Topographic statements of certain areas concerning any anlage therefore are bound to suffer from an adequate lack of exactness. Since comparable shifting processes within the egg plasm probably are common in insects other thanWachtliella, they should be considered as a certain source of error.At 60+-3 % of egg length as measured from its posterior pole, there exists a cleavage centre, an initial region of intravitelline cleavage and of repeated mitotic waves. Adjacent to the middle of the egg follows an initial region of germ band formation (differentiation centre). By their electron microscopic appearance, both developmental centres are not characterized by specific ultra structures. The factor region at the posterior pole exclusively represents an initial region of cell wall formation during superficial cleavage.Other than any experimental marking procedure the technique of time-lapse motion pictures permits to evaluate quantitatively and without artificial interfering the shifting of presumptive segment material during morphogenetic movements of the germ band. The embryonic material of the blastoderm at the egg equator is used for building up the first abdominal segment. The prothoracal and mesothoracal material at about 60% of the egg length stays in site when the germ band becomes extended lengthwise. Closely behind the differentiation centre there is a region of maximal extension as well as of shortening of the germ band. No proliferous growth of segments (segment formation zone) has been found.
盖耶尔 - 杜辛斯卡(1959年)的实验结果表明,在极质、其中所含的嗜碱性卵质体物质以及波斯菊瓦氏线虫卵后极的周质中存在三种卵质因子,这促使人们用其他技术手段进一步研究因子区域。由于卵质因子区域可能由形态发生发育的起始区域指示,因此利用延时动态摄影技术对早期胚胎发育进行体内分析。电子显微镜研究补充了卵内质体系统的微观形态学方面,以便更好地理解卵质因子的性质和有效性。卵裂核在后期移动过程中并非仅通过纺锤体活动移动。卵裂能质的核膜由带有孔的两层单位膜或许多管状及膜状成分组成。复杂多层核膜的出现与能质的移动阶段一致,这一观察结果结合了核主动移动的可能性进行讨论。在胚盘形成前期后期,内质中含有马蹄形和多叶形的卵黄吞噬体,其核质致密。随着胚盘形成,细胞核可能会固缩,同时其膜会碎片化。这些碎片可能堆积形成环状膜。除了其中所含的卵质体物质外,极质没有显示出特定结构。它不含卵黄物质,几乎完全由基质质组成。极质内的嗜碱性卵质体物质没有被任何膜包围。它由许多核糖体样单位组成,且局限于未来极细胞的质中。卵质体物质的微观结构至少在胚带达到其最大长度之前得以保留。细胞膜通过卵膜内陷形成,卵膜向内深入卵内部。当极细胞和胚盘细胞分离时,基质质可能通过在卵膜褶皱的末端延伸处形成纤维分化而参与细胞膜的生长过程。周质和内质之间没有明显的界限。不含卵黄物质的周质似乎富含基质质,且不包含特定的超微结构。在卵裂过程中,卵的外部卵质区域以有节奏的脉动平行于其较长轴移动,最大移动幅度约为整个卵长度的6%。因此,关于任何原基的某些区域的地形描述必然会存在一定程度的不精确性。由于卵质内类似的移动过程可能在除波斯菊瓦氏线虫之外的昆虫中普遍存在,它们应被视为某种误差来源。从卵的后极测量,在卵长度的60%±3%处存在一个卵裂中心,这是卵黄内卵裂和反复有丝分裂波的起始区域。在卵的中部附近是胚带形成的起始区域(分化中心)。从电子显微镜外观来看,这两个发育中心都没有以特定的超微结构为特征。后极的因子区域仅代表表面卵裂过程中细胞壁形成的起始区域。与任何实验标记程序不同,延时动态摄影技术允许在不进行人工干扰的情况下定量评估胚带形态发生运动过程中假定节段物质的移动。卵赤道处胚盘的胚胎物质用于构建第一腹节。当胚带纵向延伸时,卵长度约60%处的前胸和中胸物质保持原位。在分化中心后方紧接着是胚带最大延伸以及缩短的区域。未发现节段的增殖生长(节段形成区)。