title: [Causal mechanisms of nuclear movement and division during early cleavage stages in the egg of a gall midge,Wachtliella persicariae L.]
abstract: Between each mitotic cycle, the cleavage nuclei ofWachtliella move over long distances, thus populating the ooplasm within a short time. Besides being shifted passively by flowing pulses of the ooplasm, thenuclei are also migrating actively. The active movements are accompanied by such oscillations of yolk particles as are known from the eggs of other insects, too. For a closer analysis of these "quivering movements" the inseminated eggs werepressed, either totally or partially, reducing their smaller diameters to ooplasmic layers of 30 μm or between 12 and 4 μm, respectively. Along with the experimental reduction of the radius of the curvature at the egg surface, there is anincreased tendency of cell membrane formation, resulting in anearly total cleavage already at the 2-nuclei-stage. Furthermore, theinitial region of cleavage (= initial region of quivering movements)may be shifted to a site free from nuclei; the initial region even may becomesplit up into two, one near each of the egg's poles. Yet, in flattened eggs, division and migration activities of the nuclei are not prevented. Untreated as well as flattened eggs have been analysed by means of time-lapse motion pictures taken either by the phase contrast or by the differential interference contrast method, using apochromatic objectives of maximum resolution, combined with an inverted microscope.According to the rhythm of the cleavage divisions,waves of irregular quivering movements spread from the initial region(s) of cleavage throughout the whole egg space. They are composed of irregular oscillations of yolk particles, probably caused by the effect of actively shortening, dynamic elements irregularly spread within the ooplasm. The presence ofcleavage nuclei obviously exerts a kind ofregulative effect: Shortly before such a wave of quivering movements reaches a metaphasic cleavage energide, regular oscillations and approximations of yolk particles are visible in the surroundings of the nucleus. The movements in question are radially adjusted towards the spindle poles, starting at the one which is reached first by the wave of quivering movements. These "radial quivering movements" are caused by abig cytaster, each originating from its spindle pole and distally reaching far into the ooplasm. Synchronous with the beginning of the shortening process of the astral rays, the cleavage nucleus passes through anaphase and telophase, and the spindle poles arepulled apart. During the then following migration of each daughter nucleus, its spindle pole-the kinetocentre of the previous spindle-is preserved and becomes the centre of a "migration cytaster". Its longest rays measure up to at least 80 μm. Their distal ends temporarily insert either in motile, or in elastically suspended, or in rigid egg components.By the recurrent short-time insertions and irregular shortening processes of the astral rays,the nucleus, displaying a strong affinity to its own kinetocentre,is pulled foreward. This movement always occurs in the direction of the biggest ooplasmic region still free from nuclei and therefore permitting the greatest spacial extension of the migration cytasters. This could explain the so-called mutual "repulsion" of the energides, leading to their even dispersion all over the egg space.In some of the eggs it has been possible toseparate the cleavage nuclei from their cytaster systems experimentally. Deprived of their nuclei such migration cytasters behave autonomously, i. e. they are actively moving within the ooplasm, possibly even retaining their division power. On the other hand, thenuclei without their cytasters have lost their mobility and therefore at first remain in their sites. But they seem to be capable ofinducing new spindle poles and migration cytasters of their own and to carry out further cleavage divisions.The migration cytasters of all cleavage energides develop by division from the very cytaster whose formation had been induced by the sperm nucleus after entering the egg. On the other hand thefemale pronucleus, remainingwithout a migration cytaster and therefore lacking migration activity, ismoved towards the male nucleus, pulled by the probably permanently inserted astral rays of the latter. Thus the final act of fertilization, i.e. nuclear fusion, comes about by the affinity between the (female pro-)nucleus and the (alien) migration cytaster (of the male nucleus).Judged by their derivation from the "polar rays" of the spindle apparatus, the astral rays with high probability are built up oftubuli, the evidence being left to electron microscopical investigations. Functional structures related to the causal mechanism of the migration cytaster are suggested and their supposed derivation from the mitotic apparatus is discussed. The existence of migration cytasters might not only represent an adaptation to the specific conditions of cleavage within spacious eggs, but also could be essential for the stretching of the spindle and the separation of the daughter nuclei during the division process of many other animal cells.
authors.affiliation: Heiligenberg-Institut für experimentelle Biologie, Heiligenberg, Baden.|Zoologisches Institut I der Universität Würzburg, Deutschland.
authors.forename: Rainer
authors.initials: R
authors.lastname: Wolf