A deep (although at the first glance naïve) question which may be addressed to embryonic development is why during this process quite definite and accurately reproduced successions of precise and complicated shapes are taking place, or why, in several cases, the result of development is highly precise in spite of an extensive variability of intermediate stages. This problem can be attacked in two different ways. One of them, up to now just slightly employed, is to formulate robust macroscopic generative laws from which the observed successions of shapes could be derived. Another one, which dominates in modern embryology, regards the development as a succession of highly precise 'micropatterns', each of them arising due to the action of specific factors, having, as a rule, nothing in common with each other. We argue that the latter view contradicts a great bulk of firmly established data and gives no satisfactory answers to the main problems of development. Therefore we intend to follow the first way. By doing this, we regard developing embryos as self-organized systems transpierced by feedbacks among which we pay special attention to those linked with mechanical stresses (MS). We formulate a hypothesis of so-called MS hyper-restoration as a common basis for the developmentally important feedback loops. We present a number of examples confirming this hypothesis and use it for reconstructing prolonged chains of developmental events. Finally, we discuss the application of the same set of assumptions to the first steps of egg development and to the internal differentiation of embryonic cells.
The movements of blastomere surfaces marked with carbon particles during cytokinesis of the Ist-IVth cleavage divisions in the eggs of the gastropodsLymnaea stagnalis, L. palustris, Physa acuta and Ph. fontinalis have been studied by time-lapse cinematographic methods. The vitelline membrane was removed with trypsin. At 2- and 4-cell stages shifts of nuclei have also been studied.Symmetrical as well as asymmetrical surface movements (in respect to the furrow plane) have been revealed. Symmetrical surface movements at the beginning of cytokinesis consist mainly in contraction of the furrow zone and in expansion of the more peripheral regions; between these there is a stationary zone. After the end of cytokinesis the furrow region expands.Considerableasymmetrical surface movements have also been observed in all four divisions. From anaphase until the end of cytokinesis each of the two sister blastomeres rotates with respect to the other in such a way, that if viewed along the spindle axis, the blastomere nearest to the observer rotates dexiotropically in a dextral species and laeotropically in a sinistral species (primary rotations). After the completion of cytokinesis the blastomeres may rotate in a reverse direction. The latter rotations are less pronounced in the IInd and IIIrd divisions and most pronounced in the IVth division. Blastomeres with the vitelline membrane intact retain a slight capacity for primary rotations. In normal conditions nuclei of the first two blastomeres shift mainly laeotropically in dextral species, but dexiotropically in sinistral species, being carried along by the reverse surface rotations.The invariable primary asymmetrical rotations of blastomeres seem to be the basis of enantiomorphism in molluscan cleavage. They are assumed to be determined by an asymmetrical structure of the contractile ring carrying out the cytokinesis.
Ventral ectodermal explants taken from early gastrula embryos of Xenopus laevis were artificially stretched either by two opposite concentrated forces or by a distributed force applied to the internal explant's layer. These modes of stretching reflect different mechanical situations taking place in the normal development. Two main types of kinematic response to the applied tensions were detected. First, by 15 min after the onset of concentrated stretching a substantial proportion of the explant's cells exhibited a concerted movement towards the closest point of the applied stretching force. We define this movement as tensotaxis. Later, under both concentrated and distributed stretching, most of the cell's trajectories became reoriented perpendicular to the stretching force, and the cells started to intercalate between each other, both horizontally and vertically. This was accompanied by extensive elongation of the outer ectodermal cells and reconstruction of cell-cell contacts. The intercalation movements led first to a considerable reduction in the stretch-induced tensions and then to the formation of peculiar bipolar "embryoid" shapes. The type and intensity of the morphomechanical responses did not depend upon the orientation of a stretching force in relation to the embryonic axes. We discuss the interactions of the passive and active components in tension-dependent cell movements and their relations to normal morphogenetic events.
This paper reviews the 90 years long controversial history of the so-called “mitogenetic radiation,” the first case of non-chemical distant interactions, reported by Gurwitsch (1923). It was soon described as ultraweak UV, emitted by a number of biological systems, and stimulating mitosis in “competent” (in this sense) cells. In the following 20 years this phenomenon attracted enormous interest of the scientific community, and gave rise to more than 700 publications around the world. Yet, this wave of research vanished after several ostensibly disproving works in late 1930-s, and was not resumed later, regardless of quite serious grounds for that. The authors discuss separately two aspects of the problem: (1) do living organisms emit ultraweak radiation in the UV range (irrespective of whether it has any biological role), and (2) are there any real effects of this ultraweak photon emission (UPE) upon cell division and/or other biological functions? Analysis of the available data permits to conclude, that UV fraction of UPE should be regarded real, while its biological effects are difficult to reproduce. This causes a paradox. A number of presently known qualities of UPE were initially discovered (predicted?) by the “early workers” on the basis of biological effects. Yet the qualities they discovered were proved later (the UV component of UPE, the sources of UPE among biological systems, etc…), while the biological effect they used for UPE “detection” remains questionable. Importance of this area for basic biology and medicine, and potential usefulness of UPE as a non-invasive research method, invite scientists to attack this problem again, applying powerful research facilities of modern science. Yet, because of complexity and uncertainty of the problem, further progress in this area demands comprehensive examination of both positive and negative works, with particular attention to their methodical details.
Gastrulation in amphibian embryos is a composition of several differently located morphogenetic movements which are perfectly coordinated with each other both in space and time. We hypothesize that this coordination is mediated by biomechanical interactions between different parts of a gastrulating embryo based upon the tendency of each part to hyper-restore the value of its mechanical stress (see Beloussov and Grabovsky, 2006). The entire process of gastrulation in amphibian embryos is considered as a chain of these mutually coupled reactions, which are largely dependent upon the geometry of a given embryo part. We divide gastrulation into several partly overlapped steps, give a theoretical interpretation for each of them, formulate the experiments for testing our interpretation and describe the experimental results which confirm our point of view. Among the predicted experimental results are: inhibition of radial cell intercalation by relaxation of tensile stresses at the blastula stage; inversion of convergent intercalation movements by relaxation of circumferential stresses at the early gastrula stage; stress-promoted reorientation of axial rudiments, and others. We also show that gastrulation is going on under a more or less constant average value of tensile stresses which may play a role as rate-limiting factors. A macromorphological biomechanical approach developed in this paper is regarded as complementary to exploring the molecular machinery of gastrulation.
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