Morphomechanical rules of embryonic development.

We start by comparing two explicatory approaches, which we call law-centered and "instructivistic". Although the latter dominates in modern biology, we find it inappropriate for treating developmental problems, especially those related to morphogenesis. As an example of a law-centered approach we suggest a simple morphomechanical rule based upon the idea of the hyper-restoration of mechanical stresses.

Morphomechanical reactions and mechanically stressed structures in amphibian embryos, as related to gastrulation and axial organs formation.

Reactions of embryonic tissues to a distributed and concentrated stretching are described and compared with the mechanics of the normal gastrulation movements. A role of mechanically stressed dynamic cell structures in the gastrulation, demarcation of notochord borders and in providing proportionality of the axial rudiments is demonstrated. A morphomechanical scheme of amphibian gastrulation is presented.

Local and global dynamics in collective movements of embryonic cells.

Several important morphogenetic processes belong to the category of collective cell movements (CCM), by which we mean coordinated rearrangements of many neighboring cells. The causes of the dynamic order established during CCM are still unclear. We performed statistical studies of rates and angular orientations of cell rearrangements in two kinds of embryonic tissues, which we categorized as "committed" (in the sense of being capable of autonomous CCM) as opposed to "naïve" tissues, which are those that require external forces in order to exhibit full scale CCM.

Stress-generating tissue deformations in Xenopus embryos: Long-range gradients and local cell displacements.

Background: Although the role of endogenous mechanical stresses in regulating morphogenetic movements and cell differentiation is now well established, many aspects of mechanical stress generation and transmission in developing embryos remain unclear and require quantitative studies.

Results: By measuring stress-bearing linear deformations (caused by differences in cell movement rates) in the outer cell layer of blastula - early tail-bud Xenopus embryos, we revealed a set of long-term tension-generating gradients of cell movement rates, modulated by short-term cell-cell displacements much increasing the rates of local deformations. Experimental relaxation of tensions distorted the gradients but preserved and even enhanced local cell-cell displacements.

Two ways for interpreting Driesch's law: "Positional information" and morphogenetic fields.

The late Lev V. Beloussov wrote a 2005 textbook The Foundations of General Embryology which is available in Russian. In 2003 he prepared an excellent, annotated translation of the part of his manuscript for this book on distinguishing positional information models from morphogenetic field models of embryogenesis, which is reproduced here verbatim.

[On the Work of the Developmental Biophysics Laboratory of the Embryology Department of Moscow State University].

The laboratory is engaged in morphomechanics—the study of self-organization of mechanical forces that create the shape and structure of the embryonic primordia. As part of its work, the laboratory described pulsating modes of mechanical stresses in hydroids, identified and mapped mechanical stresses in the tissues of amphibian embryos, and studied morphogenetic reorganization caused by the relaxation and reorientation of tensions. The role of mechanical stresses in maintaining the orderly architectonics of the embryo is shown.

Revisiting the mitogenetic effect of ultra-weak photon emission.

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.

Morphogenesis can be driven by properly parametrised mechanical feedback.

A fundamental problem of morphogenesis is whether it presents itself as a succession of links that are each driven by its own specific cause-effect relationship, or whether all of the links can be embraced by a common law that is possible to formulate in physical terms. We suggest that a common biophysical background for most, if not all, morphogenetic processes is based upon feedback between mechanical stresses (MS) that are imposed to a given part of a developing embryo by its other parts and MS that are actively generated within that part. The latter are directed toward hyper-restoration (restoration with an overshoot) of the initial MS values.

A simple model for estimating the active reactions of embryonic tissues to a deforming mechanical force.

Active reactions of embryonic tissues to mechanical forces play an important role in morphogenesis. To study these reactions, experimental models that enable to evaluate the applied forces and the deformations of the tissues are required. A model based upon the active intrusion of a living early gastrula Xenopus embryo into a tube half the embryo in diameter is described.

Morphogenesis as a macroscopic self-organizing process.

We start from reviewing different epistemological constructions used for explaining morphogenesis. Among them, we explore the explanatory power of a law-centered approach which includes top-down causation and the basic concepts of a self-organization theory. Within such a framework, we discuss the morphomechanical models based upon the presumption of feedbacks between mechanical stresses imposed onto a given embryo part from outside and those generated within the latter as a kind of active response.

Active reinforcement of externally imposed folding in amphibians embryonic tissues.

Although the folding of epithelial layers is one of the most common morphogenetic events, the underlying mechanisms of this process are still poorly understood. We aimed to determine whether an artificial bending of an embryonic cell sheet, which normally remains flat, is reinforced and stabilized by intrinsic cell transformations. We observed both reinforcement and stabilization in double explants of blastocoel roof tissue from Xenopus early gastrula embryos.

Nomothetics and idiography in developmental biology.

Successions of space-temporal structures arisen during development of multicellular organisms are the most regular, complex and reproducible ones among all taking place in the entire nature without a human's intervention. Therefore, the question whether it would be possible to embrace them by a common physicalistic law (nomothetic approach) or they can be only enumerated and described one after another (idiography) is of an overall importance for the natural sciences in general. We review several nomothetic attempts performed in XX century biology and suggest that such laws may have a structure of feedback contours between the active and passive mechanical stresses generated in developing embryos.

Neuro-mesodermal patterns in artificially deformed embryonic explants: a role for mechano-geometry in tissue differentiation.

The mutual arrangement of neural and mesodermal rudiments in artificially bent double explants of Xenopus laevis suprablastoporal areas was compared with that of intact explants. While some of the bent explants straightened or became spherical, most retained and actively reinforced the imposed curvature, creating folds on their concave sides and expanding convex surfaces. In the intact explants, the arrangement of neural and mesodermal rudiments exhibited a distinct antero-posterior polarity, with some variability.

Relocations of cell convergence sites and formation of pharyngula-like shapes in mechanically relaxed Xenopus embryos.

Influence of the relaxation of mechanical tensions upon collective cell movements, shape formation, and expression patterns of tissue-specific genes has been studied in Xenopus laevis embryos. We show that the local relaxation of tensile stresses within the suprablastoporal area (SBA) performed at the early-midgastrula stage leads to a complete arrest of normal convergent cell intercalation towards the dorsal midline. As a result, SBA either remains nondeformed or protrudes a strip of cells migrating ventralwards along one of the lateral lips of the opened blastopore.

Mechanically based generative laws of morphogenesis.

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.

Information about a form (on the dynamic laws of morphogenesis).

How a developing embryo becomes "informed" about its form?" This problem remains obscure and controversial. We argue that the "information about a form" is distributed throughout three main components: the dynamic laws, the parameters and the initial/boundary conditions. In the absence of a dynamic law two other components are "blind", that is, do not contain any unambiguous information.

Gastrulation in amphibian embryos, regarded as a succession of biomechanical feedback events.

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. 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.

Morphomechanics: goals, basic experiments and models.

Morphomechanics is a branch of developmental biology, studying the generation, space-time patterns and morphogenetic role of mechanical stresses (MS) which reside in embryonic tissues. All the morphogenetically active embryonic tissues studied in this respect have been shown to bear substantial mechanical stresses of tension or pressure. MS are indispensable for organized cell movements, expression of a number of developmentally important genes and the very viability of cells.

A common biomechanical model for the formation of stationary cell domains and propagating waves in the developing organisms.

Many important morphogenetic processes that take place in the development of an animal start from the segregation of a homogeneous layer of cells into a different number of the domains of columnar and flattened cells. In many cases, waves of cell shape transformation travel throughout embryonic tissues. A biomechanical model is presented which embraces both kinds of event.

Biophotonic patterns of optical interactions between fish eggs and embryos.

The optical (non-substantial) interactions between various biological samples have been evident in a number of cases mainly by the effects on their functional activity and developmental patterns. However, the mechanisms of these interactions have remained obscure. Effect of optical interaction has been observed on the intensity and Fourier patterns of biophoton emission of fish embryos.

Formative capacities of mechanically stressed networks: developmental and evolutionary implications.

We present a biomechanical model of morphogenesis highlighting the extensive formative capacities of stressed networks with a very simple initial geometry. They consist of a restricted number of kinematically independent elements exerting a pressure to each other and increasing thus the local curvatures. The pressure is applied as a series of periodic impulses and is opposed by a constant quasi-elastic resistance force.

Integrating self-organization theory into an advanced course on morphogenesis at Moscow State University.

A lecture course on morphogenesis for fourth-year Moscow State University Specialist Diploma students specializing in embryology is described. The main goal of the course is to give the students an extensive theoretical background based on the tenets of the modern theory of Self-Organization and to show them how important this theory is for the proper understanding of developmental events. The corresponding mathematics are bound as tightly as possible to the actual morphogenetic processes.

A Geometro-mechanical model for pulsatile morphogenesis.

A model is proposed which imitates the morphogenesis of several species of the lower invertebrate animals, the hydroid polyps and permits the derivation of the geometry (surface curvature) of each developmental stage from that of the preceding stage. The model is based upon two experimentally verified assumptions. First, neighbouring cells are assumed to compress each other laterally in a regular and species-specific pulsatile manner.