Increasing complexity: Mechanical guidance and feedback loops as a basis for self-organization in morphogenesis

The article is devoted to physical views on embryo development as a combination of structurally stable dynamics and symmetry-breaking events in the general process of self-organization. The first corresponds to the deterministic aspect of embryo development. The second type of processes is associated with sudden increase of variability in the periods of symmetry-breaking, which manifests unstable dynamics. The biological basis under these considerations includes chemokinetics (a system of inductors, repressors, and interaction with their next surrounding) and morphomechanics (i.e. mechanotransduction, mechanosensing, and related feedback loops). Although the latter research area is evolving rapidly, up to this time the role of mechanical properties of embryonic tissues and mechano-dependent processes in them are integrated in the general picture of embryo development to a lesser extent than biochemical signaling. For this reason, the present article is mostly devoted to experimental data on morphomechanics in the process of embryo development, also including analysis of its limitations and possible contradictions. The general system of feedback-loops and system dynamics delineated in this review is in large part a repetition of the views of Lev Beloussov, who was one of the founders of the whole areas of morphomechanics and morphodynamics, and to whose memory this article is dedicated.