DEVELOPMENT
2291Here, I discuss selected examples of elongation in embryogenesis to identify common and unique mechanisms, useful questions for further work, and new systems that offer opportunities for answering these questions. Fiber-wound, hydraulic mechanisms of elongation highlight the importance of biomechanical linkages of otherwise unrelated cellular behaviors during elongation. Little-studied examples of elongation by cell intercalation offer opportunities to study new aspects of this mode of elongation. Elongation by oriented cell division highlights the problem of mitotic spindle orientation and the maintenance of cell-packing patterns in anisotropic force environments. The balance of internal celladhesion and external traction forces emerges as a key issue in the formation of elongate structures from compact ones by directed migration.
IntroductionThe elongation of tissues plays a major role in embryogenesis and organogenesis. Well-known examples include convergent extension movements that are driven by cell intercalation during vertebrate gastrulation, in the ascidian notochord, during Drosophila germ band extension, and during echinoderm gut elongation (Keller, 2002). However, most work has focused on the early development of a few systems, and convergent extension is only one mechanism of elongation. Here, other mechanisms of elongation, and less wellknown examples of convergent extension by cell intercalation that are instructive and have not received adequate attention, are discussed with the goal of identifying important, unexplored questions. The major conclusions are that specific outcomes of morphogenesis emerge from the global biomechanical integration of local, cellular force-generating processes. This integration follows the mechanical principles that are built in to each particular type of morphogenic machine. The challenge ahead is to integrate genetic and molecular manipulations with cell biological and biomechanical analyses to learn how genes encode the forces, and the cell and tissue material properties (Koehl, 1990) that transmit these forces, to generate the organized patterns essential for the heritable reproduction of form.
Fiber-wound, hydraulic systemsFiber-wound, hydraulic systems either maintain their shape or change it as a function of two interacting components: an external winding of restraining, tension-resisting fibers and an internal compression-resisting fluid under pressure (Koehl et al., 2000). The geometry of the fiber-windings determines the morphological changes that occur when the internal pressure is increased. Two examples will be discussed here. The first is the vertebrate notochord, in which a swelling of vacuoles increases the pressure, and the mechanical output is determined by the angle of orientation of extracellular matrix (ECM) fibrils. The second is the nematode embryo, in which the fiber-windings themselves contract to cause an increase in pressure.
Elongation and straightening of the notochordAfter participating in convergent extension during gastru...