Measuring the multi-scale integration of mechanical forces during morphogenesis

The elaborate changes in morphology of an organism during development are the result of mechanical contributions that are a mixture of those generated locally and those that influence from a distance. We would like to know how chemical and mechanical information is transmitted and transduced, how work is done to achieve robust morphogenesis and why it sometimes fails. We introduce a scheme for separating the influence of two classes of forces. Active intrinsic forces integrate up levels of scale to shape tissues. Counter-currently, extrinsic forces exert influence from higher levels downwards and feed back directly and indirectly upon the intrinsic behaviours. We identify the measurable signatures of different kinds of forces and identify the frontiers where work is most needed.