Claw asymmetry in lobsters: case study in developmental neuroethology.

An enduring debate in the study of development is the relative contribution of genetic and epigenetic factors in the genesis of an organism, that is, the nature vs. nurture debate. The behavior of the paired claws in the lobster offers promising material for pursuing this debate because of the way they develop. The paired claws and their closer muscles are initially symmetrical; both are slender in appearance and have a mixture of fast and slow fibers in their closer muscles. During a critical period of development, they become determined into a major (crusher) and minor (cutter) claw and during subsequent development acquire their final form and behavior: The crusher becomes a stout, molar-toothed claw capable of closing only slowly because its closer muscle has 100% slow fibers while the cutter becomes a slender, incisor-toothed claw capable of closing rapidly because its closer muscle has 90% fast fibers. Our initial hypothesis was that the more active claw became the crusher and its less active counterpart the cutter. Presumably, nerve activity would influence muscle transformation, which in turn would influence the exoskeleton to which they attach and hence claw morphology. Curtailing nerve activity to the claw prevented crusher development, while reflex activation of a claw promoted its development; both results support the notion that nerve activity directly regulates claw form and function. This is not, however, the case, for when both claws were reflexly exercised neither formed a crusher, signifying rather that bilateral differences in predominantly mechanoreceptive input to the paired claws somehow lateralized the claw ganglion [central nervous system (CNS)] into a crusher and cutter side. The side experiencing the greater activity becomes the crusher side while the contralateral side becomes the cutter and is also inhibited from ever becoming a crusher. This initial lateralization in the CNS is expressed, via as yet unknown pathways, at the periphery in claw morphology, muscle composition, and behavior. The critical period defines a time when the CNS is susceptible to being lateralized into a crusher and cutter side. Such lateralization is dependent upon experience of the environment in the form of mechanoreceptive input. In the absence of such experience, the CNS is not lateralized and paired cutter claws develop. Thus, while the critical period for crusher determination is genetically determined the actual trigger is influenced by experience.