Claw transformation and regeneration in adult snapping shrimp: test of the inhibition hypothesis for maintaining bilateral assymetry

In the paired asymmetric claws of adult snapping shrimp, Alpheus heterochelis, the minor, or pincer, claw may transform into a major, or snapper, claw if the existing snapper claw is damaged or lost, implying that an intact snapper claw normally inhibits the contralateral pincer claw from advancing to a snapper. We find that the pincer-to-snapper advancement in external form occurs almost immediately after the snapper is lost even as late as the premolt stage. The transforming claw in turn inhibits the newly regenerating pincer claw from becoming a snapper, but if the dactyl of the transforming claw is cut, then snapper-based inhibition is removed and the contralateral claw may regenerate as a snapper, resulting in shrimp with paired snapper claws. However, damaging an established snapper claw will not allow another snapper claw to regenerate at the pincer site, implying that less inhibition is required to restrict a newly regenerating claw to a pincer than to arrest an existing pincer claw. Inhibition may be manifested largely in terms of quantity of innervation. Hence the greater innervation of the snapper side over the pincer side would inhibit the pincer side, accounting for the regeneration of paired claws in their previous configuration following loss of both claws. Loss of the paired claws in two consecutive molts retards their development so that both claws often appear as pincers, but in succeeding molts one usually differentiates into a snapper and bilateral asymmetry is restored. In contrast, shrimp with paired snapper claws retain this configuration over several molts unless one or both of the claws are lost; in that case, regeneration restores bilateral asymmetry. Thus, bilateral asymmetry of the paired claws of adult shrimp is governed by a strong intrinsic lateralizing mechanism in which the snapper claw inhibits the pincer from advancing to another snapper.