Heterochrony and the development of the escape response: Prehatching movements in the rainbow trout Oncorhynchus mykiss

Teleost fishes produce coordinated escape responses (C-starts) at hatching. This implies that essential swimming morphologies and motor behaviors develop during the incubation interval while the embryo is in the chorion. We examined prehatching motor behaviors in rainbow trout Oncorhycus mykiss (considered morphologically mature at hatching) and compared this species with zebrafish Danio rerio (considered morphologically immature) and assessed two hypotheses concerning the development of escape behavior. (1) Escape behavior is associated with the formation of key elements of the musculoskeletal and nervous systems; thus, the escape response appears early in ontogeny, when these elements form. (2) Escape behavior is not directly associated with the formation of underlying morphological elements; instead, it appears at hatching (i.e. when needed). We find that rainbow trout, like zebrafish, respond to touch early in the incubation interval, but do not demonstrate a complete C-start (including the second, propulsive stage) until shortly before hatching. At hatching, rainbow trout and zebrafish are similar in the degree of development of the chondocranium, paired fins and visceral arches (which comprise the larval jaw and gill support); however, rainbow trout have incipient rays in their unpaired fins (dorsal, anal and caudal), whereas zebrafish retain the embryonic fin fold. Although rainbow trout are more mature in axial swimming morphology at hatching, the essential neural and musculoskeletal systems that produce a coordinated escape response are functional at hatching in both species. This finding supports the evolutionary hypothesis that an effective escape response is critical for the survival of newly hatched teleost fishes.