Developmental plasticity, modularity, and heterochrony during the phylotypic stage of the zebra fish, Danio rerio.

We studied early embryonic development of zebra fish and tested if changes in the external raising conditions could elicit phenotypic changes during the phylotypic stage which, classically, is considered as a conserved embryonic stage. In particular, we tested for internal constraints, plasticity, and heterochrony during the early embryonic development. Our tested hypotheses predict (i) no change associated with developmental stability/internal constraints, (ii) change of the rate of development associated with developmental flexibility, and (iii) heterochronic disruption of developmental pattern associated with a modular organization of the embryo. We measured 14 traits of embryos raised in different conditions (temperature, salinity, oxygen concentration). The results of our study show that zebra fish embryos respond flexibly to changes in external parameters even during the conserved "phylotypic stage." It also showed that internal constraints canalize early development when exposed to moderate external challenges. Hypoxic conditions, however, elicited a heterochronic delay of the onset of the development of the Anlagen of the eye and the otic vesicle from the remaining embryo. Therefore, we concluded that the eye and the otic vesicle are modules that may develop, to a certain degree, independently of the rest of the embryo. Because these modules become recognizable only under specific raising conditions, we suggest that the modularization acts as buffering mechanism against extreme developmental deviations. Our results provide support to the idea that modularity is present during the phylotypic stage, but it is not effective under normal conditions.