Environmental factors affecting reproduction and development in ascidians and other protochordates

Protochordate reproduction and development are influenced by many kinds of environmental factors. For example, spawning, sexual and asexual reproduction, larval behaviour, and life-cycle transitions (metamorphosis) are key processes known to be affected by environmental factors. This review must be restricted primarily to only one group of protochordates, the ascidians or "sea squirts", because information on the reproductive ecology of hemichordates and cephalochordates is limited to only a few studies. Topics discussed in the present review include (i) environmental factors that regulate larval settlement, (ii) how pelagic embryos avoid damage to DNA caused by UV radiation, (iii) the effect of water temperature and food availability on sexual reproduction in colonial ascidians, (iv) environmental regulation of asexual budding, (v) environmental regulation of metamorphosis, and (vi) the possible role of the environment in the evolution of direct-developing ascidians. A novel role for HSP90 and nitric oxide signaling in the integration of environmental factors with cell signaling pathways in ascidians is discussed near the end of this review. Throughout this review, the multiple roles of environmental stress on ascidian reproduction and development are emphasized.