Risk and the evolution of cell-cycle durations of embryos

Embryos at low risk evolve slower development rates. In seven independent evolutionary contrasts for marine invertebrates (two in asteroids, three in gastropods, one each in phoronids and brachiopods) the more protected embryos had longer cell cycles from first to second cleavage than less protected planktonic embryos. Protected embryos had longer cell cycles even when protected eggs were smaller than planktonic eggs. In an eighth contrast, among tunicates, the embryonic cell cycle was unrelated to brooding and nearly proportional to egg size, but the literature provides examples of especially slow development in some brooding tunicates. The faster development of planktonic embryos is consistent with published estimates of greater mortality rates for planktonic larvae than for embryos in broods or egg masses. Examples from the literature for annelids, arthropods, holothuroids, and chordates also demonstrated longer embryonic cell cycles for more protected embryos with no consistent effect of egg size on cell-cycle duration. Longer cell cycles presumably reduce the benefits of protecting offspring because of longer exposure to whatever hazards remain, but slow development may permit compensating benefits. Hypothesized benefits of longer cell cycles include less maternal investment in rate-limiting materials, more or different transcription, and correction of errors. Such trade-offs are independent of feeding and growth and are influenced by parental protection.