Earliest ciliary swimming effects vertical transport of planktonic embryos in turbulence and shear flow

Eggs released by broadcast-spawning marine invertebrates are often negatively buoyant. Blastulae and gastrulae of these species are commonly motile, with passive stability that leads to upward swimming in still water. The earliest occurrence of swimming in developing embryos of diverse invertebrates may therefore permit vertical migration in nature. I used turbulent and laminar shear flows to investigate: (1) the speed and direction of transport of non-motile and newly swimming stages of the echinoids Dendraster excentricus and Strongylocentrotus purpuratus in turbulence, and (2) the limit of stable vertical orientation in swimming blastulae of D. excentricus. Swimming contributed significantly to the rate of upward transport of D. excentricus in turbulence experiments where the kinetic energy dissipation rate (ε) was ∼10(-2) cm(2) s(-3). However, swimming significantly reduced the rate of upward transport of S. purpuratus blastulae in turbulence, suggesting that passively stable swimmers of this species were turned from the vertical, crossed flow-lines, and migrated into downwelling. Observations of swimming in laminar shear indicate that D. excentricus swimming blastulae maintain a vertical orientation until shear approaches 0.26 s(-1), equivalent to sub-microscale shear in turbulence where ε is ∼10(-3) cm(2) s(-3). Swimming speeds of D. excentricus showed an unexpected dependence on shear, indicating that greater shear (within limits) can enhance speed of ciliary swimming. In D. excentricus, swimming by newly hatched blastulae should support upward migration in turbulence characteristic of coastal surface waters, whereas species differences in passive stability and swimming responses to shear may lead to differences in vertical transport and subsequent dispersal.