Dev Growth Differ 1988 Oct 01;305:543-552. doi: 10.1111/j.1440-169X.1988.00543.x.

Timers in Early Development of Sea Urchin Embryos: (sea urchin/timing mechanism/temperature dependency/mid-blastula transition/cytoplasmic clock).

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To elucidate the timing mechanisms in the early development of sea urchin embryos, we measured the times of initiation of the first four cleavages, of ciliary movement, of primary mesenchyme cell ingression, and of gastrulation at four temperatures ranging from 11 to 20°C. The times of cleavage and of initiation of ciliary movement showed similar temperature dependency, indicating that these events may be controlled by a common timer (the first timer). Although batches of eggs often showed variation in the period between fertilization and the first cleavage, their subsequent cleavages were more regular. This indicates that the first timer may not start at fertilization. The ingression of mesenchyme cells and the onset of gastrulation showed similar temperature dependency that was higher than that of other events, suggesting the existence of a second timer. Temperature shift experiments indicate that the second timer starts at the mid-blastula (the 8-9th cleavage) stage when divisions of blastomeres become asynchronous.

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