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Fig. 6. Evolutionary modifications in echinoderm skeletogenesis. Only
embryonic and larval stages are illustrated. Green cells indicate
non-skeletogenic mesoderm, red cells indicate skeletogenic mesoderm, and heavy
black lines represent skeletal rods. (A) The ancestral echinoderm
exhibited indirect development and had an adult skeleton. The embryo had an
ancestral program of mesoderm specification but lacked a skeleton. (B)
In the ancestral echinoid, the adult program of biomineralization was imported
into the late embryo. This pattern of skeletogenesis is still seen in modern
cidaroid sea urchins. (C) In modern euechinoids, a second heterochronic
change occurred, shifting the skeletogenic program into the early embryo. This
change was associated with the invention of micromeres and an
early-ingressing, skeletogenic mesenchyme (PMCs). It required the
establishment of new regulatory links between the ancestral skeletogenic GRN
and an even more ancient system of early patterning mediated by
β-catenin. It was also associated with the invention of a PMC-derived
signal that suppresses the skeletogenic potential of NSM cells (white
arrows).