Contractile connective tissue in crinoids

Active movements in animals are usually attributed to cellular protein engines, e.g., the actin-myosin system of muscle cells. Here we report the first evidence of an extracellular contractile connective tissue, which we have found in sea lilies and feather stars (Echinodermata, Crinoida). These marine animals have arm muscles that are antagonized, not by other muscles, but by ligaments consisting of extracellular fibrils interspersed with neuron-like cell processes. Contractile cells are lacking, yet these arm ligaments actively contracted upon stimulation. The ligaments stayed in a contracted condition even after the stimulus had stopped. The stresses generated were lower than those of typical skeletal muscles. Additional data from crinoid cirri, which lack muscles entirely, corroborate the hypothesis that the connective tissue of the ligaments is contractile.