Retinoids and pattern formation in a hydroid.

The retinoids (retinol, retinal, retinoic acid) cause alterations in the pattern of limb elements in vertebrates (Summerbell & Harvey, 1983). As shown here, retinoids also influence pattern specification in hydroid polyps (Hydractinia echinata) in a way suggesting interference with the generation and transmission of signals responsible for the dimension and spacing of structures. A pulse-type application of low doses (e.g. retinoic acid 10(-6) to 10(-10) M, 4 h) causes metamorphosing primary polyps to develop more tentacles but fewer stolons per unit circumference, to shorten the length of the hydranth while the stolon elongates, and to bud secondary hydranths at high frequency 2-3 days after treatment (Fig. 3). Dose-response curves display optimum peaks. It is argued that the increase in budding rate is due to a reduction of the range of spacing signals emitted by the primary hydranth. In regenerating hydranths, low doses (10(-10) to 10(-9) M) improve the rate of head formation, whilst medium doses (10(-8) to 10(-6) M) result in more tentacles being regenerated. However, prolonged treatment with high doses (10(-6) to 10(-5) M) causes the animals to reduce all head structures and to transform eventually into stolons, in contravention of the rule of distal transformation that they normally obey (Fig. 8). The effects of the retinoids are counteracted by a putative morphogen, the endogenous inhibitor isolated from Hydra by Berking (1977). The Hydra-derived 'head-activator' displayed no stimulating effect on the number of tentacles and buds formed.