Model systems for environmental signaling.

Studies of environmental signaling in animals have focused primarily on organisms with relatively constrained responses, both temporally and phenotypically. In this regard, existing model animals (e.g., "worms and flies") are particularly extreme. Such animals have relatively little capacity to alter their morphology in response to environmental signals. Hence, they exhibit little phenotypic plasticity. On the other hand, basal metazoans exhibit relatively unconstrained responses to environmental signals and may thus provide more general insight, insofar as these constraints are likely traits derived during animal evolution. Such enhanced phenotypic plasticity may result from greater sensitivity to environmental signals, or greater abundance of suitable target cells, or both. Examination of what is known of the components of environmental signaling pathways in cnidarians reveals many similarities to well-studied model animals. In addition to these elements, however, macroscopic basal metazoans (e.g., sponges and cnidarians) typically exhibit a system-level capability for integrating environmental information. In cnidarians, the gastrovascular system acts in this fashion, generating local patterns of signaling (e.g., pressure, shear, and reactive oxygen species) via its organism-wide functioning. Contractile regions of tissue containing concentrations of mitochondrion-rich, epitheliomuscular cells may be particularly important in this regard, serving in both a functional and a signaling context. While the evolution of animal circulatory systems is usually considered in terms of alleviating surface-to-volume constraints, such systems also have the advantage of enhancing the capacity of larger organisms to respond quickly and efficiently to environmental signals. More general features of animals that correlate with relatively unconstrained responses to environmental signals (e.g., active stem cells at all stages of the life cycle) are also enumerated and discussed.