The origin of the tetrapod limb: from expeditions to enhancers.

More than three centuries ago natural philosophers, and later anatomists, recognized a fundamental organization to the skeleton of tetrapod limbs. Composed of three segments, stylopod, zeugopod, and autopod, this pattern has served as the basis for a remarkably broad adaptive radiation from wings and flippers to hands and digging organs. A central area of inquiry has been tracing the origins of the elements of this Bauplan in the fins of diverse fish. Can equivalents of the three segments, and the developmental processes that pattern them, be seen in fish fins? In addition, if so, how do these data inform theories of the transformation of fins into limbs? Answers to these questions come from linking discoveries in paleontology with those of developmental biology and genetics. Burgeoning discoveries in the regulatory biology of developmental genes and in the genomics of diverse species offer novel data to investigate these classical questions.