Early development of the neural plate, neural crest and facial region of marsupials

Marsupial mammals have a distinctive reproductive strategy. The young are born after an exceptionally short period of organogenesis and are consequently extremely altricial. Yet because they must be functionally independent in an essentially embryonic condition, the marsupial neonate exhibits a unique suite of adaptations. In particular, certain bones of the facial region, most cranial musculature and a few additional structures are accelerated in their development. In contrast, central nervous system structures, especially the forebrain, are markedly premature at birth, resembling an embryonic d 11 or 12 mouse. This review examines the developmental processes that are modified to produce these evolutionary changes. The focus is on the early development of the neural plate, neural crest and facial region in the marsupial, Monodelphis domestica, compared with patterns reported for rodents. Neural crest begins differentiation and migration at the neural plate stage, which results in large accumulations of neural crest in the facial region at an early stage of development. The early accumulation of neural crest provides the material for the accelerated development of oral and facial structures. The first arch region is massive in the early embryo, and the development of the olfactory placode and frontonasal region is advanced relative to the forebrain region. The development of the forebrain is delayed in marsupials relative to the hindbrain or facial region. These observations illustrate how development may be modified to produce evolutionary changes that distinguish taxa. Further, they suggest that development is not necessarily highly conserved, but instead may be quite plastic.