Development of visceral smooth muscle.

The development of the smooth musculature of viscera has attracted the interest of only relatively few investigators, and thus the field appears somewhat underexplored. The major emphasis on histochemical evidence--at the expense of ultrastructural and functional studies--may have limited the progress in this area. Mature tissue is formed through the differentiation of precursors into muscle cells and through the organization of these cells into a complex tissue where distribution and orientation of muscle cells, deployment of abundant extracellular materials and addition of other cellular elements (interstitial cells, fibroblasts, nerves, blood vessels) are characteristic and specific features. The precursor cells are found at sites where a muscle develops, and they derive predominantly from the mesoderm, but also from the neuroectoderm and from the endoderm. The process starts at different times in different organs. The earliest stages of differentiation are characterized by the precursor cells aggregating and becoming elongated; their longitudinal axis lies in a position similar to the one they will have in the mature muscle. Both the cytological and the histochemical differentiation follow distinct patterns in various muscles, with characteristic temporal sequences in the appearance of key features. This process must impart distinct functional properties to a muscle cell at each stage of its development. However, the chronological correspondence between ultrastructural and histochemical development is poorly understood. Histochemical studies have detected gradients of maturation of the muscle cells, for example, across the thickness of the gizzard musculature and along the length of the small intestine; ultrastructural studies have not yet confirmed the existence of these gradients. Muscle growth is accounted for by muscle cell enlargement (without nucleus duplication) and an increase in muscle cell number by mitosis of pre-existing differentiated muscle cells. De-differentiation and division of muscle cells, migration of muscle cells and late development of muscle cell precursors have all also been considered as possible mechanisms for muscle growth. Several authors have described the presence of precursor cells within developing smooth muscles, and they have described late differentiation of some muscle cells or waves of differentiation that would give rise to phenotypic heterogeneity of the mature muscle cell population. In contrast, other studies, mainly by electron microscopy, have suggested that, within large visceral muscles, the muscle cells differentiate synchronously. There are interesting data on the influence of adjacent tissues on the development of a smooth muscle, but the interplay of these and other factors has not been fully investigated. Smooth muscles contract from early in their development, hence mechanical factors are likely to influence development: on the one hand, passive stresses imposed on the muscle by other tissues, such as adjacent muscles or the contents of the viscera and, on the other hand, active forces generated by the muscle itself. The very attraction of visceral smooth muscles in the study of cellular morphogenesis--an attraction that has not yet been highlighted or exploited in scientific studies, either descriptively or experimentally--is that, onto a single type of cell, a large range of factors interact, such as the genetic expression, chemical influences (from other muscles, endocrine glands, nerves, other intramuscular cells) and mechanical factors.