The Organization of the Muscular System of Metridium senile
I. The muscular system of Metridium consists of fields of relatively short muscle-fibres. In extension these may exceed I mm. in length but are only about 0.5µ thick. They can shorten to about a fifth of the extended length. The fibres consist almost entirely of densely staining material. They form a connected network. At least in some cases the cells seem to be in contact rather than to form syncytial connexions. 2. Deformation of the body-wall is in part controlled by the contractility of the muscle-fibres and in part by the properties of the mesogloea. Longitudinal contraction of the body-wall is accompanied by great thickening of the substance of the mesogloea. That part of the mesogloea which carries the circular muscle-fibres of the body-wall does not thicken. It buckles, thereby throwing the muscular layer into folds. Buckling occurs during the shortening of almost every actinian tissue. The familiar folding seen in cross-sections of the retractors is a special case of excessive buckling which is permanent. 3. A natural limit to the extension of anemone tissue is reached when the muscle-layer is completely unbuckled. If contraction proceeds to a maximum, there is a second order of buckling by which the whole body-wall is thrown into folds. Con-traction ca n then proceed no further. 4. The function of the muscle-fields is analysed. The youngest cycles of mesenteries (‘imperfect microcnemes’) supply the longitudinal musculature of the column (parietal muscle). The older ‘imperfect retractor-bearers’ have only feeble parietal musculature, but possess a retractor muscle connecting the oral with the pedal disk. The perfect mesenteries have a similar organization to the imperfect retractor-bearers, and parti-cularly in the non-directive perfect mesenteries there is a well-developed sheet of radial (exocoelic) muscle whose reflex contraction opens the mouth. The vertical endocoelic muscle-fibres of all non-directive mesenteries fan out on to the pedal disk. On the exocoelic side, the parieto-basilarfans out from the pedal disk to the body-wall. As usual, the muscle-fields of the directives are developed on opposite sides from those of the non-directives. 5. The muscular plan of the pedal disk is compared with the tube foot of Asterias as described by J. E. Smith. There is a significant functional similarit y in the opera-tion of vertical, oblique, and radial muscles (basilars) bearing on the adhesive disk. The circular layer of the actinian foot has no analogue in the tube foot. It is primarily concerned with locomotion and not with adhesion. 6. The functional organization of the oral disk and tentacles is discussed. It differs from the rest of the body in the retention of ectodermal longitudinal muscle. This layer is responsible for the special movements executed in feeding. The significance of its physiological separation from the endodermal system is noted.