The mechanism of ciliary movement. —V. The effect ions the duration of beat
In a previous paper (Gray, 1924) it was shown that the whole series of events which accompany continuous ciliary movement is divisible into at least three phases: (i) A reaction which is sensitive to monovalent cations (particularly the hydrogen-ion); any interference with this mechanism involves a change in the rate of beat of the cilia, and ultimately leads to a corresponding change in the oxygen consumption of the cells, (ii) A mechanism which is brought into operation by the presence of an activating acid substance, (iii) A reaction of an oxidative nature which removes some factor which is the direct result of activity and which, if allowed to accumulate, will inhibit the activity of the whole ciliary apparatus. The present paper deals primarily with the first of these processes. For this purpose the lateral epithelium on the gills of Mytilusedulis form a convenient material, since under appropriate conditions the cilia of these cells can (for many hours after excision of the gill) either be maintained in an active state of movement or be kept quite motionless. If the gills of a healthy animal be excised and examined at once under the microscope, the lateral cilia are seen to be in active movement, and exhibit a very beautiful metachronial rhythm which passes up and down the two sides of the gill-filaments. If these gills are now thoroughly washed with sea-water the lateral cilia sooner or later come to rest; the period which elapses before quiescence varies considerably with different individuals. It is usually about 15 mins., but it may be considerably longer in the case of well-fed mussels. It is to be concluded that the amount of available energy in the lateral epithelium is usually sufficient to maintain activity in sea-water for a strictly limited period. This is in marked contrast to the frontal epithelium, which remains active in sea-water for very long periods; in this case it may be assumed that the supply of energy is still being maintained from the glycoprotein (Gray 1924).