It is then at once clear at what a disadvantage the Ascons are placed as compared with other sponges having canal systems of the second or third types. Their chamber and oscular currents can differ but slightly, the difference being obtained merely by narrowing the lumen of the distal extremity of the body to form the oscular rim. Further, the choanocytes are acting on a volume of water which they can only imperfectly control, and it is no doubt due to the necessity of limiting the volume of water which the choanocytes have to set in motion that the members of the Ascon family are so restricted in size. The oscular rim is only a special case of a device adopted by sponges at the very outset of their career, and retained and perfected when they have reached their greatest heights; the volume of water passing per second over every cross-section of the path of the current is of course the same, therefore by narrowing the cross-sectional area of the path at any point, the velocity of the current is proportionally increased at that point. The lining of the oscular rim is of pinacocytes; they determine a smooth surface, offering little frictional resistance to the current, while choanocytes in the same position would have been a hindrance, not only by setting up friction, but by causing irregularities in the motion.
Canal systems of the second type show a double advance upon that of the Ascons, namely, subdivision of the gastral cavity and much greater length of the smooth walled exhalant passage. The choanocytes have now a task more equal to their strength, and, further, there is now a very great inequality between the total sectional areas of the flagellated chambers and that of the oscular tube.
Canal systems of the third type with tubular chambers are an improvement on those of the second, in that the area of choanocytes is increased by the pouching of the chamber-layer without corresponding increase in the size of the sponge. However, the area of choanocytes represents expenditure of energy, and the next problem to be solved is how to retain the improved current and at the same time to cut down expense. The first step is to change the form of the chamber from tubular to spherical. Now the energy of all the choanocytes is concentrated on the same small volume of water. The area of choanocytes is less, but the end result is as good as before. At the wide mouth of the spherical chamber there is nevertheless still a cause of loss of energy in the form of eddies, and it is as an obviation of these that one must regard the aphodi and prosodi with which higher members of the Demospongiae are provided. The correctness of this view receives support, apart from mechanical principles, from the fact that the mass of the body of any one of these sponges is greater relatively to the total flagellated area than in those sponges with eurypylous chambers; that is to say, a few aphodal and diplodal chambers are as efficient as many of the eurypylous type.
It is manifest that the current is the bearer of the supply of food; but it requires more care to discover (1) what is the nature of the food; (2) by which of the cells bathed by the current the food is captured and by which digested. The answer to the latter question has long been sought by experimenters,[[275]] who supplied the living sponge with finely powdered coloured matters, such as carmine, indigo, charcoal, suspended in water. The results received conflicting interpretations until it became recognised that it was essential to take into account the length of time during which the sponge had been fed before its tissues were subjected to microscopic examination. Vosmaer and Pekelharing obtained the following facts: Spongilla lacustris and Sycon ciliatum, when killed after feeding for from half an hour to two hours with milk or carmine, contain these substances in abundance in the bodies of the choanocytes and to a slight degree in the deeper cells of the dermal tissue; after feeding for twenty-four hours the proportions are reversed, and if a period of existence in water uncharged with carmine intervenes between the long feed and death then the chambers are completely free from carmine. These are perhaps the most conclusive experiments yet described, and they show that the choanocytes ingest solid particles and that the amoeboid cells of the dermal layer receive the ingested matter from them. In all probability it is fair to argue from these facts that solid particles of matter suitable to form food for the sponge are similarly dealt with by it and undergo digestion in the dermal cells.
Choanocytes are the feeding organs par excellence; but the pinacocytes perform a small share of the function of ingestion, and in the higher sponges where the dermal tissue has acquired a great bulk the share is perhaps increased.
In the above experiments is implied the tacit assumption that sponges take their food in the form of finely divided solids. Haeckel[[276]] states his opinion that they feed on solid particles derived from decaying organisms, but that possibly decaying substances in solution may eke out their diet. Loisel, in 1898,[[277]] made a new departure in the field of experiment by feeding sponges with coloured solutions, and obtained valuable results. Thus solutions, if presented to the sponge in a state of extreme dilution, are subjected to choice, some being absorbed, some rejected. When absorbed they are accumulated in vacuoles within both dermal and gastral cells, mixed solutions are separated into their constituents and collected into separate vacuoles. In the vacuoles the solutions may undergo change; Congo red becomes violet, the colour which it assumes when treated with acid, and similarly blue litmus turns red. The contents of the vacuoles, sometimes modified, sometimes not, are poured out into the intercellular gelatinous matrix of the dermal layer, whence they are removed partly by amoeboid cells, partly, so Loisel thinks, by the action of the matrix itself. It adds to the value of these observations to learn that Loisel kept a Spongilla supplied with filtered spring-water, to which was added the filtered juice obtained from another crushed sponge. This Spongilla lived and budded, and was in good health at the end of ten days.
Movement.—Sponges are capable of locomotion only in the young stage; in the adult the only signs of movement are the exhalant current, and in some cases movements of contraction sufficiently marked to be visible to the naked eye. Meresjkowsky was one of the early observers of these movements. He mentions that he stimulated a certain corticate Monaxonid sponge by means of a needle point: a definite response to each prick inside the oscular rim was given by the speedy contraction of the osculum.[[278]]
Pigments and Spicules.—Various reasons lead one to conclude that the spicules have some function other than that of support and defence, probably connected with metabolism. For the spicules are cast off, sometimes in large numbers, to be replaced rapidly by new ones, a process for which it is difficult to find an adequate explanation if the spicules are regarded as merely skeletal and defensive.[[279]] Potts remarks upon the striking profusion with which spicules are secreted by developing Spongillids from water in which the percentage of silica present must have been exceedingly small. The young sponges climbed up the strands of spicules as they formed them, leaving the lower parts behind and adding to the upper ends.
Of the physiology of the pigments of sponges not much is yet known: a useful summary of facts will be found in Von Fürth's text-book.[[280]]
Spongin.—Von Fürth[[280]] points out that this term is really a collective one, seeing that the identity of the organic skeletal substance of all sponge species is hardly to be assumed. Spongin is remarkable for containing iodine. The amount of iodine present in different sponges varies widely, reaching in certain tropical species of the Aplysinidae and Spongidae the high figure 8 to 14 per cent. Seaweeds which are specially rich in iodine contain only 1.5 to 1.6 per cent.