framework; it reconciles with the laws of physics the combination of a framework with a fluid or semi-fluid aggregate condition, while variations in the fluidity of the framework are compatible with a stiffening of the protoplasm almost to the pitch of rigidity, as seen, for example, in nervous tissue; and, finally, it explains many characteristic structural peculiarities of protoplasm, such as the superficial layer of radiately arranged alveoli, the spherical form of vacuoles, the continuous wall or pellicle which limits both the vacuoles and the protoplasm as a whole, and many other points not intelligible on the theory of a sponge-like structure. Bütschli has succeeded, moreover, in producing artificial foams of minute structure, which not only mimic the appearance of protoplasm, but can be made to exhibit streaming and amoeboid movements very similar to those of simple protoplasmic organisms. Incidentally these experiments have shown that many of the apparent granulation’s and “microsomes” are an optical effect produced by the nodes of the minute framework. In his most recent works Bütschli has extended his theory of alveolar structure to many other substances, and has tried to prove that it is a universal characteristic of colloid bodies, a view strongly combated, however, by Fischer. While it cannot be claimed that Bütschli’s theory furnishes in any way a complete explanation of life, leaving untouched, as it does, the fundamental question of assimilation and metabolism, he at least draws attention to a very important class of facts, which, if demonstrated to be of universal occurrence, must be reckoned with in future treatment of the protoplasm question, and would form an indispensable preliminary to all speculations upon the mechanism of the living substance.
In opposition to the above-mentioned monomorphic theories of protoplasm, all of which agree in assuming the existence of some fundamental type of structure in all living substance, attempts have been made at various times to show that the structural appearances seen in protoplasm are in reality artificial products, due to precipitation or coagulation caused by reagents used in the study or preparation of living objects. These views have been developed by Fischer, who by experimenting upon various proteids with histological fixatives, has shown that it is possible to produce in them a granular, reticular or alveolar structure, according to treatment, and, further, that granules so produced may be differentially stained according to their size and absorptive powers. Fischer therefore suggests that many structural appearances seen in protoplasm may be purely artificial, but does not extend this view to all such structures, which would indeed be impossible, in view of the frequency with which reticular or alveolar structures have been observed during life. He suggests, however, that such structures may be temporary results of vital precipitation of proteids within the organism, and that protoplasm may have at different times a granular reticular or alveolar structure, or may be homogeneous. Fischer’s conception of living protoplasm is therefore that of a polymorphic substance, and a similar view is held at the present time by Flemming, Wilson and others. Strassburger also regards protoplasm as composed of two portions: a motile kinoplasm which is fibrillar, and a nutritive trophoplasm which is alveolar, in structure.
The chemical investigation of protoplasm labours at the outset under the disadvantage that it cannot deal with the living substance as a whole, since no analysis can be performed upon it without destroying the life. Protoplasm consists, to the extent of about 60% of its total mass, of a mixture of various nucleo-proteids—that is to say, of those substances which, in molecular structure and chemical composition, are the most complex bodies known. In association with them are always found varying amounts of fats, carbohydrates, and other bodies, and such compounds are always present in the living substance to a greater or less degree as products of both upward and downward metabolism. Protoplasm also contains a large but variable percentage of water, the amount of which present in any given case affects largely its fluid or viscid aggregate condition. Especial interest attaches to the remarkable class of bodies known as ferments or enzymes, which when prepared and isolated from the living body are capable of effecting in other substances chemical changes of a kind regarded as specifically vital. It is from their study, and from that of the complex proteids found in the living body, that the greatest advances towards an explanation of the properties of living matter may be expected at the present time.
The question may be raised how far it is probable that there is one universal living substance which could conceivably be isolated or prepared in a pure state, and which would then exhibit the phenomena characteristic of vital activity. It is sufficiently obvious, in the first place, that protoplasm, as we know it, exhibits infinite diversity of character, and that no two samples of protoplasm are absolutely similar in all respects. Chemical differences must be assumed to exist not only between the vital fabrics of allied species of organisms, but even between those of individuals of the same species. Kassowitz regards this variability as compatible with the assumption of a gigantic protoplasmic molecule in which endless variations arise by changes in the combinations of a vast number of atoms and atom complexes. It is difficult to conceive, however, of any single substance, however complex in its chemical constitution, which could perform all the functions of life. To postulate a universal living substance is to proceed along a path which leads inevitably to the assumption of biophores, plastidules or other similar units, since the ultimate living particles must then be imagined as endowed at the outset with many, if not all, of the fundamental properties and characteristic actions of living bodies. Such a conception has as its logical result a vitalistic standpoint, which may or may not embody the correct mental attitude with regard to the study of life, but which at any rate tends to check any further advance towards an explanation or analysis of elementary vital phenomena. We may rather, with Kölliker, Verworn and others, ascribe the activities of protoplasm to the mutual interaction of many substances, no single one of which can be considered as living in itself, but only in so far as it forms an indispensable constituent of a living body. From this point of view life is to be regarded, not as the property of a single definite substance, but as the expression of the ever-changing relations existing between the many substances which make up the complex and variable congeries known to us as protoplasm.
Authorities.—For exhaustive historical summaries of the protoplasm question, with full bibliographical references, the reader may be referred to the following works, especially the first five: Bütschli, Investigations on Microscopic Foams and Protoplasm (London, 1894); Untersuchungen über Strukturen (Leipzig, 1898); “Meine Ansicht über die Struktur des Protolplasmas und einige ihrer Kritiker,” Arch. f. Entwickelungsmechanik d. Org. (1901); xi. 499–584, pl. xx.; Delage, La Structure du protoplasm et les théories sur lhérédité (Paris, 1895); Wilson, The Cell (2nd ed., London, 1900); Fischer, Fixirung, Färbung, und Bau des Protoplasmas (Leipzig, 1899); Kassowitz, Allgemeine Biologie (Vienna, 1899); G. Mann, Protoplasm, its Definition, Chemistry and Structure (Oxford, 1906), p. 59. (E. A. M.)
PROTOZOA (Gr. πρῶτος, first, and ζῷον, living thing), the name given by modern zoologists to the animalcules, for the most part microscopic, which were termed by the older naturalists Infusoria, from the manner in which they appear in infusions containing decaying animal and vegetable matter. The name Infusoria is now, however, restricted to one of the four classes which comprise the Protozoa proper. The name Protozoa was coined as far back as 1820 as an equivalent for the German word Urthiere, meaning animals of primitive or archaic nature, the forms of animal life which may be supposed to have been the first that appeared upon our globe. The great naturalist C. T. von Siebold was, however, the first to give a scientific definition to the group. Von Siebold pointed out that in the Protozoa the individual was always a single vital unit or cell, in contrast with the higher division of the animal kingdom, the Metazoa, in which the body is generally, though not universally, regarded as composed of many such units. To put the matter briefly and somewhat technically: the Protozoa are unicellular animals, the Metazoa multicellular animals; in the Protozoa the cell is complete in itself, both morphologically and physiologically, and is capable of maintaining a separate and independent existence in suitable surroundings, like any other organism; in the Metazoa the cells are differentiated for the performance of distinct functions and combined together to form the various tissues of which the body is built up, and the individual cells of the Metazoan body are not capable of maintaining a separate existence apart from their fellows. This is the sense in which the term Protozoa is used by zoologists, whereby certain forms of animal life, which were formerly ranked as Protozoa, such as sponges and rotifers, are now definitely excluded from the group and classed as Metazoa.
The animal kingdom may be divided, therefore, into two sub-kingdoms, the Protozoa and the Metazoa, the first-named characterized by their essentially unicellular nature. This is a criterion by which it is easy to define the Protozoa from a purely