Chapter II

PROTOZOA (CONTINUED): SPONTANEOUS GENERATION—CHARACTERS OF PROTOZOA—CLASSIFICATION

The Question of Spontaneous Generation

From the first discovery of the Protozoa, their life-history has been the subject of the highest interest: yet it is only within our own times that we can say that the questions of their origin and development have been thoroughly worked out. If animal or vegetable matter of any kind be macerated in water, filtered, or even distilled, various forms of Protista make their appearance; and frequently, as putrefaction advances, form after form makes its appearance, becomes abundant, and then disappears to be replaced by other species. The questions suggested by such phenomena are these: (1) Do the Protista arise spontaneously, that is, by the direct organisation into living beings of the chemical substances present, as a crystal is organised from a solution: (2) Are the forms of the Protista constant from one generation to another, as are ordinary birds, beasts, and fishes?

The question of the "spontaneous generation" of the Protista was readily answered in the affirmative by men who believed that Lice bred directly from the filth of human skins and clothes;^[57] and that Blow-flies, to say nothing of Honey-bees, arose in rotten flesh: but the bold aphorism of Harvey "omne vivum ex ovo" at once gained the ear of the best-inspired men of science, and set them to work in search of the "eggs" that gave rise to the organisms of putrefaction. Redi (ob. 1699) showed that Blow-flies never arise save when other Blow-flies gain access to meat and deposit their very visible eggs thereon. Leeuwenhoek, his contemporary, in the latter half of the seventeenth century adduced strong reasons for ascribing the origin of the organisms of putrefaction to invisible air-borne eggs. L. Joblot and H. Baker in the succeeding half-century investigated the matter, and showed that putrefaction was no necessary antecedent of the appearance of these beings: that, as well as being air-borne, the germs might sometimes have adhered to the materials used for making the infusion; and that no organisms were found if the infusions were boiled long enough, and corked when still boiling. These views were strenuously opposed by Needham in England, by Wrisberg in Germany, and by Buffon, the great French naturalist and philosopher, whose genius, unballasted by an adequate knowledge of facts, often played him sad tricks. Spallanzani made a detailed study of what we should now term the "bionomical" or "oecological" conditions of Protistic life and reproduction in a manner worthy of modern scientific research, and not attained by some who took the opposite side within living recollection. He showed that infusions kept sufficiently long at the boiling-point in hermetically sealed vessels developed no Protistic life. As he had shown that active Protists are killed at much lower temperatures, he inferred that the germs must have much higher powers of resistance; and, by modifying the details of his experiments, he was able to meet various objections of Needham.

Despite this good work, the advocates of spontaneous generation were never really silenced; and the widespread belief in the inconstancy of species in Protista added a certain amount of credibility to their cause. In 1845 Pineau put forward these views most strongly; and from 1858 to 1864 they were supported by the elder Pouchet. Louis Pasteur, who began life as a chemist, was led from a study of alcoholic fermentation to that of the organisms of fermentation and of putrefaction and disease. He showed that in infusions boiled sufficiently long and sealed while boiling, or kept at the boiling-point in a sealed vessel, no life manifested itself: objections raised on the score of the lack of access of fresh air were met by the arrangement, so commonly used in "pure cultures" at the present day, of a flask with a tube attached plugged with a little cotton-wool, or even merely bent repeatedly into a zigzag. The former attachment filtered off all germs or floating solid particles from the air, the latter brought about the settling of such particles in the elbows or on the sides of the tube: in neither case did living organisms appear, even after the lapse of months. Other observers succeeded in showing that the forms and characters of species were as constant as in Higher Animals and Plants, allowing, of course, for such regular metamorphoses as occur in Insects, or alternations of generations paralleled in Tapeworms and Polypes. The regular sequences of such alternations and metamorphoses constitute, indeed, a strong support of the "germ-theory"—the view that all Protista arise from pre-existing germs. It is to the Rev. W. H. Dallinger and the late Dr. Charles Drysdale that we owe the first complete records of such complex life-histories in the Protozoa as are presented by the minute Flagellates which infest putrefying liquids (see below, p. 116 f.). The still lower Schizomycetes, the "microbes" of common speech, have also been proved by the labours of Ferdinand Cohn, von Koch, and their numerous disciples, to have the same specific constancy in consecutive generations, as we now know, thanks to the methods first devised by De Bary for the study of Fungi, and improved and elaborated by von Koch and his school of bacteriologists.

And so to-day the principle "omne vivum ex vivo" is universally accepted by men of science. Of the ultimate origin of organic life from inorganic life we have not the faintest inkling. If it took place in the remote past, it has not been accomplished to the knowledge of man in the history of scientific experience, and does not seem likely to be fulfilled in the immediate or even in the proximate future.^[58]

PROTOZOA

Organisms of various metabolism, formed of a single cell or apocyte, or of a colony of scarcely differentiated cells, whose organs are formed by differentiations of the protoplasm, and its secretions and accretions; not composed of differentiated multicellular tissues or organs.^[59]

This definition, as we have seen, excludes Metazoa (including Mesozoa, Vol. II. p. 92) sharply from Protozoa, but leaves an uncertain boundary on the botanical side; and, as systematists share with nations the desire to extend their sphere of influence, we shall here follow the lead of other zoologists and include many beings that every botanist would claim for his own realm. Our present knowledge of the Protozoa has indeed been largely extended by botanists,^[60] while the study of protoplasmic physiology has only passed from their fostering care into the domain of General Biology within the last decade. The study of the Protozoa is little more than two centuries old, dating from the school of microscopists of whom the Dutchman Leeuwenhoek is the chief representative: and we English may feel a just pride in the fact that his most important publications are to be found in the early records of our own Royal Society.

Baker, in the eighteenth century, and the younger Wallich, Carter, Dallinger and Drysdale, Archer, Saville Kent, Lankester, and Huxley, in the last half-century, are our most illustrious names. In France, Joblot, almost as an amateur, like our own Baker, flourished in the early part of the eighteenth century. Dujardin in the middle of the same century by his study of protoplasm, or sarcode as he termed it, did a great work in laying the foundations of our present ideas, while Balbiani, Georges Pouchet, Fabre-Domergue, Maupas, Léger, and Labbé in France, have worthily continued and extended the Gallic traditions of exact observation and careful deduction. Otto Friedrich Müller, the Dane, in the eighteenth century, was a pioneer in the exact study and description of a large number of forms of these, as of other microscopic forms of life. The Swiss collaborators, Claparède and Lachmann, in the middle of the nineteenth century, added many facts and many descriptions; and illustrated them by most valuable figures of the highest merit from every point of view. Germany, with her large population of students and her numerous universities, has given many names to our list; among these, Ehrenberg and von Stein have added the largest number of species to the roll. Ehrenberg about 1840 described, indeed, an enormous number of forms with much care, and in detail far too elaborate for the powers of the microscope of that date: so that he was led into errors, many and grave, which he never admitted down to the close of a long and honoured life. Max Schultze did much good work on the Protozoa, as well as on the tissues of the Metazoa, and largely advanced our conceptions of protoplasm. His work was continued in Germany by Ernst Haeckel, who systematised our knowledge of the Radiolaria, Greeff, Richard Hertwig, Fritz Schaudinn, and especially Bütschli, who contributed to Bronn's Thier-Reich a monograph of monumental conception and scope, and of admirable execution, on which we have freely drawn. Cienkowsky, a Russian, and James-Clark and Leidy, both Americans, have made contributions of high quality.

Lankester's article in the Encyclopædia Britannica was of epoch-making quality in its philosophical breadth of thought.

Delage and Hérouard have given a full account of the Protozoa in their Traité de Zoologie Concrète, vol. i. (1896); and A. Lang's monograph in his Vergleichende Anatomie, 2nd ed. (1901), contains an admirable analysis of their general structure, habits, and life-cycles, together with full descriptions of well-selected types. Calkins has monographed "The Protozoa" in the Columbia University Biological series (1901). These works of Bütschli, Delage, Lang, and Calkins contain full bibliographies. Doflein has published a most valuable systematic review of the Protozoa parasitic on animals under the title Die Protozoen als Parasiten und Krankheitserreger (1901); and Schaudinn's Archiv für Protistenkunde, commenced only four years ago, already forms an indispensable collection of facts and views.

The protoplasm of the Protozoa (see p. 5 f.) varies greatly in consistency and in differentiation. Its outer layer may be granular and scarcely altered in Proteomyxa, the true Myxomycetes, Filosa, Heliozoa, Radiolaria, Foraminifera, etc.; it is clear and glassy in the Lobose Rhizopods and the Acrasieae; it is continuous with a firm but delicate superficial pellicle of membranous character in most Flagellates and Infusoria; and this pellicle may again be consolidated and locally thickened in some members of both groups so as to form a coat of mail, even with definite spines and hardened polygonal plates (Coleps, Fig. 54, p. 150). Again, it may form transitory or more or less permanent pseudopodia,^[61] (1) blunt or tapering and distinct, with a hyaline outer layer, or (2) granular and pointed, radiating and more or less permanent, or (3) branching and fusing where they meet into networks or perforated membranes. Cilia or flagella are motile thread-like processes of active protoplasm which perforate the pellicle; they may be combined into flattened platelets or firm rods, or transformed into coarse bristles or fine motionless sense-hairs. The skeletons of the Protozoa, foreign to the cytoplasm, will be treated of under the several groups.

Most of the fresh-water and brackish forms (and some marine ones) possess one or more contractile vacuoles, often in relation to a more or less complex system of spaces or canals in Flagellates and Ciliates.

The Geographical Distribution of Protozoa is remarkable for the wide, nay cosmopolitan, distribution of the terrestrial and fresh-water forms;^[62] they owe this to their power of forming cysts, within which they resist drought, and can be disseminated as "dust." Very few of them can multiply at a temperature approaching freezing-point; the Dinoflagellates can, however, and therefore present Alpine and Arctic forms. The majority breed most freely at summer temperatures; and, occurring in small pools, can thus achieve their full development in such as are heated by the sun during the long Arctic day as readily as in the Tropics. In infusions of decaying matter, the first to appear are those that feed on bacteria, the essential organisms of putrefaction. These, again, are quickly followed and preyed upon by carnivorous species, which prefer liquids less highly charged with organic matters, and do not appear until the liquid, hitherto cloudy, has begun to clear. Thus we have rather to do with "habitat" than with "geographical distribution," just as with the fresh-water Turbellaria and the Rotifers (vol. ii. pp. 4 f., 226 f.). We can distinguish in fresh-water, as in marine Protista, "littoral" species living near the bank, among the weeds; "plankton," floating at or near the surface; "zonal" species dwelling at various depths; and "bottom-dwellers," mostly "limicolous" (or "sapropelic," as Lauterborn terms them), and to be found among the bottom ooze. Many species are "epiphytic" or "epizoic," dwelling on plants or animals, and sometimes choice enough in their preference of a single genus or species as host. Others again are "moss-dwellers," living among the root-hairs of mosses and the like, or even "terrestrial" and inhabiting damp earth. The Sporozoa are internal parasites of animals, and so are many Flagellates, while many Proteomyxa are parasitic in plant-cells. The Foraminifera (with the exception of most Allogromidiaceae) are marine, and so are the Radiolaria; while most Heliozoa inhabit fresh water. Concerning the distribution in time we shall speak under the last two groups, the only ones whose skeletons have left fossil remains.

Classification.—The classification of the Protozoa is no easy task. We omit here, for obvious reasons, the unmistakable Plant Protists that have a holophytic or saprophytic nutrition; that are, with the exception of a short period of locomotion in the young reproductive cells, permanently surrounded with a wall of cellulose or fungus-cellulose, and that multiply and grow freely in this encysted state: to these consequently we relegate the Chytridieae, which so closely allied to the Proteomyxa and the Phycomycetous Fungi; and the Confervaceae, which in the resting state form tubular or flattened aggregates and are allied to the green Flagellates; besides the Schizophyta. At the opposite pole stand the Infusoria in the strict sense, with the most highly differentiated organisation found in our group, culminating in the possession of a nuclear apparatus with nuclei of two kinds, and exhibiting a peculiar form of conjugation, in which the nuclear apparatus is reorganised. The Sporozoa are clearly marked off as parasites in living animals, which mostly begin life as sickle-shaped cells and have always at least two alternating modes of brood-formation, the first giving rise to aplanospores, wherein is formed the second brood of sickle-shaped, wriggling zoospores. The remainder, comprising the Sarcodina, or Rhizopoda in the old wide sense (including all that move by pseudopodia during the great part of their active life), and the Flagellata in the widest sense, are not easy to split up; for many of the former have flagellate reproductive cells, and many of the latter can emit pseudopodia with or without the simultaneous retraction of their flagella. The Radiolaria are well defined by the presence in the body plasm of a central capsule marking it off into a central and a peripheral portion, the former containing the nucleus, the latter emitting the pseudopodia. Again, on the other hand, we find that we can separate as Flagellata in the strict sense the not very natural assemblage of those Protista that have flagella as their principle organs of movement or nutrition during the greater part of their active life. The remaining groups (which with the Radiolaria compose the Sarcodina of Bütschli), are the most difficult to treat. The Rhizopoda, as we shall limit them, are naked or possess a simple shell, never of calcium carbonate, have pseudopodia that never radiate abundantly nor branch freely, nor coalesce to form plasmatic networks, nor possess an axial rod of firmer substance. The Foraminifera have a shell, usually of calcium carbonate, their pseudopodia are freely reticulated, at least towards the base; and (with the exception of a few simple forms) all are marine. The Mycetozoa are clearly united by their tendency to aggregate more or less completely into complex resting-groups (fructifications), and by reproducing by unicellular zoospores, flagellate or amoeboid, which are not known to pair. The Heliozoa resemble the Radiolaria in their fine radiating pseudopodia, but have an axial filament always present in each, and lack the central capsule; and are, for the most part, fresh-water forms. Finally, the Proteomyxa forms a sort of lumber-room for beings which are intermediate between the Heliozoa, Rhizopoda, and Flagellata, usually passing through an amoeboid stage, and for the most part reproducing by brood-formation. Zoospores that possess flagella are certainly known to occur in some forms of Foraminifera, Rhizopoda, Heliozoa, and Radiolaria, though not by any means in all of each group.^[63]

A. Pseudopodia the principal means of locomotion and feeding; flagella absent or transitory I. Sarcodina

(1) Plastogamy only leading to an increase in size, never to the formation of "fructifications."

(a) Pseudopodia never freely coalescing into a network nor fine to the base Rhizopoda.

(*) Ectoplasm clear, free from granules; pseudopodia, usually blunt Rhizopoda Lobosa

(**) Ectoplasm finely granular; pseudopodia slender, branching, but not forming a network, passing into the body by basal dilatation Rhizopoda Filosa

(b) Pseudopodia branching freely and coalescing to form networks; ectoplasm granular; test usually calcareous or sandy Foraminifera

(c) Pseudopodia fine to the very base; radiating, rarely coalescing.

(i.) Pseudopodia with a central filament Heliozoa

(ii.) Pseudopodia without a central filament.

(*) Body divided into a central and a peripheral part by a "central capsule" Radiolaria

(**) Body without a central capsule Proteomyxa

(2) Cells aggregating or fusing into plasmodia before forming a complex "fructification" Mycetozoa

B. Cells usually moving by "euglenoid" wriggling or by excretion of a trail of viscid matter; reproduction by alternating modes of brood-formation, rarely by Spencerian fission II. Sporozoa

C. Flagella (rarely numerous) the chief or only means of motion and feeding III. Flagellata

D. Cilia the chief organs of motion, in the young state at least; nuclei of two kinds IV. Infusoria