Infusoria.

We are now brought face to face with animals which possess considerable variation of structure, Infusorial animalcules, as they are termed. It was Ehrenberg who attributed to them a highly complex organisation, but later observations negatived these views and showed them to be animals formed of one or more cells, or colonies of so-called individuals. It is true that this cell or united protoplasm may show a wonderful amount of differentiation, what with its nucleus and vacuole, mouth and gullet, its variously-arranged cilia or flagella, its contractile fibres, its separation into an outer denser and a more fluid inner protoplasm, and its horny cup and stalks.

In these few lines we have a condensed summary of the special qualities of minute forms of life that afford much interesting work for the microscope.

Fig. 332.—Acineta, magnified 600 diameters (Warne).

Among those widespread, and in some respects heterogeneous, forms of life associated under the comprehensive title of Infusoria, we encounter types that not only differ very widely from one another, but which occupy a different rank or position, so to speak, with regard to the relation they bear to each other, and also to the outlying representatives of the series—differences that permeate throughout the ranks of this extensive group. Furthermore, a considerable number of Infusorial animalcules foreshadow or typify, in a corresponding degree, the separate or associated cell elements out of which higher tissue structures—metazoic organisms—are built up. We may take the well-known example Euglena viridis ([Plate III]., No. 67), or Paramecium (No. 74), and their allies; these would appear to be the prototypes of Turbellaria. Another more lowly organised group of the Ciliata exhibits a distinct and highly-interesting affinity to the Opalinidæ. There are many other species (Acineta, [Plate III]., No. 68, for instance), which at first sight would seem to stand by themselves and present no marked agreement with any metazoic type. Indeed, the function of these and other polypites consists simply in seizing food and conveying it through perforations at the extremity of each separate tentaculum to its interior. In Acineta certain of the tentacles only are suctorial, and these, being the inner ones, fulfil the ingestive function, while the peripheral series are prehensile. This stalked club-shaped body ([Fig. 332]), which fixes itself to seaweeds or Bryozoa, is seen to have a nucleus, and also clear vesicles in the body-substance; its embryos are ciliated. It is an object of considerable interest even among curious marine animalcules; one or two species inhabit fresh water. The spiral-mouthed Spirostomum are among the largest of the class, and in sunlight are visible to the naked eye as slender golden threads of about ¹⁄₁₀th of an inch in length. The mouth slit, extending half the length of the body, is bordered on one side by cilia. The body is cylindrical and the surface covered with rows of cilia. Its multiplication takes place by transverse fission through the middle.

Flagellate Infusoria.—The characteristic of this group, as its name implies, is the possession of one or more flagella or whip-like appendages, at the base of which is an opening in the denser surface layer of protoplasm, and in the interior a nucleus and one or more contractile vacuoles, and not infrequently a brilliant red spot of pigment known to microscopists as the eye-spot. The Monads, which constitute the simplest members of the group, are commonly found in fresh-water pools and vegetable infusions. The typical form consists simply of a spherical or oval cell provided with a flagellum. The Volvox was formerly placed in this group, but as it contains chlorophyll it is properly claimed by the botanist. The collared group possesses cup-like collars, and these frequently secrete horny receptacles or cups, and form elegant tree-like colonies.

The mail-coated group are of very varied form, the body being often prolonged into spiny processes. They have two long flagella which fit into grooves purposely provided. But the most interesting and remarkable are the phosphorescent animalcules (Noctiluca), whose beautiful bluish-green luminosity on the surface of the sea has attracted attention from very early periods. It was, however, not until the first half of the present century that the luminosity was discovered to be due to the presence of multitudes of these minute jelly-like spheres.

Fig. 333.—Noctiluca miliaris; magnified 150 diameters.

Fig. 334.—Pyrocystis; magnified 150 diameters.

The body of the Noctiluca ([Fig. 333]) is a nearly globular-shaped cyst, enclosed in a tough membranous wall, from a grooved opening in which a striated muscular flagellum or proboscis is projected forth, and it is by means of this the animal swims away even in rough seas. A fine whip-like flagellum is also located in the same groove. At the apex of the funnel there is a mass of protoplasm which extends itself as a widely-meshed, highly-vacuolated network to the inner wall of the cyst, whence it is believed the phosphorescent light emanates. It multiplies by self-division, first becoming encysted after withdrawing its flagellum, and then breaking up into numerous ciliated helmet-shaped swarm spores. Frequently two organisms fuse into one and then divide into spores.

Noctiluca mainly confines itself to the shallower seas, but there are related forms met with in the warmer open seas; these belong to the genus Pyrocystis ([Fig. 334]). In one variety the body is perfectly spherical and without the big flagellum or proboscis. Professor Butschli, however, regards this species as an encysted or resting phase of the commoner and better-known form.

The late Mr. Philip Gosse, F.R.S., was the first microscopist to describe the Noctiluca. After careful observation, he wrote in his “Naturalist’s Rambles” as follows:—“I had an opportunity of becoming acquainted with the minute animals to which a great portion of the luminousness of the sea is attributed. One of my large glass vases of sea-water I had observed to become suddenly at night, when tapped with the finger, studded with minute but brilliant sparks at various points on the surface of the water. I set the jar in the window, and was not long in discovering, without the aid of a lens, a goodly number of the tiny jelly-like globules of Noctiluca miliaris swimming about in various directions. They swam with an even gliding motion, much resembling that of the Volvox globator of our fresh-water pools. They congregated in little groups, and a shake of the vessel sent them darting down from the surface. It was not easy to keep them in view when seen, owing rather to their extreme delicacy and colourless transparency than to their minuteness. They were, in fact, distinctly appreciable by the naked eye, measuring from ¹⁄₅₀th to ¹⁄₃₀th of an inch in diameter.”

Among the numerous fresh-water members of the flagellate infusoria, there is one which especially calls for notice, Codosiga, discovered by the late Professor H. J. Clark. This minute body bears a delicate funnel-shaped protoplasmic expansion or collar, common to the several members of this organic series. The flagellum is placed at the base of the oral opening, and within the circumscribed area of the collar, which is of such extreme tenuity that its true form and nature can only be determined by a very careful adjustment of the achromatic condenser and accessory apparatus employed, together with a wide-angled objective. It is seen to greater advantage by supplying the animal with very fine particles of colouring matter. In this way it is found that the infundibuliform cup consists of protoplasm, through which the flagellum is protruded and withdrawn into the general substance of the Monad’s body (Fig. 335). As many as twenty or more zooids are attached to the extremity of a slender footstalk. The length of the body, exclusive of the collar, is ¹⁄₂₅₀₀th to the ¹⁄₁₂₀₀th of an inch. The habitat of these bodies is fresh water. Mr. Saville Kent in 1869 discovered some of these interesting infusoria in the London Docks.

“The more exact significance of the special organ, the collar, is manifest by the circulatory currents or cyclosis induced, and there can be no room for doubt that this structure finds its precise homologue in the pseudopodia of the foraminiferous group of the Rhizopoda, in which a similar circulation or cyclosis of the constituent sarcode is exhibited. The whole of this highly-interesting flagellate order, a comparatively small one as yet, are remarkable for their pale glaucous green or florescent hue, such colour assisting materially in their recognition, even when the magnifying power employed is insufficient for the detection of the very characteristic collar with its enclosed flagellum.”[66]

Fig. 335.—Codosiga umbellata; a few colonies of Zooids diverging from the parent foot-stalk with flagella extended, magnified 650 diameters.

Ciliata.—Types of Ciliata obtained from hay infusions are very numerous. Ehrenberg’s animalcules were mainly of a large size, and of those belonging to the higher order of the Ciliata, pertaining to such genera as Paramecium, Colpoda, Cyclidium, Oxytricha, and Vorticella. These, however, represent but an insignificant minority of the hosts of flagellate forms which abound in our humid climate, and in hay infusions in particular. In such infusions, watched from day to day and produced from hay obtained from different localities, the number of types developed in regular sequence is found to be perfectly marvellous, commencing with the Monas proper, Amphimonas and Heteromita; while Bacteria, in their motile and quiescent forms, are invariably present and furnish an abundant supply of material for the microscope.[67]

Vorticellidæ constitute one of the most numerous families of the ciliate infusoria. All its members are at once recognised by their normal stationary condition, and by the structure of their oral system. In but few of the genera is there any marked divergence from this formula, and when any exists it is made manifest by an increase in development of some one of its elements at the expense of another. For instance, in the genus Spirochona, the external edge of the encircling border or peristome is suppressed, while the inner portion is abnormally developed into a transparent and highly elevated spiral membrane. The bell-animalcules usually possess stalks, and are either solitary or form branching colonies. Conichilus vorticella ([Plate III]., No. 80) is a well-known member of the colony stock, all the zooids of which are united on a slender branching pedicle, which consists of a central contractile cord enclosed within a tubular hyaline sheath. There are many other shrub-like colonies all variously modified in form and character. The Epistylis opercularia, or nodding-bell animalcule, is an interesting member of a numerous host of solitary short-stalked forms ([Fig. 337]). When the animal is disturbed, the heads drop down towards the stalk. This animalcule has been found to form a colony; and another, Carchesium, whose tiny branched tree-like colonies resemble little white globular masses of moulds, are seen at once to drop down towards the base of the colony with a jerky movement if the cell be touched. By a process of encysting, all the Vorticellæ and many of the more highly-organised ciliata have the means of what may be termed self-preservation. Should the water dry up in which they have been living, the little animal encases itself in mud at the bottom of the pool. Should this be baked by the sun not the least injury arises, for at this stage it crumbles into dust, and is carried by the wind to long distances, but the first shower of rain calls it back to active life, and soon after it is seen to issue forth as a free swimming bud.

Fig. 336.—Vorticella microstoma.

Thuricola valvata ([Plate III]., No. 72) possesses a hinge-like process which closes up like a door when the animal contracts itself into its case. This very effectually protects it from assault. Both portions of the valve are capable of extension. Another group of ciliate infusoria also possess a limited number of cilia, but these, although restricted to the under surface of their bodies, have an unrestricted range of motion. The group are all free swimmers, belonging to the genus Oxytricha. They possess two separate alimentary orifices, neither of which are situated at the extremities or encased by a dense integument. Their locomotive organs consist either of setæ, vibratile cilia, or non-vibratile styles or uncini, variously situated, and all serving to make these infusorial animals very active ([Plate III]., Nos. 73 and 77). A typical species is the mussel-animalcule (Stylonychia, [Fig. 338]), common in all infusions and pools of water. Its body is oval and flattened, and about ¹⁄₁₀₀th of an inch in length. At one end a funnel-shaped depression or mouth, with a ciliated margin, leads to the inner part of the body, in which are two oval bodies, a nucleus and a contractile vacuole, which is seen to contract rhythmically. The creature can also stalk along by means of its cilia or setæ, and set up currents to the mouth. [Plate III]., Nos. 70, 71, 72, 73, and 74, are types of these interesting bodies.

Fig. 337.—Nodding-bell animalcule (Epistyles operculata) × 250 (Warne).

Fig. 338.—Mussel-animalcule (Stylonychia mytilus) under surface.

a. Mouth; b. Contractile vacuole; c. Nucleus. (Magnified 150 diameters.

Dr. Balbini believes a true sexual generation occurs among these organisms, but, with the exception of the Paramecium, this has not been seen to take place; even Gruber’s more recent investigations appear to be inconclusive on this point. Conjugation, however, it is said takes place among some attached forms, as in the Stentors. These have been seen to put forth a bud from the body base, and soon after become free swimming bodies. The trumpet-animalcule (Stentor), a conspicuous member of the ciliata, is comparatively large, being about the ¹⁄₂₅th of an inch in length when extended to the full size. It is usually found attached to the under sides of duckweed, and is continually changing its form from that of a small knob when contracted, to the trumpet shape seen in [Fig. 339], No. 6, when fully extended, and from which it derives its name. The long cilia projected from the upper part form a spiral within the margin of the open mouth leading to the digestive sac. A contractile vacuole lies to the right of the oral opening. New individuals are produced by the process of budding, and in the form of ciliated embryos from the nucleus. Stentors are commonly met with in fresh water, and are usually of a brilliant green colour. These little bodies will bear cutting up: if only a fragment of the nucleus be included in the section, the injury is soon repaired.

Rotifera, or Wheel-animalcules ([Fig. 339]).—In this group we have a higher type of animal, with a more complex organisation than those previously noticed. The great majority inhabit fresh water, and are readily developed in hay infusions, in bog-moss, in house-top gutters, everywhere if looked for after a shower of rain. The rotating organs from which these fascinating animalcula derive their name consist of two disc-like bodies whose margins are fringed with rows of cilia, which create currents toward the oral aperture, and which have given rise to the optical delusion of rotating wheels. The disposition of the cilia is so arranged as to bring food to the rotifer and conduct it to the mastax or digesting apparatus—a muscular bulb moved by a series of muscles—the gastric glands and stomach. The great transparency of the whole structure permits of the animal economy being easily studied. The body is covered with a horny envelope of two layers, and is divided into segmental divisions, which slide into each other telescopic fashion. Consequently, as the water dries up, the animal is for a long time rendered indestructible and capable of resisting varying temperatures and the action of caustic reagents.

Rotifers are oviparous, and their eggs are conspicuous and of three kinds. The common soft-shelled eggs produce females, the smaller and more spherical produce males. The ephippial, or summer eggs, are often beset with spines or bosses; these have only a membranous covering, and are hatched soon after they are laid, or before leaving the ova sac. The male rotifer is but a third of the length of the female, often without cilia, and appears to have no alimentary tract; indeed, the only internal organ is a large sperm sac. Rotifers have been divided by Dr. Hudson and the late Mr. Gosse in their charming work on these very interesting “Wheel-animalcules” into four orders, according to their powers of locomotion, as follows:—(1) Rhizota, the rooted; (2) Bdelloida, the leech-like, that swim and creep like a leech; (3) Ploïma, the sea-worthy, that only swim with their ciliary wreath; (4) Scirtopoda, the skippers, that swim with their cilia and skip with arthropodous limbs. These, again, are subdivided into families. With such hardy creatures as Philodina, Adineta, Brachionus, &c., creatures to whom extremes of cold, heat, and drought are the ordinary conditions of life, nothing can be easier to keep going throughout the year. Mr. C. F. Rousselet, who has so thoroughly succeeded in mounting Rotifers with their cilia fully extended, recently exhibited at one of the evening meetings of the Royal Microscopical Society, London, no less than four hundred specimens in a natural and perfect condition, the nervous system being seen more clearly from its successful staining throughout the body than in the living rotifer.

Fig. 339.

1. Rotifer vulgaris with its cilia; b. rotating; c. horn; d. œsophagus; f. outer case; g. ova, foot protruding through outer case. 2. Same in the contracted state and at rest, showing the segmentation of the body and development of young. 3. Pitcher-shaped Brachionus, furnished with two horny projections; a. mastax; b. shell; c. cilia, rotating disc; d. foot. 4. Baker’s Brachionus, with six horny setæ; these are retracted when the cilia are in action; the letters relate to the same internal organs as in the former; the ova sac seen filled with eggs. 5 and 6. B. ovalis, closed, and with cilia displayed.

There is also a family of Rotatoria with a single rotatory organ, disposed around the margin of the case. This comprises at present a very small group. The Œcistes is a member of the family ([Plate III]., No. 69). A single ciliary wreath leads to the alimentary canal, and a pharyngeal bulb or mastax comprises the apparatus of nutrition. The visual organs are red, as in other rotifers, and the ovarium contains several ova, shown in No. 69. The envelope is a gelatinous transparent sheath, into which the animalcule can withdraw itself, its attachment to the bottom being by the end of the foot-like tail. The most interesting among this genus are the Floscularians. These creatures may undoubtedly be described as among the most beautiful and interesting of infusorial animals.

The Stephanoceros, “crowned animalcule,” as it is termed, is about ¹⁄₃₆th of an inch in length, and enclosed in a transparent cylindrical flexible case, beyond which it protrudes five long arms in a graceful manner. These, touching at their points, give a form from which it derives its name. These arms are furnished with several rows of short cilia, which seize the food brought within their grasp until it can be swallowed. In addition to the rotatory organs, they have short flexible processes, or cornu, attached to the outside of one or more of their lobes. The water vascular system consists of two canals arising from a small pyriform contractile vesicle, situated below the stomach. The ova, after leaving the ova sac, remain quiescent until their cilia are developed. Floscularians, like Melicertans, have a certain affinity in form with Vorticellians and Stentors, and also with Campanulariæ, among polypes. Their cilia are less regular when in action than in other Rotatoria. When they retreat into their transparent cells they appear to fold themselves up. Their internal structure can be seen through the external case, and ova are observed enclosed in an ova sac; when thrown off they remain quiescent until the formation of their cilia. The whole family furnish interesting objects for microscopic investigation.

Melicerta ringens (“beaded Melicerta”).—Of all the Melicerta, or “horny floscularia,” this is the most beautiful. Its crystalline body is enclosed in a pellucid covering, wider at the top than the bottom, of a dark yellow or reddish-brown colour, which gradually becomes encrusted by zones of a variety of shapes, cemented together with a peculiar secretion that hardens in water. It derives its name from these pellets, which have the appearance of rows of beads. Mr. Gosse furnished an excellent account of the architectural instincts of Melicerta ringens: “An animalcule so minute as to be with difficulty appreciable by the naked eye, inhabiting a tube composed of pellets, which it forms and lays one by one. It is a mason who not only builds up his mansion brick by brick, but makes his bricks as he goes on, from substances which he collects around him, shaping them in a mould which he carries on his body.

“The pellets composing the case are very regularly placed in position; in a fine specimen, about the ¹⁄₃₀th of an inch in length, when fully expanded, as many as fifteen longitudinal rows of pellets were counted, which gave about thirty-two rows in all. As it exposes itself more and more, suddenly two large rounded discs are expanded, around which, at the same instant, a wreath of cilia is seen performing surprising motions.

“On mixing carmine with the water, the course of the ciliary current is readily traced, and forms a fine spectacle. The particles are hurled round the margin of the disc, until they pass off in front through the great sinus, between the larger petals. If the pigment be abundant, the cloudy torrent for the most part rushes off, and prevents our seeing what takes place; but if the atoms be few, we see them swiftly glide along the facial surface, following the irregularities of outline with beautiful precision, dash round the projecting chin like a fleet of boats doubling a bold headland, and lodge themselves, one after another, in the little cup-like receptacle beneath. Mr. Gosse, believing that the pellets of the case might be prepared in the cup-like receptacle, watched the animal, and presently had the satisfaction of seeing it bend its head forward, as anticipated, and after a second or two raise it again; the little cup having in the meantime lost its contents. It immediately began to fill again; and when it was full, and the contents were consolidated by rotation, aided probably by the admixture of a salivary secretion, it was again bent down to the margin of the case, and emptied of its pellet. This process he saw repeated many times in succession, until a goodly array of dark-red pellets were laid upon the yellowish-brown ones, but very irregularly. After a certain number were deposited in one part, the animal would suddenly turn itself round in its case, and deposit some in another part. It took from two-and-a-half to three-and-a-half minutes to make and deposit a pellet.”

Melicerta may be found in clear pools, mill-ponds, and other places through which a current of water gently flows. If a portion of water-weed be brought home and placed in a small glass zoophyte-trough, and carefully examined with a magnifying power of about fifty diameters, a few delicate-looking projections of a reddish-brown colour will probably be seen adhering to the plant; these are the tubular cases of Melicerta, which, after a short period of rest, will be seen to be animals of ¹⁄₁₂th of an inch or more in length.