# Littell's Living Age/Volume 128/Issue 1657/On the Border Territory between the Animal and the Vegetable Kingdoms

From Macmillan's Magazine.
ON THE BORDER TERRITORY BETWEEN THE ANIMAL AND THE VEGETABLE KINGDOMS.
BY T. H. HUXLEY.
In the whole history of science there is nothing more remarkable than the rapidity of the growth of biological knowledge within the last half-century, and the extent of the modification which has thereby been effected in some of the fundamental conceptions of the naturalist.
In the second edition of the "Règne Animal," published in 1828, Cuvier devotes a special section to the "Division of Organized Beings into Animals and Vegetables," in which the question is treated with that comprehensiveness of knowledge and clear critical judgment which characterize his writings, and justify us in regarding them as representative expressions of the most extensive, if not the profoundest, knowledge of his time. He tells us that living beings have been sub-divided from the earliest times into animated beings, which possess sense and motion, and inanimated beings, which are devoid of these functions, and simply vegetate.
Although the roots of plants direct themselves towards moisture, and their leaves towards air and light; although the parts of some plants exhibit oscillating movements without any perceptible cause, and the leaves of others retract when touched, yet none of these movements justify the ascription to plants of perception or of will.
From the mobility of animals, Cuvier, with his characteristic partiality for teleological reasoning, deduces the necessity of the existence in them of an alimentary cavity or reservoir of food, whence their nutrition may be drawn by the vessels, which are a sort of internal roots; and in the presence of this alimentary cavity he naturally sees the primary and the most important distinction between animals and plants.
Following out his teleological argument, Cuvier remarks that the organization of this cavity and its appurtenances must needs vary according to the nature of the aliment, and the operations which it has to undergo, before it can he converted into substances fitted for absorption; while the atmosphere and the earth supply plants with juices ready prepared, and which can be absorbed immediately.
As the animal body required to be independent of heat and of the atmosphere, there were no means by which the motion of its fluids could be produced by internal causes. Hence arose the second great distinctive character of animals, or the circulatory system, which is less important than the digestive, since it was unnecessary, and therefore is absent, in the more simple animals.
Animals further needed muscles for locomotion and nerves for sensibility. Hence, says Cuvier, it was necessary that the chemical composition of the animal body should be more complicated than that of the plant; and it is so, inasmuch as an additional substance, nitrogen, enters into it as an essential element, while in plants nitrogen is only accidentally joined with the three other fundamental constituents of organic beings — carbon, hydrogen, and oxygen. Indeed, he afterwards affirms that nitrogen is peculiar to animals; and herein he places the third distinction between the animal and the plant.
The soil and the atmosphere supply plants with water, composed of hydrogen and oxygen; air, consisting of nitrogen and oxygen; and carbonic acid, containing carbon and oxygen. They retain the hydrogen and the carbon, exhale the superfluous oxygen, and absorb little or no nitrogen. The essential character of vegetable life is the exhalation of oxygen, which is effected through the agency of light.
Animals, on the contrary, derive their nourishment either directly or indirectly from plants. They get rid of the superfluous hydrogen and carbon, and accumulate nitrogen.
The relations of plants and animals to the atmosphere are therefore inverse. The plant withdraws water and carbonic acid from the atmosphere, the animal contributes both to it. Respiration — that is, the absorption of oxygen and the exhalation of carbonic acid — is the specially ​animal function of animals, and constitutes their fourth distinctive character.
Thus wrote Cuvier in 1828. But in the fourth and fifth decades of this century, the greatest and most rapid revolution which biological science has ever undergone was effected by the application of the modern microscope to the investigation of organic structure; by the introduction of exact and easily manageable methods of conducting the chemical analysis of organic compounds; and finally, by the employment of instruments of precision for the physical measurement of the forces which are at work in the living economy.
That the semi-fluid contents (which we now term protoplasm) of the cells of certain plants, such as the Charæ, are in constant and regular motion, was made out by Bonaventura Corti a century ago but the fact, important as it was, fell into oblivion, and had to be rediscovered by Treviranus in 1807. Robert Brown noted the more complex motions of the protoplasm in the cells of Tradescuntia in 1831; and now such movements of the living substance of plants are well-known to be some of the most widely prevalent phenomena of vegetable life.
Agardh, and other of the botanists of Cuvier's generation, who occupied themselves with the lower plants, had observed that, under particular circumstances, the contents of the cells of certain water-weeds were set free and moved about with considerable velocity, and with all the appearances of spontaneity, as locomotive bodies, which, from their similarity to animals of simple organization, were called "zoospores."
Even as late as 1845, however, a botanist of Schleiden's eminence deals very sceptically with these statements; and his scepticism was the more justified, since Ehrenberg, in his elaborate and comprehensive work on the infusoria, had declared the greater number of what are now recognized as locomotive plants to be animals.
At the present day, innumerable plants and free plant-cells are known to pass the whole or part of their lives in an actively locomotive condition, in no wise distinguishable from that of one of the simpler animals; and, while in this condition, their movements are, to all appearance, as spontaneous — as much the product of volition — as those of such animals.
Hence the teleological argument for Cuvier's first diagnostic character — the presence in animals of an alimentary cavity, or internal pocket, in which they can carry about their nutriment, has broken down so far, at least, as his mode of stating it goes. And with the advance of microscopic anatomy the universality of the fact itself among animals has ceased to be predicable. Many animals of even complex structure, which live parasitically within others, are wholly devoid of an alimentary cavity. Their food is provided for them, not only ready cooked but ready digested, and the alimentary canal, become superfluous, has disappeared. Again, the males of most rotifers have no digestive apparatus; as a German naturalist has remarked, they devote themselves entirely to the Minnedienst, and are to be reckoned among the few realizations of the Byronic ideal of a lover. Finally, amidst the lowest forms of animal life, the speck of gelatinous protoplasm, which constitutes the whole body, has no permanent digestive cavity or mouth, but takes in its food anywhere; and digests, so to speak, all over its body.
But although Cuvier's leading diagnosis of the animal from the plant will not stand a strict test, it remains one of the most constant of the distinctive characters of animals. And if we substitute for the possession of an alimentary cavity, the power of taking solid nutriment into the body and there digesting it, the definition so changed will cover all animals, except certain parasites, and the few and exceptional cases of non-parasitic animals which do not feed at all. On the other hand, the definition thus amended will exclude all ordinary vegetable organisms.
Cuvier himself practically gives up his second distinctive mark when he admits that it is wanting in the simpler animals.
The third distinction is based on a completely erroneous conception of the chemical differences and resemblances between the constituents of animal and vegetable organisms, for which Cuvier is not ​responsible, as it was current among contemporary chemists.
It is now established that nitrogen is as essential a constituent of vegetable as of animal living matter; and that the latter is, chemically speaking, just as complicated as the former. Starchy substances, cellulose and sugar, once supposed to be exclusively confined to plants, are now known to be regular and normal products of animals. Amylaceous and saccharine substances are largely manufactured, even by the highest animals; cellulose is widespread as a constituent of the skeletons of the lower animals; and it is probable that amyloid substances are universally present in the animal organism, though not in the precise form of starch.
Moreover, although it remains true that there is an inverse relation between the green plant in sunshine and the animal, in so far as, under these circumstances, the green plant decomposes carbonic acid and exhales oxygen, while the animal absorbs oxygen and exhales carbonic acid; yet the exact investigations of the modern chemical investigator of the physiological processes of plants have clearly demonstrated the fallacy of attempting to draw any general distinction between animals and vegetables on this ground. In fact the difference vanishes with the sunshine, even in the case of the green plant; which, in the dark, absorbs oxygen and gives out carbonic acid like any animal. While those plants, such as the fungi, which contain no chlorophyll and are not green, are always, so far as respiration is concerned, in the exact position of animals. They absorb oxygen and give out carbonic acid.
Thus, by the progress of knowledge, Cuvier's fourth distinction between the animal and the plant has been as completely invalidated as the third and second; and even the first can be retained only in a modified form and subject to exceptions.
But has the advance of biology simply tended to break down old distinctions, without establishing new ones?
With a qualification, to be considered presently, the answer to this question is undoubtedly in the affirmative. The famous researches of Schwann and Schleiden in 1837 and the following years, founded the modern science of histology, or that branch of anatomy which deals with the ultimate visible structure of organisms, as revealed by the microscope; and from that day to this the rapid improvement of methods of investigation, and the energy of a host of accurate observers, have given greater and greater breadth and firmness to Schwann's great generalization, that a fundamental unity of structure obtains in animals and plants; and that however diverse may be the fabrics, or tissues, of which their bodies are composed, all these varied structures result from the metamorphoses of morphological units (termed cells, in a more general sense than that in which the word "cells" was at first employed), which are not only similar in animals and in plants respectively, but present a close fundamental resemblance when those of animals and those of plants are compared together.
The contractility which is the fundamental condition of locomotion, has not only been discovered to exist far more widely among plants than was formerly imagined, but, in plants, the act of contraction has been found to be accompanied, as Dr. Burdon Sanderson's interesting investigations have shown, by a disturbance of the electrical state of the contractile substance comparable to that which was found by Du Bois Reymond to be a concomitant of the activity of ordinary muscle in animals.
Again, I know of no test by which the reaction of the leaves of the sundew and of other plants to stimuli, so fully and carefully studied by Mr. Darwin, can be distinguished from those acts of contraction following upon stimuli, which are called "reflex" in animals.
On each lobe of the bibbed leaf of Venus's fly-trap (Dionæa muscipula) are three delicate filaments which stand out at right angles from the surface of the leaf. Touch one of them with the end of a fine human hair and the lobes of the leaf instantly close together in virtue of an act of contraction of part of their substance, just as ​the body of a snail contracts into its shell when one of its "horns" is irritated.
The reflex action of the snail is the result of the presence of a nervous system in that animal. A molecular change takes place in the nerve of the tentacle, is propagated to the muscles by which the body is retracted, and causing them to contract, the act of retraction is brought about. Of course the similarity of the acts does not necessarily involve the conclusion that the mechanism by which they are effected is the same; but it suggests a suspicion of their identity which needs careful testing.
The results of recent inquiries into the structure of the nervous system of animals converge towards the conclusion that the nerve-fibres, which we have hitherto regarded as ultimate elements of nervous tissue, are not such, but are simply the visible aggregations of vastly more attenuated filaments, the diameter of which dwindles down to the limits of our present microscopic vision, greatly as these have been extended by modern improvements of the microscope; and that a nerve is, in its essence, nothing but a linear tract of specially modified protoplasm between two points of an organism — one of which is able to affect the other by means of the communication so established. Hence it is conceivable that even the simplest living being may possess a nervous system. And the question whether plants are provided with a nervous system or not, thus acquires a new aspect, and presents the histologist and physiologist with a problem of extreme difficulty, which must be attacked from a new point of view and by the aid of methods which have yet to be invented.
Thus it must be admitted that plants may be contractile and locomotive; that, while locomotive, their movements may have as much appearance of spontaneity as those of the lowest animals; and that many exhibit actions comparable to those which are brought about by the agency of a nervous system in animals. And it must be allowed to be possible that further research may reveal the existence of something comparable to a nervous system in plants. So that I know not where we can hope to find any absolute distinction between animals and plants, unless we return to their mode of nutrition, and inquire whether certain differences of a more occult character than those imagined to exist by Cuvier, and which certainly hold good for the vast majority of animals and plants, are of universal application.
A bean may be supplied with water in which salts of ammonia and certain other mineral salts are dissolved in due proportion; with atmospheric air containing its ordinary minute dose of carbonic acid; and with nothing else but sunlight and heat. Under these circumstances, unnatural as they are, with proper management, the bean will thrust forth its radicle and its plumule; the former will grow down into roots, the latter grow up into the stem and leaves of a vigorous bean-plant; and this plant will, in due time, flower and produce its crop of beans, just as if it were grown in the garden or in the field.
The weight of the nitrogenous protein compounds, of the oily, starchy, saccharine and woody substances contained in the full-grown plant and its seed, will be vastly greater than the weight of the same substances contained in the bean from which it sprang. But nothing has been supplied to the bean save water, carbonic acid, ammonia, potash, lime, iron, and the like, in combination with phosphoric, sulphuric and other acids. Neither protein, nor fat, nor starch, nor sugar, nor any substance in the slightest degree resembling them have formed part of the food of the bean. But the weights of the carbon, hydrogen, oxygen, nitrogen, phosphorus, sulphur, and other elementary bodies contained in the bean-plant, and in the seeds which it produces, are exactly equivalent to the weights of the same elements which have disappeared from the materials supplied to the bean during its growth. Whence it follows that the bean has taken in only the raw materials of its fabric and has manufactured them into bean-stuffs.
The bean has been able to perform this great chemical feat by the help of its green colouring matter, or chlorophyll, which, under the influence of sunlight, has the marvellous power of decomposing carbonic acid, setting free the oxygen and laying hold of the carbon which it contains. In fact the bean obtains two of the absolutely indispensable elements of its substance from two distinct sources; the watery solution, in which its roots are plunged, contains nitrogen but no carbon; the air, to which the leaves are exposed, contains carbon, but its nitrogen is in the state of a free gas, in which condition the bean can make no use of it and the chlorophyll is the apparatus by which the carbon is extracted from the atmospheric carbonic acid — the leaves being the chief ​laboratories in which this operation is effected.
The great majority of conspicuous plants are, as everybody knows, green; and this arises from the abundance of their chlorophyll. The few which contain no chlorophyll and are colourless, are unable to extract the carbon which they require from atmospheric carbonic acid, and lead a parasitic existence upon other plants; but it by no means follows, often as the statement has been repeated, that the manufacturing power of plants depends on their chlorophyll, and its interaction with the rays of the sun. On the contrary, it is easily demonstrated, as Pasteur first proved, that the lowest fungi, devoid of chlorophyll, or of any substitute for it, as they are, nevertheless possess the characteristic manufacturing powers of plants in a very high degree. Only it is necessary that they should be supplied with a different kind of raw material; as they cannot extract carbon from carbonic acid, they must be furnished with something else that contains carbon. Tartaric acid is such a substance; and if a single spore of the commonest and most troublesome of moulds — Penicillium — be sown in a saucer full of water, in which tartrate of ammonia, with a small percentage of phosphates and sulphates is contained, and kept warm, whether in the dark or exposed to light, it will, in a short time, give rise to a thick crust of mould, which contains many million times the weight of the original spore, in protein compounds and cellulose. Thus we have a very wide basis of fact for the generalization that plants are essentially characterized by their manufacturing capacity by their power of working up mere mineral matters into complex organic compounds.
Contrariwise, there is a no less wide foundation for the generalization that animals, as Cuvier puts it, depend directly or indirectly upon plants for the materials of their bodies; that is, either they are herbivorous, or they eat other animals which are herbivorous.
But for what constituents of their bodies are animals thus dependent upon plants? Certainly not for their horny matter; nor for chondrin, the proximate chemical element of cartilage; nor for gelatine; nor for syntonin, the constituent of muscle; nor for their nervous or biliary substances; nor for their amyloid matters; nor, necessarily, for their fats.
It can be experimentally demonstrated that animals can make these for themselves. But that which they cannot make, but must, in all known cases, obtain directly or indirectly from plants, is the peculiar nitrogenous matter protein. Thus the plant is the ideal prolétaire of the living world, the worker who produces; the animal, the ideal aristocrat, who mostly occupies himself in consuming, after the manner of that noble representative of the line of Zähdarm, whose epitaph is written in "Sartor Resartus."
Here is our last hope of finding a sharp line of demarcation between plants and animals; for, as I have already hinted, there is a border territory between the two kingdoms, a sort of no-man's land, the inhabitants of which certainly cannot be discriminated and brought to their proper allegiance in any other way.
Some months ago, Professor Tyndall asked me to examine a drop of infusion of hay, placed under an excellent and powerful microscope, and to tell him what I thought some organisms visible in it were. I looked and observed, in the first place, multitudes of Bacteria moving about with their ordinary intermittent spasmodic wriggles. As to the vegetable nature of these there is now no doubt. Not only does the close resemblance of the Bacteria to unquestionable plants, such as the Oscillatoriæ, and lower forms of fungi, justify this conclusion, but the manufacturing test settles the question at once. It is only needful to add a minute drop of fluid containing Bacteria, to water in which tartrate, phosphate, and sulphate of ammonia are dissolved; and, in a very short space of time, the clear fluid becomes milky by reason of their prodigious multiplication, which, of course, implies the manufacture of living bacterium-stuff out of these merely saline matters.
But other active organisms, very much larger than the Bacteria, attaining in fact the comparatively gigantic dimensions of one three-thousandth part of an inch or more, incessantly crossed the field of view. Each of these had a body shaped like a pear, the small end being slightly incurved and produced into a long curved filament, or cilium, of extreme tenuity. Behind this, from the concave side of the incurvation, proceeded another long cilium, so delicate as to be discernible only by the use of the highest powers and careful management of the light. In the centre of the pear-shaped body a clear round space could occasionally be discerned, but not always; and careful watching showed that this clear vacuity appeared gradually, and then shut up and disappeared suddenly, at regular intervals. Such a ​structure is of common occurrence among the lowest plants and animals, and is known as a contractile vacuole.
The little creature thus described sometimes propelled itself with great activity, with a curious rolling motion, by the lashing of the front cilium, while the second cilium trailed behind; sometimes it anchored itself by the hinder cilium and was spun round by the working of the other, its motions resembling those of an anchor buoy in a heavy sea. Sometimes, when two were in full career towards one another, each would appear dexterously to get out of the other's way; sometimes a crowd would assemble and jostle one another, with as much semblance of individual effort as a spectator on the Grands Mulets might observe with a telescope among the specks representing men in the valley of Chamounix.
The spectacle, though always surprising, was not new to me. So my reply to the question put to me was, that these organisms were what biologists call monads, and though they might be animals, it was also possible that they might, like the Bacteria, be plants. My friend received my verdict with an expression which showed a sad want of respect for authority. He would as soon believe that a sheep was a plant. Naturally piqued by this want of faith, I have thought a good deal over the matter; and as I still rest in the lame conclusion I originally expressed, and must even now confess that I cannot certainly say whether this creature is an animal or a plant, I think it may be well to state the grounds of my hesitation at length. But, in the first place, in order that I may conveniently distinguish this "monad" from the multitude of other things which go by the same designation, I must give it a name of its own. I think (though for reasons which need not be stated at present, I am not quite sure) that it is identical with the species Monas lens, as defined by the eminent French microscopist Dujardin, though his magnifying power was probably insufficient to enable him to see that it is curiously like a much larger form of monad which he has named Heteromita. I shall, therefore, call it not Monas, but Heteromita lens.
I have been unable to devote to my Heteromita the prolonged study needful to work out its whole history, which would involve weeks, or it may be months of unremitting attention. But I the less regret this circumstance, as some remarkable observations recently published by Messrs. Dallinger and Drysdale on certain monads, relate, in part, to a form so similar to my Heteromita lens, that the history of the one may be used to illustrate that of the other. These most patient and painstaking observers, who employed the highest attainable powers of the microscope and, relieving one another, kept watch day and night over the same individual monads, have been enabled to trace out the whole history of their Heteromita; which they found in infusions of the heads of fishes of the cod tribe.
Of the four monads described and figured by these investigators one, as I have said, very closely resembles Heteromita lens in every particular, except that it has a separately distinguishable central particle or "nucleus," which is not certainly to be made out in Heteromita lens; and that nothing is said by Messrs. Dallinger and Drysdale of the existence of a contractile vacuole in this monad, though they describe it in another.
Their Heteromita, however, multiplied rapidly by fission. Sometimes a transverse constriction appeared; the hinder half developed a new cilium, and the hinder cilium gradually split from its base to its free end, until it was divided into two; a process which, considering the fact that this fine filament cannot be much more than .00001 of an inch in diameter, is wonderful enough. The constriction of the body extended inwards until the two portions were united by a narrow isthmus; finally they separated, and each swam away by itself, a complete Heteromita, provided with its two cilia. Sometimes the constriction took a longitudinal direction, with the same ultimate result. In each case the process occupied not more than six or seven minutes. At this rate, a single Heteromita would give rise to a thousand like itself in the course of an hour, to about a million in two hours, and to a number greater than the generally assumed number of human beings now living in the world in three hours; or, if we give each Heteromita an hour's enjoyment of individual existence, the same result will be obtained in about a day. The apparent suddenness of the appearance of multitudes of such organisms as these in any nutritive fluid to which one obtains access, is thus easily explained.
During these processes of multiplication ​by fission the Heteromita remains active; but sometimes another mode of fission occurs. The body becomes rounded and quiescent, or nearly so; and while in this resting state divides into two portions, each of which is rapidly converted into an active Heteromita.
A still more remarkable phenomenon is that kind of multiplication which is preceded by the union of two monads, by a process which is termed conjugation. Two active Heteromitæ become applied to one another, and then slowly and gradually coalesce into one body. The two nuclei run into one; and the mass resulting from the conjugation of the two Heteromitæ, thus fused together, has a triangular form. The two pairs of cilia are to be seen, for some time, at two of the angles, which answer to the small ends of the conjoined monads; but they ultimately vanish and the twin organism, in which all visible traces of organization have disappeared, falls into a state of rest. Sudden wave-like movements of its substance next occur: and, in a short time, the apices of the triangular mass burst, and give exit to a dense yellowish, glairy fluid filled with minute granules. This process, which, it will be observed, involves the actual confluence and mixture of the substance of two distinct organisms, is effected in the space of about two hours.
The authors whom I quote say that they "cannot express the excessive minuteness of the granules in question, and they estimate their diameter at less than .000005 of an inch. Under the highest powers of the microscope at present applicable such specks are hardly discernible. Nevertheless, particles of this size are massive when compared to physical molecules; whence there is no reason to doubt that each, small as it is, may have a molecular structure sufficiently complex to give rise to the phenomena of life. And, as a matter of fact, by patient watching of the place at which these infinitesimal living particles were discharged, our observers assured themselves of their growth and development into new monads. These, in about four hours from their being set free, had attained a sixth of the length of the parent, with the characteristic cilia, though at first they were quite motionless; and in four hours more they had attained the dimensions and exhibited all the activity of the adult. These inconceivably minute particles are therefore the germs of the Heteromita; and from the dimensions of these germs it is easily shown that the body formed by conjugagation may, at a low estimate, have given exit to thirty thousand of them; a result of a matrimonial process whereby the contracting parties, without a metaphor, "become one flesh," enough to make a Malthusian despair of the future of the universe.
I am not aware that the investigators from whom I have borrowed this history have endeavoured to ascertain whether their monads take solid nutriment or not; so that though they help us very much to fill up the blanks in the history of my Heteromita, their observations throw no light on the problem we are trying to solve — is it an animal or is it a plant?
Undoubtedly it is possible to bring forward very strong arguments in favour of regarding Heteromita as a plant.
For example, there is a fungus, an obscure and almost microscopic mould, termed Peronospora infestans. Like many other fungi, the Peronosporæ are parasitic upon other plants; and this particular Peronospora happens to have attained much notoriety and political importance, in a way not without a parallel in the career of notorious politicians, namely, by reason of the frightful mischief it has done to mankind. For it is this fungus which is the cause of the potato-disease; and, therefore Peronospora infestans (doubtless of exclusively Saxon origin, though not accurately known to be so) brought about the Irish famine. The plants afflicted with the malady are found to be infested by a mould, consisting of fine tubular filaments, termed hyphæ, ​which burrow through the substance of the potato-plant, and appropriate to themselves the substance of their host; while, at the same time, directly or indirectly, they set up chemical changes by which even its woody frame-work becomes blackened, sodden, and withered.
In structure, however, the Peronospora is as much a mould as the common Penicillium; and just as the Penicillium multiplies by the breaking up of its hyphæ into separate rounded bodies, the spores; so, in the Peronosj5ora, certain of the hyphæ grow out into the air through the interstices of the superficial cells of the potato-plant, and develop spores. Each of these hyphæ usually gives off several branches. The ends of the branches dilate and become closed sacs, which eventually drop off as spores. The spores falling on some part of the same potato-plant, or carried by the wind to another, may at once germinate, throwing out tubular prolongations which become hyphæ, and burrow into the substance of the plant attacked. Rut more commonly, the contents of the spore divide into six or eight separate portions. The coat of the spore gives way, and each portion then emerges as an independent organism, which has the shape of a bean, rather narrower at one end than the other, convex on one side, and depressed or concave on the opposite. From the depression, two long and delicate cilia proceed, one shorter than the other, and directed forwards. Close to the origin of those cilia, in the substance of the body, is a regularly pulsating contractile vacuole. The shorter cilium vibrates actively, and effects the locomotion of the organism, while the other trails behind; the whole body rolling on its axis with its pointed end forwards.
The eminent botanist, De Bary, who was not thinking of our problem, tells us, in describing the movements of these "Zoospores," that, as they swim about, "foreign bodies are carefully avoided, and the whole movement has a deceptive likeness to the voluntary changes of place which are observed in microscopic animals."
After swarming about in this way in the moisture on the surface of a leaf or stem (which, film though it may be, is an ocean to such a fish) for half an hour, more or less, the movement of the zoospore becomes slower, and is limited to a slow turning upon its axis; without change of place. It then becomes quite quiet, the cilia disappear, it assumes a spherical form, and surrounds itself with a distinct, though delicate, membranous coat. A protuberance then grows out from one side of the sphere, and, rapidly increasing in length, assumes the character of a hypha. The latter penetrates into the substance of the potato-plant, either by entering a stomate or by boring through the wall of an epidermic cell, and ramifies, as a mycelium, in the substance of the plant, destroying the tissues with which it comes in contact. As these processes of multiplication take place very rapidly, millions of spores are soon set free from a single infested plant; and from their minuteness they are readily transported by the gentlest breeze. Since again, the zoospores set free from each spore, in virtue of their powers of locomotion, swiftly disperse themselves over the surface, it is no wonder that the infection, once started, soon spreads from field to field, and extends its ravages over a whole country.
However, it does not enter into my present plan to treat of the potato-disease, instructively as its history bears upon that of other epidemics; and I have selected the case of the Peronospora simply because it affords an example of an organism, which, in one stage of its existence, is truly a "monad," indistinguishable by any important character from our Heteromita, and extraordinarily like it in some respects. And yet this monad can be traced, step by step, through the series of metamorphoses which I have described, until it assumes the features of an organism, which is as much a plant as an oak or an elm is.
Moreover it would be possible to pursue the analogy further. Under certain circumstances, a process of conjugation takes place in the Peronospora. Two separate portions of its protoplasm become fused together, surround themselves with a thick coat, and give rise to a sort of vegetable egg called an oospore. After a period of rest, the contents of the oospore break up into a number of zoospores like those already described, each of which, after a period of activity, germinates in the ordinary way. This process obviously corresponds with the conjugation and subsequent setting free of germs in the Heteromita.
But it may be said that the Peronospora is, after all, a questionable sort of plant; that it seems to be wanting in the manufacturing power, selected as the main distinctive character of vegetable life; or at any rate, that there is no proof that it does not get its protein-matter ready made from the potato-plant.
Let us, therefore, take a case which is not open to these objections.
There are some small plants known to botanists as members of the genus Coleochæte, which, without being truly parasitic, grow upon certain water-weeds, as lichens grow upon trees. The little plant has the form of an elegant green star, the branching arms of which are divided into cells. Its greenness is due to its chlorophyll, and it undoubtedly has the manufacturing power in full degree, decomposing carbonic acid and setting free oxygen under the influence of sunlight.
But the protoplasmic contents of some of the cells of which the plant is made up occasionally divide, by a method similar to that which effects the division of the contents of the Poronosptora spore; and the severed portions are then set free as active monad-like zoospores. Each is oval and is provided at one extremity with two long active cilia. Propelled by these, ​it swims about for a longer or shorter time, but at length comes to a state of rest and gradually grows into a Coleochæte.
Moreover, as in the Peronospora, conjugation may take place and result in an oospore; the contents of which divide and are set free as monadiform germs.
If the whole history of the zoospores of Peronospora and Coleochæte were unknown, they would undoubtedly be classed among "monads" with the same right as Heteromita; why then may not Heteromita be a plant, even though the cycle of forms through which it passes shows no terms quite so complex as those which occur in Peronospora and Coleochæte? And, in fact, there are some green organisms, in every respect characteristically plants, such as Chlamydomonas, and the common Volvox, or so-called "globe animalcule," which run through a cycle of forms of just the same simple character as those of Heteromita.
The name of Chlamydomonas is applied to certain microscopic green bodies, each of which consists of a protoplasmic central substance invested by a structureless sac. The latter contains cellulose, as in ordinary plants; and the chlorophyll which gives the green colour enables the Chlamydomonas to decompose carbonic acid and fix carbon, as they do. Two long cilia protrude through the cell-wall, and effect the rapid locomotion of this "monad," which, in all respects except its mobility, is characteristically a plant.
Under ordinary circumstances the Chlamydomonas multiplies by simple fission, each splitting into two or into four parts, which separate and become independent organisms. Sometimes, however, the Chlamydomonas divides into eight parts, each of which is provided with four, instead of two cilia. These zoospores conjugate in pairs, and give rise to quiescent bodies, which multiply by division, and eventually pass into the active state.
Thus, so far as outward form and the general character of the cycle of modifications through which the organism passes in the course of its life are concerned, the resemblance between Chlamydomonas and Heteromita is of the closest description. And on the face of the matter there is no ground for refusing to admit that Heteromita may be related to Chlamydomonas, as the colourless fungus is to the green alga. Volvox may be com pared to a hollow sphere, the wall of which is made up of coherent chlamydomonads; and which progresses with a rotating motion effected by the paddling of the multitudinous pairs of cilia which project from its surface. Each Volvox-monad has a contractile vacuole like that of Heteromita lens; and moreover possesses a red pigment spot like the simplest form of eye known among animals.
The methods of fissive multiplication and of conjugation observed in the monads of this locomotive globe are essentially similar to those observed in Chlamydomonas; and though a hard battle has been fought over it, Volvox is now finally surrendered to the botanists.
Thus there is really no reason why Heteromita may not be a plant; and this conclusion would be very satisfactory, if it were not equally easy to show that there is really no reason why it should not be an animal.
For there are numerous organisms presenting the closest resemblance to Heteromita, and, like it, grouped under the general name of "monads," which, nevertheless, can be observed to take in solid nutriment, and which therefore have a virtual, if not an actual, mouth and digestive cavity, and thus come under Cuvier's definition of an animal. Numerous forms of such animals have been described by Ehrenberg, Dujardin, H. James Clark and other writers on the infusoria.
Indeed, in another infusion of hay in which my Heteromita lens occurred, there were innumerable infusorial animalcules belonging to the well-known species Colpoda cucullus.
Full-sized specimens of this animalcule attain a length of between one three-hundredth or .0025 of an inch, so that it may have ten times the length and a thousand times the mass of a Heteromita. In shape it is not altogether unlike Heteromita. The small end, however, is not produced into one long cilium but the general surface of the body is covered with small actively vibrating ciliary organs, which are only longest at the small end. At the point which answers to that from which the two cilia arise in Heteromita, there is a conical depression, the mouth; and in young specimens a tapering filament, which reminds one of the posterior cilium of Heteromita, projects from this region.
The body consists of a soft granular protoplasmic substance, the middle of which is occupied by a large oval mass ​called the "nucleus;" while, at its hinder end, is a "contractile vacuole," conspicuous by its regular rhythmic appearances and disappearances. Obviously, although the Colpoda is not a monad, it differs from one only in subordinate details. Moreover, under certain conditions, it becomes quiescent, incloses itself in a delicate case or cyst, and then divides into two, four, or more portions, which are eventually set free and swim about as active Colpodæ.
But this creature is an unmistakable animal, and full-sized Colpodæ may be fed as easily as one feeds chickens. It is only needful to diffuse very finely ground carmine through the water in which they live, and, in a very short time, the bodies of the Colpodæ are stuffed with the deeply coloured granules of the pigment.
And if this were not sufficient evidence of the animality of Colpoda, there comes the fact that it is even more similar to another well-known ‘animalcule, Paramæcium, than it is to a monad. But Paramæcium is so huge a creature compared with those hitherto discussed — it reaches one one-hundred-and-twentieth of an inch or more in length — that there is no difficulty in making out its organization in detail; and in proving that it is not only an animal, but that it is an animal which possesses a somewhat complicated organization. For example, the surface layer of its body is different in structure from the deeper parts. There are two contractile vacuoles, from each of which radiates a system of vessel-like canals; and not only is there a conical depression continuous with a tube, which serve, as mouth and gullet, but the food ingested takes a definite course and refuse is rejected from a definite region. Nothing is easier than to feed these animals and to watch the particles of indigo or carmine accumulate at the lower end of the gullet. From this they gradually project, surrounded by a ball of water, which at length passes with a jerk, oddly simulating a gulp, into the pulpy central substance of the body, there to circulate up one side and down the other, until its contents are digested and assimilated. Nevertheless, this complex animal multiplies by division, as the monad does, and, like the monad, undergoes conjugation. It stands in the same relation to Heteromita on the animal side, as Coleochæte does on the plant side. Start from either, and such an insensible series of gradations leads to the monad that it is impossible to say at any stage of the progress — here the line between the animal and the plant must be drawn.
There is reason to think that certain organisms which pass through a monad stage of existence, such as the Myxomycetes, are, at one time of their lives, dependent upon external sources for their protein-matter, or are animals; and at another period manufacture it, or are plants. And seeing that the whole progress of modern investigation is in favour of the doctrine of continuity, it is a fair and probable speculation — though only a speculation — that, as there are some plants which can manufacture protein out of such apparently intractable mineral matters as carbonic acid, water, nitrate of ammonia, and metallic salts; while others need to be supplied with their carbon and nitrogen in the somewhat less raw form of tartrate of ammonia and allied compounds; so there may be yet others, as is possibly the case with the true parasitic plants, which can only manage to put together materials still better prepared — still more nearly approximated to protein — until we arrive at such organisms as the Psorospermiæ and the Panhistophyton, which are as much animal as vegetable in structure, but are animal in their dependence on other organisms for their food.
The singular circumstance observed by Meyer, that the Torula of yeast, though an indubitable plant, still flourishes most vigorously when supplied with the complex nitrogenous substance, pepsin; the probability that the Peronospora is nourished directly by the protoplasm of the potato-plant; and the wonderful facts which have recently been brought to light respecting insectivorous plants, all favour this view; and tend to the conclusion that the difference between animal and plant is one of degree rather than of kind; and that the problem whether, in a given case, an organism is an animal or a plant, may be essentially insoluble.
* ↑ Darwin, "Insectivorous Plants," p. 289.
* ↑ I purposely assume that the air with which the bean is supplied in the case stated contains no ammoniacal salts.
* ↑ "Researches in the Life-history of a Cercomonad: a Leason in Biogenesis," and "Further Researches in the Life-history of the Monads." — Monthly Microscopical Journal, 1873.
* ↑ Excellently described by Stein, almost all of whose statements I have verified.