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ON THE BORDER TERRITORY BETWEEN THE ANIMAL AND THE VEGETABLE KINGDOMS [1876] 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 Organised Beings into Animals and Vegetables," in which the question is treated with that comprehensiveness of knowledge and clear critical judgment which characterise 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 subdivided 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 organisation 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 be 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 he 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 measurement of the physical 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 _Charoe_ 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 _Tradescantia_ 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 organisation, were called "zoospores."
Even as late as 1845, however, a botanist of Schleiden's eminence dealt 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 recognised 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 realisations 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 researches of the modern chemical investigators 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.
[1] On the other hand, 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.
[Footnote 1: There is every reason to believe that living plants, like living animals, always respire, and, in respiring, absorb oxygen and give off carbonic acid; but, that in green plants exposed to daylight or to the electric light, the quantity of oxygen evolved in consequence of the decomposition of carbonic acid by a special apparatus which green plants possess exceeds that absorbed in the concurrent respiratory process.
] 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 generalisation, 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 metamorphosis 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 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 bilobed leaf of Venus's fly-trap (_Dionoea muscipula_) are three delicate filaments which stand out at right angle 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[2] 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.
[Footnote 2: Darwin, _Insectivorous Plants_, p. 289. ] The reflex action of the snail is the result of the presence of a nervous system in the 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 seeds, 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, has 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; for it is only the green parts of the plant which, under the influence of sunlight, have 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;[3] and the chlorophyll[4] 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.
[Footnote 3: I purposely assume that the air with which the bean is supplied in the case stated contains no ammoniacal salts.
] [Footnote 4: The recent researches of Pringsheim have raised a host of questions as to the exact share taken by chlorophyll in the chemical operations which are effected by the green parts of plants.
It may be that the chlorophyll is only a constant concomitant of the actual deoxidising apparatus.
] 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 saucerful 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 generalisation that plants are essentially characterised 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 generalisation 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.
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