The Mutual Relations of Forms in Organized Beings

Georges Cuvier

Principle by which this determination is effected.

Fortunately, comparative anatomy possesses a principle, which, when properly developed, enables us to surmount all the obstacles. This principle consists in the mutual relation of forms in organised beings, by means of which, each species may be determined, with perfect certainty, by any fragment of any of its parts.

Every organised being forms a whole—a peculiar system of its own, the parts of which mutually correspond, and concur in producing the same definitive action, by a reciprocal reaction. None of these parts can change in form, without the others also changing; and consequently, each of them, taken separately, indicates and ascertains all the others.

Thus, if the intestines of an animal are so organised as to be fitted for the digestion of flesh only, and that flesh recent, it is necessary that its jaws be so constructed as to fit them for devouring live prey; its claws for seizing and tearing it; its teeth for cutting and dividing it; the whole system of its organs of motion, for pursuing and overtaking it; and its organs of sense for discovering it at a distance. It is even requisite that nature have placed in its brain the instinct necessary for teaching it to conceal itself, and to lay snares for its victims.

Such are the general conditions which nature imposes upon the structure of carnivorous animals; and which every animal of this description must indispensably combine in its constitution, for without them its race could not subsist. But subordinate to these general conditions, there exist others, having relation to the size, the species, and the haunts of the prey for which the animal is adapted; and from each of these particular conditions, there result modifications of detail in the forms which arise from the general conditions. Thus not only the class, but the order, the genus, and even the species, are found expressed in the form of each part.

In fact, in order that the jaw may be able to seize, it must have a certain form of condyle; that the resistance, the moving power, and the fulcrum, should have a certain relative position in regard to each other; and that the temporal muscles should be of a certain size; the hollow or depression, too, in which these muscles are lodged, must have a certain depth; and the zygomatic arch, under which they pass, must not only have a certain degree of convexity, but it must be sufficiently strong to support the action of the masseter.

In order that the animal may be able to carry off its prey, it must have a certain degree of vigour in the muscles which elevate the head; whence there results a determinate form in the vertebrae from which these muscles take their rise, and in the occiput into which they are inserted.

In order that the teeth may be able to cut flesh, they must be sharp-edged, and must be so in a greater or less degree, according as they have flesh more or less exclusively to cut. Their base will be solid, according to the quantity and size of the bones which they have to break. The whole of these circumstances must necessarily influence the development and form of all the parts which contribute to move the jaws.

In order that the paws may be able to seize the prey, there must be a certain degree of mobility in the toes, and a certain degree of strength in the claws, from which there will result determinate forms in all the phalanges, and a corresponding distribution of muscles and tendons. The fore-arm, or cubitus, must possess a certain facility of turning, from which there will also result determinate forms in the bones of which it is composed. But the bones of the cubitus being articulated to the humerus, a change in the proportions of the former, will necessarily induce a corresponding change in the latter. The shoulder-bones must have a certain degree of firmness in such animals as make use of their fore-legs for seizing, and from this there must also result a certain peculiarity in their form. The play of all these parts will require certain proportions in all their muscles, and the impressions made by these muscles so proportioned, will determine still more particularly the forms of the bones.

It is easy to see that similar conclusions may be drawn with regard to the posterior extremities which contribute to the rapidity of the general motions; with regard to the composition of the trunk, and the forms of the vertebrae, which exert an influence upon the facility and flexibility of these motions; and, lastly, with regard to the forms of the bones of the nose, of the orbit, and of the ear, the connection of which with the perfection of the senses of smell, sight, and hearing, is evident. In a word, the form of the tooth regulates the forms of the condyle, of the scapula, and of the claws, in the same manner as the equation of a curve regulates all its properties; and as, by taking each property separately for the base of a particular equation, we find both the ordinary equation, and all the other properties whatever; so, the claw, the scapula, the condyle, the femur, and all the other bones taken separately, give the tooth, or are reciprocally given by it; and thus, by commencing with any one of these bones, a person who possesses an accurate knowledge of the laws of organic economy, may reconstruct the whole animal.

This principle seems sufficiently evident, in the general acceptation in which it is here taken, and does not require any fuller demonstration; but when it comes to be applied, there will be found many cases where our theoretical knowledge of the relations of forms will not be sufficient, unless it be supported by observation and experience.

For example, we are well aware, that hoofed animals must all be herbivorous, since they have no means of seizing prey. It is also evident that, having no other use to make of their fore-legs than to support their body, they do not require a shoulder so vigorously organised as that of carnivorous animals; they have, therefore, no acromion or clavicle, and their shoulder-blades are narrow. Having also no occasion to turn their fore-arm, their radius is united to the ulna by ossification, or at least articulated by a ginglimus or hinge-joint, and not by arthrodia or ball and socket, to the humerus. Their food being herbaceous, will require teeth furnished with flat surfaces, for bruising seeds and plants. The crown of the teeth must also be unequal, and, for this purpose, must be composed of parts alternately consisting of bone and of enamel. Teeth of this structure necessarily require horizontal motions to enable them to triturate the food; and hence the condyle of the jaw cannot be so strictly confined within its articulating cavity as in the carnivorous animals, but must be flattened, and thus correspond with a more or less flattened surface of the temporal bones. Further, the temporal fossa, which will only have a small muscle to contain, will be narrower, and not so shallow, as that of carnivorous animals. All these circumstances are deducible from each other, according to their greater or less generality, and in such a manner that some of them are essential and exclusively peculiar to hoofed animals, while others, although equally necessary in these animals, are not entirely peculiar to them, but may occur in other animals also, where the rest of the conditions will permit their existence. [p.422]

If we proceed to consider the orders or subdivisions of the class of hoofed animals, and examine what modifications the general conditions undergo, or rather what particular conditions are conjoined with them, according to the respective characters of these orders, the reasons of these subordinate conditions begin to appear less obvious. We can still easily conceive, in general, the necessity of a more complicated system of digestive organs in those species which have a more imperfect masticatory system; and hence we may presume, that these latter must be rather ruminating animals, in which there is wanting such or such an order of teeth; and may also deduce from the same consideration, the necessity of a certain form of the oesophagus, and of corresponding forms in the vertebrae of the neck, etc. But I doubt whether it would have been discovered, independently of actual observation, that the ruminating animals should all have cloven hoofs, and that they should be the only animals having them; that there should be horns on the forehead in this class alone; or that such of them as have sharp canine teeth, should, in general, have no horns.

However, since these relations are constant, we may be assured that they have a sufficient cause; but as we are not acquainted with that cause, we must supply the defect of theory by means of observation, and in this way establish empirical laws which become nearly as certain as those deduced from rational principles, when rounded upon observations, the authenticity of which is proved by frequent repetition. Hence, at the present day, any one who observes only the print of a cloven foot, may conclude that the animal which left this impression ruminates; and this conclusion is quite as certain as any other in physics, or in moral philosophy. This simple footmark, therefore, indicates at once to the observer the forms of the teeth, of the jaws, of the vertebrae, of all the bones of the legs, thighs, shoulders, and pelvis of the animal which had passed. It is a surer mark than all those of Zadig. That there are secret reasons, however, for all these relations, is what observation alone is sufficient to show, independently of any general principles of philosophy.

In fact, when we construct a table of these relations, we remark not only a specific constancy, if the expression may be allowed, between a particular form of a particular organ, and some other form of a different organ; but we also perceive a classic constancy of conformation, and a corresponding gradation, in the development of these two organs, which demonstrate their mutual influence, almost as well as the most perfect deduction of reason. [p.423]

For example, the dentary system of the hoofed animals, which are not ruminant, is in general more perfect than that of the cloven-footed or ruminating animals, because the former have either incisors, or canine teeth, and almost always both in each jaw; and the structure of their foot is in general more complicated, because they have more toes or claws, or their phalanges less enveloped in the hoof,—or a greater number of distinct bones in the metacarpus and metatarsus—or more numerous tarsal bones—or a fibula more distinct from the tibia—or, lastly, that all these circumstances are often united in the same species of animals.

It is impossible to assign reasons for these relations; but we are certain that they are not the effects of chance, because, whenever a cloven-footed animal manifests, in the arrangement of its teeth some tendency to approach the animals we now speak of, also manifests a similar tendency in the arrangement of its feet. Thus the camels, which have canine teeth, and even two or four incisors in the upper jaw, have an additional bone in the tarsus, because their scaphoid bone is not united to the cuboid, and they have very small hoofs, with corresponding phalanges. The musk animals, whose canine teeth are much developed, have a distinct fibula along the whole length of their tibia; while the ether cloven-footed animals have only, in place of a fibula, a small bone articulated at the lower end of the tibia. There is, therefore, a constant harmony between two organs apparently having no connection; and the gradations of their forms preserve an uninterrupted correspondence, even in those cases in which we cannot account for their relations.

Now, by thus adopting the method of observation as a supplementary means, when theory is no longer able to direct our views, we arrive at astonishing results. The smallest articulating surface of bone, or the smallest apophysis, has a determinate character, relative to the class, the order, the genus, and the species to which it belonged; insomuch, that when one possesses merely a well preserved extremity of a bone, he can, by careful examination, and the aid of a tolerable analogical knowledge, and of accurate comparison, determine all these things with as much certainty as if he had the entire animal before him. I have often made trial of this method upon portions of known animals, before reposing full confidence upon it, in regard to fossil remains; and it has always proved so completely satisfactory that I have no longer any doubts regarding the certainty of the results which it has afforded me.

It is true, that I have enjoyed all the advantages which were necessary for the undertaking; and that my favourable situation, in the Museum of Natural History at Paris, and assiduous research for nearly thirty years, have procured me skeletons of all the genera and sub-genera of quadrupeds, and even of many species in some genera, and of several varieties of some species. With such means, it was easy for me to multiply my comparisons, and to verify in all their details the applications which I have made of the various laws deducible from such circumstances as have been stated.

We cannot here enter into a more lengthened detail of this method, and must refer to the large work on Comparative Anatomy, in which all its rules will be found. In the mean time, an intelligent reader may gather a great number of these from the work upon Fossil Bones, if he take the trouble of attending to all the applications of them which we have there made. He will see, that it is by this method alone that we are guided, and that it has almost always sufficed for referring each bone to its species, when it was a living species—to its genus, when it was an unknown species—to its order, when it was a new genus—and to its class, when it belonged to an order not hitherto established—and to assign it, in the three last cases, the proper characters for distinguishing it from the nearest resembling orders, genera, and species. Before the commencement of our researches, naturalists had done no more than this with regard to animals, which they had the opportunity of examining in their entire state. Yet, in this manner, we have determined and classed the remains of more than a hundred and fifty mammiferous and oviparous quadrupeds.