Form and Function: A Contribution to the History of Animal Morphology

by E. S. Russell

CHAPTER VIII

TRANSCENDENTAL ANATOMY IN ENGLAND—RICHARD OWEN

Richard Owen is the epigonos of transcendental morphology; in him its guiding ideas find clear expression, and in his writings are no half-truths struggling for utterance.

Fig. 4.—Ideal Typical Vertebra. (After Owen.)

But he was, though a staunch transcendentalist, an eclectic of the older ideas current in his time; for he picked out what was best in the older systems—Cuvier's teleology, Geoffroy's principle of connections, Oken's idea of the serial repetition of parts. In particular, he assimilated the teaching of Cuvier, the great opponent of the transcendentalists, and reconciled it in part with his own transcendentalism. His main theoretical views are to be found in his volume On the Archetype and Homologies of the Vertebrate Skeleton (London, 1848). The master-idea of the book is that the vertebrate skeleton consists of a series of comparable segments, each of which Owen calls a vertebra.

Fig 5.—Natural Typical Vertebra; Thorax of a Bird. (After Owen.)

His definition of a vertebra is, "one of those segments of the endo-skeleton which constitute the axis of the body, and the protecting canals of the nervous and vascular trunks" (p. 81). The parts of a typical vertebra are shown in [Fig. 4], which is copied from Owen's Fig. 14.

In Fig. 5 (page 103) is shown an actual vertebra, as Owen conceives it, the "vertebra" being that of a bird.

A segment of sternum is included as the "hæmal spine" of the vertebra (hs); the vertebral rib is the "pleurapophysis" (pl); the sternal rib the "hæmapophysis" (h); the uncinate process of the vertebral rib is known as the "diverging appendage" (a). The whole vertebrate skeleton is composed of a series of vertebræ which show these typical parts. We arrive thus at the conception of an "Archetype" of the vertebrate skeleton, such as is represented in [Fig. 6.]

The archetype is only a scheme of what is usually constant in the vertebrate skeleton, and both the number and the arrangement of the bones in any real Vertebrate are subject to variation. "It has been abundantly proved," Owen writes, towards the end of his volume, "that the idea of a natural segment (vertebra) of the endoskeleton does not necessarily involve the presence of a particular number of pieces, or even a determinate and unchangeable arrangement of them. The great object of my present labour has been to deduce ... the relative value and constancy of the different vertebral elements, and to trace the kind and extent of their variations within the limits of a plain and obvious maintenance of a typical character" (p. 146).

It goes without saying that Owen considered the skull to be formed of vertebræ—the vertebral theory of the skull was, in his system, a deduction from the vertebral theory of the skeleton. He recognised four cranial vertebræ; the arrangement of them, and the relation of their constituent bones to the parts of the typical vertebra are shown in the table appearing on page 106. So far as their first three elements are concerned, these vertebræ are practically identical with the vertebræ distinguished in the classical vertebral theory of the skull, as enunciated by Oken. A divergence appears with the determination of the other elements of the vertebræ. The upper and lower jaws are associated with the nasal and frontal vertebræ respectively, not however as limbs of the head, but as constituent elements of these vertebræ. In the same way the hyoid apparatus is part and parcel of the parietal vertebra, and the pectoral girdle and fore-limbs part of the occipital vertebra.

Fig. 6.—The Archetype of the Vertebrate Skeleton. (After Owen.)

Cranial Vertebræ.[164] (After Owen, 1848, p. 165.)

Vertebræ.	Occipital.	Parietal.	Frontal.	Nasal.
Centra.	Basioccipital.	Basisphenoid.	Presphenoid.	Vomer.
Neurapopbyses.	Exoccipital.	Alisphenoid.	Orbitosphenoid.	Prefrontal.
Neural Spines.	Supraoccipital.	Parietal.	Frontal.	Nasal.
Parapopbyses.	Paroccipital.	Mastoid.	Postfrontal.	None.
Pleurapophyses.	Scapular.	Stylohyal.	Tympanic.	Palatal.
Hæmapophyses.	Coracoid.	Ceratohyal.	Articular.	Maxillary.
Hæmal Spines.	Episternum.	Basihyal.	Dentary.	Premaxillary.
Diverging Appendage.	Fore-limb or Fin.	Branchio-stegals.	Operculum.	Pterygoid and Zygoma.

Owen's reasons for considering the pectoral girdle and the fore-limb part of the occipital vertebra are as follows. In fish the pectoral girdle is slung to the skull by means of the post-temporal bone (supra-scapula, according to Owen) which abuts on the occipital arch. In Lepidosiren, whose skeleton resembles the archetype in many ways, the pectoral girdle is likewise attached to the occipital segment.

In most other Vertebrates the pectoral girdle has shifted backwards along the vertebral column, by a "metastasis" (Geoffroy) similar to that by which the pelvic fins in many fish have shifted up close to the pectoral girdle. The scapula (with supra-scapula) is the pleurapophysis, the coracoid the hæmapophysis, of the occipital vertebra. The clavicle is homologised with the slender bone in fish now known as the post-clavicle, which shows a connection with the first or atlas vertebra of the vertebral column, forming, according to Owen, the hæmapophysis of the atlas. Owen considers it no objection to this view that in other Vertebrates the clavicle is anterior to the coracoid—"its anterior position to the coracoid in the air-breathing Vertebrata is no valid argument against the determination, since in these we have shown that the true scapular arch is displaced backwards" (On the Nature of Limbs, p. 63, London, 1849). In the pelvic girdle the ilium corresponds to the scapula, the ischium to the coracoid, the pubis to the clavicle. Hence the ilium is a pleurapophysis, the ischium and pubis are both hæmapophyses. The fore-limb is the developed "appendage" of the occipital vertebra, the hind-limb the developed "appendage" of the pelvic vertebra. They are serially homologous with, for example, the uncinate processes of the ribs in birds (see Figs. 5 and 6). The fore-limb is a simple filament in Lepidosiren, and presents few joints in Proteus and Amphiuma; in other air-breathing Vertebrates it shows a more complete development, the humerus, radius and ulna, and the bones of the wrist and hand becoming differentiated out.

As the fore-limb is equivalent to a single bone of the archetype, it is said to be, in its developed state, "teleologically compound" (p. 103).

Since in the archetype every vertebra has its appendage, more than two pairs of locomotory limbs might have been developed. "Any given appendage might have been the seat of such developments as convert that of the pelvic arch into a locomotive limb; and the true insight into the general homology of limbs leads us to recognise many potential pairs in the typical endoskeleton. The possible and conceivable modifications of the vertebrate archetype are far from having been exhausted in the forms which have hitherto been recognised, from the primæval fishes of the palæozoic ocean of this planet up to the present time" (p. 102). It is not of the essence of the vertebrate type to be tetrapodal.

In determining homologies Owen remained true to Geoffroy's principle of connections. Speaking of an attempt which had been made to determine homologies by the mode of development, he writes, "There exists doubtless a close general resemblance in the mode of development of homologous parts; but this is subject to modification, like the forms, proportions, functions, and very substance of such parts, without their essential homological relationships being thereby obliterated. These relationships are mainly, if not wholly, determined by the relative position and connection of the parts, and may exist independently of form, proportions, substance, function and similarity of development. But the connections must be sought for at every period of development, and the changes of relative position, if any, during growth, must be compared with the connections which the part presents in the classes where vegetative repetition is greatest and adaptive modification least" (p. 6). It is interesting to note that in Owen's opinion comparative anatomy explains embryology. Thus the scapula, which is the pleurapophysis of the occipital vertebra, is vertical on its first appearance in the embryo of tetrapoda, and lies close up to the head (On the Nature of Limbs, p. 49)—the embryo shows a greater resemblance to the archetype than the adult. "We perceive a return to it, as it were, in the early phases of development of the highest organised of the actually existing species, or we ought rather to say that development starts from the old point; and thus, in regard to the scapula, we can explain the constancy of its first appearance close to the head, whether in the human embryo or in that of the swan, also its vertical position to the axis of the spinal column, by its general homology as the rib or 'pleurapophysis' of the occipital vertebra" (Limbs, p. 56).

We owe to Owen the first clear distinction between "homologous" and "analogous" organs; it was he who first proposed the terms "homologue" and "analogue," which he defined as follows:—"Analogue. A part or organ in one animal which has the same function as another part or organ in a different animal." "Homologue. The same organ in different animals under every variety of form and function."[165]

He introduced also useful distinctions between Special, General, and Serial Homology. "The relations of homology," he writes, "are of three kinds: the first is that above defined, viz., the correspondency of a part or organ, determined by its relative position and connections, with a part or organ in a different animal; the determination of which homology indicates that such animals are constructed on a common type; when, for example, the correspondence of the basilar process of the human occipital bone with the distinct bone called 'basi-occipital' in a fish or crocodile is shown, the special homology of that process is determined. A higher relation of homology is that in which a part or series of parts stands to the fundamental or general type, and its enunciation involves and implies a knowledge of the type on which a natural group of animals, the Vertebrate, for example, is constructed. Thus when the basilar process of the human occipital bone is determined to be the 'centrum' or 'body' of the last cranial vertebra, its general homology is enunciated.

"If it be admitted that the general type of the vertebrate endoskeleton is rightly represented by the idea of a series of essentially similar segments succeeding each other longitudinally from one end of the body to the other, such segments being for the most part composed of pieces similar in number and arrangement, and though sometimes extremely modified for special functions, yet never so as to wholly mask their typical character—then any given part of one segment may be repeated in the rest of the series, just as one bone may be reproduced in the skeletons of different species, and this kind of repetition or representative relation in the segments of the same skeleton I call 'serial homology'" (p. 7). As an example of serial homology we might take the centra of the vertebræ—the vomer, the presphenoid, the basisphenoid, the basioccipital and the series of centra in the spinal column. Such serially repeated parts are called homotypes (p. 8).

Not all the bones of the vertebrate skeleton are included in the archetype as constituents of the vertebræ. Thus the branchial and pharyngeal arches are accounted part of the splanchnoskeleton, as belonging to the same category as the heart bone of some ruminants, and the ossicles of the stomach in the lobster (p. 70). The ossicles of the ear in mammals are "peculiar mammalian productions in relation to the exalted functions of a special organ of sense" (p. 140, f.n.). This recognition of a possible development of new organs to meet new functions shows unmistakably the influence of Cuvier. Owen was indeed well aware of the importance of the functional aspect of living things, and he often adopted the teleological point of view. As a true morphologist, however, he held that the principle of adaptation does not suffice to explain the existence of special homologies. The ossification of the bones of the skull from separate centres may be purposive in Eutheria, in that it prevents injury to the skull at birth; but how explain on teleological principles the similar ossification from separate centres in marsupials, birds and reptiles? How explain above all the fact that the centres are the same in number and relative position in all these groups? Surely we must accept the idea of an archetype "on which it has pleased the divine Architect to build up certain of his diversified living works" (p. 73).

In his study of centres of ossification, Owen made in point of theory a distinct advance on his predecessors. We saw that Geoffroy recognised the importance of studying the ossification of the skeleton, and that Cuvier accepted such embryological evidence as an aid in determining homologies. Owen pointed out that it was necessary to distinguish between centres of ossification which were teleological in import and such as were purely indicative of homological relationships. Many bones, single in the adult, arise from separate centres of ossification, but we must distinguish between "those centres of ossification that have homological relations, and those that have only teleological ones; i.e., between the separate points of ossification of a human bone which typify vertebral elements, often permanently distinct bones in the lower animals; and the separate points which, without such signification, facilitate the progress of osteogeny, and have for their obvious final cause the well-being of the growing animal" (p. 105). There is, for example, a teleological reason why in mammals and leaping Amphibia (e.g., frogs), the long bones should ossify first at their ends, for the brain is thus protected from concussion; in reptiles that creep there is less danger of concussion, and the long bones ossify in the middle (p. 105). But there is no teleological reason why the coracoid process of the scapula should in all mammals develop from a separate centre. The coracoid is however a real vertebral element (hæmapophysis), and in monotremes, birds and reptiles it is in the adult a large and separate bone. Its ossification from a separate centre in mammals has therefore a homological significance. The scapula in mammals is an example of what Owen calls a "homologically compound" bone. All those bones which are formed by a coalescence of parts answering to distinct elements of the typical vertebra are "homologically compound" (p. 105). On the other hand, "All those bones which represent single vertebral elements are 'teleologically compound' when developed from more than one centre, whether such centres subsequently coalesce, or remain distinct, or even become the subject of individual adaptive modifications, with special joints, muscles, etc., for particular offices" (p. 106). The limb-skeleton, corresponding as it does to a single bone of the archetype, is the typical example of a teleologically compound bone. Owen in his definition of teleological compoundness has combined two kinds of adaptation—(1) temporary adaptation of bones to the exigencies of development, birth and growth (e.g., development of long bones from separate centres); (2) definitive adaptation of a skeletal part to the functions which it has to perform (e.g., teleological structure of limbs). Such adaptations are, so to speak, grafted on the archetype.

Owen's general views on the nature of living things merit some attention. Organic forms, according to Owen, result from the antagonistic working of two principles, of which one brings about a vegetative repetition of structure, while the other, a teleological principle, shapes the living thing to its functions. The former principle is illustrated in the archetype of the vertebrate skeleton, in the segmentation of the Articulates, in the almost mathematical symmetry of Echinoderms, and the actually crystalline spicules of sponges. It is the same principle which causes repetition of the forms of crystals in the inorganic world. "The repetition of similar segments in a vertebral column, and of similar elements in a vertebral segment, is analogous to the repetition of similar crystals as the result of polarising force in the growth of an inorganic body" (p. 171). This "general polarising force" it is which mainly produces the similarity of forms, the repetition of parts, and generally the signs of the unity of organisation. The adaptive or "special organising force" or ἰδέα, on the other hand, produces the diversity of organic beings. In every species these two forces are at work, and the extent to which the general polarising or "vegetative-repetition-force" is subdued by the teleological is an index of the grade of the species.

This view is analogous to the Geoffroyan conception that the diversity of form is limited by the unity of plan. Owen thus ranges himself with Geoffroy against Cuvier, who considered that diversity of form is limited only by the principle of the adaptation of parts.

[164] Owen introduced most of the names of bones now current.

[165] Lectures on Invertebrate Animals, pp. 374, 379, 1843.

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