Anatomy of Vertebrates

Sir Richard Owen, the great naturalist, was born July 20, 1804, at

Lancaster, England, and received his early education at the grammar

school of that town. Thence he went to Edinburgh University. In

1826 he was admitted a member of the English College of Surgeons,

and in 1829 was lecturing at St. Bartholomew's Hospital, London,

where he had completed his studies. His "Memoir on the Pearly

Nautillus," published in 1832, placed him, says Huxley, "at a bound

in the front rank of anatomical monographers," and for sixty-two

years the flow of his contributions to scientific literature never

ceased. In 1856 he was appointed to take charge of the natural

history departments of the British Museum, and before long set

forth views as to the inadequacy of the existing accommodation,

which led ultimately to the foundation of the buildings now devoted

to this purpose in South Kensington. Owen died on December 18,

1892. His great book, "Comparative Anatomy and Physiology of the

Vertebrates," was completed in 1868, and since Cuvier's

"Comparative Anatomy," is the most monumental treatise on the

subject by any one man. Although much of the classification adopted

by Owen has not been accepted by other zoologists, yet the work

contains an immense amount of information, most of which was gained

from Owen's own personal observations and dissections.

_I.--Biological Questions of 1830_

At the close of my studies at the Jardin des Plantes, Paris, in 1831, I

returned strongly moved to lines of research bearing upon the then

prevailing phases of thought on some biological questions.

The great master in whose dissecting rooms I was privileged to work held

that species were not permanent as a fact established inductively on a

wide basis of observation, by which comparative osteology had been

created. Camper and Hunter suspected the species might be transitory;

but Cuvier, in defining the characters of his anaplotherium and

palæotherium, etc., proved the fact. Of the relation of past to present

species, Cuvier had not an adequate basis for a decided opinion.

Observation of changes in the relative position of land and sea

suggested to him one condition of the advent of new species on an island

or continent where old species had died out. This view he illustrates by

a hypothetical case of such succession, but expressly states: "I do not

assert that a new creation was necessary to produce the species now

existing, but only that they did not exist in the same regions, and must

have come from elsewhere." Geoffrey Saint Hilaire opposed to Cuvier's

inductive treatment of the question the following expression of belief:

"I have no doubt that existing animals are directly descended from the

animals of the antediluvian world," but added, "it is my belief that the

season has not yet arrived for a really satisfying knowledge of

geology."

The main collateral questions argued in their debates appeared to me to

be the following:

Unity of plan or final purpose, as a governing condition of organic

development?

Series of species, uninterrupted or broken by intervals?

Extinction, cataclysmal or regulated?

Development, by epigenesis or evolution?

Primary life, by miracle or secondary law?

Cuvier held the work of organisation to be guided and governed by final

purpose or adaptation. Geoffrey denied the evidence of design and

contended for the principle which he called "unity of composition," as

the law of organisation. Most of his illustrations were open to the

demonstration of inaccuracy; and the language by which disciples of the

kindred school of Schelling illustrated in the animal structure the

transcendental idea of the whole in every part seemed little better than

mystical jargon. With Cuvier, answerable parts occurred in the

zoological scale because they had to perform similar functions.

As, however, my observations and comparisons accumulated, they enforced

a reconsideration of Cuvier's conclusions. To demonstrate the evidence

of the community of organisation I found the artifice of an archetype

vertebrate animal essential; and from the demonstration of its

principle, which I then satisfied myself was associated with and

dominated by that of "adaptation to purpose," the step was inevitable to

the conception of the operation of a secondary cause of the entire

series of species, such cause being the servant of predetermining

intelligent will.

But besides "derivation" or "filiation" another principle influencing

organisation became recognisable, to which I gave the name of

"irrelative repetition," or "vegetative repetition." The demonstrated

constitution of the vertebrate endoskeleton as a series of essentially

similar segments appeared to me to illustrate the law of irrelative

repetition.

These results of inductive research swayed me in rejecting direct or

miraculous creation, and in recognising a "natural law or secondary

cause" as operative in the production of species "in orderly succession

and progression."

_II.--Succession of Species, Broken or Linked?_

To the hypothesis that existing are modifications of extinct species,

Cuvier replied that traces of modification were due from the fossil

world. "You ought," he said, "to be able to show the intermediate forms

between the palæotherium and existing hoofed quadrupeds."

The progress of palæontology since 1830 has brought to light many

missing links unknown to the founder of the science. The discovery of

the remains of the hipparion supplied one of the links required by

Cuvier, and it is significant that the remains of such three-toed horses

are found only in deposits of that tertiary period which intervene

between the older palæotherian one and the newer strata in which the

modern horse first appears to have lost its lateral hooflets.

The molar series of the horse includes six large complex grinders

individually recognisable by developmental characters. The

representative of the first premolar is minute and soon shod. Its

homologue in palæotherium is functionally developed and retained, that

type-dentition being adhered to. In hipparion this tooth is smaller than

in palæotherium, but functional and permanent. The transitory and

singularly small and simple denticle in the horse exemplifies the

rudiment of an ancestral structure in the same degree as do the hoofless

splint-bones; just as the spurious hoofs dangling therefrom in hipparion

are retained rudiments of the functionally developed lateral hoofs in

the broader foot of palæotherium.

Other missing links of this series of species have also been supplied.

How then is the origin of these intermediate gradations to be

interpreted? If the alternative--species by miracle or by law--be

applied to palæotherium, paloplotherium, anchitherium, hipparion, equus,

I accept the latter without misgiving, and recognise such law as

continuously operative throughout tertiary time.

In respect to its law of operation we may suppose Lamarck to say, "as

the surface of the earth consolidated, the larger and more produced

mid-hoof of the old three-toed pachyderius took a greater share in

sustaining the animal's weight; and more blood being required to meet

the greater demand of the more active mid-toe, it grew; whilst, the

side-toes, losing their share of nourishment and becoming more and more

withdrawn from use, shrank"--and so on. Mr. Darwin, I conceive, would

modify this by saying that some individuals of palæotherium happening to

be born with a larger and longer middle toe, and with shorter and

smaller side-toes, such variety was better adapted to prevailing altered

conditions of the earth's surface than the parental form; and so on,

until finally the extreme equine modifications of foot came to be

"naturally selected." But the hypothesis of appetency and volition, as

of natural selection, are less applicable, less intelligible, in

connection with the changes in the teeth.

I must further observe that to say the palæotherium has graduated into

equus by "natural selection" is an explanation of the process of the

same kind and value as that by which the secretion of bile was

attributed to the "appetency" of the liver for the elements of bile.

One's surprise is that such explanatory devices should not have died out

with the "archeus faber," the "nisus formations," and other

self-deceiving, world-beguiling simulacra of science, with the last

century; and that a resuscitation should have had any success in the

present.

What, then, are the facts on which any reasonable or intelligible

conception can be formed of the mode of operation of the derivative law

exemplified in the series linking on palæotherium to equus? A very

significant one is the following. A modern horse occasionally comes into

the world with the supplementary ancestral hoofs. From Valerius Maximus,

who attributes the variety to Bucephalus downwards, such "polydactyle"

horses have been noted as monsters and marvels. In one of the latest

examples, the inner splint-bone, answering to the second metacarpal of

the pentadactyle foot, supported phalanges and a terminal hoof

resembling the corresponding one in hipparion. And the pairing of horses

with the meterpodials bearing, according to type, phalanges and hoofs

might restore the race of hipparions.

Now, the fact suggesting such possibility teaches that the change would

be sudden and considerable; it opposes the idea that species are

transmuted by minute and slow degrees. It also shows that a species

might originate independently of the operation of any external

influence; that change of structure would precede that of use and

habit; that appetency, impulse, ambient medium, fortuitous fitness of

surrounding circumstances, or a personified "selecting nature" would

have had no share in the transmutative act.

Thus I have been led to recognise species as exemplifying the continuous

operation of natural law, or secondary cause; and that not only

successively but progressively; "from the first embodiment of the

vertebrate idea under its old ichthyic vestment until it became arrayed

in the glorious garb of the human form."

_III.--Extinction--Cataclysmal or Regulated_

If the species of palæothere, paloplothere, anchithere, hipparion, and

horse be severally deemed due to remotely and successively repeated acts

of creation; the successive going out of such species must have been as

miraculous as their coming in. Accordingly, in Cuvier's "Discourse on

Revolutions of the Earth's Surface" we have a section of "Proofs that

these revolutions have been numerous," and another of "Proofs that these

revolutions have been sudden." But as the discoveries of palæontologists

have supplied the links between the species held to have perished by the

cataclysms, so each successive parcel of geological truth has tended to

dissipate the belief in the unusually sudden and violent nature of the

changes recognisable in the earth's surface. In specially directing my

attention to this moot point, whilst engaged in investigations of fossil

remains, I was led to recognise one cause of extinction as being due to

defeat in the contest which the individual of each species had to

maintain against the surrounding agencies which might militate against

its existence. This principle has received a large and most instructive

accession of illustrations from the labours of Charles Darwin; but he

aims to apply it not only to the extinction but to the origin of

species.

Although I fail to recognise proof of the latter bearing of the battle

of life, the concurrence of so much evidence in favour of extinction by

law is, in like measure, corroborative of the truth of the ascription of

the origin of species to a secondary cause.

What spectacle can be more beautiful than that of the inhabitants of the

calm expanse of water of an atoll encircled by its ring of coral rock!

Leaving locomotive frequenters of the calcarious basin out of the

question, we may ask, Was direct creation after the dying out of its

result as a "rugose coral" repeated to constitute the succeeding and

superseding "tabulate coral"? Must we also invoke the miraculous power

to initiate every distinct species of both rugosa and tabulata? These

grand old groups have had their day and are utterly gone. When we

endeavour to conceive or realise such mode of origin, not of them only

but of their manifold successors, the miracle, by the very

multiplication of its manifestations, becomes incredible--inconsistent

with any worthy conception of an all-seeing, all-provident Omnipotence.

Being unable to accept the volitional hypothesis (of Lamarck) or the

selective force exerted by outward circumstances (Darwin), I deem an

innate tendency to deviate from parental type, operating through periods

of adequate duration, to be the most probable way of operation of the

secondary law whereby species have been derived one from another.

According to my derivative hypothesis a change takes place first in the

structure of the animal, and this, when sufficiently advanced, may lead

to modifications of habits. But species owe as little to the accidental

concurrence of environing circumstances as kosmos depends upon a

fortuitous concourse of atoms. A purposive route of development and

change of correlation and inter-dependence, manifesting intelligent

will, is as determinable in the succession of races as in the

development and organisation of the individual.

Derivation holds that every species changes in time, by virtue of

inherent tendencies thereto. Natural selection holds that no such change

can take place without the influence of altered external circumstances

educing or eliciting such change.

Derivation sees among the effects of the innate tendency to change,

irrespective of altered surrounding circumstances, a manifestation of

creative power in the variety and beauty of the results; and, in the

ultimate forthcoming of a being susceptible of appreciating such beauty,

evidence of the preordaining of such relation of power to the

appreciation. Natural selection acknowledges that if power or beauty, in

itself, should be a purpose in creation, it would be absolutely fatal to

it as a hypothesis.

Natural selection sees grandeur in the "view of life, with its several

powers, having been originally breathed by the Creator into a few forms

or into one." Derivation sees, therein, a narrow invocation of a special

miracle and an unworthy limitation of creative power, the grandeur of

which is manifested daily, hourly, in calling into life many forms, by

conversion of physical and chemical into vital modes of force, under as

many diversified conditions of the requisite elements to be so combined.

Natural selection leaves the subsequent origin and succession of species

to the fortuitous concurrence of outward conditions; derivation

recognises a purpose in the defined and preordained course, due to

innate capacity or power of change, by which homogeneously-created

protozoa have risen to the higher forms of plants and animals.

The hypothesis of derivation rests upon conclusions from four great

series of inductively established facts, together with a probable result

of facts of a fifth class; the hypothesis of natural selection totters

on the extension of a conjectural condition explanatory of extinction to

the origination of species, inapplicable in that extension to the

majority of organisms, and not known or observed to apply to the origin

of any species.

_IV.--Epigenesis or Evolution?_

The derivative origin of species, then, being at present the most

admissible one, and the retrospective survey of such species showing

convergence, as time recedes, to more simplified or generalised

organisations, the result to which the suggested train of thought

inevitably leads is very analogous in each instance. If to kosmos or the

mundane system have been allotted powers equivalent to the development

of the several grades of life, may not the demonstrated series of

conversions of force have also included that into the vital form?

In the last century, physiologists were divided as to the principle

guiding the work of organic development.

The "evolutionists" contended that the new being preexisted in a

complete state of formation, needing only to be vivified by impregnation

in order to commence the series of expansions or disencasings,

culminating in the independent individual.

The "epigenesists" held that both the germ and its subsequent organs

were built up of juxtaposed molecules according to the operation of a

developmental force, or "nisus formations."

At the present day the question may seem hardly worth the paper on which

it is referred to. Nevertheless, "pre-existence of germs" and evolution

are logically inseparable from the idea of species by primary

miraculously-created individuals. Cuvier, therefore, maintained both as

firmly as did Haller. In the debates of 1830 I remained the thrall of

that dogma in regard to the origin of single-celled organisms whether in

or out of body. Every result of formfaction, I believed, with most

physiologists, to be the genetic outcome of a pre-existing "cell." The

first was due to miraculous interposition and suspension of ordinary

laws; it contained potentially all future possible cells.

Cell-development exemplified evolution of pre-existing germs, the

progeny of the primary cell. They progagated themselves by

self-division, or by "proliferation" of minutes granules or atoms,

which, when properly nourished, again multiplied by self-division, and

grew to the likeness of the parent cells.

It seems to me more consistent with the present phase of dynamical

science and the observed graduations of living things to suppose the

sarcode or the "protogenal" jelly-speck should be formable through the

concurrence of conditions favouring such combination of their elements,

and involving a change of force productive of their contractions and

extensions, molecular attractions, and repulsions--and the sarcode has

so become, from the period when its irrelative repetitions resulted in

the vast indefinite masses of the "eozoon," exemplifying the earliest

process of "formification" or organic crystallisation--than that all

existing sarcodes or "protogenes" are the result of genetic descent from

a germ or cell due to a primary act of miraculous interposition.

I prefer, while indulging in such speculations, to consider the various

daily nomogeneously developed forms of protozoal or protistal jellies,

sarcodes, and single-celled organisms, to have been as many roots from

which the higher grades have ramified than that the origin of the whole

organic creation is to be referred, as the Egyptian priests did that of

the universe, to a single egg.

Amber or steel, when magnetised, seem to exercise "selection"; they do

not attract all substances alike. A speck of protogenal jelly or

sarcode, if alive, shows analogous relations to certain substances; but

the soft yielding tissue allows the part next the attractive matter to

move thereto, and then, by retraction, to draw such matter into the

sarcodal mass, which overspreads, dissolves, and assimilates it. The

term "living" in the one case is correlative with the term "magnetic" in

the other. A man perceives ripe fruit; he stretches out his hand,

plucks, masticates, swallows, and digests it.

The question then arises whether the difference between such series of

actions in the man and the attractive and assimilative movement of the

amæba be greater or less than the difference between these acts of the

amæba and the attracting and retaining acts of the magnet.

The question, I think, may be put with some confidence as to the quality

of the ultimate reply whether the amæbal phenomena are so much more

different, or so essentially different, from the magnetic phenomena than

they are from the mammalian phenomena, as to necessitate the invocation

of a special miracle for their manifestation. It is analogically

conceivable that the same cause which has endowed His world with power

convertible into magnetic, electric, thermotic and other forms or modes

of force, has also added the conditions of conversion into the vital

force.

From protozoa or protista to plants and animals the graduation is closer

than from magnetised iron to vitalised sarcode. From reflex acts of the

nervous system animals rise to sentient and volitional ones. And with

the ascent are associated brain-cells progressively increasing in size

and complexity. Thought relates to the "brain" of man as does

electricity to the nervous "battery" of the torpedo; both are forms of

force and the results of action of their respective organs.

Each sensation affects a cerebral fibre, and, in so affecting it, gives

it the faculty of repeating the action, wherein memory consists and

sensation in a dream.

If the hypothesis of an abstract entity produces psychological phenomena

by playing upon the brain as a musician upon his instrument be rejected,

and these phenomena be held to be the result of cerebral actions, an

objection is made that the latter view is "materialistic" and adverse to

the notion of an independent, indivisible, "immaterial," mental

principle or soul.

But in the endeavour to comprehend clearly and explain the functions of

the combination of forces called "brain," the physiologist is hindered

and troubled by the views of the nature of those cerebral forces which

the needs of dogmatic theology have imposed on mankind. How long

physiologists would have entertained the notion of a "life," or "vital

principle," as a distinct entity if freed from this baneful influence

may be questioned; but it can be truly affirmed that physiology has now

established and does accept the truth of that statement of Locke--"the

life, whether of a material or immaterial substance, is not the

substance itself, but an affection of it."

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