they show that the horse deviates widely from the general structure of
mammals; and that the horse type is, in many respects, an extreme
modification of the general mammalian plan. The least modified mammals,
in fact, have the radius and ulna, the tibia and fibula, distinct and
separate. They have five distinct and complete digits on each foot, and
no one of these digits is very much larger than the rest. Moreover, in
the least modified mammals, the total number of the teeth is very
generally forty-four, while in horses, the usual number is forty, and in
the absence of the canines, it may be reduced to thirty-six; the incisor
teeth are devoid of the fold seen in those of the horse: the grinders
regularly diminish in size from the middle of the series to its front
end; while their crowns are short, early attain their full length, and
exhibit simple ridges or tubercles, in place of the complex foldings of
the horse's grinders.
Hence the general principles of the hypothesis of evolution lead to the
conclusion that the horse must have been derived from some quadruped
which possessed five complete digits on each foot; which had the bones
of the fore-arm and of the leg complete and separate; and which
possessed forty-four teeth, among which the crowns of the incisors and
grinders had a simple structure; while the latter gradually increased in
size from before backwards, at any rate in the anterior part of the
series, and had short crowns.
And if the horse has been thus evolved, and the remains of the different
stages of its evolution have been preserved, they ought to present us
with a series of forms in which the number of the digits becomes
reduced; the bones of the fore-arm and leg gradually take on the equine
condition; and the form and arrangement of the teeth successively
approximate to those which obtain in existing horses.
Let us turn to the facts, and see how far they fulfil these requirements
of the doctrine of evolution.
In Europe abundant remains of horses are found in the Quaternary and
later Tertiary strata as far as the Pliocene formation. But these
horses, which are so common in the cave-deposits and in the gravels of
Europe, are in all essential respects like existing horses. And that is
true of all the horses of the latter part of the Pliocene epoch. But, in
deposits which belong to the earlier Pliocene and later Miocene epochs,
and which occur in Britain, in France, in Germany, in Greece, in India,
we find animals which are extremely like horses--which, in fact, are so
similar to horses, that you may follow descriptions given in works upon
the anatomy of the horse upon the skeletons of these animals--but which
differ in some important particulars. For example, the structure of
their fore and hind limbs is somewhat different. The bones which, in the
horse, are represented by two splints, imperfect below, are as long as
the middle metacarpal and metatarsal bones; and, attached to the
extremity of each, is a digit with three joints of the same general
character as those of the middle digit, only very much smaller. These
small digits are so disposed that they could have had but very little
functional importance, and they must have been rather of the nature of
the dew-claws, such as are to be found in many ruminant animals. The
_Hipparion_, as the extinct European three-toed horse is called, in
fact, presents a foot similar to that of the American _Protohippus_
(Fig. 9), except that, in the _Hipparion_, the smaller digits are
situated farther back, and are of smaller proportional size, than in the
_Protohippus_.
The ulna is slightly more distinct than in the horse; and the whole
length of it, as a very slender shaft, intimately united with the
radius, is completely traceable. The fibula appears to be in the same
condition as in the horse. The teeth of the _Hipparion_ are essentially
similar to those of the horse, but the pattern of the grinders is in
some respects a little more complex, and there is a depression on the
face of the skull in front of the orbit, which is not seen in existing
horses.
In the earlier Miocene, and perhaps the later Eocene deposits of some
parts of Europe, another extinct animal has been discovered, which
Cuvier, who first described some fragments of it, considered to be a
_Palæotherium_. But as further discoveries threw new light upon its
structure, it was recognised as a distinct genus, under the name of
_Anchitherium_.
In its general characters, the skeleton of _Anchitherium_ is very
similar to that of the horse. In fact, Lartet and De Blainville called
it _Palæotherium equinum_ or _hippoides_; and De Christol, in 1847, said
that it differed from _Hipparion_ in little more than the characters of
its teeth, and gave it the name of _Hipparitherium_. Each foot possesses
three complete toes; while the lateral toes are much larger in
proportion to the middle toe than in _Hipparion_, and doubtless rested
on the ground in ordinary locomotion.
The ulna is complete and quite distinct from the radius, though firmly
united with the latter. The fibula seems also to have been complete. Its
lower end, though intimately united with that of the tibia, is clearly
marked off from the latter bone.
There are forty-four teeth. The incisors have no strong pit. The canines
seem to have been well developed in both sexes. The first of the seven
grinders, which, as I have said, is frequently absent, and, when it does
exist, is small in the horse, is a good-sized and permanent tooth, while
the grinder which follows it is but little larger than the hinder ones.
The crowns of the grinders are short, and though the fundamental pattern
of the horse-tooth is discernible, the front and back ridges are less
curved, the accessory pillars are wanting, and the valleys, much
shallower, are not filled up with cement.
Seven years ago, when I happened to be looking critically into the
bearing of palæontological facts upon the doctrine of evolution, it
appeared to me that the _Anchitherium_, the _Hipparion_, and the modern
horses, constitute a series in which the modifications of structure
coincide with the order of chronological occurrence, in the manner in
which they must coincide, if the modern horses really are the result of
the gradual metamorphosis, in the course of the Tertiary epoch, of a
less specialised ancestral form. And I found by correspondence with the
late eminent French anatomist and palæontologist, M. Lartet, that he had
arrived at the same conclusion from the same data.
That the _Anchitherium_ type had become metamorphosed into the
_Hipparion_ type, and the latter into the _Equine_ type, in the course
of that period of time which is represented by the latter half of the
Tertiary deposits, seemed to me to be the only explanation of the facts
for which there was even a shadow of probability.[3]
And, hence, I have ever since held that these facts afford evidence of
the occurrence of evolution, which, in the sense already defined, may be
termed demonstrative.
All who have occupied themselves with the structure of _Anchitherium_,
from Cuvier onwards, have acknowledged its many points of likeness to a
well-known genus of extinct Eocene mammals, _Palæotherium_. Indeed, as
we have seen, Cuvier regarded his remains of _Anchitherium_ as those of
a species of _Palæotherium_. Hence, in attempting to trace the pedigree
of the horse beyond the Miocene epoch and the Anchitheroid form, I
naturally sought among the various species of Palæotheroid animals for
its nearest ally, and I was led to conclude that the _Palæotherium
minus_ (_Plagiolophus_) represented the next step more nearly than any
form then known.
I think that this opinion was fully justifiable; but the progress of
investigation has thrown an unexpected light on the question, and has
brought us much nearer than could have been anticipated to a knowledge
of the true series of the progenitors of the horse.
You are all aware that, when your country was first discovered by
Europeans, there were no traces of the existence of the horse in any
part of the American continent. The accounts of the conquest of Mexico
dwell upon the astonishment of the natives of that country when they
first became acquainted with that astounding phenomenon--a man seated
upon a horse. Nevertheless, the investigations of American geologists
have proved that the remains of horses occur in the most superficial
deposits of both North and South America, just as they do in Europe.
Therefore, for some reason or other--no feasible suggestion on that
subject, so far as I know, has been made--the horse must have died out
on this continent at some period preceding the discovery of America. Of
late years there has been discovered in your Western Territories that
marvellous accumulation of deposits, admirably adapted for the
preservation of organic remains, to which I referred the other evening,
and which furnishes us with a consecutive series of records of the fauna
of the older half of the Tertiary epoch, for which we have no parallel
in Europe. They have yielded fossils in an excellent state of
conservation and in unexampled number and variety. The researches of
Leidy and others have shown that forms allied to the _Hipparion_ and the
_Anchitherium_ are to be found among these remains. But it is only
recently that the admirably conceived and most thoroughly and patiently
worked-out investigations of Professor Marsh have given us a just idea
of the vast fossil wealth, and of the scientific importance, of these
deposits. I have had the advantage of glancing over the collections in
Yale Museum; and I can truly say that, so far as my knowledge extends,
there is no collection from any one region and series of strata
comparable, for extent, or for the care with which the remains have been
got together, or for their scientific importance, to the series of
fossils which he has deposited there. This vast collection has yielded
evidence bearing upon the question of the pedigree of the horse of the
most striking character. It tends to show that we must look to America,
rather than to Europe, for the original seat of the equine series; and
that the archaic forms and successive modifications of the horse's
ancestry are far better preserved here than in Europe.
Professor Marsh's kindness has enabled me to put before you a diagram,
every figure in which is an actual representation of some specimen which
is to be seen at Yale at this present time (Fig. 9).
The succession of forms which he has brought together carries us from
the top to the bottom of the Tertiaries. Firstly, there is the true
horse. Next we have the American Pliocene form of the horse
(_Pliohippus_); in the conformation of its limbs it presents some very
slight deviations from the ordinary horse, and the crowns of the
grinding teeth are shorter. Then comes the _Protohippus_, which
represents the European _Hipparion_, having one large digit and two
small ones on each foot, and the general characters of the fore-arm and
leg to which I have referred. But it is more valuable than the European
_Hipparion_, for the reason that it is devoid of some of the
peculiarities of that form--peculiarities which tend to show that the
European _Hipparion_ is rather a member of a collateral branch, than a
form in the direct line of succession. Next, in the backward order in
time, is the _Miohippus_, which corresponds pretty nearly with the
_Anchitherium_ of Europe. It presents three complete toes--one large
median and two smaller lateral ones; and there is a rudiment of that
digit, which answers to the little finger of the human hand.
[Illustration: FIG. 9.]
The European record of the pedigree of the horse stops here; in the
American Tertiaries, on the contrary, the series of ancestral equine
forms is continued into the Eocene formations. An older Miocene form,
termed _Mesohippus_, has three toes in front, with a large splint-like
rudiment representing the little finger; and three toes behind. The
radius and ulna, the tibia and the fibula, are distinct, and the short
crowned molar teeth are anchitherold in pattern.
But the most important discovery of all is the _Orohippus_, which comes
from the Eocene formation, and is the oldest member of the equine series
as yet known. Here we find four complete toes on the front limb, three
toes on the hind-limb, a well-developed ulna, a well-developed fibula,
and short-crowned grinders of simple pattern.
Thus, thanks to these important researches, it has become evident that,
so far as our present knowledge extends, the history of the horse-type
is exactly and precisely that which could have been predicted from a
knowledge of the principles of evolution. And the knowledge we now
possess justifies us completely in the anticipation, that when the still
lower Eocene deposits, and those which belong to the cretaceous epoch,
have yielded up their remains of ancestral equine animals, we shall
find, first, a form with four complete toes and a rudiment of the
innermost or first digit in front, with probably a rudiment of the fifth
digit in the hind foot;[4] while, in still older forms, the series of
the digits will be more and more complete, until we come to the
five-toed animals, in which, if the doctrine of evolution is well
founded, the whole series must have taken its orgin.
That is what I mean by demonstrative evidence of evolution. An inductive
hypothesis is said to be demonstrated when the facts are shown to be in
entire accordance with it. If that is not scientific proof, there are no
merely inductive conclusions which can be said to be proved. And the
doctrine of evolution, at the present time, rests upon exactly as secure
a foundation as the Copernican theory of the motions of the heavenly
bodies did at the time of its promulgation. Its logical basis is
precisely of the same character--the coincidence of the observed facts
with theoretical requirements.
The only way of escape, if it be a way of escape, from the conclusions
which I have just indicated, is the supposition that all these different
equine forms have been created separately at separate epochs of time;
and, I repeat, that of such an hypothesis as this there neither is, nor
can be, any scientific evidence; and, assuredly so far as I know, there
is none which is supported, or pretends to be supported, by evidence or
authority of any other kind. I can but think that the time will come
when such suggestions as these, such obvious attempts to escape the
force of demonstration, will be put upon the same footing as the
supposition made by some writers, who are I believe not completely
extinct at present, that fossils are mere simulacra, are no indications
of the former existence of the animals to which they seem to belong; but
that they are either sports of Nature, or special creations,
intended--as I heard suggested the other day--to test our faith.
In fact, the whole evidence is in favour of evolution, and there is none
against it. And I say this, although perfectly well aware of the seeming
difficulties which have been built up upon what appears to the
uninformed to be a solid foundation. I meet constantly with the argument
that the doctrine of evolution cannot be well founded, because it
requires the lapse of a very vast period of time; while the duration of
life upon the earth thus implied is inconsistent with the conclusions
arrived at by the astronomer and the physicist. I may venture to say
that I am familiar with those conclusions, inasmuch as some years ago,
when President of the Geological Society of London, I took the liberty
of criticising them, and of showing in what respects, as it appeared to
me, they lacked complete and thorough demonstration. But, putting that
point aside, suppose that, as the astronomers, or some of them, and some
physical philosophers, tell us, it is impossible that life could have
endured upon the earth for as long a period as is required by the
doctrine of evolution--supposing that to be proved--I desire to be
informed, what is the foundation for the statement that evolution does
require so great a time? The biologist knows nothing whatever of the
amount of time which may be required for the process of evolution. It is
a matter of fact that the equine forms which I have described to you
occur, in the order stated, in the Tertiary formations. But I have not
the slightest means of guessing whether it took a million of years, or
ten millions, or a hundred millions, or a thousand millions of years, to
give rise to that series of changes. A biologist has no means of
arriving at any conclusion as to the amount of time which may be needed
for a certain quantity of organic change. He takes his time from the
geologist. The geologist, considering the rate at which deposits are
formed and the rate at which denudation goes on upon the surface of the
earth, arrives at more or less justifiable conclusions as to the time
which is required for the deposit of a certain thickness of rocks; and
if he tells me that the Tertiary formations required 500,000,000 years
for their deposit, I suppose he has good ground for what he says, and I
take that as a measure of the duration of the evolution of the horse
from the _Orohippus_ up to its present condition. And, if he is right,
undoubtedly evolution is a very slow process and requires a great deal
of time. But suppose, now, that an astronomer or a physicist--for
instance, my friend Sir William Thomson--tells me that my geological
authority is quite wrong; and that he has weighty evidence to show that
life could not possibly have existed upon the surface of the earth
500,000,000 years ago, because the earth would have then been too hot to
allow of life, my reply is: "That is not my affair; settle that with the
geologist, and when you have come to an agreement among yourselves I
will adopt your conclusion." We take our time from the geologists and
physicists; and it is monstrous that having taken our time from the
physical philosopher's clock, the physical philosopher should turn round
upon us, and say we are too fast or too slow. What we desire to know is,
is it a fact that evolution took place? As to the amount of time which
evolution may have occupied, we are in the hands of the physicist and
the astronomer, whose business it is to deal with those questions.
I have now, ladies and gentlemen, arrived at the conclusion of the task
which I set before myself when I undertook to deliver these lectures. My
purpose has been, not to enable those among you who have paid no
attention to these subjects before, to leave this room in a condition to
decide upon the validity or the invalidity of the hypothesis of
evolution; but I have desired to put before you the principles upon
which all hypotheses respecting the history of Nature must be judged;
and furthermore, to make apparent the nature of the evidence and the
amount of cogency which is to be expected and may be obtained from it.
To this end, I have not hesitated to regard you as genuine students and
persons desirous of knowing the truth. I have not shrunk from taking you
through long discussions, that I fear may have sometimed tried your
patience; and I have inflicted upon you details which were
indispensable, but which may well have been wearisome. But I shall
rejoice--I shall consider that I have done you the greatest service
which it was in my power to do--if I have thus convinced you that the
great question which we have been discussing is not one to be dealt with
by rhetorical flourishes, or by loose and superficial talk; but that it
requires the keen attention of the trained intellect and the patience of
the accurate observer.
ON THE PHYSICAL BASIS OF LIFE
[1868]
In order to make the title of this discourse generally intelligible, I
have translated the term "Protoplasm," which is the scientific name of
the substance of which I am about to speak, by the words "the physical
basis of life." I suppose that, to many, the idea that there is such a
thing as a physical basis, or matter, of life may be novel--so widely
spread is the conception of life as a something which works through
matter, but is independent of it; and even those who are aware that
matter and life are inseparably connected, may not be prepared for the
conclusion plainly suggested by the phrase, "_the_ physical basis or
matter of life," that there is some one kind of matter which is common
to all living beings, and that their endless diversities are bound
together by a physical, as well as an ideal, unity. In fact, when first
apprehended, such a doctrine as this appears almost shocking to common
sense.
What, truly, can seem to be more obviously different from one another,
in faculty, in form, and in substance, than the various kinds of living
beings? What community of faculty can there be between the
brightly-coloured lichen, which so nearly resembles a mere mineral
incrustation of the bare rock on which it grows, and the painter, to
whom it is instinct with beauty, or the botanist, whom it feeds with
knowledge?
Again, think of the microscopic fungus--a mere infinitesimal ovoid
particle, which finds space and duration enough to multiply into
countless millions in the body of a living fly; and then of the wealth
of foliage, the luxuriance of flower and fruit, which lies between this
bald sketch of a plant and the giant pine of California, towering to the
dimensions of a cathedral spire, or the Indian fig, which covers acres
with its profound shadow, and endures while nations and empires come and
go around its vast circumference. Or, turning to the other half of the
world of life, picture to yourselves the great Finner whale, hugest of
beasts that live, or have lived, disporting his eighty or ninety feet of
bone, muscle, and blubber, with easy roll, among waves in which the
stoutest ship that ever left dockyard would flounder hopelessly; and
contrast him with the invisible animalcules--mere gelatinous specks,
multitudes of which could, in fact, dance upon the point of a needle
with the same ease as the angels of the Schoolmen could, in imagination.
With these images before your minds, you may well ask, what community of
form, or structure, is there between the animalcule and the whale; or
between the fungus and the fig-tree? And, _a fortiori_, between all
four?
Finally, if we regard substance, or material composition, what hidden
bond can connect the flower which a girl wears in her hair and the blood
which courses through her youthful veins; or, what is there in common
between the dense and resisting mass of the oak, or the strong fabric of
the tortoise, and those broad disks of glassy jelly which may be seen
pulsating through the waters of a calm sea, but which drain away to
mere films in the hand which raises them out of their element?
Such objections as these must, I think, arise in the mind of every one
who ponders, for the first time, upon the conception of a single
physical basis of life underlying all the diversities of vital
existence; but I propose to demonstrate to you that, notwithstanding
these apparent difficulties, a threefold unity--namely, a unity of
power or faculty, a unity of form, and a unity of substantial
composition--does pervade the whole living world.
No very abstruse argumentation is needed, in the first place, to prove
that the powers, or faculties, of all kinds of living matter, diverse as
they may be in degree, are substantially similar in kind.
Goethe has condensed a survey of all powers of mankind into the
well-known epigram:--
"Warum treibt sich das Volk so und schreit?
Es will sich ernähren
Kinder zeugen, und die nähren so gut es vermag.
* * * * *
Weiter bringt es kein Mensch, stell' er
sich wie er auch will."