CHAPTER I

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EVOLUTION AND ADAPTATION

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[Illustration: Publisher's logo (The Macmillan Company)]

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EVOLUTION AND ADAPTATION

by

THOMAS HUNT MORGAN, Ph.D.

New York

The Macmillan Company

London: Macmillan & Co., Ltd.

1908

All rights reserved

Copyright, 1903,

by The Macmillan Company.

Set up and electrotyped. Published October, 1903. Reprinted January,

1908.

Norwood Press

J. S. Cushing Co.—Berwick & Smith Co.

Norwood, Mass., U.S.A.

------------------------------------------------------------------------

TO

Professor William Keith Brooks

AS A TOKEN OF SINCERE ADMIRATION AND RESPECT

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PREFACE

The adaptation of animals and plants to the conditions under which they

live has always excited the interest, and also the imagination, of

philosophers and scientists; for this relation between the organism and

its environment is one of the most characteristic features of living

things. The question at once suggests itself: How has such a relation

been brought about? Is it due to something inherent in the living matter

itself, or is it something that has been, as it were, superimposed upon

it? An example may make my meaning clearer. No one will suppose that

there is anything inherent in iron and other metals that would cause

them to produce an engine if left to themselves. The particular

arrangement of the pieces has been superimposed upon the metals, so that

they now fulfil a purpose, or use. Have the materials of which organisms

are composed been given a definite arrangement, so that they fulfil the

purpose of maintaining the existence of the organism; and if so, how has

this been accomplished? It is the object of the following pages to

discuss this question in all its bearings, and to give, as far as

possible, an idea of the present state of biological thought concerning

the problem. I trust that the reader will not be disappointed if he

finds in the sequel that many of the most fundamental questions in

regard to adaptation are still unsettled.

In attempting to state the problem as clearly as possible, I fear that

it may appear that at times I have “taken sides,” when I should only

have been justified in stating the different aspects of the question.

But this will do little harm provided the issue has been sharply drawn.

Indeed, it seems to me that the only scientific value, that a discussion

of what the French call “les grands problèmes de la Biologie” has, is to

get a clearer understanding of the relation of what is known to what is

unknown or only surmised.

In some quarters speculation concerning the origin of the adaptation of

living things is frowned upon, but I have failed to observe that the

critics themselves refrain entirely from theorizing. They shut one door

only to open another, which also leads out into the dark. To deny the

right to speculative thought would be to deny the right to use one of

the best tools of research.

Yet it must be admitted that all speculation is not equally valuable.

The advance of science in the last hundred years has shown that the kind

of speculation that has real worth is that which leads the way to

further research and possible discovery. Speculation that leads to this

end must be recognized as legitimate. It becomes useless when it deals

with problems that cannot be put to the actual test of observation or

experiment. It is in this spirit that I have approached the topics

discussed in the following pages.

The unsophisticated man believes that all other animals exist to

minister to his welfare; and from this point of view their adaptations

are thought of solely in their relation to himself. A step in advance

was taken when the idea was conceived that adaptations are for the good

of the organisms themselves. It seemed a further advance when the

conclusion was reached that the _origin_ of adaptations could be

accounted for, as the result of the benefit that they conferred on their

possessor. This view was the outcome of the acceptation of the theory of

evolution, combined with Darwin’s theory of natural selection. It is the

view held by most biologists at the present time; but I venture to

prophesy that if any one will undertake to question modern zoologists

and botanists concerning their relation to the Darwinian theory, he will

find that, while professing _in a general way_ to hold this theory, most

biologists have many reservations and doubts, which they either keep to

themselves or, at any rate, do not allow to interfere either with their

teaching of the Darwinian doctrine or with the applications that they

may make of it in their writings. The claim of the opponents of the

theory that Darwinism has become a dogma contains more truth than the

nominal followers of this school find pleasant to hear; but let us not,

therefore, too hastily conclude that Darwin’s theory is without value in

relation to one side of the problem of adaptation; for, while we can

profitably reject, as I believe, much of the theory of natural

selection, and more especially the idea that adaptations have arisen

because of their usefulness, yet the fact that living things must be

adapted more or less well to their environment in order to remain in

existence may, after all, account for the widespread _occurrence_ of

adaptation in animals and plants. It is this point of view that will be

developed in the following pages.

I am fully aware of the danger in attempting to cover so wide a field as

that of “Evolution and Adaptation,” and I cannot hope to escape the

criticism that is certain to be directed against a specialist who

ventures nowadays beyond the immediate field of his own researches; yet,

in my own defence, I may state that the whole point of view underlying

the position here taken is the immediate outcome of my work on

regeneration. One of the general questions that I have always kept

before me in my study of regenerative phenomena is how such a useful

acquirement as the power to replace lost parts has arisen, and whether

the Darwinian hypothesis is adequate to explain the result. The

conclusion that I have reached is that the theory is entirely inadequate

to account for the _origin_ of the power to regenerate; and it seemed to

me, therefore, desirable to reëxamine the whole question of adaptation,

for might it not prove true here, also, that the theory of natural

selection was inapplicable? This was my starting-point. The results of

my examination are given in the following pages.

I am deeply indebted to Professor G. H. Parker and to Professor E. G.

Conklin for advice and friendly criticism; and in connection with the

revision of the proof I am under many obligations to Professor Joseph W.

Warren and to Professor E. A. Andrews. Without their generous help I

should scarcely have ventured into a field so full of pitfalls.

Bryn Mawr, Penn., June 10, 1903.

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CONTENTS

CHAPTER I

PAGE

The Problem of Adaptation 1

– Structural Adaptations 1

– Adaptations for the Good of the 19

Species

– Organs of Little Use to the 22

Individual

– Changes in the Organism that are 25

of No Use to the Individual or to

the Race

– Comparison with Inorganic 26

Phenomena

CHAPTER II

The Theory of Evolution 30

– Evidence in Favor of the 32

Transmutation Theory

– – Evidence from Classification 32

and from Comparative Anatomy

– – The Geological Evidence 39

– – Evidence from Direct 43

Observation and Experiment

– – Modern Criticism of the Theory 44

of Evolution

CHAPTER III

The Theory of Evolution (continued) 58

– The Evidence from Embryology 58

– – The Recapitulation Theory 58

– Conclusions 84

CHAPTER IV

Darwin’s Theories of Artificial and 91

of Natural Selection

– The Principle of Selection 91

– Variation and Competition in 104

Nature

– The Theory of Natural Selection 116

CHAPTER V

The Theory of Natural Selection 129

(continued)

– Objections to the Theory of 129

Natural Selection

– Sterility between Species 147

– Weismann’s Germinal Selection 154

CHAPTER VI

Darwin’s Theory of Sexual Selection 167

– Sexual Selection 167

– General Criticism of the Theory 213

of Sexual Selection

CHAPTER VII

The Inheritance of Acquired 222

Characters

– Lamarck’s Theory 222

– Darwin’s Hypothesis of Pangenesis 233

– The Neo-Lamarckian School 240

CHAPTER VIII

Continuous and Discontinuous 261

Variation and Heredity

– Continuous Variation 261

– Heredity and Continuous Variation 270

– Discontinuous Variation 272

– Mendel’s Law 278

– The Mutation Theory of De Vries 287

– Conclusions 297

CHAPTER IX

Evolution as the Result of External 300

and Internal Factors

– The Effect of External Influences 300

– Responsive Changes in the 319

Organism that adapt it to the New

Environment

– Nägeli’s Perfecting Principle 325

CHAPTER X

The Origin of the Different Kinds 340

of Adaptations

– Form and Symmetry 340

– Mutual Adaptation of Colonial 350

Forms

– Degeneration 352

– Protective Coloration 357

– Sexual Dimorphism and Trimorphism 360

– Length of Life as an Adaptation 370

– Organs of Extreme Perfection 371

– Secondary Sexual Organs as 372

Adaptations

– Individual Adjustments as 375

Adaptations

– Color Changes as Individual 375

Adaptations

– Increase of Organs through Use 376

and Decrease through Disuse

– Reactions of the Organism to 377

Poisons, etc.

– Regeneration 379

CHAPTER XI

Tropisms and Instincts as 382

Adaptations

CHAPTER XII

Sex as an Adaptation 414

– The Different Kinds of Sexual 414

Individuals

– The Determination of Sex 422

– Sex as a Phenomenon of Adaptation 439

CHAPTER XIII

Summary and General Conclusions 452

INDEX 465

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EVOLUTION AND ADAPTATION

CHAPTER I

THE PROBLEM OF ADAPTATION

Between an organism and its environment there takes place a constant

interchange of energy and of material. This is, in general, also true

for all bodies whether living or lifeless; but in the living organism

this relation is a peculiar one; first, because the plant or the animal

is so constructed that it is suited to a particular set of physical

conditions, and, second, because it may so respond to a change in the

outer world that it further adjusts itself to changing conditions,

_i.e._ the response may be of such a kind that it better insures the

existence of the individual, or of the race. The two ideas contained in

the foregoing statement cover, in a general way, what we mean by the

adaptation of living things. The following examples will serve to

illustrate some of the very diverse phenomena that are generally

included under this head.

Structural Adaptations

The most striking cases of adaptations are those in which a special, in

the sense of an unusual, relation exists between the individual and its

surroundings. For example, the foreleg of the mole is admirably suited

for digging underground. A similar modification is found in an entirely

different group of the animal kingdom, namely, in the mole-cricket, in

which the first legs are also well suited for digging. By their use the

mole-cricket makes a burrow near the surface of the ground, similar to,

but of course much smaller than, that made by the mole. In both of these

cases the adaptation is the more obvious, because, while the leg of the

mole is formed on the same general plan as that of other vertebrates,

and the leg of the mole-cricket has the same fundamental structure as

that of other insects, yet in both cases the details of structure and

the general proportions have been so altered, that the leg is fitted for

entirely different purposes from that to which the legs of other

vertebrates and of other insects are put. The wing of the bat is another

excellent case of a special adaptation. It is a modified fore-limb

having a strong membrane stretched between the fingers, which are

greatly elongated. Here we find a structure, which in other mammals is

used as an organ for supporting the body, and for progression on the

ground, changed into one for flying in the air.

The tails of mammals show a number of different adaptations. The tail is

prehensile in some of the monkeys; and not only can the monkey direct

its tail toward a branch in order to grasp it, but the tail can be

wrapped around the branch and hold on so firmly that the monkey can

swing freely, hanging by its tail alone. The animal has thus a sort of

fifth hand, one as it were in the middle line of the body, which can be

used as a hold-fast, while the fingered hands are put to other uses. In

the squirrels the bushy tail serves as a protection during the winter

for those parts of the body not so thickly covered by hair. The tail of

the horse is used to brush away the flies that settle on the hind parts

of the body. In other mammals, the dog, the cat, and the rat, for

example, the tail is of less obvious use, although the suggestion has

been made that it may serve as a sort of rudder when the animal is

running rapidly. In several other cases, as in the rabbit and in the

higher apes, the tail is very short, and is of no apparent use; and in

man it has completely disappeared.

A peculiar case of adaptation is the so-called basket on the third pair

of legs of the worker honey-bee. A depression of the outer surface of

the tibia is arched over by stiff hairs. The pollen collected from the

stamens of flowers is stowed away in this receptacle by means of the

other pairs of legs. The structure is unique, and is not found in any

other insects except the bees. It is, moreover, present only in the

worker bees, and is absent in the queen and the males.

The preceding cases, in which the adapted parts are used for the

ordinary purposes of life of the individual, are not essentially

different from the cases in which the organ is used to protect the

animal from its enemies. The bad taste of certain insects is supposed to

protect them from being eaten by birds. Cases like this of passive

protection grade off in turn into those in which, by some reflex or

voluntary act, the animal protects itself. The bad-smelling horns of the

caterpillar of the black swallow-tailed butterfly (_Papilio polyxenes_)

are thrust out when the animal is touched, and it is believed that they

serve to protect the caterpillar from attack. The fœtid secretion of the

glands of the skunk is believed to serve as a protection to the animal,

although the presence of the nauseous odor may lead finally to the

extermination of the skunk by man. The sting of bees and of wasps serves

to protect the individual from attack. The sting was originally an

ovipositor, and used in laying the eggs. It has, secondarily, been

changed into an organ of offence.

The special instincts and reflex acts furnish a striking group of

adaptations. The building of the spider’s web is one of the most

remarkable cases of this kind. The construction of the web cannot be the

result of imitation, since, in many instances, the young are born in the

spring of the year following the death of the parents. Each species of

spider has its own type of web, and each web has as characteristic a

form as has the spider itself. It is also important to find that a

certain type of web may be characteristic of an entire family of

spiders. Since, in many cases, the web is the means of securing the

insects used for food, it fulfils a purpose necessary for the welfare of

the spider.

The making of the nests by birds appears to be also in large part an

instinctive act; although some writers are inclined to think that memory

of the nest in which the young birds lived plays a part in their

actions, and imitation of the old birds at the time of nest-building

may, perhaps, also enter into the result. It has been stated that the

first nest built by young birds is less perfect than that built by older

birds, but this may be due to the bird’s learning something themselves

in building their nests, _i.e._ to the perfecting of the instinct in the

individual that makes use of it. In any case much remains that must be

purely instinctive. The construction of the comb by bees appears to be

largely, perhaps entirely, an instinctive act. That this is the case was

shown by isolating young workers as soon as they emerged from the cell,

and before they could have had any experience in seeing comb built. When

given some wax they set to work to make a comb, and made the

characteristic six-sided structures like those made by the bees in a

hive. The formation of so remarkable a structure as the comb is worthy

of admiration, for, with the greatest economy of material, a most

perfect storeroom for the preservation of the honey is secured. This

adaptation appears almost in the nature of foresight, for the store of

honey is used not only to feed the young, but may be drawn on by the

bees themselves in time of need. It is true that a comparison with other

kinds of bees makes it probable that the comb was first made for the

eggs and larvæ, and only later became used as a storehouse, but so far

as its form is concerned there is the same economy of constructive

materials in either case.

The behavior of young birds, more especially those that take care of

themselves from the moment they leave the egg, furnishes a number of

cases of instincts that are protective. If, for example, a flock of

young pheasants is suddenly disturbed, the birds at once squat down on

the ground, and remain perfectly quiet until the danger is past. Their

resemblance to the ground is so perfect that they are almost invisible

so long as they remain quiet. If, instead of remaining still, they were

to attempt to run away when disturbed, they would be much more easily

seen.

Certain solitary wasps (_Ammophila_) have the habit of stinging

caterpillars and spiders, and dragging them to their nests, where they

are stored away for the future use of the young that hatch from the eggs

laid by the wasp on the body of the prey. As a result of the sting which

the wasp administers to the caterpillar, the latter is paralyzed, and

cannot escape from the hole in which it is stored, where it serves as

food for the young wasp that emerges from the egg. It was originally

claimed by Forel that the wasp stings the caterpillar in such a way that

the central nervous system is always pierced, and many subsequent

naturalists have marvelled at the perfection of such a wonderful

instinct. But the recent results of the Peckhams have made it clear that

the act of the wasp is not carried out with the precision previously

supposed, although it is true that the wasp pierces the caterpillar on

the lower surface where the ventral chain of ganglia lies. The habit of

this wasp is not very dissimilar from that shown by many other kinds of

wasps that sting their captive in order to quiet it. We need not imagine

in this case that the act carries with it the consciousness that the

caterpillar, quieted in this way, will be unable to escape before the

young wasps have hatched.

The resemblance in color of many animals to their natural backgrounds

has in recent years excited the interest and imagination of many

naturalists. The name of protective coloration has been given to this

group of phenomena. The following cases which have less the appearance

of purely imaginative writing may serve by way of illustration. A

striking example is that of the ptarmigan which has a pure white coat in

winter, and a brown coat in summer. The white winter plumage renders the

animal less conspicuous against the background of snow, while in summer

the plumage is said to closely resemble the lichen-covered ground on

which the bird rests. The snowy owl is a northern bird, whose color is

supposed to make it less conspicuous, and may serve either as a

protection against enemies, or may allow the owl to approach its prey

unseen. It should not pass unnoticed, however, that there are white

birds in other parts of the world, where their white color cannot be of

any use to them as a protection. The white cockatoos, for example, are

tropical birds, living amongst green foliage, where their color must

make them conspicuous, rather than the reverse.

The polar bear is the only member of the family that is white, and while

this can scarcely be said to protect it from enemies, because it is

improbable that it has anything to fear from the other animals of the

ice-fields, yet it may be claimed that the color is an adaptation to

allow the animal to approach unseen its prey.

In the desert many animals are sand-colored, as seen for instance in the

tawny color of the lion, the giraffe, the antelopes, and of many birds

that live on or near the ground.

It has been pointed out that in the tropics and temperate zones there

are many greenish and yellowish birds whose colors harmonize with the

green and yellow of the trees amongst which they live; but on the other

hand we must not forget that in all climes there are numbers of birds

brilliantly colored, and many of these do not appear to be protected in

any special way. The tanagers, humming-birds, parrots, Chinese

pheasants, birds of paradise, etc., are extremely conspicuous, and so

far as we can see they must be much exposed on account of the color of

their plumage. Whether, therefore, we are justified in picking out

certain cases as examples of adaptation, because of an agreement in

color between the organism and its surroundings, and in neglecting all

others, is, as has been already said, a point to be further examined.

Not only among mammals and birds have many cases of protective

coloration been described by writers dealing with this subject, but in

nearly every group of the animal kingdom similar cases have been

recognized. The green and brown color of lizards may protect them, the

green color of many frogs is supposed to conceal them as they sit

amongst the plants on the edge of a stream or pond. The gray-brown color

of the toad has been described as a resemblance to the dry ground, while

the brilliant green of several tree-frogs conceals them very effectively

amongst the leaves. Many fishes are brilliantly colored, and it has even

been suggested that those living amongst corals and sea-anemonies have

acquired their colors as a protection, but Darwin states that they

appeared to him very conspicuous even in their highly colored

environment.

Amongst insects innumerable cases of adaptive coloration have been

described. In fact this is the favorite group for illustrating the

marvels of protective coloration. A few examples will here serve our

purpose. The oft-cited case of the butterfly _Kallima_ is, apparently, a

striking instance of protective resemblance. When at rest the wings are

held together over the back, as in nearly all butterflies, so that only

the under surface is exposed. This surface has an unquestionably close

resemblance to a brown leaf. It is said on no less authority than that

of Wallace that when this butterfly alights on a bush it is almost

impossible to distinguish between it and a dead leaf. The special point

in the resemblance to which attention is most often called is the

distinct line running obliquely across the wings which looks like the

midrib of a leaf. Whether the need of such a close resemblance to a leaf

is requisite for the life of this butterfly, we do not know, of course,

and so long as we do not have this information there is danger that the

case may prove too much, for, if it should turn out that this remarkable

case is accidental the view in regard to the resemblance may be

endangered.

Amongst caterpillars there are many cases of remarkable resemblances in

color between the animal and its surroundings. The green color of many

of those forms that remain on the leaves of the food-plant during the

day will give, even to the most casual observer, the impression that the

color is for the purpose of concealment; and that it does serve to

conceal the animal there can be no doubt. But even from the point of

view of those who maintain that this color has been acquired because of

its protective value it must be admitted that the color is insufficient,

because some of these same green caterpillars are marvellously armed

with an array of spines which are also supposed to be a protection

against enemies. Equally well protected are the brown and mottled

geometrid caterpillars. These have, moreover, the striking and unusual

habit of fixing themselves by the posterior pairs of false legs, and

standing still and rigid in an oblique position on the twigs to which

they are affixed. So close is their resemblance to a short twig, that

even when their exact position is known it is very difficult to

distinguish them.

Grasshoppers that alight on the ground are, in many cases, so similar to

the surface of the ground that unless their exact location is known they

easily escape attention, while the green color of the katydid, a member

of the same group of orthoptera, protects it from view in the green

foliage of the trees where it lives. The veinlike wings certainly

suggest a resemblance to a leaf, but whether there is any necessity for

so close an imitation may be questioned.

There can be little doubt in some of these cases that the color of the

animal may be a protection to it, but as has been hinted already, it is

another question whether it acquired these colors because of their

usefulness. Nevertheless, if the color is useful to its possessor, it is

an adaptation in our sense of the word, without regard to the way in

which it has been acquired. Even, for instance, if the resemblance were

purely the outcome of chance in the sense that the color appeared

without relation to the surroundings, it would still be an adaptation if

it were of use to the animal under the ordinary conditions of life.

In the lower groups numerous cases in which animals resemble their

surroundings could be given. Such cases are known in crustacea, worms,

mollusks, hydroids, etc., and the possible value of these resemblances

may be admitted in many instances.

It is rather curious that so few cases of adaptive coloration have been

described for plants. No one supposes that the slate color of the lichen

is connected with the color of the rocks on which it grows, in the sense

that the resemblance is of any use to the lichen. Nor does the color of

the marine red algæ serve in any way to protect the plants so far as is

known. The green color of nearly all the higher plants is obviously

connected with the substance, chlorophyl, that is essential for the

processes of assimilation, and has no relation to external objects. But

when we come to the colors of flowers we meet with curious cases of

adaptation, at least according to the generally accepted point of view.

For it is believed by many naturalists that the color of the corolla of

flowering plants is connected with the visits of insects to the flowers,

and these visits are in many cases essential for the cross-fertilization

of the flowers. This adaptation is one useful to the species, rather

than the individual, and belongs to another category.

The leaf of the Venus’s fly-trap, which suddenly closes together from

the sides when a fly or other light body comes to rest on it, is

certainly a remarkable adaptation. A copious secretion of a digestive

fluid is poured out on the surface of the leaf, and the products of

digestion are absorbed. There can be no question that this contrivance

is of some use to the plant. In other insectivorous plants, the pitcher

plants, the leaves are transformed into pitchers. In Nepenthes a

digestive fluid is secreted from the walls. A line of glands secreting a

sweet fluid serves to attract insects to the top of the pitcher, whence

they may wander or fall into the fluid inside, and there being drowned,

they are digested. A lidlike cover projecting over the opening of the

pitcher is supposed to be of use to keep out the rain.

In _Utricularia_, a submerged water-plant, the tips of the leaves are

changed into small bladders, each having a small entrance closed by an

elastic valve opening inwards. Small snails and crustaceans can pass

into this opening, to which they are guided by small outgrowths; but

once in the cup they cannot get out again, and, in fact, small animals

are generally found in the bladders where they die and their substance

is absorbed by forked hairs projecting into the interior of the bladder.

The cactus is a plant that is well suited to a dry climate. Its leaves

have completely disappeared, and the stem has become swollen into a

water-reservoir. “It has been estimated that the amount of water

evaporated by a melon cactus is reduced to one six-hundredth of that

given off by any equally heavy climbing-plant.”

[Illustration:

Fig. 1.—The fertilization of _Aristolochia Clematitis_.

A, portion of stem with flowers in axil of leaf in different stages.

B and C, longitudinal sections of two flowers, before and after

fertilization. (After Sachs.)]

Sachs gives the following account of the fertilization process in

_Aristolochia Clematitis_, which he refers to as a conspicuous and

peculiar adaptation. In Figure 1 A a group of flowers is shown, and in

Figure 1 B and C a single flower is split open to show the interior. In

B a small fly has entered, and has brought in upon its back some pollen

that has stuck to it in another flower. The fly has entered through the

long neck which is beset with hairs which are turned inwards so that the

fly can enter but cannot get out. In roaming about, the pollen that is

sticking to its back will be rubbed against the stigmatic surface. “As

soon as this has taken place the anthers, which have been closed

hitherto, dehisc and become freely accessible,” as a result in the

change in the stigma and of the collapse of the hairs at the base of the

enlargement which has widened. The fly can now crawl under the anthers,

and, if it does so, new pollen may stick to its back. At this time the

hairs in the throat dry up, and the fly can leave its prison house,

Figure 1 C. If the fly now enters another flower this is fertilized by

repeating the process. The unfertilized flowers stand erect with widely

open mouths. As soon as they have been fertilized they bend down, as

seen in Figure 1 A, and at the same time the terminal flap bends over

the open mouth of the throat, “stopping the entrance to the flies, which

have now nothing more to do here.”

Adjustments of the Individual to Changes in the Environment

The most familiar cases of adjustments of the individual to the

environment are those that we recognize in our own bodies. After violent

exercise we breathe more rapidly, and take deeper inspirations. Since

during exercise our blood loses more oxygen and takes in more carbon

dioxide from the muscles, it is clear that one result of more rapid

breathing is to get more oxygen into the blood and more carbon dioxide

out of it. The process of sweating, that also follows exercise, may be

also looked upon as an adaptive process, since by evaporation the skin

is kept cooler, and, in consequence, the blood, which at this time flows

in larger quantities to the skin, is cooled also.

More permanent adaptive changes than these also take place as the result

of prolonged use of certain parts. If the muscles work against powerful

resistance, they become larger after several days or weeks, and are

capable of doing more work than at first. Conversely, when any group of

muscles is not used, it becomes smaller than the normal and capable of

doing less work. It would be a nice point to decide whether this latter

change is also an adaptation. If so it is one in a somewhat different

sense from that usually employed. The result is of no direct advantage

to the animal, except possibly in saving a certain amount of food, but

since the same change will take place when an abundance of food is

consumed, the result is, under these conditions, of no use.

The thickening of the skin on those parts of the body where continued

pressure is brought to bear on it is a change in a useful direction. The

thickening on the soles of the feet and on the palms of the hands is a

case in point. Not only is the skin thicker at birth in these parts, but

it becomes thicker through use. In other parts of the body also, the

skin hardens and becomes thicker if pressure is brought to bear on it.

We may regard this as a general property of the skin, which is present

even in those parts where, under ordinary circumstances, it can rarely

or never be brought into use.

Even as complicated and as much used an organ as the eye can become

adaptively improved. It is said that the lateral region of the field of

vision can be trained to perceive more accurately; and every one who has

used a microscope is familiar with the fact that if one eye is

habitually used it becomes capable of seeing more distinctly and better

than the other eye. This seems to be due, in part at least, to the

greater contraction of the iris.

Another phenomenon, which, I think, must be looked upon as an

adaptation, is the immunity to certain poisons that can be gradually

brought about by slowly increasing the amount introduced into the body.

Nicotine is a most virulent poison, and yet by slowly increasing the

dose an animal can be brought into a condition in which an amount of

nicotine, fatal to an ordinary individual, can be administered without

any ill effects at all resulting.

The same phenomenon has been observed in the case of other poisons, not

only in case of other alkaloids, such as morphine and cocaine, but also

in the case of caffein, alcohol, and even arsenic. There is a curious

phenomenon in regard to arsenic, which appears to be well established,

viz., that a person who has gradually increased the dose to an amount

great enough to kill ten ordinary men, will die if he suddenly ceases

altogether to take arsenic. He can, however, be gradually brought back

to a condition in which arsenic is not necessary for his existence, if

the dose is gradually decreased. It is a curious case of adaptation that

we meet with here, since the man becomes so thoroughly adjusted to a

poison that if he is suddenly brought back to the normal condition of

the race he will die.

Immunity to the poison of venomous snakes can also be acquired by slowly

increasing the amount given to an animal. It is possible to make a

person so immune to the poison of venomous snakes that he would become,

in a sense, adapted to live amongst them without danger to himself. It

is to be noted, moreover, that this result could be reached only by

quite artificial means, for, under natural conditions it is

inconceivable that the nicely graded series of doses of increasing

strength necessary to bring about the immunity could ever be acquired.

Hence we find here a case of response in an adaptive direction that

could not have been the outcome of experience in the past. It is

important to emphasize this capacity of organisms to adapt themselves to

certain conditions entirely new to them.

These cases lead at once to cases of immunity to certain bacterial

diseases. An animal may become immune to a particular disease in several

ways. First, by having the disease itself, which renders it immune for a

longer or a shorter period afterwards; or, second, by having a mild form

of the disease as in the case of smallpox, where immunity is brought

about by vaccination, _i.e._ by giving the individual a mild form of

smallpox; or, third, by introducing into the blood an antidote, in the

form, for example, of antitoxin, which has been made by another animal

itself immune to the disease. The first two classes of immunity may be

looked upon as adaptations which are of the highest importance to the

organism; the last case can scarcely be looked upon as an adaptive

process, since the injurious effect of the poison may as well be

neutralized outside of the body by mixing it with the antitoxin. We may

suppose, then, that in the body a similar process goes on, so that the

animal itself takes no active part in the result.

When we consider that there are a number of bacterial diseases, in each

of which a different poison is made by the bacteria, we cannot but ask

ourselves if the animal really makes a counter-poison for each disease,

or whether a single substance may not be manufactured that counteracts

all alike? That the latter is not the case is shown by the fact that an

animal made immune to one disease is not immune to others. When we

recall that the animal has also the capacity to react in one way or

another to a large number of organic and inorganic poisons, to which it

or its ancestors can have had little or no previous experience, we may

well marvel at this wonderful regulative power.

The healing of wounds, which takes place in all animals, forms another

class of adaptive processes. The immense usefulness of this power is

obvious when it is remembered how exposed most animals are to injuries.

By repairing the injury the animal can better carry on its normal

functions. Moreover, the presence of the wound would give injurious

bacteria a ready means of entering the body. In fact, an intact skin is

one of the best preventives to the entrance of bacteria.

Not only have most organisms the power of repairing injuries, but many

animals have also the closely related power of regenerating new parts if

the old ones are lost. If a crab loses its leg, a new one is

regenerated. If a fresh-water worm (_Lumbriculus_) is cut into pieces,

each piece makes a new head at its anterior end and a new tail at the

posterior end. In this way as many new worms are produced as there are

pieces. And while in a strict sense it cannot be claimed that this power

of regeneration is of any use to the original worm, since the original

worm, as such, no longer exists, yet since it has not died but has

simply changed over into several new worms, the process is of use

inasmuch as by this means the pieces can remain in existence.

We need not discuss here the relative importance to different animals of

this power of regeneration, but it may be stated, that, while in some

cases it may be necessary to replace the lost part if the animal is to

remain in existence, as when a new head is formed on an earthworm after

the old one was cut off, in other cases the replacement of the lost part

appears to be of minor importance, as in the case of the leg of the

crab. While we are not, for the moment, concerned with the relative

importance of the different adaptations, this question is one of much

importance in other connections and will be considered later.

The protective coloration of some animals, which is the direct result of

a change in color of the animal in response to the surroundings,

furnishes us with some most striking cases of adaptive coloration. A

change of this sort has been recorded in a number of fishes, more

especially in the flounders. The individuals found living on a dark

background are darker than those living on a lighter background; and

when the color of the background is changed it has been observed that

the color of the fish also changes in the same direction. I have

observed a change of this sort from dark to light, or from light to

dark, in the common minnow (_Fundulus_) in accordance with a change of

its background, and the same sort of change appears to take place in

many other fishes.

The change from green to brown and from brown to green in certain tree

frogs and in the lizard (_Anolis_), which is popularly supposed to take

place according to whether the background is green or brown, is not

after all, it appears, connected with the color of the background, but

depends on certain other responses of the animals that have not yet been

satisfactorily made out. If it be claimed that in summer the animal

would generally be warm, and therefore, often green, and that this color

would protect it at this time of year when the surroundings are green,

and in winter brown, when this color is the prevailing one in temperate

regions, then it might appear that the change is of use to the animal;

but if it is true that the same change takes place in some of the

lizards that live in the tropics, where the prevailing color is always

green, it would appear that the result may have no direct relation with

the surroundings. It has been shown in a number of well-authenticated

cases that the pupæ of certain butterflies vary in color within certain

limits in response to the color of the background. When the caterpillar

fixes itself to some surface, and there throws off the outer skin, and

acquires a new one, the color of the latter is influenced by the

background. The result is a better protection to the pupa. The change is

not brought about through the ocelli or eyes, but through the general

surface of the skin, for the same change takes place when the eyes have

been previously covered with a dark pigment.

The growth of plants toward the light may be looked upon as an adaptive

process, since only in the light can they find the conditions necessary

for their life. The extraordinary elongation of shoots and young plants

when grown in the dark may also be considered an adaptation for finding

the light, since in this way a plant, deeply embedded in the ground, may

ultimately reach the surface. Thus while the actual process of

elongation in the dark is not in itself of any use, yet under the

ordinary conditions of its life, this response may be of great benefit

to the plant.

The closing together of the leaves of some plants has been supposed

to protect them from too rapid radiation of heat, and incidentally

this purpose may be fulfilled; but since some tropical plants also

close their leaves during the night, it can hardly be maintained

that the closing has been acquired for this purpose. It has been

suggested that the opening of certain flowers under certain

conditions of light is connected with the visits of insects that

bring about cross-fertilization.

The preceding examples will suffice to give a general idea of what is

meant by adaptation in organisms. That the term includes a large number

of phenomena of very different kinds is apparent. When we have examined

these phenomena further we shall find, I think, that it will be

necessary to put some of them into different categories and treat them

differently. It is probably incorrect to suppose that all processes

useful to the organism have been acquired in the same way, nevertheless,

for the present the term adaptation is sufficiently general, even if

vague, to cover these different groups of cases.

It may be asked, in what respects are these structures and processes of

adaptation different from the ordinary structures and changes that go on

in the organism? Why is the leg of the mole more of an adaptation than

that of a dog? The one is of as much use as the other to its possessor.

What reason can we give for citing the poison of the snake, and not

mentioning in the same connection the other glands of the body? In fact,

the poison gland of the snake is supposed to be a modified superior

labial gland. Why, in short, are not the processes of digestion,

excretion, secretion, the beating of the heart, the ordinary reflex acts

of the nervous system, and the action of the sense-organs, as truly

adaptations as the special cases that have been selected for

illustration. The answer is simply that we are more impressed by those

cases of adaptation that are more unusual, as when an animal departs in

the use of certain structures from the rest of the group to which it

belongs. For example, if all mammals lived underground, ourselves

included, and the fore-legs or arms were used for burrowing, we should

not think this unusual; but if we found an animal using all four legs to

support the body and for purposes of progression, we should, most

likely, think this was an excellent illustration of adaptation.

In other instances the condition is somewhat different. The color of

certain animals may unquestionably be of use to them in concealing them

from their enemies. In other cases the color may not serve this purpose,

or any purpose at all. Thus while in the former case we speak of the

color as an adaptation to the surroundings, in the latter we do not

think of it as having any connection at all with the environment. Even

in the same animal the color of different parts of the body may appear

under this twofold relation. For example, the green color of the skin of

the frog renders it less conspicuous amongst the green plants on the

edge of the stream, but the brilliant orange and black pigment in the

body-cavity cannot be regarded as of any use to the animal.

Adaptations for the Good of the Species

Aside from the class of adaptations that are for the good of the

individual, there is another class connected solely with the

preservation of the race. The organs for reproduction are the most

important examples of this kind. These organs are of no use to the

individual for maintaining its own existence, and, in fact, their

presence may even be deleterious to the animal. The instincts connected

with the use of these organs may lead inevitably to the death of the

individual, as in the case of the California salmon, which, on entering

fresh water in order to deposit its eggs, dies after performing this

act.

The presence of the organs of reproduction in the individual is

obviously connected with the propagation of other individuals. Indeed in

many organisms the life of the individual appears to have for its

purpose the continuation of the race. In a large number of animals the

individual dies after it has deposited its eggs. The most striking case

is that of the May-flies, whose life, as mature individuals, may last

for only a few hours. The eggs are set free by the bursting of the

abdomen, and the insect dies. The male bee also dies after union with

the queen. In some annelids, the body is also said to burst when the

eggs are set free; and in other forms those parts of the body containing

the eggs break off, and, after setting free the eggs, die. These are

extreme cases of what is seen in many animals, namely the replacement of

the old individuals by a new generation; and while in general there is

only a loose connection between the death of the individual and the

consummation of its reproductive power, yet the two run a course so

nearly parallel that several writers have attempted to explain this

connection as one of racial adaptation.

It has also been pointed out that in those higher animals that take care

of their young after birth, the life of the individual does not end with

the period of birth of the young, but extends at least throughout the

time necessary to care for the young. It has even been suggested that

this lengthening of the life period has been acquired on account of its

use to the species. When, however, as in the case of the vertebrates,

the young are born at intervals either in great numbers at a birth, as

in fishes and amphibia, or in lots of twos, threes, or fours, as in many

birds and mammals, or even only one at a time, as in a few birds and in

man, it will be evident that the relation cannot be so simple as has

been supposed. It cannot be assumed in these forms that the end of the

life of the individual is in any way connected with the ripening of the

last eggs, for, on the contrary, hundreds, or even many thousands, of

potential eggs may be present in the ovaries when the animal is

overtaken by old age, and its power of reproduction lost.

In regard to several of the lower animals, we find, in a number of cases

where there are accurate data, that the individual goes on year after

year producing young. Whether they ever grow old, in the sense of losing

their power of reproduction, has not been definitely determined, but

there is, so far as I know, no evidence to show that such a process

takes place, and these animals appear to have the power of reproducing

themselves indefinitely.

The phenomenon of old age (apart from its possible connection with the

cessation of the power of reproduction), which leads to the death of the

individual, has been looked upon by a few writers as an adaptation of

the individual for the good of the species. It has been pointed out by

these writers that the longer an individual lives, the more likely it is

to become damaged, and if along with this its powers of reproduction

diminish, as compared with younger individuals, then it stands in the

way and takes food that might be used by other, younger individuals,

that are better able to carry on the propagation of the race. It is

assumed, therefore, that the life of the individual has been shortened

for the benefit of the race. Whether such a thing is probable is a

question that will also be discussed later. We are chiefly concerned

here only in recording the different groups of phenomena that have been

regarded by biologists as adaptations.

The so-called secondary sexual characters such as the brighter colors of

the males, ornaments of different kinds, crests, color-pattern, tail

feathers, etc., organs of offence and of defence used in fighting

members of the same species, present a rather unique group of

adaptations. These characters are supposed to be of use to the

individual in conquering its rivals, or in attracting the females. They

may be considered as useful to the individual in allowing it to

propagate at the expense of its rivals, but whether the race is thereby

benefited is a question that will be carefully considered later.

The colors of flowers, that is supposed to attract insects, have been

already mentioned. The sweet fluid, or nectar, secreted by many flowers

is sought by insects, which on entering the flowers bring about

cross-fertilization. Thus while the nectar seems to be of no immediate

service to the plant itself, it is useful to the species in bringing

about the fertilization of the flowers. The odors of flowers also serve

to attract insects, and their presence is one of the means by which

insects find the flowers. This also is of advantage to the race.

Organs of Little Use to the Individual

In every organism there are parts of the body whose presence cannot be

of vital importance to the individual. We may leave out of consideration

the reproductive organs, since their presence, as has just been stated,

is connected with the continuation of the race. The rudimentary organs,

so-called, furnish many examples of structures whose presence may be of

little or of no use to the individual; in fact, as in the case of the

appendix in man, the organs may be a source of great danger to the

individual. In this respect the organism is a structure not perfectly

adapted to its conditions of life, since it contains within itself parts

that are of little or of no use, which may even lead to its destruction,

and may often expose it to unnecessary danger. Nevertheless such parts

are surprisingly infrequent, and their presence is usually accounted for

on the supposition that in the past these organs have been of use, and

have only secondarily come to play an insignificant part in the

functions of the organism. Another example of the same thing is found in

the rudimentary eyes of animals living in the dark, such as the mole and

several cave animals, fishes, amphibia, and insects.

There are still other organs, which cannot be looked upon as

rudimentary, yet whose presence can scarcely be considered as essential

to the life of the individual. It is with this class that we are here

chiefly concerned. For instance, the electric organs in some of the rays

and fish can hardly protect the animal from enemies, even when as highly

developed as in the torpedo; and we do not know of any other essential

service that they can perform. Whether the same may be also said of the

phosphorescent organs of many animals is perhaps open in some cases to

doubt, but there can be little question that the light produced by most

of the small marine organisms, such as noctiluca, jellyfish,

ctenophores, copepods, pyrosoma, etc., cannot be of use to these animals

in protecting them from attack. In the case of certain bacteria it seems

quite evident that the production of light can be of no use as such to

them. The production of light may be only a sort of by-product of

changes going on in the organism, and have no relation to outside

conditions. In certain cases, as in the glowworm, it has been supposed

that the display may serve to bring the sexes together; but since the

phosphorescent organs are also present in the larval stages of the

glowworm, and since even the egg itself is said to be phosphorescent, it

is improbable, in these stages at least, that the presence of the light

is of service to the organism.

It has been pointed out that the colors of certain animals may serve to

conceal them and may be regarded as an adaptation; but it is also true

that in many cases the color of the whole animal or the color of special

parts can be of little if any direct use. While it is difficult to show

that the wonderful patterns and magnificent coloration of many of the

larger animals are not of service to the animal, however sceptical we

may be on the subject, yet in the case of many microscopical forms that

are equally brilliantly colored there can be little doubt that the

coloration can be of no special service to them. If it be admitted that

in these small forms the color and the color patterns are not

protective, we should at least be on our guard in ascribing off-hand to

larger forms a protective value in their coloration, unless there is

actual proof that it serves some purpose.

We also see in other cases that the presence of color need not be

connected with any use that it bears as such to the animal. For

instance, the beautiful colors on the inside of the shells of many

marine snails and of bivalve mollusks, can be of no use to the animal

that makes the shell, because as long as the animal is alive this color

cannot be seen from the outside. This being the case let us not jump too

readily to the conclusion that when other shells are colored on the

outer surface that this must be of use to the mollusk.

In regard to the colors of plants, there are many cases of brilliant

coloration, which so far as we can see can be of no service to the

organism. In such forms as the lichens and the toadstools, many of which

are brilliantly colored, it is very doubtful if the color, as such, is

of any use to the plant. The splendid coloring of the leaves in the

autumn is certainly of no service to the trees.

It should not pass unnoticed in this connection that the stems and the

trunks of shrubs and of trees and also many kinds of fruits and nuts are

sometimes highly colored. It is true that some of the latter have been

supposed to owe their color to its usefulness in attracting birds and

other animals which, feeding on the fruit, swallow the seeds, and these,

passing through the digestive tract and falling to the ground, may

germinate. The dissemination of the seeds of such plants is supposed to

be brought about in this way; and since they may be widely disseminated

it may be supposed that it is an advantage to the plant to have

attracted the attention of the fruit-eating birds. On the other hand one

of the most brilliantly colored seeds, the acorn, is too large to pass

through the digestive tracts of birds, and is, in fact, ground to pieces

in the gizzard, and in the case of several mammals that feed on the

acorns, the acorn is crushed by the teeth. It would seem, therefore,

that its coloration is injurious to it rather than the reverse, as it

leads to its destruction. It has been suggested by Darwin that since the

acorns are for a time stored up in the crop of the bird, the passenger

pigeon for example, and since the birds may be caught by hawks and

killed, the seeds in the crop thus become scattered. Consequently it may

be, after all, of use to the oak to produce colored acorns that attract

the attention of these pigeons. This suggestion seems too far-fetched to

consider seriously. In the case of the horse-chestnut the rich brown

color is equally conspicuous, but the nut is too large to be swallowed

by any of the ordinary seed-feeding birds or mammals. Shall we try to

account for its color on the grounds of the poisonous character of the

seed? Has it been acquired as a warning to those animals that have eaten

it once, and been made sick or have died in consequence? I confess to a

personal repugnance to imaginative explanations of this sort, that have

no facts of experience to support them.

Changes in the Organism that are of No Use to the Individual or to the

Race

As an example of a change in the organism that is of no use to it may be

cited the case of the turning white of the hair in old age in man and in

several other mammals. The absorption of bone at the angle of the chin

in man, is another case of a change of no immediate use to the

individual. We also find in many other changes that accompany old age,

processes going on that are of no use to the organism, and which may, in

the end, be the cause of its death. Such changes, for instance, as the

loss of the vigor of the muscles, and of the nervous system, the

weakening of the heart, and partial failure of many of the organs to

carry out their functions. These changes lead sooner or later to the

death of the animal, in consequence of the breaking down of some one

essential organ, or to disease getting an easier foothold in the body.

We have already discussed the possible relation of death as an

adaptation, but the changes just mentioned take place independently of

their relation to the death of the organism as a whole, and show that

some of the normal organic processes are not for the good of the

individual or of the race. In fact, the perversions of some of the most

deeply seated instincts of the species, as in infanticide, while the

outcome of definite processes in the organism, are of obvious

disadvantage to the individual, and the perversion of so deeply seated a

process as the maternal instinct, leading to the destruction of the

young, is manifestly disadvantageous to the race. As soon, however, as

we enter the field of so-called abnormal developments, the adaptive

relation of the organism to its environment is very obscure; and yet, as

in the case of adaptation to poisons, we see that we cannot draw any

sharp line between what we call normal and what we call abnormal

development.

Comparison with Inorganic Phenomena

The preceding examples and discussion give some idea of what is meant by

adaptation in living things. In what respects, it may be asked, do these

adaptations differ from inorganic phenomena? The first group of

inorganic bodies that challenges comparison are machines. These are so

constructed that they may be said to accomplish a definite purpose, and

the question arises whether this purpose can be profitably compared with

the purposefulness of the structure and response of organisms. That the

two cannot be profitably compared is seen at once, when we recall the

fact that the activity of the machine is of no use to it, in the sense

of preserving its integrity. The object of the machine is, in fact, to

perform some useful purpose for the organism that built it, namely, for

man. Furthermore, the activity of the machine only serves to wear it

out, and, therefore, its actions do not assist in preserving its

integrity as do some, at least, of the activities of an animal. It is

true, of course, that in a mechanical sense every action of the organism

leads also to a breaking down of its structure in the same way that a

machine is also worn out by use; but the organism possesses another

property that is absent in the machine, namely, the power of repairing

the loss that it sustains.

One of the most characteristic features of the organism is its power of

self-adjustment, or of regulation, by which it adapts itself to changes

in the environment in such a way that its integrity is maintained. Most

machines have no such regulative power, although, in a sense, the

fly-wheel of an engine regulates the speed, and a water-bath, with a

thermostat, regulates itself to a fixed temperature; but even this

comparison lacks one of the essential features of the regulation seen in

organisms, namely, in that the regulation does not protect the machine

from injury. It may be claimed, however, that the safety valve of an

engine does fulfil this purpose, since it may prevent the engine from

exploding. Here, in fact, we do find better grounds for comparison, but,

when we take into account the relation of the regulations in the

organism to all the other properties of the organism, we see that this

comparison is not very significant. The most essential difference

between a machine and an organism is the power of reproduction possessed

by the latter, which is absent in all machines. Here, however, we meet

with a somewhat paradoxical relation, since the reproductive power of

organisms cannot be looked upon as an adaptation for the continuation of

the individual, but rather for the preservation of a series of

individuals. Hence, in this respect also, we cannot profitably compare

the individual with a machine, but if we make any comparison we should

compare all the individuals that have come from a single one with a

machine. In this sense the power of reproduction is a sort of racial

regulation. A comparison of this sort is obviously empty of real

significance.

The regenerative power of the organism, by means of which it may replace

a lost part, or by means of which a piece may become a new whole, is

also something not present in machines.

In using a machine for comparison we should not leave out of sight the

fact that machines are themselves the work of organisms, and have been

made for some purpose useful to the organism. They may perform the same

purpose for which we would use our own hands, for they differ from parts

of the body mainly in that they are made of different compounds having

different properties, as the above comparisons have shown. But the

regulations of the machine have been added to it by man on account of

their usefulness to himself, and are not properties of the material of

which the machine itself is composed. This shows, I think, the

inappropriateness of making any comparison between these two entirely

different things.

If, then, we find the comparison between machines and organisms

unprofitable, can we find any other things in inorganic nature that can

be better compared with the phenomenon of adaptation of the organism?

The following phenomena have been made the subject of comparison from

time to time. The bendings, which are gradually made by rivers often

lead to a meeting of the loops, so that a direct, new communication is

established, and the course of the river is straightened out. The water

takes, therefore, a more direct course to the sea. It cannot be said,

however, to be of any advantage to the river to straighten its course.

Again, a glacier moulds itself to its bed, and gradually moves around

obstacles to a lower level, but this adaptation of the glacier to the

form of its surroundings cannot be said to be of advantage to the

glacier. On the contrary, the glacier reaches so much the sooner a lower

level where it is melted.

The unusual case of a solid being lighter than the liquid from which it

forms, as seen in the case of ice, has been looked upon as a useful

arrangement, since were the reverse the case all rivers and ponds would

become solid in winter in cold climates, and the polar regions would

become one solid block of ice. But no one will suppose for a moment that

there is any relation between the anomalous condition of the lightness

of ice, and its relation to the winter freezing of streams, ponds, etc.

It has even been suggested that this property of ice was given to it in

order that the animals living in the water might not be killed, which

would be the case if the ice sank to the bottom, but such a method of

interpreting physical phenomena would scarcely commend itself to a

physicist.

The formation of a covering of oxide over the surface of a piece of iron

delays the further process of oxidation, but who will imagine that this

property of iron has been acquired in order to prevent the iron from

being destroyed by oxygen?

If a piece is broken from a crystal, and the crystal is suspended in a

saturated solution of the same substance, new material is deposited over

its whole surface, and, as it grows larger, the broken side is completed

and the crystal assumes its characteristic form. But of what advantage

is it to the crystal whether it is complete or incomplete? In the case

of an animal it is of some importance to be able to complete itself

after injury, because it can then better obtain the food necessary to

keep it alive, or it can better escape its enemies; but this is not the

case with the crystal.

In conclusion, therefore, it is obvious that the adaptations of

organisms are something peculiar to living things, and their obvious

purpose is to maintain the integrity of the individual, or that of the

species to which the individual belongs. We are, therefore, confronted

with the question as to how this peculiarity has come to be associated

with the material out of which living things are made. In subsequent

chapters this will be fully discussed, but before we take up this topic,

it will be necessary to reach some understanding in regard to the theory

of evolution, for the whole subsequent issue will turn upon the question

of the origin of the forms of animals and plants living at the present

time.

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Evolution and Adaptation · 第 1 / 13 章