Darwin
Evolution
Chapter
IV of The Origin of Species
Natural
Selection; or the Survival of the Fittest
Natural Selection—
its power compared with man's selection—
its power on characters of trifling importance—
its power at all ages
and on both sexes—
Sexual selection—On the generality of intercrosses
between individuals of the same species—
Circumstances favourable
and unfavourable
to the results of natural selection, namely,
intercrossing,
isolation, number of individuals—
Slow action—Extinction caused by
natural selection—Divergence of character, related to the diversity
of inhabitants of any small area,
and to naturalization—Action of
natural selection, through divergence
of character and extinction,
on the descendants
from a common parent—
Explains the grouping
of all organic beings—
Advance in organization—Low forms preserved—
Convergence of character—I
definite multiplication of species
—
Summary.
How will the struggle
for existence, briefly discussed in the last chapter, act in regard to
variation? Can the principle of selection which we have seen is so potent
in the hands of man, apply under nature? I think we shall see that it can
act most efficiently. Let the endless number of slight variations and
individual differences occurring in our domestic productions, and, in a
lesser degree, in those under nature, be borne in mind; as well as the
strength of the hereditary tendency. Under domestication, it may be truly
said that the who]e organization becomes in some degree plastic. But the
variability, which we almost universally meet with in our domestic
productions, is not directly produced, as Hooker and Asa Gray have well
remarked, by man; he can neither originate varieties, nor prevent their
occurrence; he can preserve and accumulate such as do occur. Unintentionally
he exposes organic beings to new and changing conditions of life, and
variability ensues, but similar changes of conditions might and do occur
under nature. Let it also be borne mind how infinitely complex and
close-fitting arc the mutual relations of all organic beings to each other
and to their physical conditions of life; and consequently what infinitely
varied diversities of structure might be of use to each being under
changing conditions of life. Can it, then, be thought improbable seeing
that variations useful to man have undoubtedly occurred that other
variations useful in some way to each being in the great and complex
battle of life, should occur in the course of many successive generations.
If such do occur, can we doubt (remembering that many more individuals are
born than can possibly survive) that individuals having any advantage,
however slight, over others should have the best chance of surviving and
of procreating their kind? On the other hand, we may feel sure that any
variation in the least degree injurious would be rigidly destroyed. This
preservation of favourable individual differences and variations, and the
destruction of those which are injurious, I have called Natural Selection
or the Survival of the Fittest. Variations neither useful nor injurious
would not be affected by natural selection, and would be left either a
fluctuating element, as perhaps we see in certain polymorphic species, or
would ultimately become fixed, owing to the nature of the organism and the
nature of the conditions.
Several writers have
misapprehended or objected to the term Natural Selection. Some have even
imagined that natural selection induces variability, whereas it implies
only the preservation of such variations as arise and are beneficial to
the being under its conditions of life. No one objects to agriculturists
speaking of the potent effects of man's selection; and in this case the
individual differences given by nature, which man for some object selects
must of necessity first occur. Others have objected that the term
selection implies conscious choice in the animals which become modified;
and It had even been urged that, as plants have no volition, natural
selection is not applicable to them! In the literal sense of the word, no
doubt, natural selection is a false term, but who ever objected to
chemists speaking of the elective affinities of the various
elements?—and yet an acid cannot strictly be said to elect the base with
which it in preference combines. It has been said that I speak of natural
selection as an active power or Deity; but who objects to an author
speaking of the attraction of gravity as ruling the movements of the
planets? Every one knows what is meant and is implied by such metaphorical
expressions; and they are almost necessary for brevity. So again it Is
difficult to avoid personifying the word Nature; but I mean by Nature,
only the aggregate action and product of many natural laws, and by laws
the sequence of events as ascertained by us. With a little familiarity
such superficial objections will be forgotten. We shall best understand
the probable course of natural selection by taking the case of a country
undergoing some slight physical change, for instance, of climate. The
proportional numbers of Its inhabitants will almost immediately undergo a
change, and some species will probably become extinct. We may conclude,
from what we have seen of the intimate and complex manner in which the
inhabitants of each country are bound together, that any change in the
numerical proportions of the inhabitants, independently of the change of
climate itself, would seriously affect the others If the country were open
on its borders, new forms would certainly immigrate, and this would
likewise seriously disturb the relations of some of the former
inhabitants. Let it be remembered how powerful the influence of a single
introduced tree or mammal has been shown to be. But in the case of an
island, or of a country partly surrounded by barriers, into which new and
better adapted forms could not freely enter, we should then have places in
the economy of nature which would assuredly be better filled up, if some
of the original inhabitants were in some manner modified; for, had the
area been open to immigration, these same places would have been seized on
by intruders. In such cases, slight modifications, which in any way
favoured the individuals of any species, by better adapting them to their
altered conditions, would tend to be preserved; and natural selection
would have free scope for the work of improvement.
We have good reason to
believe, as shown in the first chapter, that changes in the conditions of
life give a tendency to increased variability—and in the foregoing cases
the conditions have changed, and this would manifestly be favourable to
natural selection, by affording a better chance of the occurrence of
profitable variations. Unless such occur, natural selection can do
nothing. Under the term of “variations,” it must never be forgotten
that mere individual differences are included. As man can produce a great
result with his domestic animals and plants by adding up in any given
direction individual differences, so could natural selection, but ar more
easily from having incomparably longer time for action. Nor do I believe
that any great physical change, as of climate, or any unusual degree of
isolation to check immigration, is necessary in order that new and
unoccupied places should be left, for natural selection to fill up by
improving some of the varying inhabitants with nicely balanced forces,
extremely slight modifications in the structure or habits of one species
would often give it an advantage over others; and still further
modifications of the same kind would often still further increase the
advantage, as long as the species continued under the same conditions of
life and profited by similar means of subsistence and defence. No country
can be named m which all the native inhabitants are now so perfectly
adapted to each other and to the physical conditions under which they
live, that none of them could be still better adapted or improved; for in
all countries, the natives have been so far conquered by naturalized
productions, that they have allowed some foreigners to take firm
possession of the land. And as foreigners have thus in every country
beaten some of the natives, we may safely conclude that the natives might
have been modified with advantage, so as to have better resisted the
intruders.
As man can produce, and
certainly has produced, a great result by his methodical and unconscious
means of selection, what may not natural selection effect? Man can act
only on external and visible characters: Nature, if I may be allowed to
personify the natural preservation or survival of the fittest, cares
nothing for appearances, except in so far as they are useful to any being.
She can act on every internal organ, on every shade of constitutional
difference, on the whole machinery of life. Man selects only for his own
good: Nature only for that of the being which she tends. Every selected
character Is fully exercised by her, as is implied by the fact of their
selection. Man keeps the natives of many climates in the same country; he
seldom exercises each selected character in some peculiar and fitting
manner; he feeds a long and a short beaked pigeon on the same food; he
does not exercise a long-backed or long-legged quadruped in any peculiar
manner; he exposes sheep with long and short wool to the same climate. He
does not allow the most vigorous males to struggle for the females. He
does not rigidly destroy all inferior animals, but protects during each
varying season as far as lies in his power, all his productions. He often
begins his selection by some half-monstrous form; or at least by some
modification prominent enough to catch the eye or to be plainly useful to
him. Under Nature, the slightest differences of structure or constitution
may well turn the nicely balanced scale in the struggle for life, and so
be preserved. How fleeting are the wishes and efforts of man, how short
his time! And consequently how poor will be his results, compared with
those accumulated by Nature during whole geological periods! Can we
wonder, then, that Nature's productions should be far “truer” in
character than man's productions that they should be infinitely better
adapted to the most complex conditions of life and should plainly bear the
stamp of far higher workmanship?
It may metaphorically
be said that natural selection is daily and hourly scrutinising,
throughout the world, the slightest variations; rejecting those that are
bad, preserving and adding up all that are good, silently and insensibly
working, whenever and wherever opportunity offers, at the improvement of
each organic being in relation to its organic and inorganic conditions of
life. We see nothing of these slow changes in progress, until the hand of
time has marked the lapse of ages, and then so imperfect is our view into
long-past geological ages, that we see only that the forms of life are now
different from what they formerly were. In order that any great amount of
modification should be seen in a species, a variety when once formed must
again, perhaps after a long interval of time, vary or present individual
differences of the same favourable nature as before; and these must be
again preserved, and so onwards step by step. Seeing that individual
differences of the same kind perpetually recur, this can hardly be
considered as an unwarrantable assumption. But whether it is true, we can
judge only by seeing how far the hypothesis accords with and explains the
general phenomena of nature. On the other hand, the ordinary belief that
the amount of possible variation is a strictly limited quantity is
likewise a simple assumption. Although natural selection can act only
through and for the good of each being, yet characters and structures,
which we are apt to consider as of very trifling importance, may thus be
acted on. When we see leaf-eating insects green, and bark-feeders mottled-grey;
the alpine ptarmigan white in winter, the red-grouse the colour of
heather, we must believe that these tints are of service to these birds
and insects in preserving them from danger. Grouse, if not destroyed at
some period of their lives would increase in countless numbers- they are
known to suffer largely from birds of prey; and hawks are guided by
eyesight to their prey--so much so, that on parts of the Continent persons
are warned not to keep white pigeons, as being the most liable to
destruction. Hence natural selection might be effective in giving the
proper colour to each kind of grouse, and in keeping that colour, when
once acquired, true and constant. Nor ought we to think that the
occasional destruction of an animal of any particular colour would produce
little effect: we should remember how essential it is in a flock of white
sheep to destroy a lamb with the faintest trace of black. We have seen how
the colour of the hogs, which feed on the “paint-root” in Virginia,
determines whether they shall live or die. In plants, the down on the
fruit and the colour of the flesh are considered by botanists as
characters of the most trifling importance: yet we hear from an excellent
horticulturist, Downing, that in the United States, smooth-skinned fruits
suffer far more from a beetle, a Curculio, than those with down; that
purple plums suffer far more from a certain disease than yellow plums;
whereas another disease attacks yellow-fleshed peaches far more than those
with other coloured flesh. If, with all the aids of art, these slight
differences make a great difference in cultivating the several varieties,
assuredly, in a state of nature, where the trees would have to struggle
with other trees, and with a host of enemies, such differences would
effectually settle which variety, whether a smooth or downy, a yellow or
purple fleshed fruit, should succeed.
In looking at many
small points of difference between species which, as far as our ignorance
permits us to judge, seem quite unimportant, we must not forget that
climate, food, &c., have no doubt produced some direct effect. It is
also necessary to bear in mind that, owing to the law of correlation, when
one part varies an the variations are accumulated through natural
selection, other modifications, often of the most unexpected nature, will
ensue.
As we see that those
variations which, under domestication, appear at any particular period of
life, tend to reappear in the offspring at the same period;--for instance,
in the shape, size, and flavour of the seeds of the many varieties of our
culinary and agricultural plants; in the caterpillar and cocoon stages of
the varieties of the silk-worm; in the eggs of poultry, and in the colour
of the down of their chickens; in the horns of our sheep and cattle when
nearly adult;—so in a state of nature natural selection will be enabled
to act on and modify organic beings at any age, by the accumulation of
variations profitable at that age, and by their inheritance at a
corresponding age. If it profit a plant to have its seeds more and more
widely disseminated by the wind, I can see no greater difficulty in this
being effected through natural selection, than in the cotton-planter
increasing and improving by selection the down in the pods on his
cotton-trees. Natural selection may modify and adapt the larva of an
insect to a score of contingencies, wholly different from those which
concern the mature insect, and these modifications may effect, through
correlation, the structure of the adult So, conversely, modifications in
the adult may affect the structure of the larva; but m all cases natural
selection will ensure that they shall not be injurious: for if they were
so, the species would become extinct.
Natural selection will
modify the structure of the young in relation to the parent, and of the
parent in relation to the young. In social animals it will adapt the
structure of each individual for the benefit of the whole community; if
the community profits by the selected change. What natural selection
cannot do, is to modify the structure of one species, without giving it
any advantage, for the good of another species; and though statements to
this effect may be found in works of natural history, I cannot find one
case which will bear investigation. A structure used only once in an
animal's life, if of high importance to it, might be modified to any
extent by natural selection; for instance, the great jaws possessed by
certain insects, used exclusively for opening the cocoon—or the hard tip
to the beak of unhatched birds, used for breaking the egg. It has been
asserted, that of the best short-beaked tumbler-pigeons a greater number
perish in the egg than are able to get out n it; so that fanciers assist
in the act of hatching. Now if nature had to make the beak of a full-grown
pigeon very short for the bird's own advantage, the process of
modification would be very slow, and there would be simultaneously the
most rigorous selection of all the young birds within the egg, which had
the most powerful and hardest beaks, for all with weak beaks would
inevitably perish; or, more delicate and more easily broken shells might
be selected, the thickness of the shell being known to vary like every
other structure. It may be well here to remark that with all beings there
must be much fortuitous destruction, which can have little or no influence
on the course of natural selection. For instance a vast number of eggs or
seeds are annually devoured, and these could be modified through natural
selection only if they varied in some manner which protected them from
their enemies. Yet many of these eggs or seeds would perhaps, if not
destroyed, have yielded individuals better adapted to their conditions of
life than any of those which happened to survive. So again a vast number
of mature animals and plants, whether or not they be the best adapted to
their conditions, must be annually destroyed by accidental causes, which
would not be in the least degree mitigated by certain changes of structure
or constitution which would in other ways be beneficial to the species.
But let the destruction of the adults be ever so heavy, If the number
which can exist in any district be not wholly kept down by such
causes,—or again let the destruction of eggs or seeds be so great that
only a hundredth or a thousandth part are developed,—yet of those which
do survive, the best adapted individuals, supposing that there is any
variability in a favourable direction, will tend to propagate their kind
in larger numbers than the less well adapted. If the numbers be wholly
kept down by the causes just indicated, as will often have been the case,
natural selection will be powerless in certain beneficial directions; but
this is no valid objection to its efficiency at other times and in other
ways; for we are far from having any reason to suppose that many species
ever undergo modification and improvement at the same time in the same
area.
Sexual
Selection
Inasmuch as
peculiarities often appear under domestication in one sex and become
hereditarily attached to that sex, so no doubt it will be under nature.
Thus it is rendered possible for the two sexes to be modified through
natural selection in relation to different habits of life, as Is sometimes
the case; or for one sex to be modified in relation to the other sex, as
commonly occurs. This leads me to say a few words on what I have called
Sexual Selection. This form of selection depends, not on a struggle for
existence in relation to other organic beings or to external conditions,
but on a struggle between the individuals of one sex, generally the males,
for the possession of the other sex. The result is not death to the
unsuccessful competitor, but few or no offspring. Sexual selection is,
therefore less rigorous than natural selection. Generally, the most
vigorous males, those which are best fitted for their places in nature,
will leave most progeny. But in many cases, victory depends not so much on
general vigor, as on having special weapons, confined to the male sex. A
hornless stag or spurless cock would have poor chance of leaving numerous
offspring. Sexual selection, by always allowing the victor to breed, might
surely give indomitable courage, length to the spur, and strength to the
wing to strike in the spurred leg, in nearly the same manner as does the
brutal cock-fighter by the careful selection of his best cocks. How low in
the scale of nature the law of battle descends, I know not; male
alligators have been described as fighting, bellowing, and whirling round,
like Indians in a war dance, for the possession of the females; male
salmons have been observed fighting all day long; male stag-beetles
sometimes bear wounds from the huge mandibles of other males, the males of
certain hymenopterous insects have been frequently seen by that Inimitable
observer M. Fabre, fighting for a particular female who sits by, an
apparently unconcerned beholder of the struggle, and then retires with the
conqueror. The war is, perhaps, severest between the males of polygamous
animals, and these seem oftenest provided with special weapons. The males
of carnivorous animals are already well armed; though to them and to
others, special means of defence may be given through means of sexual
selection, as the mane of the lion, and the hooked jaw to the male salmon,
for the shield may be as important for victory, as the sword or spear
Amongst birds, the
contest is often of a more peaceful character All those who have attended
to the subject, believe that there is the severest rivalry between the
males of many species to attract, by singing, the females. The rock-thrush
of Guiana, birds of paradise and some others, congregate; and successive
males display with the most elaborate care, and show off in the best
manner, their gorgeous plumage; they likewise perform strange antics
before the females which, standing by as spectators, at last choose the
most attractive partner. Those who have closely attended to birds in
confinement well know that they often take individual preferences and
dislikes: Sir R. Heron has described how a pied peacock was eminently
attractive to all his hen birds. I cannot here enter on the necessary
details, but if man can in a short time give beauty and an elegant
carriage to his bantams, according to the standard of beauty, I can see no
good reason to doubt that female birds, by selecting, during thousands of
generations, the most melodious or beautiful males according to their
standard of beauty, might produce a marked effect. Some well-known laws,
with respect to the plumage of male and female birds, in comparison with
the plumage of the young, can partly be explained through the action of
sexual selection on variations occurring at different ages, and
transmitted to the males alone or to both sexes at corresponding ages; but
I have not space here to enter on this subject.
Thus it is, as I
believe, that when the males and females of any animal have the same
general habits of life, but differ in structure, colour, or ornament, such
differences have been mainly caused by sexual selection: that is, by
individual males having had, in successive generations, some slight
advantage over other males, in their weapons, means of defence, or charms,
which they have transmitted to their male offspring alone. Yet, I would
not wish to attribute a sexual differences to this agency: for we see in
our domestic animals peculiarities arising and becoming attached to the
male sex, which apparently have not been augmented through selection by
man. The tuft of hair on the breast of the wild turkey-cock cannot be of
any use, and it is doubtful whether it can be ornamental in the eyes of
the female bird;—indeed, had the tuft appeared under domestication, it
would have been called a monstrosity.
Illustrations
of the Action of Natural Selection, or the Survival of the Fittest
In order to make it
clear how, as I believe, natural selection acts, I must beg permission to
give one or two imaginary illustrations. Let us take the case of a wolf,
which preys on various animals, securing some by craft, some by strength
and some by fleetness; and let us suppose that the fleetest prey, a deer
for instance, had from any change in the country increased in numbers or
that other prey had decreased in numbers, during that season of the year
when the wolf was hardest pressed for food. Under such circumstances the
swiftest and slimmest wolves would have the best chance of surviving and
so be preserved or selected,--provided always that they retained strength
to master their prey at this or some other period of the year, when they
were compelled to prey on other animals, can see no more reason to doubt
that this would be the result, than that man should be able to improve the
fleetness of his greyhounds y careful and methodical selection, or by that
kind of unconscious selection which follows from each man trying to keep
the best dogs without any thought of modifying the breed. I may add, that,
according to Mr. Pierce, there are two varieties of the wolf inhabiting
the Catskill Mountains, in the United States, one with a light
greyhound-like form, which pursues deer, and the other more bulky, with
shorter legs, which more frequently attacks the sheep
It should be observed
that, in the above illustration, I speak of the slimmest individual
wolves, and not of any single strongly-marked variation having been
preserved. In former editions of this work I sometimes spoke as if this
latter alternative had frequently occurred. I saw the great importance of
individual differences, and this led me fully to discuss the results of
unconscious selection by man, which depends on the preservation of all the
more or less valuable individuals, and on the destruction of the worst. I
saw also, that the preservation in a state of nature of any occasional
deviation of structure, such as a monstrosity, would be a rare event and
that, If at first preserved, it would generally be lost by subsequent
Intercrossing with ordinary individuals. Nevertheless, until reading an
able and valuable article in the ‘North British Review’ (l867), I did
not appreciate how rarely single variations, whether slight or strongly
marked, could be perpetuated. The author takes the case of a pair of
animals, producing during their lifetime two hundred offspring, of which,
from various causes of destruction, only two on an average survive to
procreate their kind. This is rather an extreme estimate for most of the
higher animals, but by no means so for many of the lower organisms. He
then shows that if a single Individual were born, which varied in some
manner, giving it twice as good a chance of life as that of the other
individuals, yet the chances would be strongly against its survival.
Supposing it to survive and to breed, and that half its young inherited
the favourable variation; still, as the Reviewer goes on to show, the
young would have only a slightly better chance of surviving and breeding;
and this chance would go on decreasing in the succeeding generations. The
justice of these remarks cannot, I think, be disputed. If for instance, a
bird of some kind could procure its food more easily by having its beak
curved, and if one were born with its beak strongly curved, and which
consequently flourished, nevertheless ere would be a very poor chance of
this one individual perpetuating its kind to the exclusion of the common
form; but there can hardly be a doubt, judging by what we see taking place
under domestication, that this result would follow from the presentation
during many generations of a large number of individuals with more or less
strongly curved beaks, and from the destruction of a still larger number
with the straightest beaks.
It should not, however,
be overlooked that certain rather strongly marked variations, which no one
would rank as mere individual differences, frequently recur owing to a
similar organization being similarly acted on—of which fact numerous
instances could be given with our domestic productions. In such cases, If
the varying individual did not actually transmit to its offspring its
newly-acquired character, it would undoubtedly transmit to them, as long
as the existing conditions remained the same, a still stronger tendency to
vary in the same manner. There can also be little doubt that the tendency
to vary in the same manner has often been so strong that all the
individuals of the same species have been similarly modified without the
aid of any form of selection. Or only a third, fifth, or tenth part of the
individuals may have been thus affected, of which fact several instances
could be given. Thus Graba estimates that about one-fifth of the
guillemots in the Faroe Islands consist of a variety so well marked, that
it was formerly ranked as a distinct species under the name of Uria
lacrymans. In cases of this kind, if the variation were of a beneficial
nature, the original form would soon be supplanted by the modified form,
through the survival of the fittest.
To the effects of
intercrossing in eliminating variations of all kinds, I shall have to
recur; but it may be here remarked that most animals and plants keep to
their proper homes, and do not needlessly wander about; we see this even
with migratory birds, which almost always return to the same spot.
Consequently each newly-formed variety would generally be at first local,
as seems to be the common rule with varieties in a state of nature; so
that similarly modified individuals would soon exist in a small body
together, and would often breed together. If the new variety were
successful in its battle for life, it would slowly spread from a central
district, competing with and conquering the unchanged individuals on the
margins of an ever-increasing circle.
It may be worth while
to give another and more complex illustration of the action of natural
selection. Certain plants excrete sweet juice, apparently for the sake of
eliminating something injurious from the sap: this is effected, for
instance, by glands at the base of the stipules in some Leguminosa, and at
the backs of the leaves of the common laurel. This juice, though small in
quantity, is greedily sought by insects; but their visits do not in any
way benefit the plant . Now, let us suppose that the juice or nectar was
excreted from the inside of the flowers of a certain number of plants of
any species. Insects in seeking the nectar would get dusted with pollen,
and would often transport it from one flower to another. The flowers of
two distinct individuals of the same species would thus get crossed, and
the act of crossing, as can be fully proved, gives rise to vigorous
seedlings which consequently would have the best chance of flourishing and
surviving. The plants which produced flowers with the largest glands or
nectaries, excreting most nectar, would oftenest be visited by insects,
and would oftenest be crossed; and so in the long-run would gain the upper
hand and form a local variety. The flowers, also, which had their stamens
and pistils placed, in relation to the size and habits of the particular
insects which visited them, so as to favour in any degree the transportal
of the pollen, would likewise be favoured. We might have taken the case of
insects visiting flowers for the sake of collecting pollen instead of
nectar; and as pollen is formed for the sole purpose of fertilization, its
destruction appears to be a simple loss to the plant; yet if a little
pollen were carried, at first occasionally and then habitually, by the
pollen-devouring insects from flower to flower, and a cross thus effected,
although nine-tenths of the pollen were destroyed it might still be a
great gain to the plant to be thus robbed, and the individuals which
produced more and more pollen, and had larger anthers, would be selected.
When our plant, by the
above process long continued, had been rendered highly attractive to
insects, they would, unintentionally on their part, regularly carry pollen
from flower to flower, and that they do this effectually, I could easily
show by many striking facts. I will give only one, as likewise
illustrating one step in the separation of the sexes of plants. Some
holly-trees bear only male flowers, which have four stamens producing a
rather small quantity of pollen, and a rudimentary pistil: other
holly-trees bear only female flowers; these have a full-sized pistil, and
four stamens with shrivelled anthers, in which not a grain of pollen can
be detected. Having found a female tree exactly sixty yards from a male
tree, I put the stigmas of twenty flowers, taken from different branches
under the microscope, and on all, without exception, there were a few
pollen-grains, and on some a profusion. As the wind had set for several
days from the female to the male tree, the pollen could not thus have been
carried. The weather had been cold and boisterous and therefore not
favourable to bees, nevertheless every female flower which I examined had
been effectually fertilized by the bees which had flown from tree to tree
in search of nectar. But to return to our imaginary case: as soon as the
plant had been rendered so highly attractive to insects that pollen was
regularly carried from flower to flower, another process might commence.
No naturalist doubts the advantage of what has been called the
"physiological division of labour"; hence we may believe that it
would be advantageous to a plant to produce stamens alone in one flower or
on one whole plant, and pistils alone in another flower or on another
plant. In plants under culture and placed under new conditions of life
sometimes the male organs and sometimes the female organs become more or
less impotent; now if we suppose this to occur in ever so slight a degree
under nature, then, as pollen is already carried regularly from flower to
flower, and as a more complete separation of the sexes of our plant would
be advantageous on the principle of
the division of labour,
individuals with this tendency more and more increased would be
continually favoured or selected, until at last a complete separation of
the sexes might be effected. It would take up too much space to show the
various steps, through dimorphism and other means, by which the separation
of the sexes m plants of various kinds is apparently now in progress; but
I may add that some of the species of holly in North America, are,
according to Asa Gray, in an exactly intermediate condition, or, as he
expresses It, are more or less dioeciously polygamous.
Let us now turn to the
nectar-feeding insects; we may suppose the plant, of which we have been
slowly increasing the nectar by continued selection, to be a common plant;
and that certain insects depended in main part on its nectar for food. I
could give many facts showing how anxious bees are to save time: for
instance, their habit of cutting holes and sucking the nectar at the bases
of certain flowers, which, with a very little more trouble, they can enter
by the mouth. Bearing such facts in mind, it may be believed that under
certain circumstances individual differences in the curvature or length of
the proboscis, &c., too slight to be appreciated by us, might profit a
bee or other insect, so that certain individuals would be able to obtain
their food more quickly than others; and thus the communities to which
they belonged would flourish and throw off many swarms inheriting the same
peculiarities. The tubes of the corolla of the common red and incarnate
clovers (Trifolium pratense and incarnatum) do not on a hasty glance
appear to differ in length; yet the hive-bee can easily suck the nectar
out of the incarnate clover, but not out of the common red clover, which
is visible by humble-bees alone; so that whole fields of red clover offer
in vain an abundant supply of precious nectar to the hive-bee. That this
nectar is much liked by the hive-bee is certain; for I have repeatedly
seen, but only in the autumn, many hive-bees sucking the flowers through
holes bitten in the base of the tube by bumble-bees. The difference in the
length of the corolla in the two kinds of clover, which determines the
visits of the hive-bee, must be very trifling; for I have been assured
that when red clover has been mown, the flowers of the second crop are
somewhat smaller, and that these are visited by many hive-bees. I do not
know whether this statement Is accurate, nor whether another published
statement can be trusted, namely, that the Ligurian bee which is generally
considered a mere variety of the common hive-bee, and which freely crosses
with It, Is able to reach and suck the nectar of the red clover. Thus, in
a country where this kind of clover abounded, it might be a great
advantage to the hive-bee to have a slightly longer or differently
constructed proboscis. On the other hand, as the fertility of this clover
absolutely depends on bees visiting the flowers, if humble-bees were to
become rare in any country, it might be a great advantage to the plant to
have a shorter or more deeply divided corolla, so that the hive-bees
should be enabled to suck its flowers. Thus I can understand how a flower
and a bee might slowly become, either simultaneously or one after the
other, modified and adapted to each other In the most perfect manner, by
the continued preservation of all the individuals which presented slight
deviations of structure mutually favourable to each other.
I am well aware that
this doctrine of natural selection, exemplified in the above imaginary
instances, is open to the same objections which were first urged against
Sir Charles Lyell's noble views on the modern changes of the earth, as
illustrative of “geology”; but we now seldom hear the agencies which
we see still at work spoken of as trifling or insignificant, when used in
explaining the excavation of the deepest valleys of the formation of long
lines of inland cliffs. Natural selection acts only by the preservation
and accumulation of small inherited modifications, each profitable to the
preserved being; and as modern geology has almost banished such views as
the excavation of a great valley by a single diluvial wave, so will
natural selection banish the belief of the continued creation of new
organic beings, or of any great and sudden modification in their structure.
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