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them. Meanwhile a natural destruction, or failure of intermediate
forms to survive, has gone on.
 CHAPTER IX
SPECIES IN THE MAKING

A
SERIES of important conceptions are implied in the word
"species," as used by naturalists. Some of these we have noted
in the last chapter. There is first, as a starting-point, the conception
that a species is a number or company of individuals, all closely and
clearly alike (though presenting some minor individual variations),
and capable of sharp separation by certain "characters" from other
similar groups or companies. Then follows the addition (2) that the
species is constant if the conditions of life are not changed, or but
little changed, and that year after year it reproduces itself without
change. It has a certain stability (but not permanent immutability)
greater in some species than in others. Next we find (3) that the
species constitutes a group of individuals which have descended by
natural breeding from common parents, not differing greatly from
the present individuals. They are, in fact, one "stock." Then (4) that
the species is a group, the individuals of which pair with one another
in breeding, but do not pair with the individuals of another species,
and that this is due to various peculiar and inherent chemical,
physiological and (in higher animals) psychological characteristics of
the species.
We have now further to note that species have their special
geographical centres of origin from which most spread only a small
distance, whilst others have a wonderful power of dispersal, and
have become cosmopolitan. Moreover, we find that some species are
numerically very abundant, others very rare; that rare and abundant
species have often invaded each other's territory, and exist side by
side.
 Whilst we often find a number of species, fifty or more, so much
alike that we unite them in a single genus (as, for instance, in the
case of the cats, lions, tigers, leopards, which form the genus "Felis,"
and the hundred or more species of the hedge brambles or
blackberries, which form the genus "Rubus"), there are many
species which to-day have, as it were, lost all their relatives and
stand alone, the solitary species in a well-marked genus, or have
perhaps only one other living co-species. And sometimes (curiously
enough) that one co-species is an inhabitant of a region very remote
from that inhabited by the other. Thus the two living mammals called
tapirs (genus Tapirus) inhabit, the one the Malay region, and the
other Central America. This is explained by the fact that tapirs
formerly existed all over the land-surfaces of North Europe, North
Asia, and North America, which connect these widely-separate spots.
We find the bones and teeth of the extinct tapirs embedded in the
Tertiary deposits of the connecting regions.
Once we have gained the fundamental conceptions as to what is
meant by a "species," we are able intelligently to consider
innumerable facts of the most diverse kind as to their peculiar
structure and colours, their number, localities, their interaction and
dependence on other living things, their modifications for special
modes of life, their isolation or their ubiquity. We can discuss their
genetic relations to one another, and to extinct fossil species, which
have all been to a very large extent "accounted for" or "explained"
by Mr. Darwin's theory of the origin of species by the natural
selection of favoured races in the struggle for existence. But there is
always more to be made out—difficulties to be removed, new
instances to be studied. The classification of the genera of plants
and animals, with their included species into larger groups, helps us
to state and to remember their actual build and structure, and to
survey, as it were, the living world, from the animalcule to the man,
or from the microbe to the magnolia tree. Every one interested in
natural history should carry in his mind as complete a scheme of the
classification of animals and plants as possible.
 The older naturalists held that species were suddenly "created" as
they exist, and have propagated their like ever since. Darwin has
taught us that the present "species" have developed by a slow
process of transformation from preceding species, and these from
other predecessors, and so on to the remotest geologic ages and the
dawn of life. The agents at work have been "variation"—that is to
say, the response to the never-ceasing variation of the surrounding
world or environment—and the survival in the struggle for existence
of the fittest varieties so produced.
There is nothing surprising or extraordinary in the existence of
variation. The conditions of life and growth are never absolutely
identical in two individuals, and the wonder is not that species vary,
but that they vary so little. The living substance of animals and
plants is an extremely complex chemical substance, ever
decomposing and ever being renewed. It is the most "labile" as it is
by far the most elaborately built-up chemical body which chemists
have ever ventured to imagine. It differs, chemically, not only in
every species but in every individual and is incessantly acted upon—
influenced as we may say—by the ever-changing physical and
chemical conditions around it. At the same time it has, subject to the
permanence of essential conditions, a definite stability and limitation
to its change or variation in response to variations of its
environment. That part of the living substance which in all but the
lowest plants and animals is set aside during growth to form the
eggs and sperms by which they multiply or "reproduce" themselves,
is called the "germ-plasm," and is peculiarly sensitive to variations in
(that is a change in) the environment of the plant or animal.
New conditions of life (locality and climate)—unusual food or
reproductive activity—act often in a powerful way upon the germ-
plasm and cause it to vary—that is to say, they alter some of its
qualities, though not necessarily disturbing in any way the general
living substance of the organism so far as to produce any important
change perceptible to the human eye. In consequence, the young
produced after such disturbance of the germ-plasm are found to
differ more from their parents than in cases where no such
disturbance has been set up by the natural never-ceasing variation
of the surrounding world. This fact is well known to horticulturists
and breeders, and is made use of by them. When a gardener wishes
to obtain variations of a plant from which to select and establish a
new breed, he deliberately sets to work to disturb—to shake up, to
act upon in a tentative, experimental way—the germ-plasm of one or
more parent plants by change of soil, climate, food and often by
cross-fertilizing them with another breed or variety. In this way he to
some extent "breaks" the constitutional stability of the germ-plasm
of the plant and obtains abundant "variations" in the offspring.
These are not precisely foreseen, and show themselves in all parts of
the new generation. But some of them are what the gardener wants,
and are "selected" by him for retention, rearing and breeding.
The response of the germ-plasm of organisms to the stimulus of
new environmental conditions has been compared to that of the
well-known pattern-producing toy—the kaleidoscope. The bits of
glass, beads and silk which you see in a kaleidoscope, forming by
reflection in its mirrors a beautiful and definite pattern, are changed
by a simple vibration caused by tapping the instrument into a very
different pattern, the coloured fragments being displaced and
rearranged. The apparent change or variation is very great though
produced by slight mechanical disturbance, and the new pattern is
altogether without any special significance—the fortuitous outcome
of a small displacement of the constituent coloured fragments. We
can imagine that similarly slight disturbances of the organic
molecules of the germ-plasm may produce considerable and
important variations in it and the new growth to which it gives rise:
and, further, that these variations may prove to be either (1)
injurious, or (2) of life-saving value, or often enough (3) of no
consequence whatever although bulking largely in our human eyes
and thereby misleading our judgment of them.
There is no reason to doubt that the same sequence of events
occurs in nature apart from man's interference. Changes occur in the
earth's surface, or the organism is transported by currents of water
or air into new conditions. The germ-plasm is "disturbed," "shaken"
or "shocked" by those new conditions, and a variation, in several
structures and qualities of the offspring subsequently produced,
follows. Then also follows the selection of one of the new varieties
by survival of the fitter to the new conditions into which the
organism has been transported or have developed in the region
where it was previously established.
This process of germ-variation is obviously as necessary and
constant a feature of the living organism as is the variation in the
contour of land and sea and in the extent of the polar ice-cap—a
necessary feature of the physical conditions of the terrestrial globe.
But it is the fashion with a certain school of writers nowadays to
declare that "variation" in organisms is a "mystery" unsolved.
Another very common and almost universal error is to overlook the
fact that variation is constitutional and affects whole systems of
organs and their deeply related parts, and is not, as it is so
frequently and erroneously assumed to be, a mere local affair of
patches and scraps visible on this or that part of the surface of an
animal or plant. These superficial "marks," readily seen and noted by
the collector, are rarely of any life-saving importance: they are but
the outward and visible signs of deep-lying physiological or
constitutional change or variation. The varying organism has, like
Hamlet, "that within which passeth show" and the superficial
variations (like his "inky cloak" and other customary features of
mourning) are but "the trappings and the suits" of a deep-lying
change. Variation is not an inexplicable mystery, nor, on the other
hand, are "varieties" sufficiently dealt with and their nature
appreciated when one or two surface peculiarities are enumerated
by which the collector can recognize them. A deeper study of the
varying organism is both possible and needed.
If the gradual formation of new species from ancestral species is a
true account of the matter, we must expect to find, at any rate here
and there, if not frequently, traces of the process—for instance,
gradations, or series of intermediate forms, connecting new, well-
established species with the ancestral form or with one another. We
do find such gradations—sometimes more, sometimes less,
completely persisting over a wide tract of country, or discoverable in
the fossiliferous deposits containing the remains of extinct animals.
For instance, when we look at the butterflies of a much larger
region than our little island—namely, at those of a great continent
like Africa or South America—we find that there are species which
show gradations. Thus at a series of points, A, B, C, D, separated by
some hundreds of miles from each other, we find a corresponding
series of butterflies which are apparently closely similar species of
one genus, differing by a few spots of colour, or darker and lighter
tint, much as our Large White, Garden White, and Green-veined
White differ. But when the butterflies are caught which occur at
points intermediate between A and B, B and C, C and D, we find
intermediate varieties, and, in fact, if we get a very large number
from intermediate regions, we can, in some instances, arrange them
in line so that they constitute a graduated series of forms, each
being scarcely distinguishable from the one before or the one behind
it, yet differing clearly from one a dozen places away. In such cases
there is often evidence to show that the variety found at A breeds
with that found at B, that of B with that of C, of C with D, so that
they form an inter-breeding group, though perhaps the varieties at D
will not pair with those at A, or even with those at B. Then
sometimes we find in such a series, otherwise complete, a gap. Let
us suppose it is between the butterflies of B and C. We find the
series of gradations nearly complete, but some natural condition—
such as the encroachment of the sea, or the slow elevation of a
mountain range, or the climatic destruction of the necessary food-
plant—has "wiped out" a few forms somewhere between those of B
and C. They no longer exist. The series is no longer connected by
inter-breeding forms; those occurring from A to B and some distance
beyond are one "species" varying in the direction of the series C to
D, but abruptly broken off from the latter. The series C to D is also a
"species" with graduated varieties, but distinct; it is cut off from the
lot once in continuity with it by the destruction of the intermediate
forms inhabiting an intermediate area. Thus the one species
becomes two, and these may again break up, and, having become
thus disconnected and stabilized, they may spread over one
another's territory—fly side by side and yet remain distinct forms
which do not pair together—although originally they were varieties
spreading from a common centre, where the ancestral species lived
and multiplied.
Other similar gradational series of an interesting character have
been noticed in the case of fresh-water fossil snail-shells. In the
layers of clay and marl exposed by digging a railway cutting or a pit
we may find that the successive layers represent a continuous
deposit of 100,000 years or more, and we find sometimes that a
form of snail-shell (not a species living to-day) occurs in the lowest
stratum very different from that occurring in the highest stratum—
the lowest being short and spherical, the highest elongated and of
differing texture. In the intermediate layers, each 6 or 12 ins. thick
and occupying perhaps altogether 30 ft. of vertical thickness, we find
a graduated series of snail-shells leading almost imperceptibly from
the oldest lowest form to the latest uppermost form. Such cases are
known. But it is an exceptional thing to find these graduated series
either spread over an area of the earth's surface, or following one
another in successive strata. When they came into existence they
were rapidly superseded and destroyed as a rule, and have left only
one or two widely-separated examples of the intermediate forms.
This we should naturally expect by analogy from what we know of
the successive traces of human manufactures in the deposits on the
site of some of the great cities of the ancient world which have been
carefully excavated layer by layer. But still we have the important
fact that here and there such gradational series have been found,
and we are justified in considering a few isolated intermediate forms
(which often occur connecting two greatly-differing species) as
survivors of a former complete graduated series of intermediate
forms, which came into existence by slow modification of an
ancestral stock, and may, when the stock was widely spread over a
continental area, not merely have succeeded one another in time,
but actually coexisted in neighbouring regions.
There are many remarkable facts bearing upon the origin of
"species," the description of which fills volumes written by such men
as Darwin, Wallace, Poulton, and others, and become interesting to
every one who has gained a correct notion of what naturalists mean
by a "species." I will cite one in order to illustrate this. The bird
which we call the red grouse, or nowadays simply "grouse" (the old
Scotch name for it was "muir-fowl"), is one of twenty-four birds
(among the 400 species of birds which live in the British Islands),
including several kinds of titmouse, the goldfinch, bullfinch, song-
thrush, stonechat, jay, dipper, and others which are very closely
similar to species of birds living in Continental Europe, yet show
some definite and constant marks, such as small differences in the
colour of a group of feathers, enabling us to distinguish the British
from the Continental forms. Are these twenty-four British forms to
be regarded as distinct species?
The red grouse is placed in a genus called "Lagopus," of which
there are several species in the northern hemisphere. In Scotland
the red grouse, which is distinguished as Lagopus Scoticus, is
accompanied by a rarer species of Lagopus, which lives in high, bare
regions. This is the bird called by the Celtic name "ptarmigan"; it
differs in several points from the red grouse, and acquires white
plumage in the winter, which the latter bird does not; it is called
Lagopus mutus. Now in Norway we find also two species of grouse
or Lagopus, called "rypé" (pronounced "reeper") by the Norwegians.
One is the same bird in every respect as the Scotch ptarmigan, and
is known as "the mountain rypé." The other is very close to our red
grouse, and is called "the common or bush rypé," and by English
naturalists the "willow grouse," and by ornithologists "Lagopus
salicetus." It agrees in habits, voice, eggs, and anatomical detail
with our red grouse, but the back of the cock-bird of the red grouse
and the whole plumage of the hen-bird have a darker colour.
Moreover, the willow grouse, like the ptarmigan or mountain rypé,
turns white—acquires a white plumage—in the winter which the red
grouse does not. Are the red grouse and the willow grouse to be
regarded as distinct species? Our British red grouse lives on heather-
grown moors; the willow grouse prefers the shrubby growths of
berry-bearing plants interspersed with willows, whence its name.
Their food differs accordingly. Formerly the red grouse lived on the
moors of the South of England, and when in Pleistocene times
England was a part of the Continent of Europe the willow grouse and
the red grouse were one undivided species inhabiting all the north-
west of Europe. It is probable, though the experiment would be
almost impossible to carry out, that were the eggs of a number of
willow grouse now brought to Scotland and hatched on the moors,
they would tend to keep apart from the native red grouse, and not
inter-breed with them, in which case we should say that the Scotch
form is a "species on the make," or, even, a completed and distinct
species. On the other hand, it is possible that the two forms would
freely pair with another, and that the colour and winter coat of the
one (probably that of the Scotch form if the experiment were tried in
Scotland) would predominate, and after some generations no trace
of the other strain would be observable.
 CHAPTER X
SOME SPECIFIC CHARACTERS

A
N interesting case, showing that qualities which are life-
preserving under certain severe conditions exist in some
varieties of a species and not in others, was recorded some eight
years ago. After a very severe "blizzard" 136 common sparrows were
found benumbed on the ground by Professor Bumpus at Providence,
United States. They were brought into a warm room and laid on the
floor. After a short time seventy-two revived and sixty-four perished.
They were compared to see if the survivors were distinguished by
any measurable character from those which died. It was found that
the survivors were smaller birds (the sexes and young birds being
separately compared) than those which died, and were lighter in
weight by one-twenty-fifth than the latter. Also, the birds which
survived had a decidedly longer breastbone than those which died.
Similarly, the late Professor Weldon found that in the young of the
common shore-crab, taken in certain parts of Plymouth harbour,
those with a little peculiarity in the shape of the front of the shell
survived when those without this peculiarity died. Many thousands
were collected and measured in this experiment. It is not necessary
to suppose that the distinguishing mark of the survivors in such
cases is "the cause" of their survival. Such marks as the breadth of
the front part of the crab's shell and the length of a bird's
breastbone very probably are but "the outward and visible signs of
an inward and (physiological) grace."
The marks, little peculiarities of colour and proportionate size, or
some peculiar knob or horn, by which the student of species
distinguishes one constant form from another, can rarely, if ever, be
shown to have in themselves an active value in aiding or saving the
life of the species of plant or animal. The mark or "character" is an
accompaniment of a chemical, nutritional, physiological condition,
and is in itself of no account. It is what is called "a correlated
character." Such, for instance, is the black colour of the skin of pigs
which in Virginia, U.S., are found, as stated by Darwin, not to be
poisoned by a marsh plant ("the paint-root," Lachnanthes tinctoria),
whilst all other coloured and colourless pigs are. The pigs which are
not black develop a disease of their hoofs which rot and fall off,
causing their death when they eat this special plant "the paint-root."
The colour does not save the pig—it cannot correctly be called the
cause of the pig's survival—but is an accompaniment of the
physiological quality which enables the pig to resist the poisonous
herb. So, too, with white-spotted animals. They are known to
breeders as being liable to diseases from which others are free.
Fantail pigeons have extra vertebræ in their tails, and pouter pigeons
have their vertebræ increased in number and size. But the vertebræ
were never thought of and "selected" by the breeders. They only
wanted a fanlike set of tail feathers in the one case, and a longer
body in the other. Some varieties of feathering maintained by pigeon
breeders lead to the growth of abundant feathers on the legs (as in
Cochin-China fowls), and it is found that these feather-legged
pigeons always have the two outer toes connected by a web of skin.
If it were a stabilized wild form we should separate it as a species on
account of its webbed toes, yet the real selection and survival in the
hands of the breeder had nothing to do with the toes or their web,
but was simply "caused" by these pigeons having feathers of
"survival or selection value" in his judgment. Male white cats with
blue eyes are deaf. If deafness were ever an advantage (a difficult
thing to imagine), you would get a species of cat with white hair and
blue eyes, and be led to distinguish the species by those characters,
not by the real cause of survival, namely, deafness. Not enough is
yet known of this curious and very important subject of correlation,
but its bearing on the significance of "specific characters" is
sufficiently indicated by what I have said.
 An interesting group of species, three of which are to be
purchased alive through London fishmongers, are the European
crayfishes, not to be confused with the rock-lobster or Langouste
(Palinurus), sometimes called "crawfish" in London, nor with the
Dublin prawn (Nephrops). The little river crayfishes are like small
lobsters, and were placed by older naturalists in one genus with the
lobsters. Now we keep the European species of crayfishes as the
genus Astacus, and the common lobster and the American lobster
have been put (by H. Milne-Edwards) into a separate genus
(Homarus). You can buy in London the "écrevisses à pattes rouges"
of French and German rivers, which is called Astacus fluviatilis, and
differs from that of the Thames and other English and European
rivers (which you can also buy) called A. pallipes ("pattes blanches"
of the French), by the bright orange-red tips of its legs, and by
having the side teeth of the horn or beak at the front of the head
larger and more distinct. The English crayfish grows to be nearly as
large as the "pattes rouges" in the Avon at Salisbury, though it has
nearly disappeared about Oxford. You can also sometimes buy in
London the big, long-clawed Astacus leptodactylus of East Europe.
There are two or three other species, named and distinguished,
which do not come into the London market.
Crayfishes, lobsters and the like have groups of plume-like gills
(corresponding in the most ancient forms to the number of the legs
and jaw-legs) overhung and hidden by the sides of the great shield
or "head" of the animal. The common lobsters and crayfishes retain
most of these in full size and activity, but have lost in the course of
geologic ages the original complete number. These plume-like gills—
each half an inch or so in length—are attached, some to the bases of
the legs and some to the sides of the body above the legs. In the
ancestral form there were thirty-two plumes on each side, twenty-
four attached to the bases of the legs, and eight placed each at
some distance above the connection of one of the eight legs with
the side of the body. It is those on the side of the body which have
suffered most diminution in the course of the development of
modern crayfishes (and lobsters) from the ancestral form provided
with the full equipment of thirty-two gill-plumes on each side. In
fact, only one well-grown gill-plume, out of the eight which should
exist on each side of the body-wall, is to be found—and that is the
one placed above the insertion of the hindermost or eighth of the
eight legs (eight when we reckon the three jaw-legs as "legs" as well
as the five walking-legs). In front of this the side or wall of the body
is bare of gill-plumes though they are present in full size on the
basal part of most of the legs. Nevertheless, when one examines
carefully with a lens the bare side of the body overhung by the
head-shield or "carapace," one finds in a specimen of the common
English "pale-footed crayfish" a very minute gill-plume high above
the articulation of the seventh leg and another above the articulation
of the sixth leg. They are small dwindled things, as though on the
way to extinction, and are the mere vestiges of what were once
well-grown gill-plumes, and still are so in the rock lobster and some
prawns. In the red-footed crayfish of the Continent (Astacus
fluviatilis) yet another minute vestige of a gill-plume is found, farther
in front, on the body-wall above the fifth leg on each side of the
animal. This furnishes a definite mark or character by which we can
distinguish the red-footed crayfish from the common English pale-
footed one. But these three rudimentary gill-plumes in the red-foot
species, and two in the pale-foot species are all that until lately were
recorded. The region of the body-wall above the fourth, third,
second, and first of the legs was declared to be devoid even of a
vestige of the branchial plumes which were there in ancestral forms,
and have been retained more or less in some exceptional prawn-like
creatures allied to the crayfish.
Zoologists take a special interest in the crayfish because it is found
to be a most convenient type for the purpose of teaching the
principles of zoology to young students, and with that end in view
was made the subject of a very beautiful little book by the great
teacher Huxley. The conclusions above stated in regard to the gills
are set forth in that book with admirable illustrative drawings, and
the striking fact of the dwindling and suppression of the various gill-
plumes is clearly explained.
Fig. 33.—The rudimentary gill-plume of a crayfish from that part of the body-wall
to which the first pair of jaw-legs (maxillipedes) is articulated. Found in the
red-footed crayfish (Astacus fluviatilis) but in no other species of Astacus. It is
one-fifteenth of an inch long. Drawn by Miss Margery Moseley in 1904. ("Quart.
Journal of Microscopical Science," vol. 26 (1904-5).)

And now we come to an interesting discovery in this matter of the

gill-plumes of crayfishes. Some fifteen years ago the daughter of my
friend and colleague—Professor Moseley—was a member of the class
of Elementary Biology at Oxford. She had to examine and identify
these and other points in the structure of the crayfish. The class was
supplied with specimens of the French red-footed crayfish "Astacus
fluviatilis," as it is more readily obtained from fishmongers than our
own "pale-foot" or "Astacus pallipes." She found in her specimen far
forward on each side of the "head" a very minute gill far away from
the others and previously unknown. The demonstrator in charge of
the class refused even to look at her discovery. So she confirmed it
by examining three other specimens—made drawings of the tiny
branched gill (as shown in Fig. 33) and their position, and sent them
to me in London. It was at once clear that she had discovered in this
much studied little animal a very interesting pair of gills (right and
left)—unknown to Huxley and the rest of the zoological world. She
proceeded to examine specimens of A. fluviatilis from various rivers
of Germany and France and always found the new gill-plume. She
also showed (I supplied her with specimens at the Natural History
Museum) that it was, on the other hand, absent from A.
leptodactylus, A. pallipes, and all the foreign species (some from
Asia) which are known, and she published an illustrated account of it
in the "Quarterly Journal of Microscopical Science." This tiny gill-
plume is placed very far forward on each side of the body, the
farthest point forward at which any gill-plume is found in any kind of
prawn, shrimp or lobster, namely in the region where the first pair of
jaw-legs is attached, so that there are three empty spaces between
it and the rudimentary gill over the fifth pair of legs, already known
in the red-footed crayfish. It is only two millimetres long—about one-
fifteenth of an inch! But its presence serves very distinctly to
separate the red-footed crayfish, Astacus fluviatilis of French and
German rivers, thus discovered to have four pairs of rudimentary
gill-plumes, from the Astacus leptodactylus of the Danube basin and
East Europe, which has only three pairs, and still more to emphasize
the difference between it and our British species, the "white-foot" or
Astacus pallipes, which has only two!
This little history is noteworthy, firstly, because it shows that a
young student may, to use an appropriate term, "wipe the eye" of an
expert observer and rightly venerated teacher (who would have
delighted in the little discovery had he been alive), as well as the
eyes of tens of thousands of students and teachers (including
myself) who have studied the red-foot crayfish year after year, and
missed the little gill. It is also interesting as showing us a good
sample of a specific mark or character which has no survival value;
that is, could not advantage the crayfish in the struggle for life. The
fact is, that this one particular very minute forward pair of gill-
plumes is like the other rudimentary gills—a survival in a reduced
condition of a pair of gill-plumes which were well-grown, useful
plumes aerating the blood, in the prawn-like ancestors of all
crayfishes, lobsters, shrimps, and prawns, and is, owing to
circumstances of nutrition and growth which we know nothing about
but can vaguely imagine, retained by the red-foot species of
crayfish, but lost by all other crayfishes, lobsters, common prawns
and shrimps, and, in fact, only retained besides by a very few out-of-
the-way kinds of marine prawns. That is the sort of thing which
frequently has to serve as "a specific character" or mark,
distinguishing one "species" from another.
A more ample discussion of the origin of species is not within the
scope of this book. But I may say that until recently the conception
that every organ, part and feature of a plant and animal must be
explained, and can only be explained, as being of life-saving value to
its possessor, and accordingly "selected" and preserved in the
struggle for existence, was held by many "Darwinians" in too
uncompromising a spirit. This conception was, really from the first,
qualified by the admission that the life-saving value and consequent
preservation of a structure must undoubtedly in some cases have
been in operation in ancestors of the existing species, and is no
longer operative in their descendants although they inherit the
structure which has now become useless. Moreover, the operation of
those subtle laws of nutrition and of form which are spoken of as the
"correlation of parts in growth and in variation" (mentioned on p.
119) was pointed out by Darwin himself as probably accounting for
many remarkable growths, structures and colour-marks which we
cannot imagine to be now, or ever to have been in past ancestry, of
a life-saving value. Nevertheless, the old "teleology," according to
which, in pre-Darwinian days, it was held that every part and feature
of an animal or plant has been specially created to fulfil a definite
pre-ordained function or useful purpose, still influenced the minds of
many naturalists. Natural selection and survival of the fittest were
reconciled with the old teleological scheme, and it was held that we
must as good Darwinians account for every structure and distinctive
feature in every animal and plant as due to its life-saving value.
Herbert Spencer's term, "the survival of the fittest," conduced to the
diffusion of this extreme view: Darwin's equivalent term, "the
preservation of favoured races," did not raise the question of greater
or less fitness.
The extreme view is now, however, giving place to the recognition
of the fact that the actual tendencies to variation—accumulated in
the living substance of the various stocks or lines of descent and
handed on during an immense succession of ages of change by
hereditary transmission—counts for more in the production of new
species and strange, divergent, even grotesque forms of both
animals and plants than had been supposed.
Undoubtedly selection or survival of the fittest mainly accounts for
the colouring and adaptive shaping of living things, and so for those
several great types of modelling which arrest the eye and have
excited the interest of inquisitive man. But there seems to be no
justification for the assumption that in all cases a variation—that is
to say, an increase or a diminution of the volume of some existing
structure in proportion to other coexisting structures in the body of a
living plant or animal—must be either favourable, that is, conducive
to survival, or injurious, that is, tending to the defeat and
destruction of its possessors or their race. On the contrary, it is the
fact that there are vast areas and conditions related to countless
myriads of living creatures in which variations of those creatures of
large and imposing kind and degree are neither advantageous nor
disadvantageous, but matters of absolute indifference, that is to say,
without any effect upon the preservation or survival of their race or
stock. Nature is far more tolerant than some of us were inclined to
assume. In certain restricted conditions of competition and in regard
to some special structures and components which are often so
minute and obscure as to be not yet detected by that recent arrival,
the investigating biologist—though sometimes, fortunately for him,
large enough to jump to his eyes—it is undeniable that there must
be a "survival" or "favouring" of individuals presenting a variation in
increase, or it may be decreased, of this or that special feature of its
"make-up" or structural components. But it is a more correct
statement of the case to say that natural selection or survival
preserves not the fittest, but the least fit possible under the
circumstances—namely, all those which, however great their
divagations and eccentricities of variation in other respects, yet at
the same time attain to a minimum standard of qualification in those
structures (or inner chemical qualities) essential for success in the
competition for safety, food and mating determined by the particular
conditions in which the competition is taking place. Consequently
forms which are meaningless so far as standards of utility or "life-
saving" are concerned, and are rightly described as grotesque,
monstrous, gigantic or dwarfed—excessive (as compared with more
familiar kinds) in hypertrophy or atrophy of their colouring and
clothing, or of out-growths such as leaves of plants and limbs, jaws
or other regions of the body of animals—are found existing in
various degrees of eccentricity in every class of both plants and
animals. Among animals such tolerated "exuberances of non-
significant growth" are more striking than in plants. The group of
fishes seems to be especially privileged in this way. They are freely
variable in the position of the fins, the suppression or exaggeration
of them, as well as of the scales on the surface of the body (e.g.
leather carp and mirror carp). Take, for example, the mackerel and
the salmon as standards of utilitarian adaptation of the body to an
active life in sea or river, and then compare with theirs the
astounding proportions of the sun-fish (Orthagoriscus) like a cherub
"all head and no body," or the almost incredible Pteraclis—with its
little body framed immovably between a huge dorsal and a huge
ventral fin (see figures on p. 130). The fin-like crest of enormous
size on the back of the great extinct lizard Dimetrodon of the
Permian age supported by long bony spines is a similarly excessive
and useless outgrowth. (This astonishing creature is shown in our
Frontispiece.) Such exuberant products may be ascribed to an
unrestrained "momentum" of growth which once set going by
fortuitous variation has been tolerated but not favoured by natural
selection. Or (as supposed by some) their excessive development
may be due to the persistence of some nutritional condition which at
first resulted in a moderate growth of the fin-like crests in question
as a serviceable structure, but has persisted and increased long after
the fin or crest has attained a sufficient size—simply because its
increase though of no life-saving value—yet was not harmful and so
did not bring its owner under the guillotine of natural selection. Such
disproportionate exuberance of growth due to innate variability,
tolerated but not specially favoured by natural selection, will account
for many strange and grotesque forms of living things. From time to
time in the long process of change, such exuberances may suddenly
become of service and be, so to speak, taken in hand by natural
selection, or they may become dangerous and lead to the
extermination of the stock in which they have been previously
tolerated.
Before my reader turns—as I hope he or she will do—to some
handbook of zoology in which the genealogical tree or classification
of the species of animals and of plants is treated at length, I will
endeavour to give some estimate of the immense numbers of
"species" which exist. As to mere individuals, it is impossible to form
any estimate, but when we reckon up the teaming population of a
meadow or forest in England, the hundreds of thousands of plants,
including the smallest mosses and grasses, as well as the larger
flowers, shrubs, and trees, the still greater number of insects,
spiders, snails, and larger animals and birds, feeding on and hiding
among them, and when we remember that in the ever-warm tropical
regions of the earth life is ten or twenty times more exuberant than
with us,—then the immensity of the living population of the land and
water of the globe becomes as difficult to realize as are the figures
in which the astronomer tells of the number and distances of the
stars. On the other hand, some idea of the number of distinct
species of animals and plants which have up to this date been
recognized and described by naturalists as at present existing, may
be formed by a statement of those which have been described in
some of the more familiar groups. About 10,000 species of mammals
have been described; about 14,000 of birds; 7000 of reptiles; 15,000
of fishes; 500,000 of six-legged insects; 14,000 of crustacea
(shrimps, lobsters, crabs); 62,000 of molluscs (snails, mussels, etc.);
15,000 of star-fishes and sea-urchins; 5000 of corals and polyps;
3000 of sponges; and 6000 of microscopic protozoa. In all about
800,000 species of animals have been recorded, and probably as
many more remain yet to be recognized and described.
The total number of described species of plants has never been
estimated, but some idea of it may be formed from the fact that
1860 species of flowering plants alone have been distinguished in
Britain, 17,000 in British India, and 22,000 in Brazil, not to mention
those of Africa and Australia! These figures do not include the vast
numbers of flowerless plants, the ferns, mosses, sea-weeds,
mushrooms, moulds, lichens, and microscopic plants.
And then we have to add to these enumerations of living species
the extinct species of successive geological ages, the remains of
which are sufficiently well preserved to admit of identification. Those
which are known are only a few thousands in number, and a mere
fragment of the vast series of species which have existed in
successive past ages of the earth. They are a few samples of the
predecessors of the existing species, and some of them were the
actual ancestors of those existing to-day. The larger number of them
have left no direct issue, but represent side branches of the "tree of
life" which have died out ages ago.
Strangely-shaped Fishes.—1. The Coffer-fish (Ostracion); 2. Pteraclis, an oceanic fish
allied to the so-called Dolphins; 3. The Sun-fish (Orthagoriscus); 4. An
Australian Blenny Patæcus.
 CHAPTER XI
HYBRIDS

T
HE subject treated in this and the next chapter is one of the
most interesting to mankind, and is surrounded by extraordinary
prejudice, sentiment, and ignorance. It is one upon which really
trustworthy information is to a very large extent absent—and difficult
to obtain. I cannot profess to supply this deficiency, but I can put
the matter before the reader.
It is a well-established fact that the various "kinds" of animals and
of plants do not breed promiscuously with one another. The
individuals of a "species" only breed with other individuals of that
"species." They do not even, as a habit, breed with the individuals of
an allied species. So nearly universal is this rule that it was for a long
time held by naturalists to be an absolute definition of "a species,"
that it is a group of individuals capable of producing fertile young by
breeding with one another and incapable of producing fertile young
by mating with individuals of another such group, which were,
therefore, held to constitute a distinct species. The practical
importance of this definition was that it could, in a large number of
instances among animals, and still more amongst plants, be made
use of as a test and decided by experiment.
It is a curious fact that popular belief amongst country-folk and
those who have opportunities of coming to a conclusion on so simple
and direct a question has never accepted this law of the limitation of
species in breeding as more than a general rule to which it has
always been supposed that frequent exceptions occur. I mention this
not in order to add that "there is always some basis of truth in these
popular beliefs," but on the contrary to point out that popular beliefs
on such matters are very frequently altogether erroneous, and
though their origin can sometimes be explained, it is rare to find that
they are due, in however small a degree, to true observation and
inference. Where the subject under consideration has the obscurity
and strong fascination for the natural man which all that relates to
the processes of life, growth, and reproduction possess, we find that
traditional fancies of the most unwarrantable kind are current, and
hold their ground with tenacity even at the present day. Some 250
years ago, and earlier—in fact, before the commencement of that
definite epoch of "the New Philosophy" marked by the foundation of
the Royal Society of London—any queer-looking animal brought from
remote lands, and any misshapen monstrosity born of cattle, sheep,
dogs, or men, was "explained," and confidently regarded as a
"hybrid," the result of a "cross" or irregular coupling of two distinct
species of animals to which the "monster" presented some fanciful
resemblance. Whole books were devoted to the description and
picturing of such supposed examples of mis-begotten progeny.
The belief in the existence of such extraordinary hybrids is still
common among so-called "well-educated" people. I have with
difficulty avoided causing annoyance and offence to a friend, a
celebrated painter, by refusing to admit that a deformed cat, of
which he gave me an account, was a hybrid between a cat and a
rabbit. A very eminent person whom I was conducting some years
ago round the galleries of the Natural History Museum, declared, as
we stood in front of the specimen of the Okapi of the Congo Forest,
that it was clearly a hybrid between the giraffe and the zebra. He
insisted that it was obvious that such was its explanation, and
pointed to its striped haunches and legs, and its cloven hoofs and
giraffe-like head. I failed to change his opinion.
It is the fact—ascertained by careful observation of natural
occurrences and by experiment—that, in spite of the almost absolute
law or general truth to the effect that the members of a species
(whether of plant or animal) only produce fertile offspring by mating
with members of that same species, yet there are rare instances
known in which individuals of two distinct but allied species have
mated and produced fertile offspring. The cases in which such
unions have resulted in the production of offspring, but in which the
offspring so produced prove to be infertile—that is, incapable of
producing offspring in their turn—are much more numerous. An
important distinction has also to be made between cases of either
fertile or infertile hybrid-production which occur spontaneously in
nature, and those in which man by separating the parent animals or
plants from their natural conditions of life, or by bringing about
impregnation (as in "pollinating" one flower with the pollen-dust of
another) succeeds in obtaining a "cross" or "hybrid," whether fertile
or infertile, not known to occur in "wild" (that is, not humanly
controlled) nature. The rarest case would be that of the production
of fertile hybrids in uncontrolled natural conditions. Such possibly
occur in the case of some fishes in which the fertilization of the eggs
takes place in water, the fertilizing microscopic sperms passing from
the males like dust into the water and thus reaching the eggs laid by
the females. Occasionally hybrids are thus produced between some
common fresh-water fishes—species of the same genus—and
between species of flat-fish, such as the turbot and the brill, though
it is difficult to be sure that the rare hybrids so produced are fertile
even if they attain to maturity. The same is true as to certain small
flowering plants having distinct regions of natural distribution and
occurrence. At the confines of the regions proper to two such allied
species, insects passing from one to the other do sometimes effect a
reciprocal fertilization of the two species, and a natural hybrid is the
result. Here, again, it is difficult to follow the subsequent history of
the hybrids, but it is believed that in some instances they are fertile,
and that the hybrid race is only gradually merged by subsequent
crossing into one or other of the parent species. Not a single
instance is on record of the production of a "natural" hybrid (that is
to say, one produced in natural conditions without man's
interference), whether fertile or infertile, between two species of the
larger animals (such as between horse and ass or zebra and ass, or
between lion and tiger or any of the species of cats, or between
species of bears) or birds (such as pheasants of various species,
including the jungle cock, the wild original of our domestic fowl, or
between various species of ducks, various species of geese, or
between various species of the grouse-birds).
Nevertheless, in conditions brought about by man—that is to say,
confinement in cages or paddocks, or at any rate removal from their
native climate and home—all the groups of species just cited
commonly and frequently produce hybrids inter se, that is, one or
more species of the horse group thus inter-breed with one another,
so will certain species of cats, certain species of bears, many species
of pheasants, also of ducks, of geese, and of grouse. In nearly every
case the hybrids so produced are infertile; they will not mate with a
similar hybrid, and even when mated with one of the parent species
rarely produce offspring, though they sometimes do so. The best
cases of the production of fertile hybrids are between species of
flowering plants brought to this country from widely separated
regions. The surprising and instructive result has been obtained that
a cross between two allied species (that is, of one and the same
"genus") which will fail altogether or "come to nothing" as infertile
hybrids—if the two species crossed are from the same or contiguous
regions—yet will yield readily vigorous fertile hybrid offspring when
the two species (always, of course, of one and the same genus)
have their native homes in widely separate parts of the world—as,
for instance, the Indian Himalaya range and the South American
Andean range.
This has been found in crossing species of rhododendrons, of
orchids, and of many other plants with which horticulturists occupy
themselves for commercial purposes. It is in some ways the reverse
of what one might expect. It would be reasonable to suppose that
allied species from the same climate and geographical region would
have more affinity and be more readily hybridized than species from
widely remote and physically differing regions. But the reverse is the
case, many thriving hybrid stocks which duly fertilize and set their
seed are now in cultivation, having been produced by the union of
parent species from "the opposite ends of the earth."
The consideration of this case throws some light on the
significance of the non-occurrence of natural hybrids and of the very
remarkable and curious fact that hybrids are so usually sterile. When
we come to think of it, the natural preliminary assumption should be
(as is that of unsophisticated humanity) that any animal or plant
might, so far as possibilities go, breed with any other; and the
questions to be answered are: (1) What advantage to a species is it
not to be able to hybridize with other species, and (2) how—that is
to say, by what structure or by what subtle chemical differences or
other features in their make-up and habit—are they prevented from
so hybridizing? Then we come on further to the question, Why
should a hybrid, once produced, fail to bear healthy eggs or sperms
according to its sex, although it grows up to full size and is to all
appearances mature? And why should hybrids between parents of
origin locally remote from one another not show this failure, but
behave like ordinary healthy organisms?
In the full solution of these inquiries we should get very near to
some of the most important secrets of the living body which have
still to be searched out. But a reply to these questions which is
probably in large measure true, and serves to help us in the further
collection and examination of facts, is as follows: First, the
production and maintenance of "species" of plants and of animals by
survival of favourable variations in the struggle for existence (Darwin
and Wallace's theory of the origin of species) requires the
maintenance of the purity of the favourable stock which survives in
the struggle. If it were continually liable to hybridization by other
species it would never establish its own distinctive features. It would
deteriorate by departing from those characteristics which have been
"naturally selected" and have rendered it a successful "species."
Thus the breeder, when he has selected a stock for propagation
which approaches the standard at which he is aiming, keeps it apart,
and does not allow it to be "crossed" by other stock. One of the
qualities "naturally selected" in "the wild" is the power of resistance
to fertilization by neighbouring species.
This power of resistance or immunity to fertilization by other
species may be attained by several different methods. Amongst
these are (1) a difference in the season of breeding or sexual
ripening; (2) the production of secretions (whether by plant or by
animal) which poison or paralyse the fertilizing sperms of allied and
locally associated species, but are harmless to those of the secreting
species; (3) the mechanical differences of size, etc., which prevent
the fertilizing material of a strange species from gaining access to
the egg-cells; (4) psychical activities (antipathies) in the case of
animals or mere attraction and repulsion by odoriferous substances,
which serve to repel a strange species, but are attractive to
individuals of the same species; (5) finally, a chemical and
physiological incompatibility between the sperms of one species and
the germs of another (as distinct from the attraction or repulsion of
the entire living individual), which, even when all other difficulties
are absent or have been overcome, may be, and frequently is,
present, so that the spermatozoon cannot penetrate the egg-cell
even when resting upon it, but may be paralysed or repelled, and in
any case is not guided and drawn into the aperture of the egg-
covering, called the micropyle, or "little entry," so as to fuse with and
fertilize the egg.
The operation of these hindrances to hybrid fertilization and
breeding have been ascertained in several different instances. It is
not always possible, and certainly not easy to ascertain, which is at
work in any and every case. But we can well conceive that one or
other of these agencies have been developed and accentuated by
survival of the fittest, so as to protect a species against fertilization
by a neighbouring species, and thus to enable it to maintain its own
"bundle of characteristics" free from the swamping effects of
"mixture" (that is, "hybridization") with another species. It is also
thus intelligible that an allied species from a distant land against
which our native species and its closer ancestry—struggling for
purity of race—have had no occasion or opportunity to develop a
repelling protection—will have no such difficulty in effecting the
fertilization of the native species as have those adjacent species
against whose intrusions the latter is specifically moulded and
selected by long generations of severe natural selection.
The failure of hybrids generally to ripen their ova and sperm so as
to reproduce themselves is a subject upon which, considering its
enormous importance and significance, singularly little has been
done in the way of investigation. Fifty years ago it was usually
taught that the mule, between the horse and the ass, so largely
produced under human superintendence for transport service, was
unable to breed owing to some deformity in the reproductive
passages. Even now no adequate study of the subject has been
made, but it appears that whilst a female mule can be, and
sometimes is, successfully mated to a horse or an ass, giving birth to
a foal, the male mule does not produce fully-formed spermatozoa.
What precisely is the nature of this failure, what the ultimate
microscopic condition of the sperm cells in infertile male mules, or in
any other infertile male hybrids, has not yet been properly worked
out by modern cytological methods. It would be a matter of vast
interest to determine what is the difference in the structure of the
sperm-cells of a fertile and of an infertile male hybrid. At present, so
far as I know, this has not been done.
So far what I have written applies to hybridization—the inter-
breeding of distinct species. A similar but by no means identical
subject is that of the inter-breeding of distinct races or varieties of
one species, and the production of "mongrels." "Mongrels" are to
races what "hybrids" are to species. To this branch of the subject
belongs the study of the effects of intermarriage between distinct
races of men.
 CHAPTER XII
THE CROSS-BREEDING OF RACES

W
E have seen that there is no simple rule as to the "mating" of
individuals of a species with individuals of another closely allied
but distinct species. Such mating very rarely comes about in natural
conditions, but man by his interference sometimes succeeds in
procuring "hybrids" between allied species. Hybrids between species
belonging to groups so different as to be distinguished by zoologists
as distinct "families" or "orders" are quite unknown under any
circumstances. Such remoteness of natural character and structure
as is indicated by the two great divisions of hoofed mammals—the
even-toed (including sheep, cattle, deer, antelopes, giraffes, pigs and
camels), and the odd-toed (including tapirs, rhinoceroses, horses,
asses and zebras) is an absolute bar to inter-breeding. So, too, the
carnivora (cats, dogs, bears and seals, and smaller kinds) are so
remote in their nature from the rabbits, hares and rats—called "the
rodents"—that no mating between members of the one and the
other of these groups has ever been observed, either in nature or
under artificial conditions.
Even when individuals of closely allied species mate with one
another it is a very rare occurrence that the hybrids so produced
ripen their ova and sperms so as to be capable of carrying on the
hybrid race, though sometimes they do ripen them and breed. The
great naturalist Alfred Wallace, in his most valuable and readable
book called "Darwinism," expressed the opinion that the apparent
failure of hybrid races to perpetuate themselves by breeding was to
a large extent due to the small number of individuals used in
experiments on this matter, and the in-and-in breeding which was
the consequence. One of the great generalizations established by
Darwin is that in-and-in breeding is, as a rule, resisted in all animals
and plants, and leads when it occurs to a dying-out of the inbred
race by resulting feebleness and infertility. A large part of Darwin's
work consisted in demonstrating the devices existing in the natural
structure and qualities of plants and animals for securing cross-
fertilization among individuals of the same species but of different
stock. Both extremes seem to be barred in nature—namely, the
inter-breeding of stocks so diverse in structure and quality as to be
what we call "distinct species," and again the inter-breeding of
individuals of the same immediate parentage or near cousinship.
What seems to be favoured by the natural structure and qualities of
the plant or the animal is that it shall only breed within a certain
group—the species—and shall within that group avoid constant self-
fertilization or fertilization by near cousins. Thus we find numerous
cases in which, though the same flower has both pollen and ovules,
and might fertilize itself, the visits of insects (specially made use of
by mechanisms in the flower) carry the pollen of one flower to the
ovules of another and to flowers on separate plants growing at a
distance. It is necessary to note that there are, nevertheless, self-
fertilizing flowers, and also self-fertilizing lower animals, the special
conditions of which require and have received careful examination
and consideration, upon which I cannot now enter.
In relation to the question of the possibility of establishing hybrids
between various species experimentally, I must (before going on to
the cognate question of "mongrels") tell of an interesting suggestion
made to me by my friend Professor Alphonse Milne-Edwards not
long before he died, and never published by him. He was director of
the Jardin des Plantes in Paris, where there is a menagerie of living
beasts as well as a botanic garden and great museum collections
and laboratories. He held it to be probable, as many physiologists
would agree, that the fertilization of the egg of one species by the
sperm of another, even a remotely related one, is ultimately
prevented by a chemical incompatibility—chemical in the sense that
the highly complex molecular constitution of such bodies as the anti-
toxins and serums with which physiologists are beginning to deal is
"chemical"—and that all the other and secondary obstacles to
fertilization can be overcome or evaded in the course of experiment.
He proposed to inject one species by "serums" extracted from the
other, in such a way as seemed most likely to bring the chemical
state of their reproductive elements into harmony, that is to say, into
a condition in which they should not be actively antagonistic but
admit of fusion and union. He proposed, by the exchange of living or
highly organized fluids (by means of injection or transfusion)
between a male and female of separate species, to assimilate the
chemical constitution of one to that of the other, and thus possibly
so to affect their reproductive elements that the one could tolerate
and fertilize the other. The suggestion is not unreasonable, but
would require a long series of experiments in which the possibility of
producing such "assimilation," even to a small extent and in respect
of less complex processes than those ultimately aimed at, would
have to be, first of all, established. My friend did not live to
commence this investigation, but it is possible that some day we
may see the obstacle to the union of ovum and sperm of species,
which are to some extent allied, removed in this way by transfusion
or injection of important fluids from the one into the other.
We must not lose sight of the fact, in the midst of these various
and diverging observations about the fertilization of the ova of one
species by sperms of another species, that there is such a thing as
"parthenogenesis," or virgin-birth. In some of the insects and lower
forms of animals the egg-cell habitually and regularly develops and
gives rise to a new individual without being fertilized at all. And in
other cases by special treatment, such as rubbing with a brush, or in
the case of marine animals by addition of certain salts to the water
in which the eggs are floating—or, again, in the case of the eggs of
the common frog by gently scratching them with a needle—the eggs
which usually and regularly require to be penetrated by and fused
with a spermatozoon or sperm-filament before they will develop,
proceed to develop into complete new individuals without the action
upon them of any spermatozoon. In such marine animals as the sea-
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