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THE MONIST
A
QUARTERLY MAGAZINE
VOL. I.
CHICAGO:
THE OPEN COURT PUBLISHING CO
1890-1891
COPYRIGHT BY
THE OPEN COURT PUBLISHING CO.
1890-1891
CONTENTS OF VOLUME I.
ARTICLES.
PAGE.
Anarchists, The Physiognomy of the. By Cesare Lombroso 336
Anarchist's Reply to Professor Lombroso, A Convicted. By Michael
Schwab 520
Architecture of Theories, The. By Charles S. Peirce 161
Criminal Anthropology, Illustrative Studies in. (1) "La Bête
Humaine" and Criminal Anthropology. (2) Criminal Anthropology
and Psychiatry. By Cesare Lombroso 177
Ethics, The Criterion of——an Objective Reality. Editor 552
Evolution, The Factors of. By Joseph Le Conte 321
Evolution, The Right of. By Moncure D. Conway 506
Feelings and the Elements of Feelings. Editor 401
Five Souls with but a Single Thought. By Carus Sterne 245
Höffding on the Relation of the Mind to the Body. By W. M. Salter 118
Immortality. By George M. Gould 372
Infusoria, The Immortality of. By Alfred Binet 21
Innovation and Inertia in the World of Psychology. By Cesare
Lombroso 344
Magic Mirror, The. By Max Dessoir 87
Mind, The Origin of. By Paul Carus 69
Mind, The Question of Duality of. By R. Meade Bache 362
Philosophy in American Colleges and Universities 148-156
Psycho-Physics, Some Questions of. A Discussion. (1) Sensations
and the Elements of Reality. By Ernst Mach 393
(2) Feelings and the Elements of Feelings. Editor 401
Psychology of Conception. By J. Sully 481
Sensations, The Analysis of the. By Ernst Mach 48
Sensations and the Elements of Reality. By Ernst Mach 393
Sex, On the Material Relations of—in Human Society. By E. D. Cope 38
Squaring of the Circle, The. By Hermann Schubert 197
Thought and Language, On. F. Max Müller 572
Truth, The Criterion of. Editor 229
Wallace on Physiological Selection, A. R. George J. Romanes 1
Welfare, The Principle of. By Harald Höffding 525
LITERARY CORRESPONDENCE.
France. By Lucien Arréat 124, 278, 421, 590
German Philosophy in the XIX Century. By F. Jodl 263
The Modern Literature of Italy since 1870. By C. Lombroso 428
The Science of Pedagogy in Germany. Chr. Ufer 597
BOOK REVIEWS.
Abbott, Francis Ellingwood. _The Way Out of Agnosticism_ 129
Bois-Reymond, Paul Du. _Ueber die Grundlagen der Erkenntnis in
den exacten Wissenschaften_ 608
Booth, General. _In Darkest England and the Way Out_ 451
Bray, Charles. _The Philosophy of Necessity_ 136
Brinton, Daniel G. _Races and Peoples_ 131
Büchner, Ludwig. _Fremdes und Eigenes aus dem geistigen Leben
der Gegenwart_ 303
Carneri, B. _Der Moderne Mensch_ 607
Carus, Paul. _The Soul of Man_ 620
Clarke, James Freeman. _Deacon Herbert's Bible Class_ 305
Coit, Stanton. _Die ethische Bewegung in der Religion_ 301
Cox, Charles F. _Protoplasm and Life_ 297
Dewey, John. _Outlines of a Critical Theory of Ethics_ 600
Dillmann, C. _Die Mathematik die Fackelträgerin einer neuen Zeit_ 617
Edinger, Ludwig. _Twelve Lectures on the Structure of the Central
Nervous System_ 604
Everett, Charles Carroll. _The Science of Thought_ 287
Forel, August. _Der Hypnotismus_ 605
Fullerton, George Stuart. _On Sameness and Identity_ 291
Geddes, Patrick, and J. Arthur Thomson. _The Evolution of Sex_ 439
Haeckel, Ernst. _Plankton-Studien_ 455
Harris, William T. _Introduction to the Study of Philosophy_ 438
Höffding, Harald. _Ethik_ 139
James, William. _The Principles of Psychology_ 284
Jastrow, Joseph. _The Time-Relations of Mental Phenomena_ 290
Jodl, Friedrich. _Geschichte der Ethik in der neueren Philosophie_ 137
Krause, Ernst. _Tuisko-Land der arischen Stämme und
Götter-Urheimat_ 612
Kroman, K. _Kurzgefasste Logik und Psychologie_ 142
Lehmann, Alfred. _Die Hypnose und die damit verwandten normalen
Zustände_ 298
Loeb, Jacques. _Der Heliotropismus der Thiere_ 300
Loeb, Jacques. _Untersuchungen zur physiologischen Morphologie
der Thiere_ 300
Lombroso, Cesare. _Der geniale Mensch_ 146
Mackenzie, John S. _An Introduction to Social Philosophy_ 601
Mantegazza, Paolo. _Physiognomy and Expression_ 447
Moll, Albert. _Hypnotism_ 604
Morgan, C. Lloyd. _Animal Life and Intelligence_ 443
Naden, Constance C. W. _Induction and Deduction_ 292
Natorp, Paul. _Einleitung in die Psychologie nach kritischer
Methode_ 143
Peet, Stephen D. _Emblematic Mounds and Animal Effigies_ 295
Post, Albert Hermann. _Ueber die Aufgaben einer allgemeinen
Rechtswissenschaft_ 457
Royer, Madame Clémence. _Nouveaux aperçus sur la Phylogenie
de l'Homme_ 297
Savage, M. J. _Life_ 296
Schopenhauer, Arthur. _Le Monde comme Volonté et comme
Representation_ 298
Sterrett, J. Macbride. _Studies in Hegel's Philosophy of Religion_ 133
Sterne, Carus. _Die allgemeine Weltanschauung_ 456
Taylor, Isaac. _The Origin of the Aryans_ 435
Ufer, Christian. _Geistesstörungen in der Schule_ 619
Wolff, Joh. _Das Bewusstsein und sein Object_ 147
PERIODICALS 157-160; 307-320; 459-480; 621-640
VOL. I. OCTOBER, 1890. NO. 1.
THE MONIST.
MR. A. R. WALLACE ON PHYSIOLOGICAL SELECTION.[1]
[1] In a private letter to the editor of this magazine
Professor Geo. J. Romanes writes: "The article refers to a
completely new departure in the theory of evolution, striking
in the principle of homogamy, the root-principle of the whole,
and in physiological selection, one of the main branches.
Yet neither principle has so far been perceived except by
Mr. Gulick.... The theory of physiological selection has
been better understood in America than in this country; and
I should like the naturalists there, who have taken such a
warm and appreciative interest in it, to see my reply to Mr.
Wallace published in an American periodical."
In 1886 I published a paper entitled "Physiological Selection: an
additional suggestion on the origin of species," (Zoölogical Journal of
the Linnean Society, Vol. XIX, p. 337). The view there expressed is,
briefly, as follows.
Given the facts of heredity and variability, the whole theory of
organic evolution becomes neither more nor less than a theory of
the causes which determine the breeding of like with like, to the
exclusion of unlike. For the more firmly that we believe in heredity
with variability as the fundamental principle of organic evolution,
the stronger must become our persuasion that segregate breeding (or
exclusive mating of like with like) must lead to divergence, while
indiscriminate breeding (or free intercrossing of all varieties) must
lead to uniformity. So long as there is free intercrossing, heredity
makes in favor of fixity of type—or, at most, can permit change only
in a single line, where successive generations undergo a continuous
improvement, which may give rise to a ladder-like series of species in
time. But in order that there should be a tree-like multiplication of
species in space, or a simultaneous divergence of type, it is essential
that free intercrossing be prevented at the origin, and throughout the
development, of each branch. In other words, it is only when assisted
by some form of segregation—which determines exclusive breeding of
like with like—that heredity can effect arborescent or polytypic, as
distinguished from catenated or monotypic, evolution. For the sake of
greater clearness, I will call segregation in this sense _homogamy_, or
the exclusive mating of individuals which belong to the same variety.
Now homogamy may be secured in a very great number of different
ways. Of these the most important, from every point of view, is
natural selection. Here the exclusive breeding of like with like is
determined by general fitness, and is effected by extermination of the
unlike—i. e., the comparatively unfit. Moreover, this process leads
to a continuous improvement in the way of adaptation, and in this
important respect it stands alone among all the forms of homogamy.
Nevertheless, we must note that, unless assisted by some other form of
homogamy, natural selection can only produce monotypic evolution; never
polytypic. Successive generations may thus continuously mount to higher
stages of adaptation on the steps supplied by their own dead selves;
but although they may thus give rise to a linear series of species in
time, they can never thus give rise to a multiplication of species
in space. In order to effect such multiplication, or _divergence_ of
types, natural selection must be supplemented by some other form of
homogamy, which can prevent intercrossing between the equally fit at
the origin, and throughout the development, of every separate branch.
Well, as I have said, these other forms of homogamy are very numerous.
First we may notice geographical isolation. When a comparatively small
portion of a species is thus separated from the rest of its kind,
intercrossing is effectually prevented between the two sections; and
inasmuch as the general average of specific characters in the isolated
section will be somewhat different from that of the other section,
heredity will determine that the two sections shall not run parallel
in their subsequent lines of evolutionary history: there will arise
an increasing divergence between them, as was first pointed out by the
mathematician Delbœuf, subsequently by the naturalist Weismann, and
more recently, with greater emphasis, by Mr. Gulick as well as myself.
Again, there is homogamy that arises as a result of sexual preference,
or, as I have called it, "psychological selection." It is a matter of
observation that the breeding of like with like is often determined
among the higher vertebrata by individuals of each variety preferring
to mate with other individuals of their own variety; and this is
homogamy.
Not to occupy space with any attempt at enumerating all the many forms
of homogamy[2] I will at once pass on to the form which constitutes the
subject-matter of the present paper—and the form which, in my opinion,
is probably of more importance than any other in the multiplication of
species. This is the form of homogamy which I have termed Physiological
Selection, or Segregation of the Fit, and Mr. Gulick—who independently
perceived the principle—has called Segregate Fecundity.
[2] This has been done in a most careful and exhaustive manner
by Mr. Gulick in his papers which have succeeded mine in the
publications of the Linnean Society.
As my object on the present occasion is to answer criticisms which
have been passed on my enunciation of this principle, I do not propose
to go into further detail by way of explanation than is necessary in
order to render intelligible both the criticisms and my reply thereto.
Moreover, this reply is only an abstract of a fuller one which has been
prepared for publication in a forthcoming book. Therefore it deals only
with the main points. Lastly, I may remark that the criticisms which
have hitherto appeared have all been derived from the same source,
viz., from Mr. A. R. Wallace; for, although many other naturalists have
expressed themselves as more or less opposed to the new theory, or
"additional suggestion on the origin of species," they have all done so
on the grounds, or for the reasons supplied by Mr. Wallace. Therefore,
in dealing with Mr. Wallace's objections, I shall be dealing with the
only objections which have thus far been advanced.
In order at once to restate the theory of physiological selection,
and to do so in a form which cannot be suspected of being in any way
influenced by Mr. Wallace's more recent criticisms, I will begin by
reproducing the main features of the theory in the words which were
employed for this purpose more than three years ago, when I supplied
an article to the _Nineteenth Century_ in answer to one by him in the
_Fortnightly Review_. Moreover, for the most part this restatement of
the theory is quoted _verbatim_ from my original paper—the differences
being due only to the conditions imposed by limits of an article.
The following, then, is quoted from the _Nineteenth Century_ for
January, 1887:
"According to the Darwinian theory [which, as elsewhere
fully explained, the present theory is in no way capable
of supplanting, but only of supplementing, and this among
other ways, by explaining why it is that some degree of
mutual infertility is so general a phenomenon as between
allied species—a phenomenon which Darwin expressly regarded
as not explicable by the theory of natural selection], it
is for the most part only those variations which happen to
have been useful that have been preserved: yet, even as thus
limited, the principle of variability is held able to furnish
sufficient material out of which to construct the whole
adaptive morphology of nature. How immense, therefore, must
be the number of unuseful variations. Yet these are all, for
the most part, still-born, or allowed to die out immediately
by intercrossing. Should such intercrossing be prevented,
however, there is no reason why unuseful variations should
not be perpetuated by heredity quite as well as useful ones
when under the nursing influence of natural selection—as,
indeed, we see to be the case in our domesticated productions.
Consequently, if from any reason a section of a species is
prevented from intercrossing with the rest of its species,
new varieties of a trivial or unuseful kind might be expected
to arise within that section. And this is just what we
find. Oceanic islands, for example, are well known to be
extraordinarily rich in peculiar species; and this can best
be explained by considering that a complete separation of the
fauna and flora on such an island permits them to develop
varietal histories of their own, without interference by
intercrossing with their originally parent forms. We see the
same principle exemplified by the influence of geographical
barriers of any kind, and also by the consequences of
migration. Therefore, given an absence of overwhelming
intercrossing, and the principle of what I term independent
variability may be trusted to evoke new species, without the
aid of natural selection. [Homogamy.]
"Were it not for the very general occurrence of some degree
of sterility between even closely allied species and were
it not also for the fact, that closely allied species are
not always—or even generally—separated from one another
by geographical barriers, we might reasonably attribute all
cases of species-formation by independent variability to the
prevention of intercrossing by geographical barriers or by
migration. But it is evident that these two facts can no more
be explained by the influence of geographical barriers, or
by migration, than they can be by the influence of natural
selection.
"Now, of all parts of those variable objects which we call
organisms, the most variable is the reproductive system; and
the variations may be either in the direction of increased
or diminished fertility. Consequently, variations in the
way of greater or less sterility frequently take place both
in plants and animals; and probably, if we had adequate
means of observing this point, we should find that there is
no one variation more common. But, of course, whenever it
arises—whether as a result of changed conditions of life, or,
as we say, spontaneously—it immediately becomes extinguished,
seeing that the individuals which it affects are less able
(if able at all) to propagate the variation. If, however, the
variation should be such that, while showing some degree of
sterility with the parent form, it continues to be as fertile
as before within the limits of the varietal form, it would
neither be swamped by intercrossing nor die out on account of
sterility.
"For example, suppose the variation in the reproductive
system is such that the season of flowering, or of pairing,
becomes either advanced or retarded. Whether this variation
be "spontaneous," or due to change of food, climate, habitat,
etc., does not signify. The only point we need attend to
is that some individuals, living on the same geographical
area as the rest of their species, have demonstrably varied
in their reproductive systems, so that they are perfectly
fertile _inter se_, while absolutely sterile with the rest
of their species. By inheritance there would thus arise a
variety living on the same geographical area as its parent
form, and yet prevented from intercrossing with that form by
a barrier quite as effectual as a thousand miles of ocean;
the only difference would be that the barrier, instead of
being geographical, is physiological. And now, of course, the
two sections of the physiologically divided species would be
able to develop independent histories of their own without
intercrossing; even though they are living together on the
same geographical area, their physiological isolation would
lead to their taking on distinct specific characters by
independent variations, [or homogamy,] just as is the case
with sections of a species when separated from each other by
geographical isolation.
"To state this suggestion in another form, it enables us to
regard many, if not most, species as the records of variations
in the reproductive systems of ancestors. When variations of
a non-useful kind occur in any of the other systems or parts
of organisms, they are, as a rule, immediately extinguished
by intercrossing. But whenever they happen to arise in the
reproductive system in the way here suggested, they must
tend to be preserved as new natural varieties, or incipient
species. At first the difference would only be in respect
of the reproductive systems; but eventually, on account of
independent variation, other differences would supervene, and
the new variety would take rank as a true species.
"The principle thus briefly sketched in some respects
resembles, and in other respects differs from, the principle
of natural selection, or survival of the fittest. For the sake
of convenience, therefore, and in order to preserve analogies
with already existing terms, I have called this principle
Physiological Selection, or Segregation of the Fit.
"Let it be noted that we are not concerned either with the
causes or the degrees of the particular kind of variation on
which this principle depends. Not with the causes, because
in this respect the theory of physiological selection is in
just the same position as that of natural selection: it is
enough for both that the needful variations are provided,
without its being incumbent on either to explain the causes
which in all cases underlie them. Neither are we concerned
with the degrees of sterility which the variation in question
may in any particular case supply. For whether the degree of
sterility with the parent form be originally great or small,
the result of it will be in the long run the same: the only
difference will be that in the latter case a greater number
of generations would be required in order to separate the
varietal from the parent form. [In other words, homogamy due
to such physiological isolation is cumulative.]
"The object of this paper being that of furnishing a general
answer to criticisms on the hypothesis of physiological
selection, I will not occupy space by detailing evidence
of that hypothesis, further than is needful for the object
just mentioned.[3] This evidence abundantly proves that the
particular kind of variation which the theory of physiological
selection requires does take place, (_a_) in individuals,
(_b_) in races, and (_c_) in species. Next, the evidence goes
on to show that the facts of organic nature are such as they
ought to be, supposing it true that this variation has played
any considerable part in the differentiation of specific
types. In particular, it is shown that the general association
between the one primary, or relatively constant, specific
distinction (mutual sterility), and the innumerable secondary,
or relatively variable, distinctions (slight morphological
changes which may effect _any_ parts of _any_ organisms),
of itself indicates that the former has been the original
condition to the occurrence of the latter, in all cases where
free intercrossing has not been otherwise prevented. For even
in cases where the secondary distinctions may be supposed to
have induced the primary,—or where morphological changes
taking place in other parts of an organic type have exercised
a reflex influence on the reproductive system, such that the
changed organism is no longer fertile with its unchanged
parent form,—even in such cases the theory of physiological
selection is available to explain the association in question.
For even in these cases, notwithstanding that the secondary
changes are historically the prior changes, they still depend
for their preservation on the principles of physiological
selection. These principles have, in all such cases,
_selected_ the particular kinds of secondary distinction
which have proved themselves capable of so reacting on the
reproductive system as to bring about the primary distinction,
and thus to protect themselves against the destructive power
of free intercrossing."
[3] The evidence, so far as yet published, may be read by
any one who cares to purchase the original paper, which can
be obtained from the Linnean Society in a separate form.
Now for Mr. Wallace's criticism of this theory, as presented in his
recently published work on "Darwinism."
Briefly put, he furnishes a numerical calculation, showing that when
"the physiological peculiarity is not correlated with any external
differences of form or color, or with inherent peculiarities of likes
or dislikes leading to any choice as to pairing," even when so large
a proportion as ten per cent. of the exceptional variety arises every
year in the midst of the species, "it is unable to increase its numbers
much above its starting-point, and remains wholly dependent on the
continued renewal of the variety for its existence beyond a few years."
This, it must be observed, is a reproduction of the criticism which I
answered in 1888; but, as Mr. Wallace ignores that answer, I must now
repeat it.
The criticism does not dispute the fact that the required variation in
the way of "selective sterility" occurs. Indeed, Mr. Wallace allows
that it certainly must be of very general occurrence as between
incipient species (or pronounced varieties in a state of nature),
seeing that it is of such general occurrence as between allied species
when fully differentiated as such. In other words, this variation in
the way of selective sterility must be recognised as a very general
_fact_, even if it be not regarded as a _condition_, or a _cause_, of
specific differentiation. Which is merely another way of saying that
the particular variation which is required by the theory in question
is admittedly a variation which does occur; and occurs, moreover, in
very frequent association with the origin of a new species. But Mr.
Wallace's objection to regarding this variation as itself a cause of
(or condition to) the origin of a new species is, as we have seen, that
the changes must always be greatly against the similar variations of
the opposite sexes meeting—i. e., of the "physiological complements"
happening to pair. Now, I have already shown, in the _Nineteenth
Century_ of three years ago, that this criticism can only apply to
species the sexes of which unite for every birth; but as Mr. Wallace
continues to ignore this important consideration, I will now present it
in somewhat more detail.
In considering any "supplementary theory" of the origin of species,
it is obviously absurd to disregard the realm of organic nature as a
whole, and to fasten attention exclusively upon the part of it where
a particular difficulty against the theory may be supposed to lie. As
will presently be shown, Mr. Wallace is entirely mistaken in supposing
that his particular difficulty does lie against the theory in any
part of organic nature; but, even if this could not have been shown,
it would not have followed that the theory of physiological selection
is inapplicable to _all_ the classes of the animal and vegetable
kingdoms, because it is taken to be inapplicable to _some_. One might
just as well argue against Mr. Darwin's theory of sexual selection
on the ground that it cannot be held to apply to the coloration and
the sculpture of shells. If either sexual selection or physiological
selection were put forward as an _exclusive_ theory of the origin of
_all_ species, this kind of argument would, of course, have been valid;
but as the matter actually stands, it is largely irrelevant.
I say _largely_ irrelevant, because I do not dispute that there is
this much force in it. If the theory of physiological selection can be
proved inapplicable to Birds and Mammals (which are the only classes
that Mr. Wallace considers in connection with it), its applicability to
all other divisions, both of the animal and vegetable kingdoms, would
be rendered doubtful; seeing that the process of species-formation
appears to have been _everywhere_ more or less associated with the
occurrence of "selective sterility"; and hence, if in any division
of organic nature it could be shown that selective sterility cannot
possibly have been a cause of specific differentiation, we might well
doubt whether it has been such a cause elsewhere—just as we may doubt
whether sexual selection has been a cause of the brilliant colors of
birds and butterflies, because we know, that it cannot have been a
cause of the equally brilliant colors of corals and flowers. But,
as far as physiological selection is concerned, no such question can
arise, as I will presently proceed to show.
First of all, however, it is desirable briefly to indicate the strength
of this theory in the parts of organic nature where Mr. Wallace's
sole criticism cannot possibly be held to apply—viz., the larger
part of the vegetable kingdom, where ovules are fertilised either by
insects or by the wind. Here the phenomena of "prepotency" are highly
suggestive—not to say, in my opinion, virtually demonstrative—of
physiological selection; seeing that, as Mr. Darwin remarks in another
connexion:
"There can be no doubt that if the pollen of all these species
(of Compositae) could be simultaneously or successively placed
on the stigma of any one species, this one would elect with
unerring certainty its own pollen. This elective capacity is
all the more wonderful; as it must have been acquired since
the many species of this great group of plants branched off
from a common progenitor."[4]
[4] _Variation_, etc., Vol. ii.
Darwin is here speaking of elective affinity in its more
fully developed form, as this so often obtains between fully
differentiated species. But we meet with all lower degrees of its
development—sometimes between "incipient species," or varieties, and
at other times between closely allied species. It is then known as
"prepotency" of the pollen belonging to the same variety, or species,
over the pollen of the other variety or species, when both sets of
pollen are applied to the same stigma. This is one form of what I have
called physiological selection, and in my view it serves to explain
why it is that hybrids between closely allied forms growing on common
areas (whether they be called "species" or "constant varieties")
are so comparatively rare in nature, even in cases where there is
no difficulty in producing hybrids artificially by an intentional
exclusion of the pollen belonging to the same form. And I allude to
these facts in the present connexion for two reasons. In the first
place, they serve to show how entirely irrelevant Mr. Wallace's
whole criticism is to the vegetable kingdom, as well as to the
majority of aquatic animals. In the next place, they serve to show
how entirely unwarranted is his statement, that "we have at present
no evidence whatever" in support of my belief that a physiological
incompatibility may affect a whole race or strain. Not only have we
the multitudinous cases of prepotency, where the incompatibility is
_partial_ (or in course of becoming, as Mr. Darwin says in the above
quotation, "acquired"); but we have also multitudinous cases where the
incompatibility has become _absolute_, both as between closely allied
species, and even as between varieties of the same species growing on
common areas—as M. Jordan has experimentally proved. Therefore in the
above remark we have but an additional example of Mr. Wallace's entire
forgetfulness, in the present connexion, of any organisms other than
those which belong to the class of Birds or of Mammals.[5]
[5] It seems scarcely worth while to add that Mr. Wallace
is doubly mistaken where he says, "Mr. Romanes's theory
of Physiological Selection—which assumes sterility or
infertility between first crosses as the fundamental fact
in the origin of species—does not accord with the general
phenomena of hybridism in nature." In the first place, as
shown above, "infertility between the first crosses" is by no
means out of accord with "the general phenomena of hybridism
in nature"—seeing that all degrees of such infertility, from
the slightest perceptible amount of prepotency up to absolute
sterility, are of the most general occurrence in nature. In
the second place, why Mr. Wallace should suppose that in my
view physiological selection can only act as regards first
crosses, and not also as regards hybrid progeny, I have no
means of surmising.
Turning, then, to the only parts of organic nature where his criticism
can even appear to apply, I have here the sufficiently easy task of
proving, that this appearance of application arises wholly and entirely
out of Mr. Wallace's misapprehension of the theory against which the
criticism is directed. In other words, he is not criticising the theory
of physiological selection at all, but merely his own travesty of it.
For, as repeatedly stated in my original paper, and again reiterated
three years ago in the _Nineteenth Century_, it constitutes no part of
my theory to deny the co-operation of other forms of segregate breeding
or homogamy. On the contrary, I have always insisted—and Mr. Gulick
has proved by calculation—that the more efficient the co-operation of
other forms of homogamy, the _greater_ must become the importance of
the physiological form. Yet, as I trust has already been made fully
apparent, the whole of Mr. Wallace's criticism (even as regards Birds
and Mammals) goes upon the supposition that Mr. Gulick and I believe
that, if physiological selection ever acts in any case _at all_, it
must necessarily act _alone_. For reasons afterwards to be given, I do
indeed believe that in some cases it may act alone (in this differing
from Mr. Gulick); but, clearly, whether or not there are any such
cases, is a question quite distinct from that touching the validity
of a criticism which attributes to our theory the absurd dogma, that
segregate breeding which arises from physiological isolation, can
never be associated with segregate breeding that may arise from any
other form of isolation. And that the whole of Mr. Wallace's criticism
collapses when once this correction has been supplied, is proved most
effectually by the curious fact that, _after having himself supplied
the correction, he reproduces our theory as an original one of his
own_. How he can have supposed that I did not entertain the possibility
of physiological selection being associated with natural selection,
"psychological selection," or any other known form of isolation
(excepting only the geographical), I am quite at a loss to understand;
seeing that from end to end of my paper I continually refer to such
association—especially as regards natural selection. And, if possible,
I am still less able to understand Mr. Wallace's carelessness in this
connection with reference to Mr. Gulick's paper; because there the
belief is repeatedly and most clearly expressed, that without such
association, "segregate fecundity" _can never act at all_—which is
precisely the theory which Mr. Wallace proceeds to elaborate on his own
account.
It is now time to show, by means of quotations, how unequivocal and
complete is Mr. Wallace's adoption of our theory:
"The simplest case to consider will be that in which two forms
or varieties of a species, occupying an extensive area, are
in process of adaptation to somewhat different modes of life
within the same area. If these two forms freely intercross
with each other, and produce mongrel offspring which are quite
fertile _inter se_, then the further differentiation of the
forms into two distinct species will be retarded, or perhaps
entirely prevented; for the offspring of the crossed unions
will be, perhaps, more vigorous on account of the cross,
although less perfectly adapted to the conditions of existence
than either of the pure breeds; and this would certainly
establish a powerful antagonistic influence to the further
differentiation of the two forms.
"Now, let us suppose that a partial sterility of the hybrids
between the two forms arises, in correlation with the
different modes of life and the slight external or internal
peculiarities that exist between them, both of which we have
seen to be real causes of infertility. The result will be
that, even if the hybrids between the two forms are still
freely produced, these hybrids will not themselves increase
so rapidly as the two pure forms; and as these latter are, by
the terms of the problem, better suited to their conditions
of life than are the hybrids between them, they will not only
increase more rapidly, but will also tend to supplant the
hybrids altogether whenever the struggle for existence becomes
exceptionally severe. Thus, the more complete the sterility of
the hybrids the more rapidly will they die out and leave the
two parent forms pure. Hence it will follow that, _if there
is greater infertility between the two forms in one part of
the area than the other, these forms will be kept more pure
wherever this greater infertility prevails_, will therefore
have an advantage at each recurring period of severe struggle
for existence, and will thus ultimately supplant the less
infertile or completely fertile forms that may exist in other
portions of the area. It thus appears that, in such a case as
here supposed, natural selection would preserve those portions
of the two breeds which were most infertile with each other,
or whose hybrid offspring were most infertile; and would,
therefore, if variations in fertility continued to arise, tend
to increase that infertility. It must particularly be noted
that this effect would result, not by the preservation of the
infertile variations on account of their infertility, but by
the inferiority of the hybrid offspring, both as being fewer
in numbers, less able to continue their race, and less adapted
to the conditions of existence than either of the pure forms.
It is this inferiority of the hybrid offspring that is the
essential point; and as the number of these hybrids will be
permanently less where the infertility is greatest, therefore
those portions of the two forms in which infertility is
greatest will have the advantage, and will ultimately survive
in the struggle for existence."
We have here a full acceptance of the theory of physiological
selection. For it is represented, as Mr. Gulick and I have represented,
that, if "two forms or varieties" occupying a common area are to
undergo further differentiation at the hands of natural selection, it
becomes a highly favoring condition to the process that some degree
of segregate fecundity should arise (if it has not already arisen)
between these two forms or varieties; seeing that "if these two forms
freely intercross with each other, and produce mongrel offspring
which are quite fertile _inter se_, then the further differentiation
of the forms into two distinct species will be retarded, or perhaps
entirely prevented." Here the importance of segregate fecundity, or
physiological selection, as a factor in the differentiation of specific
types on common areas is _fully recognised_; and the _only_ respect
in which Mr. Wallace alleges that his view of the matter differs from
the view of Mr. Gulick and myself, is in drawing special attention to
the part which is played by the infertility, or other "inferiority,"
of the mongrels. But clearly, this infertility, or other inferiority,
of the mongrels, in all cases where it occurs, _is part and parcel of
the segregate fecundity of the parent forms_. Whether the segregate
fecundity has reference to first crosses alone, or likewise to
second crosses, it is segregate fecundity all the same; and the only
difference is that for the same degree of segregate fecundity in first
crosses, the process of physiological selection will become the more
effective in proportion to the degree in which the infertility extends
also to second crosses. But I think it is very doubtful whether such
infertility (or inferiority) on the part of mongrels can react upon
the sexual system of their parent forms, so as directly to increase
whatever degree of segregate fecundity may have already arisen between
these forms. Does the high sterility of mules and mutes, for instance,
tend to diminish the degree of fertility that obtains between horses
and asses? The only way in which even an absolute degree of sterility
(or other inferiority) on the part of mongrels or hybrids may clearly
be seen to operate in this direction, is as a negative condition; not
as an active cause. In the proportion that mongrels are impotent with
one another, they will not so much compete with their parent forms
for food, etc.; and in the proportion that they are impotent with
their parent forms, they will not counteract any tendency which the
latter may continue to develop in the direction of a still further
segregation. If the mongrels are fully vigorous and fully fertile, both
_inter se_ and with their parent forms, the effect will be to retard,
if not altogether to prevent, any further progress of physiological
separation between the parent forms; because the free intercrossing of
the mongrels with one another, and also with their parent forms, will
be continually supplying progeny in which the physiological peculiarity
is either attenuated or altogether abolished. But this is quite a
different thing from supposing that infertility (or inferiority) of
the mongrels can react upon the generative system of the parent forms,
so as to increase in them the physiological peculiarity on which
their segregate breeding depends: infertility (or inferiority) of the
mongrels is but a negative condition which favors the preservation of
further degrees of this segregate breeding, if such further degrees
should be induced by any other causes.
Now, it does not appear that Mr. Wallace has clearly perceived
this important distinction, because he throughout speaks of "this
inferiority of the hybrid offspring as the essential point." Obviously,
however, the essential point is the physiological variation in the
parent forms, i. e., the original occurrence and subsequent development
of infertility between the first crosses. Granting to Mr. Wallace, for
the sake of argument, that this development could not proceed at all,
were it not for the inferiority of the mongrels; still the inferiority
of the mongrels need not be the cause of this development. Therefore
it is most incorrect to say, "it must be particularly noted that this
effect (i. e., increase of infertility between the parent forms)
would result, not by the preservation of the infertile variations on
account of their infertility, but by the inferiority of the hybrid
offspring." "This effect" must be due to causes which act upon the
generative systems of the parent forms, even though such causes might
be _counteracted_ by the _withdrawal_ of the negative condition in
question.
I trust, then, it has now been rendered sufficiently clear that, no
matter how infertile the hybrid progeny may become, and no matter at
how great a disadvantage they may thus (or otherwise) be placed in
their struggle for existence with the parent varieties, it is not
apparent that their infertility (or their extinction) can ever become
the cause of a further increase of infertility arising between their
parent forms. Consequently, although this is the cause assigned by Mr.
Wallace, when he comes to "the essential point" of showing how it is
to act so as to increase cross-sterility between the parent forms, he
naïvely substitutes the sentence which I have printed in italics—which
assumes a "greater infertility between the two forms" as arising
through any other causes that we may choose to suppose. The very thing
that his entire argument professes to explain (i. e., the rise and
development of cross-sterility between the parent varieties) is slipped
in as granted, or given by other causes than those which are said to
explain it.[6]
[6] The only conceivable way in which infertility (or other
inferiority) of hybrids could react on the sexual system of
their parent forms, is one which Mr. Wallace appears to have
missed: at all events he has nowhere stated it. This way is
as follows. Suppose A and B to be two varieties which produce
comparatively infertile hybrids. In the proportion that the
hybrids are infertile, or otherwise inferior, it must be a
disadvantage to both varieties for individuals belonging
to one to cross with individuals belonging to the other,
because by so doing they are wasting their time and their
energy in propagating comparatively poor offspring—thereby
failing to impress their characters on the next generation as
effectually as they might have done by pairing homogamously.
Hence, those individuals which do pair homogamously will
leave a larger number—or better quality—of offspring
to the next generation, than is left by those which fail
to pair homogamously. Hence, also, in the course of many
generations a selective premium will be set on the homogamous
pairing, A plus A, B plus B, whether such pairing be due
to a sexual instinct or to a sexual incompatibility. For
example, if horses and asses were to occupy the same area
for a sufficient length of time, it is conceivable that the
instinct which many horses now present of preferring asses
to their own kind would become obsolete; because the horses
or mares which have such an instinct would always fail to
leave progeny that could transmit it, while such would not be
the case with the horses and mares which preferred to pair
homogamously, and so it might be if a physiological instead
of a psychological character were concerned. But now observe,
if this consideration were adduced, I should not be concerned
to dispute it. For, even if such a principle of segregation
does obtain, to what category does the principle belong?
Clearly it does not belong to natural selection, inasmuch
as a mere failure to impress individual characters on the
next generation is not a matter of life and death in the
struggle for existence. But, no less clearly, it does belong
to physiological selection; and therefore, if it be an active
principle in nature, it is an additional cause of segregate
fecundity in first crosses. Moreover, such a principle, if
it ever acts, presupposes some considerable degree of sexual
differentiation as already given by some other cause.
Having thus endeavored to make it as clear as I can, that the causes of
segregate fecundity, both in its origin and subsequent "increase," must
be causes acting on the physiology of the segregating forms themselves,
and not the effects of these causes in the character of their mongrel
offspring; I must next comment upon the extraordinary idea which
underlies the whole of Mr. Wallace's exposition, and which in one place
he expressly states. This extraordinary idea is that the theory of
physiological selection, as held both by Mr. Gulick and myself, takes
no cognizance of the possible effects of cross-sterility in leading
to infertility or inferiority on the part of mongrel progeny. I call
this an extraordinary idea, because it appears to me most extraordinary
that Mr. Wallace can have read our papers, and then have supposed that
he was adding anything to our theory by arguing the points which he
does argue in the above quotation. When once this argument is correctly
stated, it amounts, as we have just seen, to nothing more than pointing
out how a segregate fecundity of first crosses will have a better
chance of increasing, if the mongrel progeny are infertile or inferior.
But surely this goes without saying; or, if it be said, let it be added
that physiological selection, when it thus extends to second crosses,
is really or ultimately due to physiological selection as regards the
first crosses. If the segregate fecundity of the first crosses is of
such a kind, that, besides tending to a physiological isolation of the
parent forms, it leads to inferiority of the mongrel progeny; this is
merely a further expression of the segregate fecundity in question.
Its effect is that of so far extinguishing the influence of progeny in
the subsequent history of parental segregation: therefore, its effect
is just the same as if, owing to a somewhat higher degree of segregate
fertility in the first instance (i. e., in the first crosses), a
proportionately smaller number of mongrel offspring had been produced
at all. In either case the result (physiological differentiation) is
equally due to causes acting on the sexual system of the parent forms;
and whether this effect is brought about by a suppression of progeny
as to their numbers alone, or likewise as to their efficiency, is
quite immaterial to the theory of physiological selection. Which shows
once more how wide of the mark is Mr. Wallace's statement, that "the
inferiority of the hybrid offspring is the essential point" in any
process of sexual segregation. The "essential point" must always be
the original occurrence and subsequent "preservation of the infertile
variations" arising between the parent forms, whether these variations
are only in the direction of producing a smaller number of mongrels, or
also in that of suppressing their efficiency when produced.
Upon the whole, then, it is surely the oddest of misconceptions on Mr.
Wallace's part that has led him to present the above-quoted "argument"
as a substitute for the theory of physiological selection. As far as
it goes, and as far as it is sound, it is the theory of physiological
selection pure and simple—neither adding to, nor detracting from it
one iota. Nevertheless, the "argument" has not yet gone far enough to
embody some of the other elements of the theory. Therefore I will now
continue the quotation: