PART II

SYSTEMATICS AND HISTORY

*A.* THE PRINCIPLES OF SYSTEMATICS

RATIONAL SYSTEMATICS

All systematics which deserves the predicate “rational” is founded

upon a concept or upon a proposition, by the aid of which a totality

of specific diversities may be understood. That is to say: every

system claiming to be rational gives us a clue by which we are able to

apprehend either that there cannot exist more than a certain number of

diversities of a certain nature, or that there can be an indefinite

number of them which follow a certain law with regard to the character

of their differences.

Solid geometry, which states that only five regular bodies are possible,

and points out the geometrical nature of these bodies, is a model

of what a rational system should be. The theory of conic sections

is another. Take the general equation of the second degree with two

unknowns, and study all the possible forms it can assume by a variation

of its constants, and you will understand that only four different types

of conic sections are possible--the circle, the ellipse, the hyperbola,

and the parabola.

In physics and chemistry no perfect rational systems have been

established hitherto, but there are many systems approaching the ideal

type in different departments of these sciences. The chemical type of

the monohydric saturated alcohols, for instance, is given by the formula

C_nH_{2n+1}OH, and in this formula we not only have an expression of

the law of composition which all possible alcohols are to follow,--but,

since we know empirically the law of quantitative relation between

*n* and various physical properties, we also possess in our formula a

general statement with respect to the totality of the properties of any

primary alcohol that may be discovered or prepared in the future. But

chemistry has still higher aims with regard to its systematics: all of

you know that the so-called “periodic law of the elements” was the first

step towards a principle that may some day give account of the relation

of all the physical and chemical properties of any so-called element

with its most important constant, the atomic weight, and it seems to be

reserved for the present time to form a real fundamental system of the

“elements” on the basis of the periodic law by the aid of the theory

of electrons. Such a fundamental system of the elements would teach us

that there can only be so many elements and no more, and only of such a

kind. In crystallography a similar end has been reached already by means

of certain hypothetic assumptions, and systematics has here accounted

for the limited number and fixed character of the possible forms of

crystalline symmetry.

It is not difficult to understand the general logical type of all

rational systems, and logic indeed can discover it without appealing

to concrete sciences or to geometry. Rational systematics is always

possible whenever there exists any fundamental concept or proposition

which carries with it a principle of division; or to express it somewhat

differently, which would lead to contradictions, if division were to

be tried in any but one particular manner. The so-called “genus,” as

will easily be perceived, then embraces all its “species” in such a

manner that all peculiarities of the species are represented already in

properties of the genus, only in a more general form, in a form which

is still unspecified. The genus is both richer in content and richer in

extent than are the species, though it must be added that its richness

in content is, as it were, only latent: but it may come into actuality

by itself and without any help from without.

We are dealing here with some of the most remarkable properties of the

so-called synthetic judgments *a priori* in the sense of Kant, and,

indeed, it seems that rational systematics will only be possible where

some concept of the categorical class or some proposition based upon

such concept lies at the root of the matter or at least is connected

with it in some way. In fact, all rational systems with regard to the

relations of symmetry in natural bodies deal ultimately with space; or

better, all systems in such fields are able to become rational only if

they happen to turn into questions of spatial symmetry.

All other genera and species, whether of natural bodies or of facts,

can be related only on the basis of empirical abstraction, *i.e.* can

never attain rationality: here, indeed, the genus is richer in extent

and poorer in content than are the species. The genus is transformed

into the species, not by any inherent development of latent properties,

but by a mere process of addition of characteristic points. It is

impossible to deduce the number or law or specifications of the species

from the genus. Mere “classification,” if we may reserve the honorable

name of systematics for the rational type, is possible here, a mere

statement in the form of a catalogue, useful for orientation but for

nothing more. We may classify all varieties of hats or of tables in the

same way.

BIOLOGICAL SYSTEMATICS

At this point we return from our logical excursion to our proper subject

of biology; for I am sorry to say biological systematics is at present

of our second type of systematics throughout: it is classification pure

and simple. We have a catalogue in our hands, but nothing more.

Such a statement of fact conveys not a particle of censure, casts not

the least reflection on the gifted men who created the classification of

animals or plants. It is absolutely necessary to have such a catalogue,

and indeed the catalogue of the organisms can be said to have been

improved enormously during the advance of empirical and descriptive

biological science. Any classification improves as it becomes more

“natural,” as the different possible schemes of arrangement, the

different reasons of division, agree better and better in their results;

and, in fact, there has been a great advance of organic classification

in this direction. The “natural” system has reached such perfection,

that what is related from one point of view seems nearly related also

from almost all points of view which are applicable, at least from those

which touch the most important characteristics. There has been a real

weighing of all the possible reasons of division, and that has led to a

result which seems to be to some extent final.

But, nevertheless, we do not understand the *raison d’être* of the

system of organisms; we are not at all able to say that there must be

these classes or orders or families and no others, and that they must be

such as they are.

Shall we ever be able to understand that? Or will organic systematics

always remain empirical classification? We cannot answer this question.

If we could, indeed, we should have what we desire! As simple relations

of space are certainly not the central point of any problematic rational

organic systematics even of the future, the question arises, whether

there could be found any principle of another type in the realm of

synthetic *a priori* judgments which could allow an inherent sort of

evolution of latent diversities, as do all judgments about spatial

symmetry. At the end of the second course of these lectures, which is to

be delivered next summer, we shall be able to say a few more words about

this important point.

The concept of what is called “a type,” due almost wholly to Cuvier and

Goethe, is the most important of all that classification has given to

us. Hardly second in importance is the discovery of the “correlation of

parts,” as a sort of connection which has the character of necessity

without being immediately based upon causality. Rádl seems to be

the only modern author who has laid some stress on this topic. The

harmony which we have discovered in development is also part of this

correlation. When, later on, we come to discuss analytically our well

established entelechy as the ultimate basis of individual organisation,

we shall be able to gain more satisfactory ideas with respect to the

meaning of the non-causal but necessary connection, embraced in the

concepts of type and of correlation of parts.

The type is a sort of irreducible arrangement of different parts; the

correlation deals with the degree and the quality of what may be called

the actual make of the parts, in relation to one another: all ruminants,

for instance, are cloven-footed, the so-called dental formulae are

characteristic of whole groups of mammals. Of course all such statements

are empirical and have their limits: but it is important that they are

possible.[142]

[142] Recent years have created the beginnings of a systematics based on

chemical differences of metabolism and its products: such differences in

fact have been found to go hand in hand with diversities of the type in

some cases (v. Bunge, Przibram, etc.).

It has been the chief result of comparative embryology to show that the

type as such is more clearly expressed in developmental stages than

it is in the adults, and that therefore the embryological stages of

different groups may be very much more similar to each other than are

the adults: that is the truth contained in the so-called “biogenetisches

Grundgesetz.” But the specific differences of the species are not

wanting in any case of ontogeny, in spite of such similarities in

different groups during development.

We have applied the name “systematics” or, if rationality is excluded,

“classification” to all that part of a science which deals with

diversities instead of generalities: in such a wide meaning systematics,

of course, is not to be confused with that which is commonly called so

in biology, and which describes only the exterior differences of form.

Our systematics is one of the two chief parts of biology; what are

called comparative anatomy and comparative embryology are its methods.

For it must be well understood that these branches of research are only

methods and are not sciences by themselves.

*B.* THE THEORY OF DESCENT

1. GENERALITIES

It is most generally conceded at the present time that the actually

existing state of all organisms whatsoever is the result of their

history. What does that mean? What are the foundations upon which the

assumption rests? What is the relation of systematics to history? In

raising such questions and considerations we are treading the ground

sacred to the theory of descent.

I well know that you prefer the name “theory of evolution” for what

I am speaking of: but it may be misleading in various respects. We

already know that quite a determinate meaning has been given to the word

“evolutio” as applied to individual morphogenesis, “evolutio” being

here opposed to “epigenesis.” Now there would be nothing against the

use of the word evolution in a wider sense--indeed it is often applied

nowadays to denote the fact that a something is actually “evolved” in

embryology--if only our entelechy had taken the place of the machine

of the mechanists. But that is the very point: there must be a real

“evolving” of a something, in order that the word evolution may be

justified verbally: and that is not the case in so-called phylogeny. At

least we know nothing of an evolutionary character in the problematic

pedigree of the organisms, as we shall see more fully hereafter. The

term “theory of descent” is therefore less open to objection than is the

usual English term. The word transformism, as used by the French, would

also be a very good title.

The theory of descent is the hypothetic statement that the organisms

are really allied by blood among each other, in spite of their

diversities.[143] The question about their so-called monophyletic

or polyphyletic origin is of secondary importance compared with the

statement of relationship in general.

[143] We prefer this unpretending definition of the theory of descent

to every other. As soon as one introduces into the definition the

concept of the “transmutability of species,” the term “species” would

require a special definition, and that would lead to difficulties which

it is unnecessary to deal with for our main purposes. It has been

remarked by Krašan, (*Ausichten und Gespräche über die individuelle und

specifische Gestaltung in der Natur*) and by several other writers,

that the problem of mutability or immutability of course relates to the

individuals in the first place. I should like to add to this remark that

the possibility must be admitted of the individuals being transmutable,

whilst the “species” are not transmutable at the same time, the line

of the “species” being a fixed order, through which the “individuals”

have to pass in the course of their generations. What is meant here

will become clearer, when we study the different possible aspects of

“phylogeny.”

There are two different groups of facts which have suggested the idea

of transformism: none of these facts can be said to be conclusive, but

there certainly is a great amount of probability in the whole if taken

together.

The first group of evidences which lead to the hypothesis of the real

relationship of organisms consists of facts relating to the geographical

distribution of animals and plants and to palæontology. As to geography,

it seems to me that the results of the floral and faunal study of groups

of islands are to be mentioned in the first place. If, indeed, on

each of the different islands, *A* *B* *C* and *D*, forming a group,

the species of a certain genus of animals or plants are different in a

certain respect, and show differences also compared with the species

living on the neighbouring continent, of which there is geological

evidence that the islands once formed a part, whilst there is no change

in the species on the continent itself for very wide areas, then, no

doubt, the hypothesis that all these differing species once had a common

origin, the hypothesis that there is a certain community among them all,

will serve to elucidate in some way what would seem to be very abstruse

without it. And the same is true of the facts of palaeontology. In

the geological strata, forming a continuous series, you find a set of

animals, always typical and specific for every single stratigraphical

horizon, but forming a series just as do those horizons. Would not the

whole aspect of these facts lose very much of its peculiarity if you

were to introduce the hypothesis that the animals changed with the

strata? The continuity of life, at least, would be guaranteed by such an

assumption.

The geographical and geological evidences in favour of the theory of

descent are facts taken from sciences which are not biology proper; they

are not facts of the living but only facts about the living. That is not

quite without logical importance, for it shows that not biology alone

has led to the transformism hypothesis. Were it otherwise, transformism

might be said to be a mere hypothesis *ad hoc*; but now this proves to

be not the case, though we are far from pretending that transformism

might be regarded as resting upon a real *causa vera*.

But let us study the second group of facts which support the theory of

descent. It is a group of evidences supplied by biology itself that we

meet here, there being indeed some features in biology which can be

said to gain some light, some sort of elucidation, if the theory of

descent is accepted. Of course, these facts can only be such as relate

to specific diversities, and indeed are facts of systematics; in other

words, there exists something in the very nature of the system of

organisms that renders transformism probable. The system of animals and

plants is based upon a principle which might be called the principle

of *similarities and diversities by gradation*; its categories are not

uniform but different in degree and importance, and there are different

kinds of such differences. No doubt, some light would be shed upon this

character of the system, if we were allowed to assume that the relation

between similarities and diversities, which is gradual, corresponded to

a blood-relationship, which is gradual also.

THE COVERT PRESUMPTION OF ALL THEORIES OF DESCENT

We have used very neutral and somewhat figurative words, in order to

show what might be called the logical value of the theory of descent,

in order to signify its value with respect to so-called “explanation.”

We have spoken of the “light” or the “elucidation” which it brings, of

the “peculiarity of aspect” which is destroyed by it. We have used this

terminology intentionally, for it is very important to understand that

a specific though hidden addition is made almost unconsciously to the

mere statement of the hypothesis of descent as such, whenever this

hypothesis is advocated in order to bring light or elucidation into any

field of systematic facts. And this additional hypothesis indeed must be

made from the very beginning, quite irrespective of the more detailed

problems of the law of transformism, in order that *any* sort of

so-called explanation by means of the theory of descent may be possible

at all. Whenever the theory that, in spite of their diversities, the

organisms are related by blood, is to be really useful for explanation,

it must necessarily be assumed in every case that the steps of change,

which have led the specific form *A* to become the specific form *B*,

have been such as only to change *in part* that original form *A*. That

is to say: the similarities between *A* and *B* must never have become

overshadowed by their diversities.

Only on this assumption, which indeed is a newly formed additional

subsidiary hypothesis, joined to the original hypothesis of descent in

general--a hypothesis regarding the very nature of transformism--only

on this almost hidden assumption is it possible to speak of any sort of

“explanation” which might be offered by the theory of transformism to

the facts of geography, geology, and biological systematics. Later on

we shall study more deeply the logical nature of this “explanation”; at

present it must be enough to understand this term in its quasi-popular

meaning.

What is explained by the hypothesis of descent--including the additional

hypothesis, that there always is a prevalence of the similarities

during transformism--is the fact that in palaeontology, in the groups

of island and continent faunae and florae taken as a whole, as well as

in the single categories of the system, the similarities exceed the

diversities. The *similarities* now are “explained”; that is to say,

they are understood as resting on but one principle: the similarities

are understood as being due to inheritance;[144] and now we have but one

problem instead of an indefinite number. For this reason Wigand granted

that the theory of descent affords what he calls a numerical reduction

of problems.

[144] It seems to me that my argument gives a broader logical basis

to the theory of descent than does that of G. Wolff (*Die Begründung

der Abstammungslehre*, München, 1907). Wolff starts from the concept

of organic teleology, and thus finds the only reason for accepting

the theory of transformism in the existence of so-called “rudimentary

organs”; these organs would form an obstacle to teleology if they could

not be regarded as inherited.

Understanding then what is explained by the theory of descent with its

necessary appendix, we also understand at once what is *not* elucidated

by it: the diversities of the organism remain as unintelligible as they

always were, even if we know that inheritance is responsible for what

is similar or equal. Now there can be no doubt that the diversities are

the more important point in systematics; if there were only similarities

there would be no problem of systematics, for there would be no system.

Let us be glad that there are similarities in the diversities, and that

these similarities have been explained in some way; but let us never

forget what is still awaiting its explanation. Unfortunately it has been

forgotten far too often.

THE SMALL VALUE OF PURE PHYLOGENY

And so we are led to the negative side of the theory of transformism,

after having discussed its positive half. The theory of descent as

such, without a real knowledge of the factors which are concerned in

transformism, or of the law of transformism, in other terms, leaves the

problem of systematics practically where it was, and adds really nothing

to its solution. That may seem very deplorable, but it is true.

Imagine so-called historical geology, without any knowledge of the

physical and chemical factors which are concerned in it: what would

you have except a series of facts absolutely unintelligible to you? Or

suppose that some one stated the cosmogenetic theory of Kant and Laplace

without there being any science of mechanics: what would the theory mean

to you? Or suppose that the whole history of mankind was revealed to

you, but that you had absolutely no knowledge of psychology: what would

you have but facts and facts and facts again, with not a morsel of real

explanation?

But such is the condition in which so-called phylogeny stands. If it

is based only on the pure theory of transformism, there is nothing

explained at all. It was for this reason that the philosopher Liebmann

complained of phylogeny that it furnishes nothing but a “gallery of

ancestors.” And this gallery of ancestors set up in phylogeny is not

even certain; on the contrary, it is absolutely uncertain, and very

far from being a fact. For there is no sound and rational principle

underlying phylogeny; there is mere fantastic speculation. How could it

be otherwise where all is based upon suppositions which themselves have

no leading principle at present? I should not like to be misunderstood

in my polemics against phylogeny. I fully grant you that it may be

possible in a few cases to find out the phylogenetic history of smaller

groups with some probability, if there is some palaeontological

evidence in support of pure comparative anatomy; and I also do not

hesitate to allow that such a statement would be of a certain value with

regard to a future discovery of the “laws” of descent, especially if

taken together with the few facts known about mutations. But it is quite

another thing with phylogeny on the larger scale. Far more eloquent than

any amount of polemics is the fact that vertebrates, for instance, have

already been “proved” to be descended from, firstly, the amphioxus;

secondly, the annelids; thirdly, the *Sagitta* type of worms; fourthly,

from spiders; fifthly, from *Limulus*, a group of crayfishes; and

sixthly, from echinoderm larvae. That is the extent of my acquaintance

with the literature, with which I do not pretend to be specially

familiar. Emil du Bois-Reymond said once that phylogeny of this sort is

of about as much scientific value as are the pedigrees of the heroes of

Homer, and I think we may fully endorse his opinion on this point.

HISTORY AND SYSTEMATICS

A few words should be devoted to the relations between history and

systematics in biology. Is there no contradiction between historical

development and a true and rational system which, we conceded, might

exist some day in biological sciences, even though it does not at

present? By no means. A totality of diversities is regarded from quite

different points of view if taken as the material of a system, and if

considered as realised in time. We have said that chemistry has come

very near to proper rational systematics, at least in some of its

special fields; but the compounds it deals with at the same time may

be said to have originated historically also, though not, of course,

by a process of propagation. It is evident at once that the geological

conditions of very early times prohibited the existence of certain

chemical compounds, both organic and inorganic, which are known at

present. None the less these compounds occupy their proper place in

the system. And there may be many substances theoretically known to

chemical systematics which have never yet been produced, on account

of the impossibility of arranging for their proper conditions of

appearance, and nevertheless they must be said to “exist.” “Existence,”

as understood in systematics, is independent of special space and of

special time, as is the existence of the laws of nature: we may speak of

a Platonic kind of existence here. Of course it does not contradict this

sort of ideal existence if reality proper is added to it.

Thus the problem of systematics remains, no matter whether the theory

of descent be right or wrong. There always remains the question about

the totality of diversities in life: whether it may be understood by

a general principle, and of what kind that principle would be. As,

in fact, it is most probably by history, by descent, that organic

systematics is brought about, it of course most probably will happen

some day that the analysis of the causal factors concerned in the

history will serve to discover the principle of systematics also.

Let us now glance at the different kinds of hypotheses which have been

established in order to explain how the descent of the organisms might

have been possible. We have seen that the theory of transformism alone

is not worth very much as a whole, unless at least a hypothetical

picture can be formed of the nature of the transforming factors: it is

by some such reasoning that almost every author who has defended the

theory of descent in its universality tries to account for the manner in

which organisms have acquired their present diversities.

2. THE PRINCIPLES OF DARWINISM

There is no need in our times and particularly in this country, to

explain in a full manner the theory known under the name of Darwinism.

All of you know this theory, at least in its outlines, and so we may

enter at once upon its analytic discussion. A few words only I beg

you to allow me as to the name of “Darwinism” itself. Strange to say,

Darwinism, and the opinion of Charles Darwin about the descent of

organisms, are two different things. Darwin, the very type of a man

devoted to science alone and not to personal interests,--Darwin was

anything but dogmatic, and yet Darwinism is dogmatism in one of its

purest forms. Darwin, for instance, gave the greatest latitude to the

nature of the variations which form the battleground of the struggle

for existence and natural selection; and he made great allowances for

other causal combinations also, which may come into account besides

the indirect factors of transformism. He was Lamarckian to a very

far-reaching extent. And he had no definite opinion about the origin

and the most intimate nature of life in general. These may seem to

be defects but really are advantages of his theory. He left open the

question which he could not answer, and, in fact, he may be said to be a

good illustration of what Lessing says, that it is not the possession

of truth but the searching after it, that gives happiness to man. It was

but an outcome of this mental condition that Darwin’s polemics never

left the path of true scientific discussions, that he never in all his

life abused any one who found reason to combat his hypotheses, and that

he never turned a logical problem into a question of morality.

How different is this from what many of Darwin’s followers have made out

of his doctrines, especially in Germany; how far is “Darwinism” removed

from Darwin’s own teaching and character!

It is to Darwinism of the *dogmatic* kind, however, that our next

discussions are to relate, for, thanks to its dogmatism, it has the

advantage of allowing the very sharp formulation of a few causal

factors, which *a priori* might be thought to be concerned in organic

transformism, though we are bound to say that a really searching

analysis of these factors ought to have led to their rejection from the

very beginning.

The logical structure of dogmatic Darwinism reveals two different parts,

which have nothing at all to do with one another.

NATURAL SELECTION

We shall first study that part of it which is known under the title

of natural selection, irrespective of the nature of the causes of

primary differences, or, in other words, the nature of variability.

This part may be said to belong to Darwin’s personal teachings and not

only to “Darwinism.” The offspring of a certain number of adults show

differences compared with each other; there are more individuals in the

offspring than can grow up under the given conditions, therefore there

will be a struggle for existence amongst them, which only the fittest

will survive; these survivors may be said to have been “selected” by

natural means.

It must be certain from the very beginning of analysis that natural

selection, as defined here, can only eliminate what cannot survive, what

cannot stand the environment in the broadest sense, but that natural

selection never is able to create diversities. It always acts negatively

only, never positively. And therefore it can “explain”--if you will

allow me to make use of this ambiguous word--it can “explain” only why

certain types of organic specifications, imaginable *a priori*, do

*not* actually exist, but it never explains at all the existence of the

specifications of animal and vegetable forms that are actually found. In

speaking of an “explanation” of the origin of the living specific forms

by natural selection one therefore confuses the sufficient reason for

the non-existence of what there is not, with the sufficient reason for

the existence of what there is. To say that a man has explained some

organic character by natural selection is, in the words of Nägeli, the

same as if some one who is asked the question, “Why is this tree covered

with these leaves,” were to answer “Because the gardener did not cut

them away.” Of course that would explain why there are no more leaves

than those actually there, but it never would account for the existence

and nature of the existing leaves as such. Or do we understand in the

least why there are white bears in the Polar Regions if we are told that

bears of other colours could not survive?

In denying any real explanatory value to the concept of natural

selection I am far from denying the action of natural selection. On

the contrary, natural selection, to some degree, is *self-evident*;

at least as far as it simply states that what is incompatible with

permanent existence cannot exist permanently, it being granted that

the originating of organic individuals is not in itself a guarantee of

permanency. Chemical compounds, indeed, which decompose very rapidly

under the conditions existing at the time when they originated may also

be said to have been eliminated by “natural selection.” It is another

question, of course, whether in fact all eliminations among organic

diversities are exclusively due to the action of natural selection in

the proper Darwinian sense. It has been pointed out already by several

critics of Darwinism and most clearly by Gustav Wolff, that there are

many cases in which an advantage with regard to situation will greatly

outweigh any advantage in organisation or physiology. In a railway

accident, for instance, the passengers that survive are not those who

have the strongest bones, but those who occupied the best seats; and

the eliminating effect of epidemics is determined at least as much by

localities, *e.g.* special houses or special streets, as by the degree

of immunity. But, certainly, natural selection is a *causa vera* in many

other cases.

We now may sum up our discussion of the first half of Darwinism.

Natural selection is a negative, an eliminating factor in transformism;

its action is self-evident to a very large degree, for it simply

states that things do not exist if their continuance under the given

conditions is impossible. To consider natural selection as a positive

factor in descent would be to confound the sufficient reason for the

non-existence of what is not, with the sufficient reason of what is.

Natural selection has a certain important logical bearing on

systematics, as a science of the future, which has scarcely ever been

alluded to. Systematics of course has to deal with the totality of the

possible, not only of the actual diversities; it therefore must remember

that more forms may be possible than are actual, the word “possible”

having reference in this connection to originating, not to surviving.

Moreover, systematics is concerned not only with what has been

eliminated by selection, but also with all that might have originated

from the eliminated types. By such reasoning natural selection gains a

very important aspect--but a logical aspect only.

FLUCTUATING VARIATION THE ALLEGED CAUSE OF ORGANIC DIVERSITY

The second doctrine of dogmatic Darwinism states that all the given

diversities among the organisms that natural selection has to work

upon are offered to natural selection by so-called fluctuating

variation; that is, by variation as studied by means of statistics.

This sort of variation, indeed, is maintained to be indefinite in

direction and amount, at least by the most conservative Darwinians; it

has occasionally been called a real differential; in any case it is

looked upon as being throughout contingent with regard to some unity

or totality; which, of course, is not to mean that it has not had a

sufficient reason for occurring.

It could hardly be said to be beyond the realm of possibility that such

differences among organic species as only relate to degree or quantity

and perhaps to numerical conditions also, might have been “selected” out

of given contingent variations, if but one postulate could be regarded

as fulfilled. This postulate may appropriately be stated as the fixation

of new averages of variation by inheritance. Let the average value of

a variation, with regard to a given property of a given species be *n*

and let the value *n* + *m*--*m* being variable--which is represented in

fewer individuals of course than is *n*, be such as to offer advantages

in the struggle for existence; then the individuals marked by *n* + *m*

will have the greater chance of surviving. Our postulate now states

that, in order that a permanent increase of the average value of the

variation in question may be reached, *n* + *m* in any of its variable

forms must be able to become the average value of the second generation,

as *n* was the average value of the first. Out of the second generation

again it would be the few individuals marked by *n* + *m* + *o*, which

would be selected; *n* + *m* + *o* would be the new average; afterwards

*n* + *m* + *o* + *p* would be selected, would become the new average,

and so on. A black variety for instance might be selected by such a

series of processes out of a grey-coloured one without difficulty.

But our postulate is not beyond all doubt: certain experiments, at

least, which have been carried out about the summation of variations

of the true fluctuating type by any kind of selection seem to show

that there may be a real progress for a few generations, but that this

progress is always followed by a reversion. Of course our experience

is by no means complete on this subject, and, indeed, it may be shown

in the future that positive transforming effects of fluctuating

variability, in connection with selective principles, are possible in

the case of new quantitative differences (in the widest sense), but we

are not entitled to say so at present.

And this is the only condition on which we can give credit to the second

doctrine of dogmatic Darwinism. Its second principle, indeed, proves

to be absolutely inadequate to explain the origin of any other kind of

specific properties whatever.

I cannot enter here into the whole subject of Darwinian criticism.[145]

Our aims are of a positive character, they desiderate construction

and only use destruction where it is not to be avoided. So I shall

only mention that dogmatic Darwinism has been found to be unable to

explain every kind of mutual adaptations, *e.g.* those existing between

plants and insects; that it can never account for the origin of those

properties that are indifferent to the life of their bearer, being mere

features of organisation as an arrangement of parts; that it fails in

the face of all portions of organisation which are composed of many

different parts--like the eye--and nevertheless are functional units

in any passive or active way; and that, last not least, it has been

found to be quite inadequate to explain the first origin of all newly

formed constituents of organisation even if they are not indifferent:

for how could any rudiment of an organ, which is not functioning at all,

not only be useful to its bearer, but be useful in such a degree as to

decide about life or death?

[145] See Wigand, *Der Darwinismus und die Naturforschung Newton’s und

Cuvier’s*, Braunschweig, 1874-7; Nägeli, *Mechanisch-physiologische

Theorie der Abstammungslehre*, München, 1884; G. Wolff, *Beiträge zur

Kritik der Darwin’schen Lehre*, 2nd ed. Leipzig, 1898; etc.

It is only for one special feature that I should like to show, by

a more full analysis, that dogmatic Darwinism does not satisfy the

requirements of the case. The special strength of Darwinism is said to

lie in its explaining everything that is useful in and for organisms;

the competitive factor it introduces does indeed seem to secure at least

a relative sort of adaptedness between the organism and its needs. But

in spite of that, we shall now see that Darwinism fails absolutely to

explain those most intimate organic phenomena which may be said to be

the most useful of all.

Darwinism in its dogmatic form is not able to explain the origin of any

sort of organic restitution; it is altogether impossible to account for

the restitutive power of organisms by the simple means of fluctuating

variation and natural selection in the struggle for existence. Here we

have the logical *experimentum crucis* of Darwinism.

Let us try to study in the Darwinian style the origin of the

regenerative faculty, as shown in the restitution of the leg of a

newt. All individuals of a given species of the newt, say *Triton

taeniatus*, are endowed with this faculty; all of them therefore must

have originated from ancestors which acquired it at some time or other.

But this necessary supposition implies that all of these ancestors must

have lost their legs in some way, and not only one, but all four of

them, as they could not have acquired the restitutive faculty otherwise.

We are thus met at the very beginning of our argument by what must be

called a real absurdity, which is hardly lessened by the assumption that

regeneration was acquired not by all four legs together, but by one

after the other. But it is absolutely inevitable to assume that *all*

the ancestors of our *Triton* must have lost one leg, or more correctly,

that only those of them survived which had lost one! Otherwise not all

newts at the present day could possess the faculty of regeneration! But

a second absurdity follows the first one; out of the ancestors of our

newt, which survived the others by reason of having lost one of their

legs, there were selected only those which showed at least a very small

amount of healing of their wound. It must be granted that such a step

in the process of selection, taken by itself, would not at all seem to

be impossible; since healing of wounds protects the animals against

infection. But the process continues. In every succeeding stage of it

there must have survived only those individuals which formed just a

little more of granulative tissue than did the rest: though *neither*

they themselves *nor* the rest could use the leg, which indeed was

not present! That is the second absurdity we meet in our attempt at a

Darwinian explanation of the faculty of regeneration; but I believe the

first one alone was sufficient.

If we were to study the “selection” of the faculty of one of the

isolated blastomeres of the egg of the sea-urchin to form a whole larva

only of smaller size, the absurdities would increase. At the very

beginning we should encounter the absurdity, that of all the individuals

there survived only those which were not whole but half; for *all*

sea-urchins are capable of the ontogenetical restitution in question,

*all* of their ancestors therefore must have acquired it, and they

could do that only *if* they became halved at first by some accident

during early embryology. But we shall not insist any further on this

instance, for it would not be fair to turn into ridicule a theory which

bears the name of a man who is not at all responsible for its dogmatic

form. Indeed, we are speaking against Darwinism of the most dogmatic

form only, not against Darwin himself. He never analysed the phenomena

of regeneration or of embryonic restitution--they lay in a field very

unfamiliar to him and to his time. I venture to say that if he had taken

them into consideration, he would have agreed with us in stating that

his theory was not at all able to cover them; for he was prepared to

make great concessions, to Lamarckism for instance, in other branches of

biology, and he did not pretend, to know what life itself is.

Darwin was not a decided materialist, though materialism has made

great capital out of his doctrines, especially in Germany. His book,

as is well known, is entitled “The Origin of *Species*,” that is of

organic *diversities*, and he himself possibly might have regarded all

restitution as belonging to the original properties of life, anterior to

the originating of diversities. Personally he might possibly be called

even a vitalist. Thus dogmatic “Darwinism” in fact is driven into all

the absurdities mentioned above, whilst the “doctrine of Darwin” can

only be said to be wrong on account of its failing to explain mutual

adaptation, the origin of new organs, and some other features in organic

diversities; the original properties of life were left unexplained by it

intentionally.

DARWINISM FAILS ALL ALONG THE LINE

The result of our discussion then must be this: selection has proved to

be a negative factor only, and fluctuating variation as the only way

in which new properties of the organisms might have arisen has proved

to fail in the most marked manner, except perhaps for a few merely

quantitative instances. Such a result betokens the complete collapse

of dogmatic Darwinism as a general theory of descent: the most typical

features of all organisms remain as unexplained as ever.

What then shall we put in the place of pure Darwinism? Let us first try

a method of explanation which was also adopted occasionally by Darwin

himself: let us study that form of transformation theories which is

commonly known under the title of Lamarckism.

3. THE PRINCIPLES OF LAMARCKISM.

As the word “Darwinism” does not signify the proper theoretical system

of Charles Darwin, so Lamarckism as commonly understood nowadays is a

good deal removed from the original views of Jean Baptiste Lamarck.

Lamarckism is generally regarded as reducing all organic diversities to

differences in the needs of individual life, but Lamarck himself, as

must be emphasised from the very beginning, did not at all maintain the

opinion that the great characteristics of the types were only due to

such accidental factors. He supposed a sort of law of organisation to

be at the root of systematics, as developed in history, and the needs

of life were only responsible, according to him, for splitting the

given types of organisation into their ultimate branches. Thus Lamarck,

to a great extent at any rate, belongs to a group of authors that we

shall have to study afterwards: authors who regard an unknown law of

phylogenetic development as the real basis of transformism. Modern

so-called Neo-Lamarckism, on the other hand, has indeed conceded the

principle of needs to be the sole principle of transformism. Let us then

study Lamarckism in its dogmatic modern form.

ADAPTATION AS THE STARTING-POINT

All facts of morphological adaptations--facts which we have analysed

already from a different point of view, as being among the most typical

phenomena of organic regulation--form the starting-point of this

theory, and it must be granted that they form a very solid foundation,

for they are facts. The theory only has to enlarge hypothetically the

realm of these facts, or rather the realm of the law that governs them.

Indeed, it is assumed by Lamarckism that the organism is endowed with

the faculty of responding to *any* change of the environment which may

change its function by a morphologically expressed alteration of its

functional state and form, which is adapted to the state of conditions

imposed from without. Of course, as stated in this most general form,

the assumption is not true, but it is true within certain limits, as

we know; and there seems to be no reason why we should not believe

that there are many more cases of adaptation than we actually know at

present, or that, in former phylogenetic times, the organisms were more

capable of active adaptation than they are now. So to a certain extent,

at least, Lamarckism can be said to rest upon a *causa vera*.

It is important to notice that this *causa vera* would imply vitalistic

causality when taken in the wide meaning which Lamarckism allows to

it: indeed, the power of active adaptation to indefinite changes would

imply a sort of causal connection that is nowhere known except in the

organism. Lamarck himself is not very clear about this point, he seems

to be afraid of certain types of uncritical vitalism in vogue in his

days; but modern writers have most clearly seen what the logical

assumptions of pure Lamarckism are. Next to Cope, August Pauly[146] may

be said to be the most conscious representative of a sort of so-called

psychological vitalism, which indeed Lamarckism as a general and

all-embracing theory must have as its basis.

[146] *Darwinismus und Lamarckismus*, München, 1905.

THE ACTIVE STORING OF CONTINGENT VARIATIONS AS A HYPOTHETIC PRINCIPLE

This point will come out more fully, if now we turn to study a certain

group of principles, upon which dogmatic Lamarckism rests: I say

principles and not facts, for there are no facts but only hypothetic

assumptions in this group of statements. We do know a little about

adaptations, at least to a certain extent, and it was only about the

sphere of the validity of a law, which was known to be at work in

certain cases, that hypothetical additions were made. In the second

group of the foundations of Lamarckism we know absolutely nothing;

accidental variations of form are supposed to occur, and the organism is

said to possess the faculty of keeping and storing these variations and

of handing them down to the next generation, if they happen to satisfy

any of its needs.

But these needs are not of the actual type, brought forth by a change of

the functional state of the individual, as in the case of adaptations:

they are of a somewhat mysterious nature. A glance at the theory of the

origin of the movements which are called acts of volition in the human

child may serve to elucidate what is meant.

Acts of volition are said thus to originate in random movements of the

new-born infant: certain of these accidental motions which happen to

relieve some pain or to afford some pleasure are “remembered,” and are

used another time quite consciously to bring forth what is liked or to

remove what is disliked. So much for the present on a very difficult

subject, which will occupy us next year at much greater length. It is

clear that at least three fundamental phenomena are concerned in this

theory of the origin of acts of volition: the liking and disliking, the

keeping in mind, and the volition itself. The real act of volition,

indeed, is always based upon a connection of all these factors, these

factors now being connected in such a way that even their kind of

connection may be said to be a fourth fundamental principle. In order

that the particular effect may be obtained which is wanted because it is

liked, the possible ways leading to it, which appeared among the random

movements in the very beginning, are now regarded as “means” and may now

be said to be “used.” But that is as much as to say that the “means”

are judged with respect to their usefulness for the actual purpose, and

therefore *judgment* is the fourth foundation of the act of volition.

In fact, Pauly does not hesitate to attribute judgment, along with the

other psychological elements, to the organisms whilst undergoing their

transformation. There has been formed, for instance, by accidental

variation some pigment which by its chemical nature brings the organism

into a closer connection with the light of the medium; the individual

likes that, keeps the pigment for itself and produces it again in the

next generation; and indeed it will safeguard any sort of improvement

which chance may effect in this primitive “eye.” Such a view is said

to hold well with respect to the origin of every new organ. And this

psychological argument is also said to afford the real explanation of

adaptation proper. Adaptation also is regarded not as a truly primary

faculty of the organism, but as a retention or provoking of metabolic

states which occurred by accident originally and were then found to

be useful; now they are reproduced either in every single case of

individual morphogenesis, without regard to actual requirements, or

else only in response to such: in the first case they are “inherited,”

in the second they only occur as regulations. Thus the process of

judgment, together with all the other elemental factors of psychical

life concerned in it, has been made to account for adaptation proper.

The whole theory has accordingly become very uniform and simple.

CRITICISM OF THE “INHERITANCE OF ACQUIRED CHARACTERS” ASSUMED BY

LAMARCKISM

In addressing ourselves to the criticism of Neo-Lamarckism we shall

neglect as far as possible all the different psychological principles

concerned in it--which in any case would need rather a great amount of

epistemological sifting--and shall keep to those hypothetic facts which

are supposed to be such as may be actually observed in nature.

All of you know that the so-called inheritance of acquired characters

lies at the root of Lamarckism; and from this hypothesis our critical

analysis is to start, disregarding a larger or smaller number of

psychological principles that are brought into the field.

The name of “acquired characters” may *a priori* be given to three

different types of phenomena: firstly, variations including mutations;

secondly, disease or injuries; and thirdly, the results of the actual

process of adaptation of every kind.

In the first of these groups, the true problem of the inheritance of

“acquired” characters appears only with certain restrictions. All

variations and mutations are indeed “acquired” by one generation so

far as the earlier generation did not possess them, but mutations, at

least, cannot be said to be acquired by the actual adult personality:

they are innate in it from its very beginning, and therefore may better

be called congenital.[147] Congenital properties of the mutation type

are, in fact, known to be inherited: their inheritance does not present

any problem of its own, but is included in the changes of the hereditary

condition to which they are due altogether.[148] All properties of the

variation type, on the other hand, having been studied statistically,

are known to be inherited, to a certain small extent, as we have seen

already whilst studying Darwinism, though they are possibly always

liable to reversion. Modern science, as we know,[149] regards them as

due to changes of nutrition, in the most general meaning of the word.

Under such a view variations might indeed be said to belong to the

acquired group of organic specifications; their inheritance, as will

be seen later on, would hardly be quite a pure instance of what we are

searching for. In no case can true variations claim to be of great

importance in problems of transformism.

[147] This would not be true, if the varieties of plants produced by

Blaringhem, Klebs, and MacDougal by means of *external* agents were

really “mutations” (comp. page 238, note 3).

[148] Of course, the inheritance of mutations would imply a certain sort

of “inheritance of acquired characters,” on the condition stated in the

preceding note. But, probably, the germs of the next generation might

be regarded here as being directly affected by the external agent, in a

manner that will briefly be mentioned later on in the text.

[149] Comp. page 238, note 2.

But what is known about the inheritance of those properties which

beyond any doubt may be said to have originated in the adult individual

as such, and of which lesions and adaptations proper, as shown for

instance among amphibious plants, are instances of the two most

typical groups?[150] Weismann did good service by putting an end to

the scientific credulity which prevailed with regard to this subject.

Weismann was led by his theory of the germ plasm to deny the inheritance

of acquired characters of the typical kinds. He could not imagine how

the effect of any agent upon the adult, be it of the merely passive or

of the adaptive kind, could have such an influence upon the germ as to

force it to produce the same effect in spite of the absence of that

agent. In fact, that is what the inheritance of acquired characters

would render necessary, and a very strange phenomenon it would be,

no doubt. But, of course, taken alone, it could never be a decisive

argument against such inheritance. I fully agree, that science is

obliged to explain new facts by what is known already, as long as it is

possible; but if it is no longer possible, the theory of course has to

be changed, and not the facts. On this principle one would not neglect

the fact of an inheritance of acquired properties, but on the contrary

one perhaps might use it as a new evidence of vitalism.

[150] Certain English authors have applied the term “modification” to

all kinds of organic properties acquired from without, whether they are

adapted or not.

But are there any facts?

At this point we come to speak about the second group of Weismann’s

reasonings. He not only saw the difficulty of understanding inheritance

of acquired characters on the principles of the science of his time,

but he also criticised the supposed facts; and scarcely any of them

stood the test of his criticism. Indeed, it must fairly be granted that

not one case is known which really proves the inheritance of acquired

characters, and that injuries certainly are never found to be inherited.

In spite of that, I do not believe that we are entitled to deny the

possibility of the inheritance of a certain group of acquired characters

in an absolute and dogmatic manner, for there are a few facts which seem

at least to tend in the direction of such an inheritance, and which seem

to show that it might be discovered perhaps one day, if the experimental

conditions were changed.

I am not referring here to the few cases in which bacteria were made

colourless or non-virulent by outside factors, or in which certain

fungi were forced to permanent agamic reproduction by abnormal external

conditions and were shown to retain their “acquired properties”

after the external conditions had been restored. In these cases only

reproduction by simple division occurred, and that does not imply the

true problem of inheritance. Nor am I referring to the few cases of

non-adaptive “modifications” found by Standfuss and Fischer, in which

butterflies that had assumed an abnormal kind of pigmentation under

the influence of abnormal temperature acting upon the pupa, were seen

to form this same kind of pigmentation in the next generation under

normal conditions of temperature. These cases, though important in

themselves, are capable perhaps of a rather simple explanation, as in

fact has been suggested. Some necessary means both of inheritance and of

morphogenesis, the former being present in the propagation cells, may

be said to have been changed or destroyed by heat, and therefore, what

seems to be inherited after the change of the body only, would actually

be the effect of a direct influence of the temperature upon the germ

itself.[151] Let me be clearly understood: I do not say that it is so,

but it may be so. What seems to me to be more important than everything

and to have a direct bearing on the real discovery of the inheritance

of acquired characters in the future, is this. In some instances

plants which had been forced from without to undergo certain typical

morphological adaptations, or at least changes through many generations,

though they did not keep the acquired characters permanently in spite

of the conditions being changed to another type, were yet found to lose

the acquired adaptations not suddenly but only in the course of three

or more generations. A certain fern, *Adiantum*, is known to assume a

very typical modification of form and structure, if grown on serpentine;

now Sadebeck,[152] while cultivating this serpentine modification of

*Adiantum* on ordinary ground, found that the first generation grown in

the ordinary conditions loses only a little of its typical serpentine

character, and that the next generation loses a little more, so that

it is not before the fifth generation that all the characters of the

serpentine modification have disappeared. There are a few more cases

of a similar type relating to plants grown in the plains or on the

mountains. There also it was found to take time, or rather to take the

course of *several* generations, until what was required by the new

conditions was reached. Of course these cases are very very few compared

with those in which a *sudden* change of the adaptive character,

corresponding to the actual conditions, sets in; but it is enough that

they do exist.

[151] Of course the inheritance of specific values from the results of

fluctuating variations, leading to new averages of variability (see

p. 265), may also be understood in this manner, the conditions of

nourishment acting upon the adult and upon its germs equally well.

[152] *Berichte üb. d. Sitzung. d. Ges. f. Bot.*, Hamburg, 1887, 3 Heft.

Would it not be possible at least that adaptations which last

for thousands of generations or more might in fact change the

adaptive character into a congenital one? Then we not only should

have inheritance of acquired characters, but should have a sort

of explanation at the same time for the remarkable fact that

certain histological structures of a very adapted kind are formed

ontogenetically before any function exists, as is known to be the

case with the structures in the bones of vertebrates, for instance.

Experiments are going on at Paris, and perhaps in other places of

scientific research also, which, it is hoped, will show that animals

reared in absolute darkness for many generations will lose their

perfectly formed eyes, and that animals from the dark with very

rudimentary eyes will be endowed with properly functioning ones, after

they have been reared in the light for generations. Such a result indeed

would account for the many animals, of the most different groups, which

live in dark caves and possess only rudiments of eyes: functional

adaptation is no longer necessary, so-called atrophy by inactivity sets

in, and the results “acquired” by it are inherited.[153]

[153] Quite recently Kammerer (*Arch. Entw. Mech.* 25, 1907, p. 7) has

published very important experiments on the inheritance of “acquired”

modifications with regard to the peculiarities of reproduction in

*Salamandra atra* and *S. maculosa*. It seems rather improbable--though

not absolutely impossible--that the germ cells were directly affected by

the external modifying agent in this case.

But enough of possibilities. Let us be content at present to know at

least a few real instances with regard to the slowness of the process of

what might be said to be “re-adaptation” in some plants. This process

shows us a way by which our problem may some day be solved; it allows

us to introduce inheritance of acquired characters as a legitimate

hypothesis at least, which not only will explain many of the diversities

in systematics historically, but also can be called, though not a *causa

vera*, yet certainly more than a mere fiction.

OTHER PRINCIPLES WANTED

We have only dealt with the probability of the inheritance of

morphological or physiological[154] adaptation. If that could really be

considered as one of the factors concerned in the theory of descent,

many, if not all of those congenital diversities among organic species

which are of the type of a true structural correspondence to their

future functional life, might be regarded as explained, that is,

as reduced to one and the same principle. But nothing more than an

explanation of *this* kind of diversities is effected by our principle,

and very much more remains to be done, for organic diversities not only

consist in specifications and differences as to histology, but are to a

much more important degree, differences of organisation proper, that is,

of the arrangement of parts, in the widest sense of the word.[155]

[154] We have not spoken about the hypothetic inheritance of pure

physiological adaptations, for it is clear without further discussion

that innate specific immunity, for instance, being a specific

“adaptedness” (*see* p. 186) *might* be due to the inheritance of the

results of active immunity as an adaptation, just as adaptive congenital

structures *might* be due to such an inheritance.

[155] C. E. v. Baer clearly discriminated between the type, the degree

of organisation, and the histological structure. All these three topics

indeed have to be taken into account separately; the third alone is of

the adaptive type. All of them may be independent of each other: the

Amoeba may be as adapted histologically as is a high vertebrate, but it

is of much lower type; and in its own type it is of a lower degree of

organisation than Radiolaria are.

Would it be possible to interpret the origin of this sort of systematic

diversities by a reasoning similar to that by which we have understood,

at least hypothetically, congenital adaptedness?

Dogmatic Lamarckism, we know, uses two principles as its foundations;

one of them, adaptation and its inheritance, we have studied with what

may be called a partly positive result. The other is the supposed

faculty of the organism to keep, to store, and to transfer those

variations or mutations of a not properly adaptive sort which, though

originating by chance, happen to satisfy some needs of the organism.

CRITICISM OF THE HYPOTHESIS OF STORING AND HANDING DOWN CONTINGENT

VARIATIONS

Strange to say, this second hypothesis of dogmatic Lamarckism, invented

with the express purpose of defeating Darwinism and taking the place of

its fluctuating variability, which was found not to do justice to the

facts--this second hypothesis of dogmatic Lamarckism is liable to just

the same objections as dogmatic Darwinism itself.

As it is important to understand well the real logical nature of

our objections to both of the great transformistic theories, we

think it well to interrupt our argument for a moment, in order to

consider a certain point which, though very important in itself,

seems of only secondary importance to us in our present discussion.

Dogmatic Darwinism--I do not say the doctrine of Charles Darwin--is

materialistic at bottom, and indeed has been used by many to complete

their materialistic view of the universe on its organic side. The word

“materialism” must not necessarily be taken here in its metaphysical

sense, though most materialists are dogmatic metaphysicians. It also

can be understood as forming part of a phenomenological point of view.

Materialism as a doctrine of science means simply this: that whether

“nature” be reality or phenomenon, in any case there is but one ultimate

principle at its base, a principle relating to the movements of

particles of matter. It is this point of view which dogmatic Darwinism

strengthens; on the theory of natural selection and fluctuating

variations, due to accidental differences of nutrition, organisms are

merely arrangements of particles of matter, nothing else; and moreover,

their kinds of arrangement are understood, at least in principle.

Lamarckism, on the other hand, is not materialistic, but most markedly

vitalistic--psychistic even; it takes life for granted when it begins

its explanations.

You may tell me that Darwin did the same, that he expressly states that

his theory has nothing to do with the origin of life; that the title

of his work is “The Origin of *Species*.” It would certainly be right

to say so, at least with reference to Darwin personally; but in spite

of that, it must be granted that Darwin’s doctrine contains a certain

germ of materialism which has been fully developed by the Darwinian

dogmatists, while Lamarckism is antimaterialistic by its very nature.

Now it is very important, I think, to notice that this difference

between the two theories is unable to disguise one main point which is

common to both: and it is to this point, and to this point only, that

our chief objections against both these theories converge at present.

The *contingency* of the typical organic form is maintained by Darwinism

as well as by Lamarckism: both theories, therefore, break down for

almost the same reasons. The term “contingency” can signify very

different relations, having but little in common; but it is sufficient

for our present purpose to observe that there may be distinguished

roughly two main classes of contingencies, which may provisionally be

called the “contingency of being,” and the “contingency of occurring.”

It is with the contingency of being that criticism of Darwinism and

Lamarckism of the dogmatic type has to deal. Darwinism dealt with

variations occurring at random; the organic form was the result of

a fixation of only one kind of such variations, all others being

extinguished by selection. In other terms, the specific organised

form, as understood by Darwinism, was a unit only to the extent that

all its properties related to one and the same body, but for the rest

it was a mere aggregation or summation. It may be objected to this

statement, that by being inherited in its specificity the Darwinian

form proved to be a unit in a higher sense of the word, even in the

opinion of dogmatic Darwinians; and this objection, perhaps, holds good

as far as inheritance is concerned. But on the other hand, it must

never be forgotten that the word “unit” had quite a vague and empty

meaning even then, as indeed everything the organism is made up of

is regarded as being in itself due to a contingent primary process,

which has no relation to its fellow-processes. “Unit,” indeed, in spite

of inheritance--which, by the way, is alleged also to be a merely

materialistic event--means to Darwinians no more when applied to the

organism than it does when applied to mountains or islands, where of

course a sort of “unit” also exists in some sense, as far as one and the

same body comes into account, but where every single character of this

unit, in every single feature of form or of quality, is the result of

factors or agents each of which is independent of every other.

To this sort of contingency of being, as maintained by Darwinians,

criticism has objected, as we know, that it is quite an impossible

basis of a theory of descent, since it would explain neither the first

origin of an organ, nor any sort of harmony among parts or among whole

individuals, nor any sort of restitution processes.

Now Lamarckism of the dogmatic kind, as will easily be seen, only

differs from Darwinism in this respect, that what according to the

latter happens to the organism passively by means of selection, is

according to the former performed actively by the organism, by means of

a “judgment”--by the retention and handing down of chance variations.

The specificity of the form as a whole is contingent also according

to Lamarckism. And, indeed, criticism must reject this contingency of

being in exactly the same way as it rejected the contingency of form

maintained by Darwinians.

As far as the inheritance of truly adaptive characters comes into

account--that is, the inheritance of characters which are due to the

active faculty of adaptation possessed by the organism, bearing a

vitalistic aspect throughout--hardly anything could be said against

Lamarckism, except that inheritance of acquired characters is still

an hypothesis of small and doubtful value at present. But that

*specific organisation proper* is due to *contingent* variations, which

accidentally have been found to satisfy some needs of the individual and

therefore have been maintained and handed down, this reasoning is quite

an impossibility of exactly the same kind as the argument of Darwinism.

The process of restitution, perfect the very first time it occurs, if it

occurs at all, is again the classical instance against this new sort of

contingency, which is assumed to be the basis of transformism. Here we

see with our eyes that the organism can do more than simply perpetuate

variations that have occurred at random and bear in themselves no

relation whatever to any sort of unit or totality. There *exists* a

faculty of a certain higher degree in the organism, and this faculty

cannot possibly have originated by the process which Lamarckians[156]

assume. But if their principle fails in one instance, it fails as a

*general* theory altogether. And now, on the other hand, as we actually

see the individual organism endowed with a morphogenetic power,

inexplicable by Lamarckism, but far exceeding the organogenetic faculty

assumed by that theory, would it not be most reasonable to conclude from

such facts, that there exists a certain organising power at the root of

the transformism of species also, a power which we do not understand,

which we see only partially manifested in the work of restitutions,

but which certainly is not even touched by any of the Lamarckian

arguments? There does indeed exist what Gustav Wolff has called primary

purposefulness (“primäre Zweckmässigkeit”), at least in restitutions,

and this is equally unexplainable by Darwinism and by the dogmatism of

the Lamarckians.

[156] I repeat once more that we are dealing here with dogmatic

“Neo-”Lamarckism exclusively. This theory indeed claims to explain *all*

features and properties of organic bodies on the basis of the feeling

of needs and storing of contingent fulfilments and on this basis alone,

just as dogmatic “Neo”-Darwinism claims to account for *all* those

phenomena on the ground of contingent variations and natural selection.

Darwin himself, as we have seen, intentionally left unexplained certain

primary features of life and therefore cannot be blamed for having

failed to explain them, though even then his theory remains wrong.

Lamarck personally considered a real primary organisatory law of

phylogeny as being of fundamental importance, and therefore he is not in

the least responsible if “Neo-Lamarckism” fails as a universal theory.

But before entering into this area of hypothesis, let us mention a few

more objections to be made to the theory of the contingency of form as

put forward by Lamarckians. In the first place, let us say a few words

about the appropriateness of the term “contingency” as used in this

connection. The forms are regarded as contingent by Lamarckians inasmuch

as the variations which afterwards serve as “means” to the “needs” of

the organism occur quite accidentally with regard to the whole organism.

It might be said that these “needs” are not contingent but subject

to an inherent destiny, but this plea is excluded by the Lamarckians

themselves, when they say that the organism experiences no need until it

has enjoyed the accidental fulfilment of the same. So the only thing in

Lamarckian transformism which is not of a contingent character would be

the psychological agent concerned in it, as being an agent endowed with

the primary power of feeling needs after it has felt fulfilment, and of

judging about what the means of future fulfilment are, in order to keep

them whenever they offer. But these are characteristics of life itself,

irrespective of all its specific forms, which alone are concerned in

transformism. Now indeed, I think, we see as clearly as possible that

Darwinism and Lamarckism, in spite of the great contrast of materialism

and psychologism, shake hands on the common ground of the contingency of

organic forms.

The whole anti-Darwinistic criticism therefore of Gustav Wolff for

instance, may also be applied to Lamarckism with only a few changes

of words. How could the origin of so complete an organ as the eye of

vertebrates be due to contingent variations? How could that account for

the harmony of the different kinds of cells in this very complicated

organ with each other and with parts of the brain? And how is it to be

understood, on the assumption of contingency, that there are two eyes of

almost equal perfection, and that there are two feet, two ears? Islands

and mountains do not show such symmetry in *their* structures.

We shall not repeat our deduction of the origin of restitutions, of

regeneration for instance, on the dogmatic Lamarckian theory. As we

have said already, it would lead to absurdities as great as in the

case of dogmatic Darwinism, and indeed we already have mentioned that

Lamarckians would hardly even attempt to explain these phenomena.

It follows that dogmatic Lamarckism fails as a general theory about

form.[157]

[157] Compare also the excellent criticism of Lamarckism lately given by

G. Wolff, *Die Begründung der Abstammungslehre*, München, 1907.

There is finally one group of facts often brought forward against

Lamarckism by Darwinian authors[158] which may be called the logical

*experimentum crucis* of this doctrine, an *experimentum* destined

to prove fatal. You know that among the polymorphic groups of bees,

termites, and ants, there exists one type of individuals, or even

several types, endowed with some very typical features of organisation,

but at the same time absolutely excluded from reproduction: how could

those morphological types have originated on the plan allowed by the

Lamarckians? Of what use would “judgment” about means that are offered

by chance and happen to satisfy needs, be to individuals which die

without offspring? Here Lamarckism becomes a simple absurdity, just as

Darwinism resulted in absurdities elsewhere.

[158] It has also very often been said by Darwinians that Lamarckism is

only able to explain those cases of adaptedness which relate to active

functioning but not mere passive adapted characters, like “mimicry” for

example. But this argument *taken by itself*, it seems to me, would not

be fatal to Neo-Lamarckism in the special form August Pauly gave to this

doctrine.

We were speaking about dogmatic Darwinism then, and it is about dogmatic

Lamarckism that we are reasoning at present; both theories must fall in

their dogmatic form, though a small part of both can be said to stand

criticism. But these two parts which survive criticism, one offered by

Lamarck, the other by Darwin, are far from being a complete theory of

transformism, even if taken together: they only cover a small area of

the field concerned in the theory of descent. Almost everything is still

to be done, and we may here formulate, briefly at least, what we expect

to be accomplished by the science of the future.

4. THE REAL RESULTS AND THE UNSOLVED PROBLEMS OF TRANSFORMISM

What has been explained to a certain extent by the two great theories

now current is only this. Systematic diversities consisting in mere

differences as to intensity or number may perhaps owe their origin to

ordinary variation. They may at least, if we are entitled to assume

that heredity in some cases is able to hand on such variations without

reversion, which, it must be again remarked, is by no means proved by

the facts at present. Natural selection may share in this process by

eliminating all those individuals that do not show the character which

happens to be useful. That is the Darwinian part of an explanation of

transformism which may be conceded as an hypothesis. On the other side,

congenital histological adaptedness may be regarded hypothetically as

due to an inheritance of adaptive characters which had been acquired by

the organism’s activity, exerted during a great number of generations.

That is the Lamarckian part in the theory of descent.

But nothing more is contributed to this theory either by the doctrine

of Darwin or by that of Lamarck. So it follows that almost everything

has still to be done; for no hypothesis at present accounts for

the foundation of all systematics, viz., for the differences in

organisation, in all that relates to the so-called types as such and

the degree of complication in these types, both of which (types and

degree of complication) are independent of histological adaptation and

adaptedness.

What then do we know about any facts that might be said to bear on

this problem? We have stated already at the end of our chapter devoted

to the analysis of heredity that what we actually know about any

deviation of inheritance proper, that is, about congenital differences

between the parents and the offspring, relating to mere tectonics,

is practically nothing: indeed, there are at our disposal only the

few facts observed by de Vries or derived from the experience of

horticulturalists and breeders. We may admit that these facts at

least prove the possibility[159] of a discontinuous variation, that

is of “mutation,” following certain lines of tectonics and leading to

*constant* results; but everything else, that is everything about a real

theory of phylogeny, must be left to the taste of each author who writes

on the theory of the Living. You may call that a very unscientific state

of affairs, but no other is possible.

[159] But nothing more. All “mutations” hitherto observed in nature

or (comp. page 238, note 3) experimentally produced relate only to

“varieties” and not to “species.” One could hardly say that the

recent investigations about the production of mutations by *external*

means have strengthened their importance for the general theory of

transformism.

And, in fact, it has been admitted by almost all who have dealt with

transformism without prepossessions that such is the state of affairs.

Lamarck himself, as we have mentioned already, was not blind to the fact

that a sort of organisatory law must be at the base of all transformism,

and it is well known that hypothetical statements about an original law

of phylogeny have been attempted by Nägeli, Kölliker, Wigand, Eimer,

and many others. But a full discussion of all these “laws” would hardly

help us much in our theoretical endeavour, as all of them, it must be

confessed, do little more than state the mere fact that some unknown

principle of organisation must have been at work in phylogeny, if we are

to accept the theory of descent at all.

It is important to notice that even such a convinced Darwinian as

Wallace, who is well known to have been an independent discoverer of the

elimination principle, admitted an exception to this principle in at

least one case--with regard to the origin of man. But one exception of

course destroys the generality of a principle.

As we ourselves feel absolutely incapable of adding anything specific

to the general statement that there *must* be an unknown principle of

transformism, if the hypothesis of descent is justified at all, we may

here close our discussion of the subject.

5. THE LOGICAL VALUE OF THE ORGANIC FORM ACCORDING TO THE DIFFERENT

TRANSFORMISTIC THEORIES

A few words only must be added about two topics: on the character of

organic forms as regarded by the different transformistic theories, and

on the relation of transformism in general to our concept of entelechy.

We have learnt that both Darwinism and Lamarckism, in their dogmatic

shape, regard the specific forms of animals and plants as being

contingent; in fact, it was to this contingency that criticism was

mainly directed. We therefore are entitled to say that to Darwinism and

Lamarckism organic forms are accidental in the very sense of the *forma

accidentalis* of the old logicians. There are indefinite forms possible,

according to these theories, and there is no law relating to these

forms. Systematics, under such a view, must lose, of course, any really

fundamental importance. “There is no rational system about organisms”:

that is the ultimate statement of Darwinism and of Lamarckism on this

doubtful question. Systematics is a mere catalogue, not at present

only, but for ever, by the very nature of the organisms. It is not

owing to the indefinite number of possible forms that both our theories

came to deny the importance of systematics, but to the want of a *law*

relating to this indefinite number: among chemical compounds indefinite

possibilities also exist in some cases, but they obey the law of the

general formula. It is very strange that Darwinians of all people are

in the forefront of systematic research in all countries: do they not

see that what they are trying to build up can only relate to accidental

phenomena? Or have they some doubts about the foundations of their own

theoretical views, in spite of the dogmatic air with which they defend

them? Or is it the so-called historical interest which attracts them?

A new question seems to arise at this point: Have not we ourselves

neglected history in favour of systematics and laws? Our next lecture,

the last of this year, will give the answer to this question.

At present we continue our study of the possible aspects of systematics.

It is not difficult to find out what meaning organic forms would assume

under any phylogenetic theory opposed to the theories of contingency.

It was their defence of contingency, that is, their lack of any law

of forms, that caused these theories to be overthrown--reduced to

absurdities even--and therefore, it follows that to assume any kind of

transformistic law is at the same time to deny the accidental character

of the forms of living beings.

There is no *forma accidentalis*. Does that mean that the *forma

essentialis* is introduced by this mere statement? And what would *that*

assert about the character of systematics?

THE ORGANIC FORM AND ENTELECHY

This problem is not as simple as it might seem to be at the first

glance, and, in fact, it is insoluble at present. It is here that the

relation of the hypothetic transformistic principle to our concept of

entelechy is concerned.

We know that entelechy, though not material in itself, uses material

means in each individual morphogenesis, handed down by the material

continuity in inheritance. What then undergoes change in phylogeny,

the means or the entelechy? And what would be the logical aspect of

systematics in either case?

Of course there would be a law in systematics in any case; and therefore

systematics in any case would be rational in principle. But if the

transformistic factor were connected with the means of morphogenesis,

one could hardly say that specific form as such was a primary essence.

Entelechy would be that essence, but entelechy in its generality and

always remaining the same in its most intimate character, as the

specific diversities would only be due to a something, which is not

form, but simply means to form. But the *harmony* revealed to us in

every typical morphogenesis, be it normal or be it regulatory, seems

to forbid us to connect transformism with the means of morphogenesis.

And therefore we shall close this discussion about the most problematic

phenomena of biology with the declaration, that we regard it as more

congruent to the general aspect of life to correlate the unknown

principle concerned in descent with entelechy itself, and not with

its means. Systematics of organisms therefore would be in fact

systematics of entelechies, and therefore organic forms would be

*formae essentiales*, entelechy being the very essence of form in its

specificity. Of course systematics would then be able to assume a truly

rational character at some future date: there might one day be found

a principle to account for the totality of possible[160] forms, a

principle based upon the analysis of entelechy.[161] As we have allowed

that Lamarckism hypothetically explains congenital adaptedness in

histology, and that Darwinism explains a few differences in quantity,

and as such properties, of course, would both be of a contingent

character, it follows that our future rational system would be combined

with certain accidental diversities. And so it might be said to be

one of the principal tasks of systematic biological science in the

future to discover the really rational system among a given totality of

diversities which cannot appear rational at the first glance, one sort

of differences, so to speak, being superimposed upon the other.

[160] The word “possible” relating to originating, of course, not to

surviving. It is here that natural selection may acquire its logical

importance alluded to above (see page 264).

[161] The discussions in the second volume of this book will show the

possible significance of such an analysis. We at present are dealing

with entelechy in a quasi-popular manner.

*C.* THE LOGIC OF HISTORY

History, in the strictest sense of the word, is the enumeration of the

things which have followed one another in order of time. History deals

with the single, with regard both to time and space. Even if its facts

are complex in themselves and proper to certain other kinds of human

study, they are nevertheless regarded by history as single. Facts, we

had better say, so far as they are regarded as single, are regarded

historically, for what relates to specific time and space is called

history.

Taken as a simple enumeration or registration, history, of course,

cannot claim to be a “science” unless we are prepared to denude that

word of all specific meaning. But that would hardly be useful. As a

matter of fact, what has actually claimed to be history, has always

been more than a mere enumeration, even in biology proper. So-called

phylogeny implies, as we have shown, that every one of its actual

forms contains some rational elements. Phylogeny always rests on

the assumption that only some of the characters of the organisms

were changed in transformism and that what remained unchanged may be

explained by the fact of inheritance.

But this, remember, was the utmost we were able to say for phylogeny.

It remains fantastic and for the most part unscientific in spite of

this small degree of rationality, as to which it is generally not very

clear itself. For nothing is known with regard to the positive factors

of transformism, and we were only able to offer the discussion of a few

possibilities in place of a real theory of the factors of descent.

In spite of that it will not be without a certain logical value to begin

our analysis of history in general by the discussion of possibilities

again. Biology proper would hardly allow us to do more: for the

simple “fact” of history is not even a “fact” in this science, but an

hypothesis, albeit one of some probability.

As discussions of mere possibilities should always rest on as broad a

basis as possible, we shall begin our analysis by raising two general

questions. To what kinds of realities may the concept of history

reasonably be applied? And what different types of “history” would be

possible *a priori*, if the word history is to signify more than a mere

enumeration?

1. THE POSSIBLE ASPECTS OF HISTORY

Of course, we could select one definite volume in space and call all the

consecutive stages which it goes through, its history: it then would be

part of its history that a cloud was formed in it, or that a bird passed

through it on the wing. But it would hardly be found very suggestive to

write the history of space-volumes. In fact, it is to *bodies* in space

that all history actually relates, at least indirectly, for even the

history of sciences is in some respect the history of men or of books.

It may suffice for our analysis to understand here the word body in its

popular sense.

Now in its relation to bodies history may have the three following

aspects, as far as anything more than a simple enumeration comes into

account. Firstly, it may relate to one and the same body, the term

body again to be understood popularly. So it is when the individual

history of the organism is traced from the egg to the adult, or when the

history of a cloud or of an island or of a volcano is written. Secondly,

the subject-matter of history may be formed by the single units of a

consecutive series of bodies following each other periodically. To this

variety of history the discoveries of Mendel and his followers would

belong in the strictest sense, but so does our hypothetical phylogeny

and a great part of the history of mankind. And lastly, there is a

rather complicated kind of sequence of which the “history” has actually

been written. History can refer to bodies which are in no direct

relation with one another, but which are each the effect of another body

that belongs to a consecutive series of body-units showing periodicity.

This sounds rather complicated; but it is only the strict expression

of what is perfectly familiar to you all. Our sentence indeed is

simply part of the definition of a history of art or of literature for

instance--or, say, of a phylogenetic history of the nests of birds. The

single pictures are the subjects of the history of art, and nobody would

deny that these pictures are the effects of their painters, and that

the painters are individuals of mankind--that is, that they are bodies

belonging to a consecutive series of body-units showing periodicity. Of

course, it is only improperly that we speak of a history of pictures or

of books or of nests. In fact, we are dealing with painters, and with

men of letters or of science, and with certain birds, and therefore

the third type of history may be reduced to the second. But it was not

without value to pursue our logical discrimination as far as possible.

So far we have always spoken of history as being more than a mere

enumeration, but we have not ascertained what this “more” signifies. It

is not very difficult to do so: in fact, there are three different types

of history, each of a different degree of importance with respect to the

understanding of reality.