THE HYPOTHESIS OF PERIODIC MUTATIONS

The prevailing belief that slow and gradual, nearly invisible changes

constitute the process of evolution in the animal and vegetable kingdom,

did not offer a strong stimulus for experimental research. No

appreciable response to any external agency was of course to be

expected. Responses were supposed to be produced, but the corresponding

outward changes would be too small to betray themselves to the

investigator.

The direct observation of the mutations of the evening-primrose has

changed the whole aspect of the problem at once. It is no longer a

matter dealing with purely hypothetical conditions. Instead of the vague

notions, uncertain hopes, and a priori conceptions, that have hitherto

confused the investigator, methods of observation have been formulated,

suitable for the attainment of definite results, the general nature of

which is already known.

To my mind the real value of the discovery [687] of the mutability of

the evening-primrose lies in its usefulness as a guide for further work.

The view that it might be an isolated case, lying outside of the usual

procedure of nature, can hardly be sustained. On such a supposition it

would be far too rare to be disclosed by the investigation of a small

number of plants from a limited area. Its appearance within the limited

field of inquiry of a single man would have been almost a miracle.

The assumption seems justified that analogous cases will be met with,

perhaps even in larger numbers, when similar methods of observation are

used in the investigation of plants of other regions. The mutable

condition may not be predicated of the evening-primroses alone. It must

be a universal phenomenon, although affecting a small proportion of the

inhabitants of any region at one time: perhaps not more than one in a

hundred species, or perhaps not more than one in a thousand, or even

fewer may be expected to exhibit it. The exact proportion is immaterial,

because the number of mutable instances among the many thousands of

species in existence must be far too large for all of them to be

submitted to close scrutiny.

It is evident from the above discussion that next in importance to the

discovery of the prototype of mutation is the formulation of methods

[688] for bringing additional instances to light. These methods may

direct effort toward two different modes of investigation. We may search

for mutable plants in nature, or we may hope to induce species to become

mutable by artificial methods. The first promises to yield results most

quickly, but the scope of the second is much greater and it may yield

results of far more importance. Indeed, if it should once become

possible to bring plants to mutate at our will and perhaps even in

arbitrarily chosen directions, there is no limit to the power we may

finally hope to gain over nature.

What is to guide us in this new line of work? Is it the minute

inspection of the features of the process in the case of the

evening-primroses? Or are we to base our hopes and our methods on

broader conceptions of nature's laws? Is it the systematic study of

species and varieties, and the biologic inquiry into their real

hereditary units? Or is the theory of descent to be our starting-point?

Are we to rest our conceptions on the experience of the breeder, or is

perhaps the geologic pedigree of all organic life to open to us better

prospects of success?

The answer to all such questions is a very simple one. All possibilities

must be considered, and no line of investigation ignored. For myself I

have based my field-researches and my [689] testing of native plants on

the hypothesis of unit-characters as deduced from Darwin's _Pangenesis_.

This conception led to the expectation of two different kinds of

variability, one slow and one sudden. The sudden ones known at the time

were considered as sports, and seemed limited to retrograde changes, or

to cases of minor importance. The idea that sudden steps might be taken

as the principal method of evolution could be derived from the

hypothesis of unit characters, but the evidence might be too remote for

a starting point for experimental investigation.

The success of my test has given proof to the contrary. Hence the

assertion that no evidence is to be considered as inadequate for the

purpose under discussion. Sometime a method of discovering, or of

producing, mutable plants may be found, but until this is done, all

facts of whatever nature or direction must be made use of. A very slight

indication may change forever the whole aspect of the problem.

The probabilities are now greatly in favor of our finding out the causes

of evolution by a close scrutiny of what really happens in nature. A

persistent study of the physiologic factors of this evolution is the

chief condition of success. To this study field-observations may

contribute as well as direct experiments, [690] microscopical

investigations as well as extended pedigree-cultures. The cooperation of

many workers is required to cover the field. Somewhere no doubt the

desired principle lies hidden, but until it is discovered, all methods

must be tried.

With this conception as the best starting point for further

investigation, we may now make a brief survey of the other phase of the

problem. We shall try to connect our observations on the

evening-primroses with the theory of descent at large.

We start with two main facts. One is the mutability of Lamarck's

primrose, and the second is the immutable condition of quite a number of

other species. Among them are some of its near allies, the common and

the small flowered evening-primrose, or _Oenothera biennis_ and _O.

muricata_.

From these facts, a very important question arises in connection with

the theory of descent. Is the mutability of our evening-primroses

temporary, or is it a permanent condition? A discussion of this problem

will give us the means of reaching a definite idea as to the scope of

our inquiries.

Let us consider the present first. If mutability is a permanent

condition, it has of course no beginning, and moreover is not due to the

[691] agency of external circumstances. Should this be granted for the

evening-primrose, it would have to be predicated for other species found

in a mutable state. Then, of course, it would be useless to investigate

the causes of mutability at large, and we should have to limit ourselves

to the testing of large numbers of plants in order to ascertain which

are mutable and which not.

If, on the other hand, mutability is not a permanent feature, it must

once have had a beginning, and this beginning itself must have had an

external cause. The amount of mutability and its possible directions may

be assumed to be due to internal causes. The determination of the moment

at which they will become active can never be the result of internal

causes. It must be assigned to some external factor, and as soon as this

is discovered the way for experimental investigation is open.

In the second place we must consider the past. On the supposition of

permanency all the ancestors of the evening-primrose must have been

mutable. By the alternative view mutability must have been a periodic

phenomenon, producing at times new qualities, and at other times leaving

the plants unchanged during long successions of generations. The present

mutable state must then have been preceded by an immutable [692]

condition, but of course thousands of mutations must have been required

to produce the evening-primroses from their most remote ancestors.

If we take the species into consideration that are not mutable at

present, we may ask how we are to harmonize them with each of the two

theories proposed. If mutability is permanent, it is manifest that the

whole pedigree of the animal and vegetable kingdom is to be considered

as built up of main mutable lines, and that the thousands of constant

species can only be taken to represent lateral branches of the

genealogic tree.

These lateral branches would have lost the capacity of mutating,

possessed by all their ancestors. And as the principle of the hypothesis

under discussion does not allow a resumption of this habit, they would

be doomed to eternal constancy until they finally die out. Loss of

mutability, under this conception, means loss of the capacity for all

further development. Only those lines of the main pedigree which have

retained this capacity would have a future; all others would die out

without any chance of progression.

If, on the other hand, mutability is not permanent, but a periodic

condition, all lines of the genealogic tree must be assumed to show

alternatively [693] mutating and constant species. Some lines may be

mutating at the present moment; others may momentarily be constant. The

mutating lines will probably sooner or later revert to the inactive

state, while the powers of development now dormant may then become

awakened on other branches.

The view of permanency represents life as being surrounded with

unavoidable death, the principle of periodicity, on the contrary,

follows the idea of resurrection, granting the possibility of future

progression for all living beings. At the same time it yields a more

hopeful prospect for experimental inquiry.

Experience must decide between the two main theories. It demonstrates

the existence of polymorphous genera, such as _Draba_ and _Viola_ and

hundreds of others. They clearly indicate a previous state of

mutability. Their systematic relation is exactly what would be expected,

if they were the result of such a period. Perhaps mutability has not

wholly ceased in them, but, might be found to survive in some of their

members. Such very rich genera however, are not the rule, but are

exceptional cases, indicating the rarity of powerful mutative changes.

On the other hand, species may remain in a state of constancy during

long, apparently during indefinite, ages.

[694] Many facts plead in favor of the constancy of species. This

principle has always been recognized by systematists. Temporarily the

current form of the theory of natural selection has assumed species to

be inconstant, ever changing and continuously being improved and adapted

to the requirements of the life-conditions. The followers of the theory

of descent believed that this conclusion was unavoidable, and were

induced to deny the manifest fact that species are constant entities.

The mutation theory gives a clew to the final combination of the two

contending ideas. Reducing the changeability of the species to distinct

and probably short periods, it at once explains how the stability of

species perfectly agrees with the principle of descent through

modification.

On the other hand, the hypothesis of mutative periods is by no means

irreconcilable with the observed facts of constancy. Such casual changes

can be proved by observations such as those upon the evening-primrose,

but it is obvious that a disproof can never be given. The principle

grants the present constancy of the vast majority of living forms, and

only claims the exceptional occurrence of definite changes.

Proofs of the constancy of species have been given in different ways.

The high degree of similarity of the individuals of most of our [695]

species has never been denied. It is observed throughout extended

localities, and during long series of years. Other proofs are afforded

by those plants which have been transported to distant localities some

time since, but do not exhibit any change as a result of this migration.

Widely dispersed plants remain the same throughout their range, provided

that they belong to a single elementary species. Many species have been

introduced from America into Europe and have spread rapidly and widely.

The Canadian horsetail (_Erigeron canadensis_), the evening-primrose and

many other instances could be given. They have not developed any special

European features after their introduction. Though exposed to other

environmental conditions and to competition with other species, they

have not succeeded in developing a new character. Such species as proved

adequate to the new environment have succeeded, while those which did

not have succumbed.

Much farther back is the separation of the species which now live both

in arctic regions and on the summits of our highest mountaintops. If we

compare the alpine flora with the arctic plants, a high degree of

similarity at once strikes us. Some forms are quite identical; others

are slightly different, manifestly representing elementary species of

the same systematic [696] type. Still others are more distant or even

belong to different genera. The latter, and even the diverging, though

nearly allied, elementary species, do not yield adequate evidence in any

direction.

They may as well have lived together in the long ages before the

separation of the now widely distant floras, or have sprung from a

common ancestor living at that time, and subsequently have changed their

habits. After excluding these unreliable instances, a good number of

species remain, which are quite the same in the arctic and alpine

regions and on the summits of distant mountain ranges. As no

transportation over such large distances can have brought them from one

locality to the other, no other explanation is left than that they have

been wholly constant and unchanged ever since the glacial period which

separated them. Obviously they must have been subjected to widely

changing conditions. The fact of their stability through all these

outward changes is the best proof that the ordinary external conditions

do not necessarily have an influence on specific evolution. They may

have such a result in some instances, in others they obviously have not.

Many arctic forms bearing the specific name of _alpinus_ justify this

conclusion. _Astragalus alpinus_, _Phleum alpinum_, _Hieracium alpinum_

and [697] others from the northern parts of Norway may be cited as

examples.

Thus Primula imperialis has been found in the Himalayas, and many other

plants of the high mountains of Java, Ceylon and northern India are

identical forms. Some species from the Cameroons and from Abyssinia have

been found on the mountains of Madagascar. Some peculiar Australian

types are represented on the summit of Kini Balu in Borneo. None of

these species, of course, are found in the intervening lowlands, and the

only possible explanation of their identity is the conception of a

common post-glacial origin, coupled with complete stability. This

stability is all the more remarkable as nearly allied but slightly

divergent forms have also been reported from almost all of these

localities. Other evidence is obtained by the comparison of ancient

plants with their living representatives. The remains in tombs of

ancient Egypt have always afforded strong support of the views of the

adherents of the theory of stability, and to my mind they still do so.

The cereals and fruits and even the flowers and leaves in the funeral

wreaths of Rameses and Amen-Hotep are the same that are still now

cultivated in Egypt. Nearly a hundred or more species have been

identified. Flowers of _Acacia_, leaves of _Mimusops_, [698] petals of

_Nymphaea_ may be cited as instances, and they are as perfectly

preserved as the best herbarium-specimens of the present time. The

petals and stamens retain their original colors, displaying them as

brightly as is consistent with their dry state.

Paleontologic evidence points to the same conclusion. Of course the

remains are incomplete, and rarely adequate for a close comparison. The

range of fluctuating variability should be examined first, but the test

of elementary species given by their constancy from seed cannot, of

course, be applied. Apart from these difficulties, paleontologists agree

in recognizing the very great age of large numbers of species. It would

require a too close survey of geologic facts to go into details on this

point. Suffice it to say that in more recent Tertiary deposits many

species have been identified with living forms. In the Miocene period

especially, the similarity of the types of phanerogamic plants with

their present offspring, becomes so striking that in a large number of

cases specific distinctions rest in greater part on theoretical

conceptions rather than on real facts. For a long time the idea

prevailed that the same species could not have existed through more than

one geologic period. Many distinctions founded on this belief have since

had to be abandoned. [699] Species of algae belonging to the

well-preserved group of the diatoms, are said to have remained unchanged

from the Carboniferous period up to the present time.

Summing up the results of this very hasty survey, we may assert that

species remain unchanged for indefinite periods, while at times they are

in the alternative condition. Then at once they produce new forms often

in large numbers, giving rise to swarms of subspecies. All facts point

to the conclusion that these periods of stability and mutability

alternate more or less regularly with one another. Of course a direct

proof of this view cannot, as yet, be given, but this conclusion is

forced upon us by a consideration of known facts bearing on the

principle of constancy and evolution.

If we are right in this general conception, we may ask further, what is

to be the exact place of our group of new evening-primroses in this

theory? In order to give an adequate answer, we must consider the whole

range of the observations from a broader point of view. First of all it

is evident that the real mutating period must be assumed to be much

longer than the time covered by my observations. Neither the beginning

nor the end have been seen. It is quite obvious that _Oenothera

lamarckiana_ was in a mutating condition when I first [700] saw it,

seventeen years ago. How long had it been so? Had it commenced to mutate

after its introduction into Europe, some time ago, or was it already

previously in this state? It is as yet impossible to decide this point.

Perhaps the mutable state is very old, and dates from the time of the

first importation of the species into Europe.

Apart from all such considerations the period of the direct

observations, and the possible duration of the mutability through even

more than a century, would constitute only a moment, if compared with

the whole geologic time. Starting from this conception the pedigree of

our mutations must be considered as only one small group. Instead of

figuring a fan of mutants for each year, we must condense all the

succeeding swarms into one single fan, as might be done also for _Draba

verna_ and other polymorphous species. In _Oenothera_ the main stem is

prolonged upwards beyond the fan; in the others the main stem is lacking

or at least undiscernable, but this feature manifestly is only of

secondary importance. We might even prefer the image of a fan, adjusted

laterally to a stem, which itself is not interrupted by this branch.

On this principle two further considerations are to be discussed. First

the structure of the [701] fan itself, and secondly the combination of

succeeding fans into a common genealogic tree.

The composition of the fan as a whole includes more than is directly

indicated by the facts concerning the birth of new species. They arise

in considerable quantities, and each of them in large numbers of

individuals, either in the same or in succeeding years. This multiple

origin must obviously have the effect of strengthening the new types,

and of heightening their chances in the struggle for life. Arising in a

single specimen they would have little chance of success, since in the

field among thousands of seeds perhaps one only survives and attains

complete development. Thousands or at least hundreds of mutated seeds

are thus required to produce one mutated individual, and then, how small

are its chances of surviving! The mutations proceed in all directions,

as I have pointed out in a former lecture. Some are useful, others might

become so if the circumstances were accidentally changed in definite

directions, or if a migration from the original locality might take

place. Many others are without any real worth, or even injurious.

Harmless or even slightly useless ones have been seen to maintain

themselves in the field during the seventeen years of my research, as

proved by _Oenothera laevifolia_ and _Oenothera_ [702] _brevistylis_.

Most of the others quickly disappear.

This failure of a large part of the productions of nature deserves to be

considered at some length. It may be elevated to a principle, and may be

made use of to explain many difficult points of the theory of descent.

If, in order to secure one good novelty, nature must produce ten or

twenty or perhaps more bad ones at the same time, the possibility of

improvements coming by pure chance must be granted at once. All

hypotheses concerning the direct causes of adaptation at once become

superfluous, and the great principle enunciated by Darwin once more

reigns supreme.

In this way too, the mutation-period of the evening-primrose is to be

considered as a prototype. Assuming it as such provisionally, it may aid

us in arranging the facts of descent so as to allow of a deeper insight

and a closer scrutiny. All swarms of elementary species are the remains

of far larger initial groups. All species containing only a few

subspecies may be supposed to have thrown off at the outset far more

numerous lateral branches, out of which however, the greater part have

been lost, being unfit for the surrounding conditions. It is the

principle of the struggle for life between elementary species, followed

by the survival of the [703] fittest, the law of the selection of

species, which we have already laid stress upon more than once.

Our second consideration is also based upon the frequent repetition of

the several mutations. Obviously a common cause must prevail. The

faculty of producing _nanella_ or _lata_ remains the same through all

the years. This faculty must be one and the same for all the hundreds of

mutative productions of the same form. When and how did it originate? At

the outset it must have been produced in a latent condition, and even

yet it must be assumed to be continuously present in this state, and

only to become active at distant intervals. But it is manifest that the

original production of the characters of _Oenothera gigas_ was a

phenomenon of far greater importance than the subsequent accidental

transition of this quality into the active state. Hence the conclusion

that at the beginning of each series of analogous mutations there must

have been one greater and more intrinsic mutation, which opened the

possibility to all its successors. This was the origination of the new

character itself, and it is easily seen that this incipient change is to

be considered as the real one. All others are only its visible

expressions.

Considering the mutative period of our evening-primrose [704] as one

unit-stride section in the great genealogic tree, this period includes

two nearly related, but not identical changes. One is the production of

new specific characters in the latent condition, and the other is the

bringing of them to light and putting them into active existence. These

two main factors are consequently to be assumed in all hypothetic

conceptions of previous mutative periods.

Are all mutations to be considered as limited to such periods? Of course

not. Stray mutations may occur as well. Our knowledge concerning this

point is inadequate for any definite statement. Swarms of variable

species are easily recognized, if the remnants are not too few. But if

only one or two new species have survived, how can we tell whether they

have originated-alone or together with others. This difficulty is still

more pronounced in regard to paleontologic facts, as the remains of

geologic swarms are often found, but the absence of numerous mutations

can hardly be proved in any case.

I have more than once found occasion to lay stress on the importance of

a distinction between progressive and retrograde mutations in previous

lectures. All improvement is, of course, by the first of these modes of

evolution, but apparent losses of organs or qualities are [705] perhaps

of still more universal occurrence. Progression and regression are seen

to go hand in hand everywhere. No large group and probably even no genus

or large species has been evolved without the joint agency of these two

great principles. In the mutation-period of the evening-primroses the

observed facts give direct support to this conclusion, since some of the

new species proved, on closer inspection, to be retrograde varieties,

while others manifestly owe their origin to progressive steps. Such

steps may be small and in a wrong direction; notwithstanding this they

may be due to the acquisition of a wholly new character and therefore

belong to the process of progression at large.

Between them however, there is a definite contrast, which possibly is in

intimate connection with the question of periodic and stray mutations.

Obviously each progressive change is dependent upon the production of a

new character, for whenever this is lacking, no such mutation is

possible. Retrograde changes, on the other hand, do not require such

elaborate preliminary work. Each character may be converted into the

latent condition, and for all we know, a special preparation for this

purpose is not at all necessary. It is readily granted that such special

preparation may occur, because the [706] great numbers in which our

dwarf variety of the _Oenothera_ are yearly produced are suggestive of

such a condition. On the other hand, the _laevifolia_ and _brevistylis_

mutations have not been repeated, at least not in a visible way.

From this discussion we may infer that it is quite possible that a large

part of the progressive changes, and a smaller part of the retrograde

mutations, are combined into groups, owing their origin to common

external agencies. The periods in which such groups occur would

constitute the mutative periods. Besides them the majority of the

retrograde changes and some progressive steps might occur separately,

each being due to some special cause. Degressive mutations, or those

which arise by the return of latent qualities to activity, would of

course belong with the latter group.

This assumption of a stray and isolated production of varieties is to a

large degree supported by experience in horticulture. Here there are no

real swarms of mutations. Sudden leaps in variability are not rare, but

then they are due to hybridization. Apart from this mixture of

characters, varieties as a rule appear separately, often with intervals

of dozens of years, and without the least suggestion of a common cause.

It is quite superfluous to go into details, as we have dealt with the

horticultural [707] mutations at sufficient length on a previous

occasion. Only the instance of the peloric toadflax might be recalled

here, because the historic and geographic evidence, combined with the

results of our pedigree-experiment, plainly show that peloric mutations

are quite independent of any periodic condition. They may occur anywhere

in the wide range of the toad-flax, and the capacity of repeatedly

producing them has lasted some centuries at least, and is perhaps even

as old as the species itself.

Leaving aside such stray mutations, we may now consider the probable

constitution of the great lines of the genealogic tree of the evening

primroses, and of the whole vegetable and animal kingdom at large. The

idea of drawing up a pedigree for the chief groups of living organisms

is originally due to Haeckel, who used this graphic method to support

the Darwinian theory of descent. Of course, Haeckel's genealogic trees

are of a purely hypothetic nature, and have no other purpose than to

convey a clear conception of the notion of descent, and of the great

lines of evolution at large. Obviously all details are subject to doubt,

and many have accordingly been changed by his successors. These changes

may be considered as partial improvements, and the somewhat picturesque

form of Haeckel's pedigree might well be replaced by [708] more simple

plans. But the changes have by no means removed the doubts, nor have

they been able to supplant the general impression of distinct groups,

united by broad lines. This feature is very essential, and it is easily

seen to correspond with the conception of swarms, as we have deduced it

from the study of the lesser groups.

Genealogic trees are the result of comparative studies; they are far

removed from the results of experimental inquiry concerning the origin

of species. What are the links which bind them together? Obviously they

must be sought in the mutative periods, which have immediately preceded

the present one. In the case of the evening-primrose the systematic

arrangement of the allied species readily guides us in the delimitations

of such periods. For manifestly the species of the large genus of

_Oenothera_ are grouped in swarms, the youngest or most recent of which

we have under observation. Its immediate predecessor must have been the

subgenus _Onagra_, which is considered by some authors as consisting of

a single systematic species, _Oenothera biennis_. Its multifarious forms

point to a common origin, not only morphologically but also

historically. Following this line backward or downward we reach another

apparent mutation-period, which includes the origin of [709] the group

called _Oenothera_, with a large number of species of the same general

type as the _Onagra-forms, Still farther downward comes the old genus

_Oenothera_ itself, with numerous subgenera diverging in sundry

characters and directions.

Proceeding still farther we might easily construct a main stem with

numerous succeeding fans of lateral branches, and thus reach, from our

new empirical point of view, the theoretical conclusion already

formulated.

Paleontologic facts readily agree with this conception. The swarms of

species and varieties are found to succeed one another like so many

stories. The same images are repeated, and the single stories seem to be

connected by the main stems, which in each tier produce the whole number

of allied forms. Only a few prevailing lines are prolonged through

numerous geologic periods; the vast majority of the lateral branches are

limited each to its own storey. It is simply the extension of the

pedigree of the evening-primroses backward through ages, with the same

construction and the same leading features. There can be no doubt that

we are quite justified in assuming that evolution has followed the same

general laws through the whole duration of life on earth. Only a moment

of their lifetime is disclosed to us, but it [710] is quite sufficient

to enable us to discern the laws and to conjecture the outlines of the

whole scheme of evolution.

A grave objection which has, often, and from the very outset, been urged

against Darwin's conception of very slow and nearly imperceptible

changes, is the enormously long time required. If evolution does not

proceed any faster than what we can see at present, and if the process

must be assumed to have gone on in the same slow manner always,

thousands of millions of years would have been needed to develop the

higher types of animals and plants from their earliest ancestors.

Now it is not at all probable that the duration of life on earth

includes such an incredibly long time. Quite on the contrary the

lifetime of the earth seems to be limited to a few millions of years.

The researches of Lord Kelvin and other eminent physicists seem to leave

no doubt on this point. Of course all estimates of this kind are only

vague and approximate, but for our present purposes they may be

considered as sufficiently exact.

In a paper published in 1862 Sir William Thomson (now Lord Kelvin) first

endeavored to show that great limitation had to be put upon the enormous

demand for time made by Lyell, Darwin and other biologists. From a

consideration [711] of the secular cooling of the earth, as deduced from

the increasing temperature in deep mines, he concluded that the entire

age of the earth must have been more than twenty and less than forty

millions of years, and probably much nearer twenty than forty. His views

have been much criticised by other physicists, but in the main they have

gained an ever-increasing support in the way of evidence. New mines of

greater depth have been bored, and their temperatures have proved that

the figures of Lord Kelvin are strikingly near the truth. George Darwin

has calculated that the separation of the moon from the earth must have

taken place some fifty-six millions of years ago. Geikie has estimated

the existence of the solid crust of the earth at the most as a hundred

million years. The first appearance of the crust must soon have been

succeeded by the formation of the seas, and a long time does not seem to

have been required to cool the seas to such a degree that life became

possible. It is very probable that life originally commenced in the

great seas, and that the forms which are now usually included in the

plankton or floating-life included the very first living beings.

According to Brooks, life must have existed in this floating condition

during long primeval epochs, and evolved nearly all the main branches of

the animal and vegetable kingdom [712] before sinking to the bottom of

the sea, and later producing the vast number of diverse forms which now

adorn the sea and land.

All these evolutions, however, must have been very rapid, especially at

the beginning, and together cannot have taken more time than the figures

given above.

The agency of the larger streams, and the deposits which they bring into

the seas, afford further evidence. The amount of dissolved salts,

especially of sodium chloride, has been made the subject of a

calculation by Joly, and the amount of lime has been estimated by Eugene

Dubois. Joly found fifty-five and Dubois thirty-six millions of years as

the probable duration of the age of the rivers, and both figures

correspond to the above dates as closely as might be expected from the

discussion of evidence so very incomplete and limited.

All in all it seems evident that the duration of life does not comply

with the demands of the conception of very slow and continuous

evolution. Now it is easily seen, that the idea of successive mutations

is quite independent of this difficulty. Even assuming that some

thousands of characters must have been acquired in order to produce the

higher animals and plants of the present time, no valid objection is

raised. The demands of the biologists and the results of [713] the

physicists are harmonized on the ground of the theory of mutation.

The steps may be surmised to have never been essentially larger than in

the mutations now going on under our eyes, and some thousands of them

may be estimated as sufficient to account for the entire organization of

the higher forms. Granting between twenty and forty millions of years

since the beginning of life, the intervals between two successive

mutations may have been centuries and even thousands of years. As yet

there has been no objection cited against this assumption, and hence we

see that the lack of harmony between the demands of biologists and the

results of the physicists disappears in the light of the theory of

mutation.

Summing up the results of this discussion, we may justifiably assert

that the conclusions derived from the observations and experiments made

with evening-primroses and other plants in the main agree satisfactorily

with the inferences drawn from paleontologic, geologic and systematic

evidence. Obviously these experiments are wonderfully supported by the

whole of our knowledge concerning evolution. For this reason the laws

discovered in the experimental garden may be considered of great

importance, and they may guide us in our further inquiries. Without

doubt many minor [714] points are in need of correction and elaboration,

but such improvements of our knowledge will gradually increase our means

of discovering new instances and, new proofs.

The conception of mutation periods producing swarms of species from time

to time, among which only a few have a chance of survival, promises to

become the basis for speculative pedigree-diagrams, as well as for

experimental investigations.

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