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PLANT-BREEDING

Garden-Cratt Series

THE HOoRTICULTURIST’S RULE-BOOK

PLANT-BREEDING

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PLANT-BREEDING

BEING FIVE LECTURES UPON THE
AMELIORATION OF DOMESTIC

PLANTS

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MACMILLAN AND CO.

AND LONDON
1895

All rights reserved

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Copyrient, 1895,

By L. H. BAILEY.

Norwood 4Bress

Jj. S. Cushing & Co. — Berwick & Smith
Norwood Mass. U.S.A.

PREFACE.

THERE is no subject associated with the care
of plants respecting which there is so much mis-
apprehension and imperfect knowledge, as that
of the origination of new forms. Most of the
scattered writing touching it treats the subject
as if all our knowledge of the matter were and
must be derived wholly from experiment. It
therefore recites examples of how this and that
new form has come to be, and has made little
attempt to discover the fundamental causes of
the genesis of the novelties. Horticulturists
commonly look upon each novelty as an isolated
fact, whilst we ought to regard each one as but
an expression of some law of the variation of
plants. It is the common notion, too, to con-
sider any type of plant to be essentially a fixed
entity, and to regard any marked departure from

the type as a phenomenon rather more to be
Vv

vl PREFACE.

wondered at than to be explained. It is evident,
however, that one cannot understand the pro-
duction of new varieties until he has grasped
some of the fundamental principles of the on-
ward progression of the vegetable kingdom.
Any attempt, therefore, to explain the origin of
garden varieties, and the methods of producing
them, must be at the same time a contribution
to the literature of the philosophy of organic
evolution.

I do not know of any explicit and sustained
attempt to account for the evolution of all gar-
den forms, and I have therefore brought together
in this volume the subject-matter of various
lectures which I have been in the habit of
giving before my students. The first and
third lectures were newly elaborated the present
summer for two addresses before the class in
biology which came together at the University
of Pennsylvania, under the -auspices of the
American Society for the Extension of Univer-
sity Teaching. The second lecture was first
presented before the Massachusetts State Board
of Agriculture, in Boston, December 1, 1891.
In April, 1892, it was republished, with a bibli-

PREFACE. Vil

ography of the subject, by the Rural Publishing
Co., under the title, “« Cross-Breeding and Hybrid-
izing.” This publication is now out of print.
I have made no attempt to collect lists or cata-
logues of varieties, but have endeavored to make
very brief statements of some of the underlying
principles of the amelioration of plants, with only
sufficient examples to fix them in the mind.

I hope that teachers of horticulture and botany
may find the book useful in their classes. When
it ig necessary to abridge the instruction or to
present it to untrained students, only Lectures
IIT. and V. may be used, for these contain the
matters of greatest demonstrative importance.

L. AH BAT LEY.

CORNELL UNIVERSITY,
IrHaca, N.Y., September 1, 1895.

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CONTENTS.

LECTURE I.

Tur Fact AND PHILOSOPHY OF VARIATION s

I. The Fact of Individuality .
The seed-individual.
The bud-individual.
Il. The Causes of Individual Differences

a. Fortuitous variation
b. Sex as a factor in the variation of plants
¢. Physical environment and variation
1. Variation in food supply
2. Variation in climate
3. Change of seed. Bud-variation
d. Struggle for life a cause of variation .

Ill. The Choice and Fixation of Variations : =

LECTURE II.

Tur PHILOSOPHY OF THE CROSSING OF PLANTS, CONSID-
ERED IN REFERENCE TO THEIR IMPROVEMENT UNDER
CULTIVATION

I. The Struggle for Life
Il. The Division of Labor ; : .

Ill. The Limits of Crossing é : : :
ix

PAGE

CONTENTS.

TY. Function of the Cross :
a. The gradual amelioration of the os
b. Change of seed and crossing
. The outright production of new varieties

V. Characteristics of Crosses .

VI. Uncertainties of Pollination
Conclusion

LECTURE III.

How Domestic VARIETIES ORIGINATE

I. Indeterminate Varieties
II. Plant-breeding

Rule
Rule

Rule
Rule
Rule

Rule
Rule

Rule
Rule

Rule 12.
Rule 13.
Rule 14.

Rule 15.

iis
2.

-

8.
9.
Rule 10.
Rule 11.

Antagonistic features
Quickest results in the most vastalste
groups .

. Breed for one thing a ihe ‘

Contradictory attributes .

Characters of the entire ie fast
important

Plants differ in her entitagy power

Less marked variations more impor-
tant :

Crossing a means, 0 an aad

Choice of parents to a cross

The ideal should be mental

Seek to produce variation in the
desired direction .

Watch for bud-varieties

Progress lies in selection .

The type is kept up to standard By
continued selection

The best final results are to he es
tained by high tillage and intelli-
gent selection

PAGE

127

CONTENTS. X1

PAGE

Ill. Specific Hxanples i ‘ ‘ ; : - 129
The dewberry and blackberry ‘ ; s 29

The apple f ; : , ‘ : - lor

Beans : : ; : - ; é . 135
Cannas . : ; : 3 4 : . 140

LECTURE IV.

BORROWED OPINIONS ; BEING EXTRACTS FROM THE WRIT-
Incs oF B. Vertot, E. A. CARRIERE, AND W. O.

FocokeE ; ; . , ‘ : ‘ , . 148
I. Verlot’s Classification of Varieties of Ornamen-
tal Plants. ; : : : k . 148
Il. Carriére’s Account of Bud-varieties . ; YS
1. General remarks upon bud-variation . . 154
2. List of bud-varieties . 2 z ; EO
Ill. Focke’s Discussion of the Characteristics of
Crosses ; : h , y : «, 2h
i. The simple primary cross . ‘ 215
Proposition 1. Similarity of Bascal off.
spring . : 215
Proposition 2. Dissimilarity of et
offspring. . at
Proposition 3. Weueatire powers of hy-
brids . ‘ 225
Proposition 4. Caumanstiye fertility of
hybrids : ‘ 228
Proposition 5. Mintitonmiaitions: in hybrid
offspring . ‘ : : : Bor
ii. The progeny of crosses. 237

1. Progeny of crosses with ‘itil own
pollen. ‘ , : : . 2388

Xli CONTENTS.

PAGE
2. Derivative hybridization of crosses
with the parent forms . . . 242
3. Hybrids of several species . i . 244
a. Triple hybrids. ; : . 244
b. Hybrids of four to six species . 246
c. Crosses of plants grown together 247
iii. Cross-breeds and hybrids . . ; . 247
LECTURE V.
POLLINATION; OR How To Cross PLANTs . : ; . 252

I. The Structure of the Flower : : ‘ . 252
Il. Manipulating the Flowers . : : , . 265

GLOSSARY. ; : ; : : F : .. 282

INDEX —.-. : : : ‘ . : - : . 286

PLANT-BREEDING.

LECTURE I.

THE FACT AND PHILOSOPHY OF VARIATION.

THERE is no one fact connected with horticult-
ure which so greatly interests all persons as the
existence of numerous varieties of plants which
seem to satisfy every need of the gardener.
Whence came all this multitude of forms? What
are the methods employed in securing them? Are
they simply isolated facts or phenomena of gar-
dening, or have they some relation to the broader
phases of the evolution of the forms of life?
These are some of the questions which occur to
every reflective mind when it contemplates an
attractive garden, but they are questions which
seem never to be answered. Whatever attempt
the gardener may make at answering them is
either befogged by an effort to define what a vari-
ety is, or else it consists in simply reciting how a
few given varieties came to be known. But there

B 1

2 FACT AND PHILOSOPHY OF VARIATION.

must be some fundamental method of arriving at
a conception of how the varieties of fruits and
flowers and other cultivated plants have origi-
nated. If there is no such method, then the
origination of these varieties must follow no law,
and the discussion of the whole subject is fruit-
less. But we have every confidence in the con-
secutive uniformity of the operations of nature,
and it were strange if some underlying principle
of the unfolding or progression of plant life does
not dominate the origin of the varied and in-
numerable varieties which, from time unknown,
have responded to the touch of the cultivator.
Let us first, therefore, make a broad survey of the
subject in a philosophical spirit, and, later, discuss
the more specific instances of the origination of
varieties.

I. THe Fact oF INDIVIDUALITY.

There is universal difference in nature. No
two living things are exact counterparts, for no
two are born into exactly the same conditions and
experiences. Every living object has individ-
uality ; that is, there is something about it which
enables the acute observer to distinguish it from
all-other objects, even of the same class or spe-
cies. Every plant in a row of lettuce is different
from every other plant, and the gardener, when

INDIVIDUALITY. 3

transplanting them, selects out, almost uncon-
sciously, some plants which please him and others
which do not. Every apple tree in an orchard of
a thousand Baldwins is unlike every other one,
perhaps in size or shape, or possibly in the vigor
of growth or the kind of fruit it bears. Persons
who buy apples for export know that fruit from
certain regions stands the shipments better than
the same variety from other regions; and if one
were to go into the orchards where these apples
are grown, he would find the owner still further
refining the problem by talking about the merits
of individual trees in his orchard. If one were
to make the effort, he would find that it is pos-
sible to distinguish differences between every two
spears of grass in a meadow, or every two heads
of wheat in a grain-field.

All this is equivalent to saying that plants are
infinitely variable. The ultimate causes of all
this variation are beyond the purpose of the
present discussion, but it must be evident, to
the reflective mind, that these differences are
the means of adapting the innumerable indi-
viduals to every little difference or advantage
in the environment in which they live. And if
the object of variation is better adaptation to
the physical conditions of life, then the same
motive must have been present in the circum-
stances which determined the birth of the indi-

4 . FACT AND PHILOSOPHY OF VARIATION.

vidual. The variation in environment, therefore,
must be the cause of much of the variation in
plants, since differences in plants were positively
injurious if it were possible for the conditions of
environment to be the same.

If no two plants are anywhere alike, then it is
not strange if now and then some departure, more
marked than common, is named and becomes a
garden variety. We have been taught to feel
that plants are essentially stable and inelastic,
and that any departure from the type is an excep-
tion and calls for immediate explanation. The
fact is, however, that plants are essentially un-
stable and plastic, and that variation between the.
individuals must everywhere be expected. This
erroneous notion of the stability of organisms
comes of our habit of studying what we call
species. We set for ourselves a type of plant or
animal, and group about it all those individuals
which are more like this type than they are like
any other, and this group we name a species.
Nowadays, the species is regarded as nothing
more than a convenient and arbitrary expression
for classifying our knowledge of the forms of
life, but the older naturalists conceived that the
species is the real entity or unit in nature, and we
have not yet wholly outgrown the habit of mind
which was born of that fallacy. Nature knows
nothing about species; she is concerned with the

INDIVIDUALITY. 5

individual, the ultimate unit. ‘This individual she
moulds and fits into the chinks of environment,
and each individual tends to become the more
unlike its birthmates the more the environments
of the various individuals are unlike. I would
impress upon you, therefore, as a fundamental
conception to the discussion of the general subject
before us, the importance of the individual plant,
rather than the importance of the species; for
thereby we put ourselves as nearly as possible in
a sympathetic attitude with nature, and, resting
upon the ultimate object of her concern, we are
able to understand what may be conceived to be
her motive in working out the problem of life.
That I may still more forcibly emphasize this
thought, let me recall to your minds the fact
that the whole tendency of contemporary civili-
zation, in sociology and religion, is to deal with
the individual person and not with the mass.
This is only an unconscious feeling after natu-
ral methods of solving the most complex of
problems, for it is exactly the means to which
every organic thing has been subjected from the
beginning.

In looking for the ultimate unit or individuality
or personality in nature, we must make a broad dis-
tinction between the animal and the plant. Every
higher animal is itself a unit; it is one. It has
a more or less definite span of life, and every part

6 FACT AND PHILOSOPHY OF VARIATION.

and organ contribute a certain indispensable part
to the life and personality of the organism. No
part is capable of propagating itself independently
of the sex-organs of the animal, nor is it capable
of developing sex-organs of its own. If any part
is removed, the animal is maimed and perhaps it
dies. The plant, on the contrary, has no definite
or distinct autonomy. Most plants live an indefi-
nite existence, dependent very closely upon the
immediate conditions in which they grow. Every
part or branch of the plant lives largely for itself,
it is capable of propagating and multiplying itself
when removed from the parent plant or the colony
of branches of which it is a member, and it de-
velops sex-organs and other individual features
of its own. If any branch is removed, the tree
or plant does not necessarily suffer; in fact, the
remaining branches usually profit by the removal,
a fact which shows that there is a competition,
or struggle for existence, between the different
branches or elements of the plant. The whole
theory and practice of pruning rest upon the fact
of the individual unlkenesses of the branches of
plants; and these unlikenesses are of the same
kind and often of the same degree as those which
exist between different plants which are grown
from seeds. That is, the branches of a Crawford
peach tree, for example, differ amongst themselves
in size, shape, vigor, productiveness, and season of

BUD-INDIVIDUALS. c

maturity, the same as any two or more separate
Crawford trees, or any number of trees of other
varieties, differ the one from the others. If any
one of these branches or buds is removed and is
grown into an independent tree, a person could
not tell—it he were ignorant of its history — if
this tree were derived from a branch or a seed.
This proves that there is no essential unlikeness
between branches and independent plants, except
the mere accident that one grows upon another
branch or plant whilst the other grows in the
ground. But the branch may be severed and
grown in the ground, and the seedling may be
pulled up and grafted on the tree, and no one
can distinguish the different origins of the two.
And then, as a matter of fact, a very large pro-
portion of our cultivated plants are not distinct
plants at all, in the sense of being different crea-
tions from seeds, but are simply the results of
the division of branches of one original plant or
branch. All the fruit trees of any one variety
are obtained from the dividing up and multiplica-
tion of the branches of the first or original tree.
You are now curious to know how this orig-
inal tree came to be, and this I hope to tell
you before I am done; but for the present, let
me impress it upon you that it is equally possi-
ble for it to have come from a seed, or to have
sprung from a branch which some person had

8 FACT AND PHILOSOPHY OF VARIATION.

noticed to be very different from the associated
branches in the tree-top. In other words, the
ultimate unit or individual in growing plants is
the bud and the bit of wood or tissue to which it
is attached ; for every bud, like every seed, pro-
duces an offspring which can be distinguished
from every other offspring whatsoever.

Il. THE CAUSES OF INDIVIDUAL DIFFERENCES.

We have now gotten back to the starting-point,
to that unit with which nature begins to make
her initial differences or individualities ; that is,
to the point where variations arise. This unit
is the bud and the seed,—one sexless, or the
offspring of one parent; the other sexual, or the
offspring of two parents. Now, inasmuch as
the horticultural variety is only a well-marked
variation which the gardener has chanced to
notice and to propagate, it follows that the only
logical method of determining how garden vari-
eties originate is to discover the means by which
plants vary or differ one from another.

There is probably no one fact of organic nature
concerning the origin of which modern philoso-
phers are so much divided as the genesis or
reasons for the beginnings of variations or dif-
ferences. It seems to be an inscrutable problem,
and it would be useless, therefore, for us to at-

FORTUITOUS VARIATION. 9

tempt to discover these ultimate forces in the
present hour. Still, we must give them sufficient
thought to enable us to satisfy our minds as to
how far these variations may be produced by
man; and, in doing this, we must discover at
least the underlying philosophy of plant variation.
It is the nature of organisms to be unlike their
parents and their birthmates. Why?

a. Fortuitous Variation.

It will probably never be possible to refer every
variation to a distinct cause, for it is probable that
some of them have no antecedent. If we con-
ceive of the forms of life as having been created
with characters exactly uniform from generation
to generation, then we should be led to look for
a distinct occasion or cause for every departure
from the type; but we know, as I have already
pointed out, that heredity by its very nature is
not so exact as to carry over every attribute, and
no other, of the parent to the offspring. Elas-
ticity, plasticity, is a part of the essential consti-
tution of all organic beings. There is probably
no inherent tendency in organisms towards any
ultimate or predetermined completion of form, as
the older naturalists supposed, but simply a laxity
or indefiniteness of constitution which is expressed
in numberless minor differences in individuals.

10 FACT AND PHILOSOPHY OF VARIATION.

That is, some variation is simply fortuitous, an
inevitable result of the inherent plasticity of
organisms, and it has no immediate inciting
cause. If we were to assume that every minor
difference is the result of some immediate cause,
then we should expect every individual plant or
animal to fill some niche, to satisfy some need, to
produce the definite effect for which the cause
stands. But it is apparent to one who contem-
plates the operations of nature that very many —
certainly more than half — of the organisms which
are born are wholly useless in the struggle for
life and very soon perish. From these fortuitous
variations nature selects, to be sure, many indi-
viduals to be the parents of other generations
because they chance to be fitted to live, but this
does not affect the methods or reasons of their
origin. It is possible that, whilst many of these
mere individual differences have no direct and
immediate cause, they may still be the result of a
devious line of antecedent causes long since so
much diffused and modified that they will remain
forever unrecognizable ; but even if so, the fact
still remains that these present differences or
variations may be purposeless, and it is quite as
well to say that they exist because it is a part
of the organic constitution of living things that
unlike produces unlike.

SEX AND VARIATION. tt

b. Sex as a Factor in the Variation of Plants.

All plants have the faculty, either potential or
expressed, of propagating themselves by means
of buds, or asexual parts. This is obviously the
cheapest and most direct possible method of propa-
gation for many-membered plants, since it requires
no special reproductive organization and energy,
and, as only one parent is concerned in it, there
is none of the risk of failure which resides in any
mode of propagation in which two parents must
find each other and form a union. There must
be some reason, therefore, for the existence of
such a costly mechanism as sex aside from its use
as a mere means of propagation. It may be said
that it exists because it is a means of more rapid
multiplication than bud-propagation, but such is
not necessarily the fact. ‘There are many plants
which produce buds as freely as they produce
seeds ; and then, if mere multiplication were the
only destiny of the plant, bud-production would
no doubt have greatly increased to have met the
demand for new generations. ‘The only reason
for the existence of sex in the vegetable world
seems to be the need for a constant rejuvenation
and modification of the offspring by uniting the
features of two individuals into one. ‘There thus
arises from every sexual union a number of new
or different forms from which nature may select

12 FACT AND PHILOSOPHY OF VARIATION.

the best, — that is, those best fitted to live in the
conditions in which they chance to be placed.
But whilst sex is undoubtedly one of the most
potent sources of present unlikenesses, it 1s not
necessarily an original cause of individual differ-
ences, since the two parties to any sexual con-
tract must be unlike before they can produce
unlike. When once the initial unlikenesses were
established, every new sexual union would pro-
duce new combinations, so that now, when every
new form, from whatever source it appears, comes
into existence, there are other intimately related
forms with which it may cross. This state of
things has existed to a greater or less degree from
the moment sex first appeared, so that the organic
world is now endlessly varied as the result of a
most complex ancestry.

The variety which sexual union has introduced
into the world performs such an important part in
the evolution of the forms of plants, and the prob-
lems which it presents are so complex, that I shall
leave the whole subject for an independent dis-
cussion (Lecture II.).

c. Physical Environment and Variation.

Every phase and condition of physical cireum-
stances, which are not absolutely prohibitive of
plant life, have plants which thrive in them.

EEE eel

ENVIRONMENT AND VARIATION. 13

Every soil and climate, every degree of humidity,
hills, swamps, and ponds,—every place is filled
with plants. Even the trunks and branches of
trees support other plants, as epiphytes and para-
sites. That is, plants have adapted themselves to
every physical environment ; or, to turn the propo-
sition around, every physical environment pro-
duces adaptive changes in plants. There are
those, like Weismann and his adherents, who
contend, from purely speculative reasons, that
these changes do not become hereditary or perma-
nent until they have influenced a certain physio-
logical substance which is assumed to reside in
the reproductive regions of the organism, and
that all those changes which have not yet reached
this germ-plasm are, therefore, lost, or die with
the organism. It is not necessary to combat this
philosophy, for we know, as a matter of common
horticultural experience, that every change or va-
riation in any organism — unless it proceeds from
mere accident or mutilation — may become heredi-
tary or be the beginning of a new variety; it is
only necessary, therefore, for the Weismannians to
assume —as they are always ready to do—that
any variation which has become fixed or permanent
has already affected the germ. Their assumption
needs only another assumption to prove it, and,
therefore, when we are considering merely plain
matters of fact and experience, we need give little

14 FACT AND PHILOSOPHY OF VARIATION.

attention to the subtleties of this Neo-Darwinian
philosophy.

Weismann teaches that “ acquired characters,”
or those variations which first appear in the life-
time of the individual because of the influences of
environment, are lost, because they have not yet
affected the reproductive substance. But if these
characters are induced by the effect of impinging
environment during two or more generations, they
may come to be so persistent that the plant can-
not throw them off, and they become, thereby, a
part of the hereditary and non-negotiable prop-
erty of the species. Now, it is apparent that in
one or another of the generations which are thus
acted upon by the environment, there must be a
beginning towards the fixing or hereditable per-
manency of the new form, and we might as well
assume that this beginning takes place in the first
generation as in the last, since there can be no
proof that it does not take place in either one.
The tendency towards fixity, if it exists at all,
undoubtedly originates at the very time that the
variation itself originates, and it is only sophistry
to assume that the form appears at one time and
the tendency towards permanence at another time.
Since plants fit themselves into their circumstances
by means of adaptive variations, we must con-
clude that all adaptive variations have the power
of persisting, upon occasion.

NATAL AND POST-NATAL VARIATION. 15

All these remarks, whilst somewhat abstruse,
have a most important bearing upon the phi-
losophy of the origin of garden varieties, because
they show, first, that changes in the conditions in
which plants grow introduce modifications in the
plants themselves, and second, that wherever any
modification occurs it is probable that it may be
fixed and perpetuated.

It is necessary, at this point, that we distin-
guish between natal and post-natal variations ;
that is, between those variations which are born
with plants, and those which appear, as a result of
environment, after the plant has begun to grow.
It is commonly assumed that the form and general
characters of the plant are already determined in
the seed, but a moment’s reflection will show that
this is far from the truth. One may sow a hun-
dred selected peas, for instance, all of which may
be alike in every discernible character. If these
are planted in a space a foot square, it will be
found, after two or three weeks, that some indi-
viduals are outstripping the others, although all
of them came up equally well and were at first
practically indistinguishable. This means that,
because of a little advantage in food or moisture,
or other circumstance, some plants have obtained
the mastery and are crowding out the less fortu-
nate ones. Here is a variation taking place before
our very eyes, and we may be able to see the exact

16 FACT AND PHILOSOPHY OF VARIATION.

cause of it. Moreover, variations which originate
in this way may pass down to the offspring
through the seeds, as in the case of “ viney” peas,
which are grown on too rich soil. All this is a
matter of the commonest observation with the
gardener, who is so accustomed to seeing great
differences arise in batches of plants, all of which
start equal and with an equal chance, that he
never thinks to comment upon the occurrence.
In fact, the theory and practice of agriculture
rest upon the fact that plants can be modified
greatly by the conditions in which they grow,
after they have become thoroughly established in
the soil. Plants may start equal, but may differ
widely at the harvest; and this difference may be
controlled to a nicety by the cultivator. Every
farmer knows, too, that the best results for the
succeeding year are to be got only when he selects
seeds from the best which he has been able to pro-
duce this year. So, given uniformity or equality
at the start, the operator moulds the individual
plants largely at his will.

Having noticed that physical environments may
modify plants, we are now ready to consider just
what changes in these circumstances of plant life
are most fruitful in the production of new forms.

1. Variation in Food Supply.— The greater
part of the changes in the physical conditions
of life hinge upon the relative supply of food.

Ee

FOOD SUPPLY AND VARIATION. Ly

Climbing plants assume their form because, by
virtue of the divergence of character, they are
enabled to fit themselves into places which other
plants cannot occupy. They rear their foliage
into the air, where food and sunlight are unappro-
priated. ‘The lower branches of the tree-top die,
and the others thereby appropriate the more food
and grow the faster. ‘The entire practice of agri-
culture is built upon the augmentation of the
food supply. For this purpose, we set the plants
in isolated positions, we till the ground, keep
down other plants or weeds, add plant food to
the soil, and prune the tree and thin the fruit.
Thomas Andrew Knight, the chief of horticul-
tural philosophers, appears to have been the first
to clearly enunciate the law that excess of food
supply is the most prolific cause of the variation
of plants. Darwin subscribes to it without re-
serve: ‘*Of all the causes which induce. varia-
bility, excess of food, whether or not changed in
nature, is probably the most powerful.” Alex-
ander Braun, an earlier writer upon the philosophy
of the organic world, said that ‘it appears rather,
on the whole, as if the unusual conditions favor-
able to a luxuriant state of development, afforded
by cultivation, awakened in the plant the inward
impulse to the display of all those variations pos-
sible within the more or less narrowly circum-
scribed limits of the species.” It is generally
Cc

18 FACT AND PHILOSOPHY OF VARIATION.

agreed by those who have given the matter much
thought, that an excess of food above the amount
normally or habitually received is one of the very
chief, if not the most dominant, causes of indi-
vidual differences in plants. Certainly every
farmer and gardener knows that the richer the
soil in available plant food, the stronger and the
more abnormal and unusual his product will be.

If, then, excess of food supply is a strong factor
in the modification of plants, and if the one fun-
damental aim of agriculture is to supply food in
excess of natural conditions, it must naturally
follow that cultivated plants should be of all
others the most variable. ‘This is notably true.
Now, the first variation which usually comes of
this liberal food supply is increase in mere big-
ness. Probably every plant which has ever been
cultivated has increased its stature or the size of
some or all of its parts. Moreover, this is gener-
ally the direct object of cultivation, —to secure
larger herbage, fruits, seeds, or flowers. Inci-
dentally, we find here an indubitable proof of the
truth of the hypothesis of evolution, for if it were
impossible for plants to vary or to assume new
characters, there would be no cultivation and no
agriculture; for there would be little object in
cultivating a product if it grew equally well in
the wild.

This variation into mere bigness is more impor-

BREAKING THE TYPE. 19

tant than it may seem at first sight. All thought-
ful horticulturists agree in beheving that the first
thing to be done in ameliorating any plant is to
‘‘break the type,” that is, to cause it to vary.
The particular direction of variation is not so
important, at first; for all experience has shown
that if once the seedlings of a plant begin to
depart from the parental type, other and various
modifications will soon follow. If a plant is once
strongly modified in size, variations in shape,
color, flavor, or other attributes are forthcoming.
This apparent accumulation of variation seems at
first to be incapable of scientific explanation, but
the reasons for it are not difficult to understand
when once they are presented.

When plants are placed in new conditions,
whether in the wild or in cultivation, then they
begin to vary, but usually only in one direction
at first, although the amount of the variation, and
sometimes the kind, is determined very largely
by the nature and the extent of the change in the
conditions. This initial variation, particularly
when plants are transferred to cultivated areas,
is generally in the direction of greater size con-
sequent upon the greater amount of food. This
initial variation is generally soon followed by
others in various directions, and from these the
cultivator may be able to establish new varieties.
We now ask ourselves why these many variations

20 FACT AND PHILOSOPHY OF VARIATION.

appear when once the type begins to modify itself.
Consider the fact that the world is now full of
plants. In untamed nature, not one more plant
can grow unless another plant dies. All plants,
therefore, are held down to narrow limits of num-
bers, and since there are so few individuals, —
in comparison with the seeds and buds which
each plant produces for the chance of multiplying
itself, — there must be, also, few kinds and degrees
of individual differences. The farther and more
freely a plant distributes itself, the greater must
be the differences between the various individuals,
because they must adapt themselves to a wider
range of conditions. All plants are held in equi-
librium, so to speak; but the plant organism is
plastic by nature and quickly responds to every
touch of environment; so, as soon as the pressure
is removed in any direction, the plant at once
springs into the breach. Recall the monotonous
vegetation of the deep forest, where the battle
of centuries has subdued all but the strongest.
Clear away the forest, and then observe the
fierce scramble for place and life amongst a multi-
tude of forms which spring in for an opportunity
to better their conditions. In a few years more,
the tender low herbs have gone. The briars and
underbrush have usurped the land. As time goes
on, one species after another perishes, and when
the place is again reforested, two or three species

EQUILIBRIUM IN NATURE. a

hold undisputed sway over the land. The pop-
lars which followed the pines have long since per-
ished and pines again dominate the forest. Or,
if the area were turned to pasture a few years
after the woods were removed, the herbs and
bushes die with the browsing, and in time the
June grass covers the whole landscape with the
mantle of conquest. So plants may be said to
be always ready to fill new places in the polity
of nature by adapting themselves to the new cir-
cumstances as they grow into them. The appear-
ing of any one marked variation, therefore, is
evidence that the plant has found a new condi-
tion, that the pressure is somewhat lifted, and
that its whole plastic organization will soon re-
spond to the new environment. It is apparent,
then, how the simplest and rudest cultivation has
been able, through the centuries, to so profoundly
modify our domestic plants that we are often
unable to recognize the forms from which they
sprung.

We must not forget to notice, at this point,
that the food supply differs amongst the various
branches of the same plant. Some branches, by
reason of position with reference to the main
trunk or with reference to air and sunlight, or,
because of a better start in the beginning as a
result of some incidental advantage, gain the
mastery over others and crowd them out. We

22 FACT AND PHILOSOPHY OF VARIATION.

have already seen that no two branches on a plant
are alike; and we are now able to understand that
sports or bud-varieties are no more inexplicable
than seed-varieties are.

Cultivation is really but an extension or intensi-
fication of nature’s methods of dealing with the
plant world. The ultimate object of both nature
and man is to supply more food. ‘The variations
which arise from the effects of mere cultivation,
therefore, are in kind very like those which nature
produces, the chief difference being that of degree.
The accustomed operations of the farmer, there-
fore, have been powerful agents in the evolution
of vegetable forms. The ways in which cultiva-
tion affords a more liberal food supply are as fol-
lows :— |

1. By isolating the individual plant. The
husbandman sets each plant by itself, and then
protects it by destroying the weeds or plants
which endeavor to crowd it out. There is a
partial exception to this in the “sowed crops,”
like the grains, and it is noticeable that variation
in these plants is usually less marked than in the
‘hoed. .cropsid

2. By giving the plant the advantage of posi-
tion, whereby it is allowed the most congenial
exposure to sun and contour of land.

3. By increasing the fertility of the soil, either
by tillage or the direct application of plant

MEANS OF AUGMENTING FOOD SUPPLY. 23

food, or both. Rich and moist soils tend to
“break” the type,—or to cause initial varia-
tions, —to produce verdant colors and loss of
saccharine and pungent qualities, to induce re-
dundant growth, and to delay maturity and
thereby to render plants tender to cold winter
climates.

4. By thinning the tops of plants and the
fruits, whereby the remaining parts receive an
amount of food in excess of the habitual allow-
ance.

5. By divergence of character in associated
plants. It is well known that a field which is
planted so thickly to corn that it cannot grow
more with profit, may still grow pumpkins be-
tween. The pumpkins and the corn are so unlike
in form that they complement each other, the one
filling the niche which the other is not fitted to
occupy. We have already seen that a copse ever
so full of bushes may still grow vines. A meadow
which is full of timothy may still grow clover
in the bottom, and land which is covered with
apple trees still grows weeds beneath. “The
more diversified the descendants from any one
species become in structure, constitution, and
habits,” writes Darwin, ‘‘ by so much will they
be better enabled to seize on many and widely
diversified places in the polity of nature, and so
be enabled to increase in numbers.”

24 ¥FACT AND PHILOSOPHY OF VARIATION.

2. Variation in Climate. —The fact that any
distinct climatic region usually has plants which
are very closely related to those of other climatic
regions in the same zone, points strongly to the
probable profound modification of plants by cli-
mate. And, furthermore, we should expect that if
the food environment modifies plants, the climatic
environment must have the same power. More-
over, there is abundant historical and experimental
proof that climate is capable of greatly modifying
the vegetable kingdom. There are those who
contradict any great effect of climate in the varia-
tion of plants, and acclimatization has been even
stoutly denied. ‘These persons make the mistake
of asking that a visible modification take place at
once upon the transfer of a plant from one climate
to another, and they also err in supposing that a
plant can adapt itself to a cold climate only by
developing a capability to withstand more cold.
Indian corn is sometimes cited as proof that
plants do not become acclimatized, for it is as
tender to frost now as ever, for all that we know.
Yet this very plant affords a most unequivocal
example of complete acclimatization, because it
has shortened its period of growth fully one-half
to enable it to escape the cold of the north.

The influence of a change of climate upon
plants, or, what may amount to the same thing,
the result of a transfer of plants to new climates,

CLIMATE AND VARIATION. 25

is so complex and so general that no detailed dis-
cussion of the subject can be made at this time.
It will answer our present purpose to briefly
designate the ways in which climate modifies
plants: —

1. Climate greatly modifies the stature of plants.
They become dwarfer in high latitudes and alti-
tudes.

2. It modifies form. Plants tend to be broader-
headed, and also more prostrate, in high latitudes
and altitudes.

3. Proportionate leafiness generally increases,
at the same time.

4. There is, also, often a gain in comparative
fruitfulness following transfer towards the poles.

5. The colors of leaves, flowers, fruits, and
seeds are greatly influenced by climate, there
being a general tendency, in plants of temperate
regions, to augmentation in intensity of colors as
they are carried towards the poles.

6. There is modification in the flavor and es-
sential ingredients of various parts, following a
change of climate.

7. There is a variation in variability itself. The
more difficult the climate in which a plant finds
itself, the more it tends to vary to meet the uncon-
genial environments. In the high north, many
plants are so variable that the marks used to iden-
tify the species in other latitudes are often lost.

26 FACT AND PHILOSOPHY OF VARIATION.

8. There may be a profound variation or modi-
fication in constitution and habit by which plants
become acclimatized, or enabled to endure a cli-
mate at first injurious to them. ‘This may occur
by a variation in the constitution of the descend-
ants, which enables them to endure directly more
untoward conditions. It generally comes about,
however, through a change in habit, by which
plants, when transferred towards the poles, shorten
their season of growth or even become annuals.
Plants become more sensitive to spring tempera-
tures in cold climates, so that they start relatively
much earlier in the season — that is, at a lower
sum-temperature — than they do in warm climates.
Any one who has passed the springtime in both
the North and South must have noticed how much
more suddenly the vegetation comes forward in
the North; and it is surprising how the spring-
sowed crops accelerate their growth in the North
over those in the South.

The characters which result from a change of
climatic environment are peculiarly within the
control of the agriculturist, for a leading factor
in his business is the transfer of plants far and
wide over the earth. So it has come that the
staple varieties of the important grains and fruits
are unlike in Europe and America and in all great
geographical areas, although all the various forms
may have sprung from one ancestor within historic

CLIMATE AND VARIATION. oT

times. A new country is stocked with varieties
from the mother country; but in the course of
a few generations it is found that the varieties
in cultivation are unlike the ones originally in-
troduced, and from which they came. As wild
plants have become separated from each other as
species in the different geographical regions, so
the cultivated plants soon begin to follow similar
lines of divergence. In the beginning of the colo-
nization of this country, for example, all the vari-
eties of apples were of European origin. But in
1817, over sixty per cent of the apples recommended
for cultivation here were of American origin, that
is, American-grown seedlings from the original
stock. At the present time, fully ninety per cent
of the popular apples of the Atlantic states are
American productions. The northern states of
the Mississippi valley, to which most of our east-
ern apples are not adapted, are now witness-
ing a similar transformation in the adaptation
and modification of the varieties introduced
from the East and from Russia. The newly
introduced Japanese plums are conceded to be
great acquisitions to our fruit-growing, but no
doubt the best results are yet to come with the
origination of domestic varieties of them. So
there is an irresistible tendency towards a di-
vergence of forms in different continental or
geographical regions, and much of the inevi-

28 FACT AND PHILOSOPHY OF VARIATION.

table result is no doubt chargeable to climatic
environment.

3. Change of seed. Bud-variation. —1 wili
pause for a moment to consider two agencies or
phenomena which are often associated with the
genesis of varieties. One of these is the fact
that the simple change of seed from one locality
to another generally gives a larger or better
product or even more marked variation. Mere
transfer of seed is not of itself, however, a cause
of variation. The change is beneficial because
it fits together characters and environments which
are not in equilibrium with each other. <A plant
which is grown for several years in one set of con-
ditions becomes fitted into them, so to speak, and
is in a comparative state of rest. When the plant
or its progeny is taken to other conditions, all the
adjustments are broken up, and in the refitting to
the new circumstances new or strange characters
are apt to appear. We shall leaye this subject
for the present, expecting to give it a fuller
treatment in the second lecture.

Bud-variation, or sport, is a name given to
those branches which are so much unlike the
normal plant in any particular that they attract
attention. Many garden varieties are simply
multiplications of such abnormal branches. ‘This
bud-variation is commonly held to be such an
unusual and inexplicable phenomenon that it is

STRUGGLE FOR LIFE AND VARIATION. 29

considered apart from all the general discussions
of variation. It is not, of course, a cause of vari-
ability, but simply an effect of some antecedent,
the same as seed-variation is. We have already
seen that all the different branches, or even joints,
of any plant are, in a very important sense, dis-
tinct individuals, since every one develops its
own organs, each is capable of reproducing’ itself
independently, and each is unlike every other
because it is acted upon differently by environ-
ment and food supply. It is not strange, there-
fore, that some of these individuals should now
and then depart very widely from the ordinary
type, and thereby attract the attention of a gar-
dener, who would forthwith make cuttings or set
grafts from the part. Every branch is a bud-
variety, just as truly as every seedling is a seed-
variety, — since no seedling is ever exactly like its
parent, —and there should be no greater mystery
connected with the sports of buds than there is
with the variations from seeds, for the causes
which produce the one may be and are equally
competent to produce the other.

d. Struggle for Infe a Cause of Variation.

We have seen that the world is full of plants.
There is room for more only as the present indi-
viduals die. Yet nearly every species produces

30 FACT AND PHILOSOPHY OF VARIATION.

a great number of seeds, and makes a most stren-
uous effort to multiply its kind. Any one plant,
if left to itself, is capable of covering the earth in
a comparatively short time. <A fierce struggle for
a chance to live is therefore inevitable. This con-
flict is most apparent to the general observer in
the springtime, when every “herb yielding seed
after his kind, and the tree yielding fruit, whose
seed was in itself, after his kind,” are sending
forth a host of sturdy offspring. The very land
seems to be pregnant with weeds and aspiring
young growths. But by midsummer the num-
bers may be less. The weaker and less fortunate
ones have perished, and the victors have waxed
stronger thereby. The annual and half of the
biennial species complete their course upon the
approach of winter, and the older perennial herbs
are becoming weak; so in the succeeding spring-
time there is again a fierce combat for the vacant
places.

One of the results of this conflict is the adjust-
ment of plants to each other. We have seen how
the climbing plant insinuates itself in amongst the
shrubberies and ties them together in an impene-
trable tangle in order that it, itself, may have a
chance to live. So the low plants of the deep
forest are such as have been plastic enough to
adapt themselves to the damp shades. Thus
plants have developed companionships or diver-

FIXATION OF VARIATIONS. 31

gences in characters, by means of which, under
the stress of circumstances, they are able to live
together. Plants have adapted themselves to
other plants as truly as they have adapted them-
selves to soil or climate; and if these latter en-
vironments are ever the sources or causes of
variation, then the first must be also. I must
look upon the struggle for existence, therefore,
as itself a cause of individual differences, since
we know that any continued pressure from with-
out awakens an adaptive response in the form of
the vegetable organism.

Ill. THrt CHoIcE AND FIXATION OF VARIA-
TIONS.

We have now seen that every lhving object is
unlike every other. In plants, even every branch
is unlike any other branch. We have endeay-
ored to discover some of the causes of these uni-
versal differences. We have found that they
are intimately associated with the welfare of the
type or species, inasmuch as they appear, for
the most part, to be the means of fitting the
plant to live in the conditions in which it is
placed. But we have also seen that there are
more individuals than can find a place to live.
How, then, does nature choose the best from the
poorest, and, having chosen them, how does she

32 FACT AND PHILOSOPHY OF VARIATION.

endeavor to fix them, or to make them more or
less stable ?

“This preservation of favorable individual dif-
ferences and variations, and the destruction of
those which are injurious, I have called Natural
Selection or the Survival of the Fittest.” This
is the philosophy which was propounded by Dar-
win, and which will carry his name to the last
generation of men. It looks simple enough.
Those forms which are best fitted to live, do live,
because they crowd out the others. Yet, this
simple principle of natural selection was the first
explanation of the process of evolution which
seemed to be capable of interpreting the complex
phenomena of the forms of organic life. For a
time, this philosophy was thought to be the one
fundamental motive of the evolution or progres-
sion of life, but we are now convinced that there
are other motives or forces at work; but it seems
to be indisputable that natural selection is the
chief force underlying the evolution of plants,
and it is the only one with which the person who
desires to breed plants need intimately concern
himself.

We must now determine what a variety is. This
is a vexed question, and one which seems never to
be: capable of an answer which is satisfactory to
the gardener. ‘Time and again, some person has
introduced what he considered to bea distinct new

WHAT IS A VARIETY ? 33

variety, only to find that other horticulturists dis-
pute him and declare that it is only some old vari-
‘ety renamed. And yet the introducer knows that
he has not renamed an old variety, but that he has
simply propagated a form which appeared or origi-
nated upon his own grounds.

Now, let us see. Nature starts out with the
individual to make a new form. Every individual
is unlike every other one. When the individual
differences are so well marked that we can readily
describe and distinguish them, and so permanent
that they pass down nearly intact to a few genera-
tions, we say that we have a variety. If the differ-
ences are still more marked, we say that we have a
species. Where the variety ends and the species
begins it may be utterly impossible to determine ;
so we mark off at a certain point and say, arbi-
trarily, that this much is variety and that much
is species. Asa Gray once said to me that ‘‘ species
are judgments.’’ Now, if there is no hard and
fast line between the variety and the species, so
there is none between the individual and the vari-
ety; for a variety is only the family of descendants
from some one individual. That is, the idea of
variety or species rests upon difference, but just
how much difference shall constitute one grade or
another is a matter of individual opinion. So,
when two gardeners cannot agree as to whether a
given introduction is a new variety or not, they

D

34 FACT AND PHILOSOPHY OF VARIATION.

are having just the same difficulty that two botan-
ists have when they cannot decide whether two
plants are two species or one.

It is apparent, then, that every individual plant
is a distinct variety, only that the differences be-
tween it and other individuals may be so slight
that they have no practical utility and cannot be
described and recorded. Just as soon as an indi-
vidual plant has characters so unlike its kin that
it has some commercial value, then the plant will
be increased by cuttings or grafts or seeds, the
brood of offspring will be given a name, and a new
variety is born.

Individuals with the same general features may
appear simultaneously in two or more places, and
two or more men may propagate, name, and intro-
duce them. When they are all brought together
and compared, it will be said that they are all the
same variety, that, according to the rules of nomen-
clature, the brood which chanced to be named first
must “stand” or be held to be the type of the
variety, and that the other names must become
synonyms. Yet some person may discover minor
differences in them and demand that the varieties
be kept distinct. So the see-saw goes on —a vari-
ety isa variety so long as it answers some purpose
in use or trade, and it is not a variety when it is so
much like some other variety that it has no merit
which the other does not possess.

———————————E ee

WHAT IS A VARIETY ? ao

As soon as a plant appears with some feature
which is more desirable than anything which has
preceded it, therefore, it may be made the begin-
ning of a new variety. Man chooses it, and then
propagates it. This is human selection. If nat-
ure did the same, it would be natural selection.

Now, how does nature preserve or fix this type?
She does not preserve it! She simply chooses it
as a beginning and gradually modifies it and shapes
it into the form which she needs. She has no per-
manent forms. ‘There is a general onward pro-
gression of every type either towards other types
or towards extinction. We have seen that nature
is constantly choosing and selecting. If she selects
an individual for the beginning of a race, then
she selects just as keenly from every offspring of
that individual, and so on to the end of time. The
process never stops. So nature fixes her forms by
keeping them moving, growing, constantly develop-
ing farther away from their beginnings.

Now, man does the same thing. <A plant in a
cabbage row pleases him. It has a solid, small
head and stout stem. He stores it away for seed.
Amongst the offspring, perhaps fifty per cent are
as good as the parent. These are saved. So the
process goes on, from season to season. In four
or five generations of plants, he finds that ninety
per cent of the seeds “come true.’ Then he
names it and introduces it. It is well advertised

36 FACT AND PHILOSOPHY OF VARIATION.

in the seed catalogues. Many people buy the
seeds. Some of these persons will grow their
own seed, and every one of them has a different
ideal in mind when selecting the seed parents.
So, in the course of a few years, it is found that
there are really several more or less different
forms going under the same name. Some person
may observe this difference and legitimately in-
troduce one or more of the forms as distinct
varieties. Some other person, however, who has
known the history of the stock and who is not
aware that varieties pass into other forms, objects
to the new names and declares that the introducer
is imposing upon the public.

This is the history of ninety-nine out of every
hundred varieties which are habitually propa-
gated by seeds, like the kitchen-garden vege-
tables and the annual flowers. Some peculiar
individual, appearing we know not why, is dis-
covered, and seeds are saved and _ selection —
perhaps unconscious selection— begins. After
a time the variety is broken up into several, or
else, if it varies only slightly into divergent
forms, the whole body or generations of the
variety move onward, gradually departing from
the initial type until it is no longer the same,
although it may still bear the same name. The
life of seed-varieties, in their pure and original
form, is very short. Even the best of them are

THE PASSING OF VARIETIES. 31

usually measured by one or two decades. They
run out or pass out by variation, into other forms.
The Trophy tomato is not the Trophy tomato
which was introduced over twenty years ago,
although it bears the old name and is a direct
descendant of the first stock.

In plants multiplied by buds — that is, by bud-
ding, grafting, cuttings, tubers, and the like —
there is less variation in the offspring than in
those propagated by seeds. Yet we have seen
that no two Baldwin apple trees—all of which
are but divisions, more or less remote, of the one
original tree —are alike, and now and then one
branch of a fruit tree may “sport,” or develop a
strange bud-variety. We know, too, that the
same variety of fruit tree takes on different
characters in different geographical regions, so
that the Greening apple is no longer the Green-
ing of Rhode Island in the West and South. So,
it is apparent that even when we divide a plant
into many parts and distribute the members far
and wide, and when there is no occasion for con-
cerning ourselves with fixing the type,—even
here there is variation. In some cases, particu-
larly in those in which we multiply the plant by
dividing abnormally developed parts, there is a
tendency to scatter or to vary in many directions,
and also a tendency to run out by degeneration.
This is admirably true of the potato, varieties of

88 FACT AND PHILOSOPHY OF VARIATION.

which, in ten years or less, become so mixed in
their characters, through rapid variation and de-
terioration, that we must return to seedling pro-
ductions for a new start.

The gist of all this is the fact that nature does
not make new varieties or species by leaps and
bounds or sudden starts, but that she gradually
produces the new out of the old, so slowly that
were a man to live a thousand years he might
note little change in the grosser features of plants.
She employs crossing, the changes in all the varied
conditions of life, and whatever other forces she
may possess, to give small differences between
individuals. ‘Then the slow and cumulative
process of selection carries the work forward
forever. Man must follow the same course, in
the main. He is only rarely the direct means of
originating variations. He finds them amongst
the normal plants of the field and garden. His
skill and science are exercised in the selection
and so-called breeding of the offspring, not in
the original genesis of the new form. It is only
in those plants which he multiplies by simple
division that he gains much direct profit by
crossing or hybridizing. It is the slow and pa-
tient care and selection, day by day, which per-
manently ameliorate and improve the vegetable
world. Nature starts the work; man may com-
plete it.

a

LECTURE II.

THE PHILOSOPHY OF THE CROSSING OF PLANTS,
CONSIDERED IN REFERENCE TO THEIR IMPROVE-
MENT UNDER CULTIVATION.

I. THE STRUGGLE FOR LIFE.

It is now understood that the specific forms or
groups of plants have been determined largely by
the survival of the fittest in a long and severe
struggle for existence. The proof that this strug-
gle everywhere exists becomes evident upon a
moment's reflection. We know that all organisms
are eminently variable. In fact, no two plants or
animals in the world are exactly alike. We also
know that very few of the whole number of seeds
which are produced in any area ever grow into
plants. If all the seeds produced by the eims
upon Boston Common in any fruitful year were to
erow into trees, this city would become a forest as
aresult. If all the seeds of the rarest orchid in
our woods were to grow, in a few generations of
plants even our farms would be overrun. If all
the rabbits which are born were to reach old age,
and all their offspring were to do the same, in less
than ten years every vestige of herbage would be

39

40 PHILOSOPHY OF CROSSING PLANTS.

swept from the country, and our farms would
become barren. ‘There is, then, a wonderful latent
potency in these species; but the same may be
said of every species of plant and animal, even of
man himself. If one species of plant would over-
run and usurp the land if it increased to the full
extent of its possibilities, what would be the result
if each of the two thousand and sixty-one plants
known to inhabit Middlesex County were to do the
same? And then fancy the result if each of the
animals, from rabbits and mice to frogs and leeches,
were to increase without check! The plagues of
Egypt would be insignificant in the comparison !
The fact is, the world is not big enough to hold
the possible first offspring of the plants and ani-
mals at this moment living upon it. Struggle for
existence, then, is inevitable, and it must be severe.
It follows as a necessity that those seeds grow or
those plants live which are best fitted to grow and
live, or which are fortunate enough to find a con-
genial foothold. It would appear, at first thought,
that much depends upon the accident of falling
into a congenial place, or one unoccupied by other
plants or animals; but, inasmuch as scores of
plants are contending for every unoccupied place,
it follows that everywhere only the fittest can
germinate or grow. In the great majority of cases,
plants grow in a certain place because they are
better fitted to grow there, to hold their own, than

STRUGGLE FOR EXISTENCE. 41

any other plants are ; and the instances are rare in
which a plant is so fortunate as to find an un-
occupied place. We are apt to think that plants
chance to grow where we find them, but the chance
is determined by law, and therefore is not chance.

Much of the capability of a plant to persist
under all this struggle depends, therefore, upon
how much it varies; for the more it varies the
more likely it is to find places of least struggle.
It grows under various conditions,—in sun and
shade, in sand and clay, by the sea-shore or upon
the hills, in the humidity of the forest or the
aridity of the plain. In some directions it very
likely finds less struggle than in others, and in
these directions it expands itself, multiplies, and
gradually dies out in other directions. So it
happens that it tends to take on new forms, or to
undergo an evolution. In the meantime, all the
intermediate forms, which are at best only indif-
ferently adapted to their conditions, tend to dis-
appear. In other words, gaps appear which we
eall “ missing links.” The weak links break and
fall away, and what was once a chain becomes a se-
ries of rings. So the “ missing links” are amongst
the best proofs of evolution.

The question now arises as to the cause of these’
numerous variations in animals and plants. Why
are no two individuals in nature exactly alike?
The question is exceedingly difficult to answer.

42 PHILOSOPHY OF CROSSING PLANTS.

It was once said that plants vary because it is
their nature to vary; that variation is a necessary
function, as much as growth or fructification.
This really removes the question beyond the reach
of philosophy ; and direct observation leads us to
think that some variation, at least, is due to ex-
ternal circumstances. (See Lecture I.) We are
now looking for the cause of variation as a part of
the scheme of evolution; and we are wondering if
the varied surroundings, or, as Darwin put it, the
“changed conditions of life,’ may not actually
induce variability. This conclusion would seem
to follow from the fact of the severe and universal
struggle in nature whereby plants are constantly
forced into new and strange conditions. But there
is undoubtedly much variation which has sprung
from more remote causes, one of which it is my
purpose to discuss here.

II. THe DIVISION oF LABOR.

In the lowest animals and plants— which are
simply single cells—the species multiples by
means of simple division or by budding. One in-
dividual, of itself, becomes two, and the two are
therefore recasts of the one. But, as organisms
multiplied and conditions became more complex,
that is, as struggle increased, there came a differen-
tiation in the parts of the individual, so that one

DIVISION OF LABOR. 43
cell or one cluster of cells performed one labor
and other cells performed other labor; and this
tendency resulted in the development of organs.
Simple division, therefore, could no longer repro-
duce the whole complex individual; and, as all
organs are necessary to the existence of life, the
organism may die if it is divided. Along with
this specialization came the differentiation into
sex ; and sex clearly has two offices: to hand over,
by some mysterious process, the complex organ-
ization of the parent to the offspring, and also to
unite the essential characters or tendencies of two
beings into one. The second office is manifestly
the greater, for, as it unites two organizations into
one, it insures that the offspring is somewhat un-
like either parent, and is therefore better fitted to
seize upon any place or condition new to its kind.
And as the generations increase, the tendency
to variation in the offspring must be constantly
greater, because the impressions of a greater num-
ber of ancestors are transmitted to it. I have said
that this office of sex to induce variation is more
important than the mere fact of reproduction of a
complex organization; for it must be borne in
mind that the complexity of organization is itself
a variation and adaptation made necessary by the
increasing struggle for existence.

If, therefore, the philosophy of sex is to promote
variation by the union of different individuals, it

As PHILOSOPHY OF CROSSING PLANTS.

must follow that the greatest variation must come
from parents considerably unlike each other in
their minor characters. Thus it comes that in-
breeding tends to weaken a type, and cross-breeding
tends to strengthen it. And at this point we meet
the particular subject which I am to present to
you. I have introduced you to this preliminary
sketch because I contend that we can understand
crossing only as we make it a part of the general
philosophy of nature. There are the vaguest
notions concerning the possibilities of crossing,
some of which I hope to correct by presenting the
subject in its relations to the general aspects of
the vegetable world.

We are now prepared to understand that crossing
is good for the species, because it constantly revi-
talizes offspring with the strongest traits of the
parents, and ever presents new combinations which
enable the individuals to stand a better chance of
securing a place in the polity of nature. The fur-
ther discussions of the subject are such as have to
do with the extent to which crossing is possible and
advisable, and the general results of the operation.

III. THe Limits or CROSSING.

If crossing is good for the species, which philoso-
phy and direct experiment abundantly show, it is
necessary at once to find out to what extent it can

LIMITS OF GROSSING. 45

be carried. Does the good increase in proportion
as the cross becomes more violent, or as the parents
are more and more unlike? Or do we soon find a
limit beyond which it is not profitable or even
possibie to go,—a point at which we say that “an
inch is as good as an ell”? If great variability is
good for the species in the struggle for existence,
and if crossing induces variability because of the
union of unlike individuals, it would seem to follow
that the more unlike the parents are, the greater
will be the variation in offspring and the more the
type will prosper; and, carrying this thought to
its logical conclusion, we should expect to find that
the most closely related plants would constantly
tend to refuse to cross, because the offspring of
them would be little variable and therefore little
adapted to the struggle for existence; while the
most widely separated plants would constantly tend
to cross more and more, because their offspring
would present the greatest possible degrees of dif-
ferences. We should expect, for instance, that a
Baldwin apple would be less likely to cross with a
Greening than it is to cross with a peach or a gourd.
And, if we should carry our thought a step farther,
we should at once see that this crossing between
different species would soon fill in all differences
between those species, and that definite specific
types would cease to exist. This would be pande-
monium, and crossing would be the cause of it!

46 PHILOSOPHY OF CROSSING PLANTS.

Now, essentially this reasoning has been ad-
vanced to combat the evolution of plants and
animals by means of natural selection; and this
proposition that intermixing must constantly tend
to obliterate all differences between plants and to
prevent the establishment of well-marked types,
has been called the “swamping effects of inter-
crossing.” It is exceedingly important that we
consider this question, for it really lies at the
foundation of the improvement of cultivated plants
by means of crossing, as well as the persistence
and evolution of varieties and species under wholly
natural conditions.

We find, however, that distinct species, as a rule,
refuse to cross; and the first question which natu-
rally arises is, What is the immediate cause of the
refusal of plants to cross? How does this refusal
express itself? It comes about in many ways.
The commonest cause is the positive refusal of
a plant to allow its ovules to be impregnated by
the pollen of another plant. The pollen will not
“take.” For instance, if we apply the pollen of a
Hubbard squash to the flower of the common field
pumpkin, there will simply be no result, —the
fruit will not form. The same is true of the pear
and the apple, the oat and the wheat, and most
very unlike species. Or the refusal may come in
the sterility of the cross or hybrid: the pollen may
“take” and seeds may be formed and the seeds

BARRIERS TO INTERCROSSING. AT

may grow, but the plants which they produce may
be wholly barren, sometimes even refusing to pro-
duce flowers as well as seeds, as in the instance of
some hybrids between the Wild Goose plum and
the peach. Sometimes the refusal to cross is due
to some difference in the time of blooming or some
incompatibility in the structure of the flowers. But
it is enough for our purpose to know that there are
certain characters in widely dissimilar plants which
prevent intercrossing, and that these characters are
just as positive and just as much influenced by
change of environment and natural selection as are
size, color, productiveness, and other characters.
Here, then, is the sufficient answer to the
proposition that intercrossing must swamp all
natural selection, and also the explanation of the
varying and often restricted limits within which
crossing is possible. That is, the checks to cross-
ing have been developed through the principle
of universal variability and natural selection, as
has been shown by Darwin and Wallace. Plants
vary in their reproductive organs and powers just
the same as they do in other directions; and when
such a variation is useful it is perpetuated, and
when hurtful it is lost. Suppose that a certain
well-marked individual of a species should find an
unusually good place in nature, and it should mul-
tiply rapidly. Crosses would be made between
its own offspring and perhaps between those off-

48 PHILOSOPHY OF CROSSING PLANTS.

spring and itself in succeeding years; and it is
fair to suppose that some of the crosses would be
particularly well adapted to the conditions in
which the parent grew, and these would constantly
tend to perpetuate themselves, while less adaptive
forms would constantly tend to disappear. Now,
the same thing would take place if this individual
or its adaptive offspring were to cross with the
main stock of the parent species; for all the off-
spring of such a cross which are intermediate in
character and therefore less adapted to the new con-
ditions would tend to disappear, and the two types
would, as a result, become more and more fixed and
the tendency to cross would constantly decrease.
The refusal to cross, therefore, becomes a posi-
tive character of separation, and the “missing
links” which result from crossing are no more
or no less inexplicable than the “missing links ”
due to simple selection; or, to put the case more
accurately, natural selection weeds out the ten-
dency to promiscuous crossing, when it is hurtful,
in just the same manner that it weeds out any
other injurious tendency. It makes no difference
in what way this tendency expresses itself, whether
in some constitutional refusal to cross, —if such
exists, — or in. infertility of offspring, or in differ-
ent: times of blooming: all equally come under the
power of natural selection. We are apt to look
upon infertility as the absence of a character, a

HYBRIDS RARELY USEFUL. 49

sort of a negative feature which is somehow not
the legitimate property of natural selection; but
such is not the case. We are perhaps led the
more to this feeling because the word infertility
is itself negative, and because we associate full pro-
ductiveness with the positive attributes of plants.
But loss of productiveness is surely no more a sub-
ject of wonder than loss of color or size, if there is
some corresponding gain to be accomplished. In
fact, we see, in numerous plants which propagate
easily by means of runners and suckers, a very low
degree of productiveness, that is, infertility.

Now, if this reasoning is sound, it leads us to
conclusions quite the reverse of those held by the
advocates of the swamping effects of intercrossing,
and these conclusions are of the most vital impor-
tance to every man who tills the soil. The logical
result is simply this: the best results of crossing
are obtained, as a rule, when the cross is made be-
tween different individuals of the same variety,
or at farthest, between different individuals of the
same species. In other words, hybrids — or crosses
between species — are rarely useful in nature, and
it follows that the more unlike the species the less
useful will be the hybrids. This, I am aware, is
counter to the notions of most horticulturists, and,
if true, must entirely overthrow our common think-
ing upon this subject. But I think that I shall be
able to show that observation and experiment lead

E

50 PHILOSOPHY OF CROSSING PLANTS.

to the same conclusion to which our philosophy
has brought us.

IV. FUNCTION OF THE CROSS.

a. The Gradual Amelioration of the Type.

At this point we must ask ourselves what we
mean by “best results.” I take this phrase to
refer to those plants which are best fitted to sur-
vive in the struggle for existence, those which are
most vigorous or most productive or most hardy,
or which possess any well-marked character or
characters which distinguish them in virility from
their fellows. We commonly associate the term
more particularly with marked vigor and produc-
tiveness; these are the characters most useful in
nature and also in cultivation, the ones which we
oftenest desire to obtain. Another type of varia-
tion which we constantly covet is something which
we can call a new character, which will lead to the
production of a new cultural variety, and we are
always looking to this as the legitimate result of
crossing. We have forgotten —if, indeed, we ever
knew —that the commoner, all-pervading, more
important function of the cross is to infuse some
new strength or power into the offspring, to improve
or to perpetuate an existing variety, rather than to
create anew one. Or, if a new one is created, it

FUNCTION OF THE CROSS. 51

comes from the gradual passing of one into another,
an inferior variety into a good one, a good one into
a superlative one. So nature employs crossing in
a process of slow or gradual improvement, one step
leading to another, and not in any bold or sudden
creation of new forms. And there is evidence to
show that something akin to this must be done to
secure the best and most permanent results under
cultivation. The notion is somehow firmly rooted
in the popular mind that new varieties can be pro-
duced with the greatest ease by crossing parents of
given attributes. There is something captivating
about the notion. It smacks of a somewhat magic
power which man evokes as he passes his wand
over the untamed forces of nature. But the wand
is often only a gilded stick, and is apt to serve no
better purpose than the drum major’s pretentious
baton !

Let me say further that crossing alone can
accomplish comparatively little. The chief power
in the evolution or progression of plants appears
to be selection, or, as Darwin puts it, the law of
“preservation of favorable individual differences
and variations, and the destruction of those which
are injurious.” Selection is the force which aug-
ments, develops, and fixes types. Man must not
only practice a judicious selection of parents from
which the cross is to come, which is in reality but
the exercise of a choice, but he must constantly

nw PHILOSOPHY OF CROSSING PLANTS.

select the best from among the crosses, in order to
maintain a high degree of usefulness and to make
any advancement; and it sometimes happens that
the selection is much more important to the culti-
vator than the crossing. I do not wish to discour-
age the crossing of plants, but I do desire to dispel
the charm which too often hangs about it.

Further discussion of this subject naturally falls
under two heads: the improvement of existing
types or varieties by means of crossing, and the
summary production of new varieties. I have
already stated that the former office is the more
important one, and the proposition is easy of proof.
It is the chief use which nature makes of crossing,
— to strengthen the type. Think, for instance, of
the great rarity of hybrids or pronounced crosses
in nature. No doubt all the authentic cases on
record could be entered in one or two volumes, but
a list of all the individual plants of the world
could not be compressed into ten thousand volumes.
There are a few genera, in which the species are
not well defined or in which some character of
inflorescence favors promiscuous crossing, in which
hybrids are conspicuous; but even here the num-
ber of individual hybrids is very small in compari-
son to the whole number of individuals. That is,
the, hybrids are rare, while the parents may be
common. This is well illustrated even in the
willows and oaks, in which, perhaps, hybrids are

eee

RARITY OF HYBRIDS. 53

better known than in any other American plants.
The great genus carex or sedge, which occurs in
great numbers and many species in almost every
locality in New England, and in which the species
are particularly adapted to intercrossing by the
character of their inflorescence, furnishes but few
undoubted hybrids. Among one hundred and
eighty-five species and prominent varieties inhabit-
ing the northeastern states, there are only eleven
hybrids recorded, and all of them are rare or local,
some of them having been collected but once.
Species of carex of remarkable similarity may grow
side by side for years, even intertangled in the
same clump, and yet produce no hybrid. These
instances prove that nature avoids hybridization,
—a conclusion at which we have already arrived
from philosophical considerations. And we have
reason to infer the same conclusion from the fact
that flowers of different species are so constructed
as not to invite intercrossing. But, on the other
hand, the fact that all higher plants habitually
propagate by means of seeds, which is far the most
expensive to the plant of all methods of propa-
gation, while at the same time most flowers are so
constructed as to prevent self-fertilization, proves
that some corresponding good must come from
crossing within the limits of the species or variety ;
and there are purely philosophical reasons, as we
have seen, which warrant a similar conclusion.

54. PHILOSOPHY OF CROSSING PLANTS.

But experiment has given us more direct proof of
our propositions, and we shall now turn our atten-
tion to the garden.

Darwin was the first to show that crossing within
the limits of the species or variety results in a con-
stant revitalizing of the offspring, and that this is
the particular ultimate function of cross-fertiliza-
tion. Kdélreuter, Sprengel, Knight, and others had
observed many, if, indeed, not all, the facts obtained
by Darwin; but they had not generalized upon
them broadly, and did not conceive their relation
to the complex life of the vegetable world. Dai-
win’s results are, concisely, these ; self-fertilization
tends to weaken the offspring; crossing between
different plants of the same variety gives stronger
and more productive offspring than arises from
self-fertilization; crossing between stocks of the
same variety grown in different places or under dif-
ferent conditions gives better offspring than cross-
ing between different plants grown in the same
place or under similar conditions; and his re-
searches have also shown that, as a rule, flowers
are so constructed as to favor cross-fertilization.
In short, he found, as he expressed it, that ‘ nature
abhors perpetual self-fertilization.” Some of his
particular results, although often quoted, will be
useful in fixing these facts in our minds. Plants
from crossed seeds of morning-glory exceeded in
height those from self-fertilized seeds as 100 exceeds

ae. a, ee

INCREASED VIGOR OF CROSS-BREEDS. 5d

76, in the first generation. Some flowers from these
plants were self-pollinated and some were crossed,
and in this second generation the crossed plants
were to the uncrossed as 100 is to 79; the opera-
tion was again repeated, and in the third generation
the figures stand 100 to 68; fourth generation, the
plants having been grown in midwinter, when none
of them did well, 100 to 86; fifth generation, 100
to T5; sixth generation, 100 to 72; seventh genera-
tion, 100 to 81; eighth generation, 100 to 85; ninth
generation, 100 to 79; tenth generation, 100 to
54. The average total gain in height of the
crossed over the uncrossed was as 100 to TT, or:
about 30 per cent. There was a corresponding
gain in fertility, or the number of seeds and seed-
pods produced. Yet, striking as the results are,
they were produced by simply crossing between
plants grown near together, and under what would
ordinarily be called uniform conditions. In order
to determine the influence of crossing with fresh
stock, plants of the same variety were obtained
from another garden, and these were crossed with
the ninth generation mentioned above. The off-
spring of this cross exceeded those of the other
crossed plants as 100 exceeds 78, in height; as 100
exceeds 57, in the number of seed-pods; and as 100
exceeds 51, in the weight of the seed-pods. In
other words, crosses between fresh stock of the same
variety were nearly 380 per cent more vigorous than

56 PHILOSOPHY OF CROSSING PLANTS.

crosses between plants grown side by side for some
time and over 44 per cent more vigorous than
plants from self-fertilized seeds. On the other
hand, experiments showed that crosses between
different flowers upon the same plant gave actu-
ally poorer results than offspring of selt-fertilized
flowers. It is evident, from all these figures, that
nature desires crosses between plants, and, if pos-
sible, between plants grown under somewhat dif-
ferent conditions. All the results are exceedingly
interesting and important; and there is every
reason to believe that, as a rule, similar results can
be obtained with all plants.

Darwin extended his investigations to many
plants, only a few of which need be discussed here.
Cabbage gave pronounced results. Crossed plants
were to self-fertilized plants in weight as 100 is
to 87. A cross was now made between these
crossed plants and a plant of the same variety
from another garden, and the difference in weight
of the resulting offspring was the difference be-
tween 100 and 22, showing a gain of over 350 per
cent, due to a cross with fresh stock. Crossed
lettuce plants exceeded uncrossed in height as
100 exceeds 82. Buckwheat gave an increase in
weight of seeds as 100 to 82, and in height of
plants as 100 to 69. Beets gave an increase in
height represented by 100 and 87. Maize, when
full grown, from crossed and uncrossed seeds,

INCREASED VIGOR OF CROSS-BREEDS. 57

gave the differences in height between 100 and 91.
Canary-grass gave similar results.

I have obtained results as well marked as these
upon a large and what might be called a commer-
cial scale. I raised the plants during the first
generation of seeds from known parentage, the
flowers from which they came having been care-
fully pollinated by hand. In some instances the
second generations were grown from hand-crossed
seeds, but in other cases the second generations
were grown from seeds simply selected from the
first-year patches. As the experiments have been
made in the field and upon a somewhat extensive
scale, it was not possible to accurately measure
the plants and the fruits from individuals in all
eases; but the results have been so marked as to
admit of no doubt as to their character. In 1889
several hand-crosses were made among egg-plants.
Three fruits matured, and the seeds from them
were grown in 1890. Some two hundred plants
were grown, and they were characterized through-
out the season by great sturdiness and vigor of
erowth. They grew more erect and taller than
other plants near by grown from commercial seeds.
They were the finest plants which I had ever seen.
It was impossible to determine productiveness,
from the fact that our seasons are too short for
ego-plants, and only the earliest flowers, in the
large varieties, perfect their fruit, and the plant

08 PHILOSOPHY OF CROSSING PLANTS.

blooms continuously through the season. In order
to determine how much a plant will bear, it must
be grown until it ceases to bloom. When frost
came, I could see little difference in productive-
ness between these crossed plants and commercial
plants. A dozen fruits were selected from various
parts of this patch, and in 1891 about twenty-five
hundred plants were grown from them. Again
the plants were remarkably robust and healthy,
with fine foliage, and they grew erect and tall, —
an indication of vigor. They were also very pro-
ductive; but, as the cross had been made between
unlike varieties, and the offsprings were therefore
unlike either parent, I could not make an accurate
comparison. But they compared well with com-
mercial egg-plants, and I am satisfied that they
would have shown themselves to be more produc-
tive than common stock could they have grown a
month or six weeks longer. Professor Munson, of
the Maine Agricultural College, grew some of this
crossed stock in 1891, and he told me that it was
better than any commercial stock in his gardens.

In extended experiments in the crossing of
pumpkins, squashes, and gourds, carried on dur-
ing several years, increase in productiveness due
to crossing has been marked in many instances.
Marked increase in productiveness has been ob-
tained from tomato crosses, even when no other
results of crossing could be seen.

a ee ee ee ee ee

ee

BENEFITS FROM CHANGE OF SEED. 59

b. Change of Seed and Crossing.

Bearing in mind these good influences of cross-
ing, let us recall another series of facts following
the simple change of seed. Almost every farmer
and gardener at the present day feel that an
occasional change of seed results in better crops,
and there are definite records to show that such
is often the case. In fact, I am convinced that
much of the rapid improvement in fruits and vege-
tables in recent years is due to the practice of
buying plants and seeds so largely of dealers, by
means of which the stock is often changed. Even
a shght change, as between farms or neighboring
villages, sometimes produces marked results, such
as more vigorous plants and often more fruitful
ones. We must not suppose, however, that be-
cause a small change gives a good result, a violent
or very pronounced change gives a better one.
There are many facts on record to show that great
changes often profoundly influence plants, and
when such influence results in lessened vigor or
lessened productiveness we call it an injurious one.
Now, this injurious influence may result even
when all the conditions in the new place are
favorable to the health and development of the
plant; it is an influence which is wholly indepen-
dent, so far as we can see, of any condition which
interferes injuriously with the simple processes of

60 PHILOSOPHY OF CROSSING PLANTS.

growth. Seeds of a native physalis or husk-tomato
were sent to me from Paraguay in 1889 by Dr.
Thomas Morong, then travelling in that country.
I grew it both in the house and out of doors, and
for two generations was unable to make it set
fruit, even though the flowers were hand-polli-
nated; yet the piants were healthy and grew
vigorously. The third generation grown out of
doors set fruit freely. This is an instance of the
fact that very great changes of conditions may
injuriously affect the plant, and an equally good
illustration of the power to overcome these condi-
tions. Now, there is great similarity between the
effects of slight and violent changes of conditions
and small and violent degrees of crossing, as both
Darwin and Wallace have pointed out, and it is
pertinent to this discussion to endeavor to dis-
cover why this similarity exists.

It is well proved that crossing is good for the
resulting offspring, because the differences be-
tween the parents carry over new combinations
of characters or at least new powers into the
crosses. It is a process of revitalization, and the
more different the stocks in desirable characters
within the limits of the variety, the greater is
the revitalization; and frequently the good is of
a more positive kind, resulting in pronounced
characters which may serve as the basis for new
varieties. In the cross, therefore, a new combina-

BENEFITS FROM CHANGE OF SEED. 61

tion of characters or a new power fit it to live
better than its parents in the conditions under
which they lived.

In the case of change of stock we find just the
reverse, which, however, amounts to the same
thing, — that the same characters or powers fit the
plant to live better in conditions new to it than
plants which have long lived in those conditions.
In either case, the good comes from the fitting
together of new characters or powers and new
environments. Plants which live during many
generations in one place become accustomed to the
place, thoroughly fitted into its conditions, and are
in what Mr. Spencer calls a state of equilibrium.
When either plant or conditions change, new ad-
justments must take place; and the plant may find
an opportunity to take advantage, to expand in
some direction in which it has before been held
back; for plants always possess greater power
than they are able to express. ‘These rhythmical
actions or functions [of the organism],” writes
Spencer, “‘and the various compound rhythms
resulting from their combinations, are in such
adjustment as to balance the actions to which
the organism is subject. There is a constant or
periodic genesis of forces which, in their kinds,
amounts, and directions, suffice to antagonize the
forces which the organism has constantly or peri-
odically to bear. If, then, there exists this state

62 PHILOSOPHY OF CROSSING PLANTS.

of moving equilibrium among a definite set of
internal actions, exposed to a definite set of ex-
ternal actions, what must result if any of the
external actions are changed? Of course there is
no longer an equilibrium. Some force which the
organism habitually generates is too great or too
small to balance some incident force; and there
arises a residuary force exerted by the environ-
ment on the organism, or by the organism on the
environment. This residuary force, this unbal-
anced force, of necessity expends itself in produc-
ing some change of state in the organism.”

The good results, therefore, are processes of
adaptation, and when adaptation is perfectly com-
plete the plant may have gained no permanent
advantage over its former condition, and new
crossing or another change may be necessary; yet
there is often a permanent gain, as when a plant
becomes visibly modified by change to another cl-
mate. Now, this adaptive change may express
itself in two ways: either by some direct influence
upon the stature, vigor, or other general character,
or indirectly upon the reproductive powers, by
which some new influence is carried to the off-
spring. If the direct influences become heredi-
tary, as observation seems to show may sometimes
occur, the two directions of modification may
amount, ultimately, to the same thing.

For the purposes of this discussion it is enough

CHANGE OF STOCK AND CROSSING. 63

to know that crossing within the variety and
change of stock within ordinary bounds are bene-
ficial, that the results in the two cases seem to
flow from essentially the same causes, and that
crossing and change of stock combined give much
better results than either one alone; and this
benefit is expressed more in increased yield and
vigor than in novel and striking variations. These
processes are much more important than any mere
eroping after new varieties, as I have already said;
not only because they are surer, but because they
are universal and necessary means of maintaining
and improving both wild and cultivated plants.
Even after one succeeds in securing and fixing
a new variety, he must employ these means to a
ereater or less extent to maintain fertility and
vigor, and to keep the variety true to its type.
In the case of some garden crops, in which many
seeds are produced in each fruit and in which
the operation of pollination is easy, actual hand-
crossing from new stock now and then may be
found to be profitable. But in most cases the
operation can be left to nature, if the new stock
is planted among the old. Upon this point Dar-
win expressed himself as follows: “It is a common
practice with horticulturists to obtain seeds from
another place having a very different soil, so as to
avoid raising plants for a long succession of gen-
erations under the same conditions; but with all

64 PHILOSOPHY OF GROSSING PLANTS.

the species which freely intercross by the aid of
insects or the wind, it would be an incomparably
better plan to obtain seeds of the required variety,
which had been raised for some generations under
as different conditions as possible, and sow them
in alternate rows with seeds matured in the old
garden. The two stocks would then intercross,
with a thorough blending of their whole organi-
zations, and with no loss of purity to the variety,
and this would yield far more favorable results
than a mere change of seed.”

ce. The Outright Production of New Varieties.

But you are waiting for a discussion of the
second of the great features of crossing, —the sum-
mary production of new varieties. This is the sub-
ject which is almost universally associated with
crossing in the popular mind, and even among
horticulturists themselves. It is the commonest
notion that the desirable characters of given
parents can be definitely combined in a pronounced
cross or hybrid. There are two or three philo-
sophical reasons which somewhat oppose this doc-
trine, and which we will do well to consider at the
outset. In the first place, nature is opposed to
hybrids, for species have been bred away from each
other in the ability to cross. If, therefore, there
is no advantage for nature to hybridize, we may

PRODUCTION OF NOVELTIES. 65

suppose that there would be little advantage for
man to do so; and there would be no advantage
for man did he not place the plant under conditions
different from nature, or desire a different set of
characters. We have seen that nature’s chief
barriers to hybridization are total refusal of species
to unite, and entire or comparative seedlessness
of offspring. We can overcome the refusal to
cross in many cases by bringing the plant under
cultivation; for the character of the species be-
comes so changed by the wholly new conditions
that its former antipathies may be overpowered.
Yet it is doubtful if such a plant will ever acquire
a complete willingness to cross. In like manner
we can overcome in a measure the comparative
seedlessness of hybrids, but it is very doubtful if
we can ever make such hybrids completely fruit-
ful. It would appear, therefore, upon theoretical
grounds, that in plants in which seeds are the parts
sought, no good can be expected, as a rule, from
hybridization; and this seems to be affirmed by
facts.1

It is evident that species which have been
differentiated or bred away from each other in
a given locality will have more opposed qualities
or powers than similar species which have arisen
quite independently in places remote from each

1See definition of hybrids, crosses, and other terms in the
Glossary.

F

66 PHILOSOPHY OF CROSSING PLANTS.

other. In the one case the species have likely
struggled with each other until each one has at-
tained to a degree of divergence which allows it
to persist; while in the other case there has been
no struggle between the species, but similar con-
ditions have brought about similar results. These
similar species which appear independently of
each other in different places are called representa-
tive species. Islands remote from each other but
similarly situated with reference to climate very
often contain representative species; and the same
may be said of other regions much like each other,
as eastern North America and Japan. Now, it
follows that, if representative species are less
opposed than others, they are more likely to
hybridize with good results; and this fact is re-
markably well illustrated in the Kieffer and allied
pears, which are hybrids between representative
species of Europe and Japan; and I am inclined
to think that the same may be found to be true
of the common or European apple and the wild
crab of the Mississippi valley. Various crabs of
the Soulard type, which I once thought to con-
stitute a distinct species, appear upon further study
to be hybrids. We will also recall that the hybrid
grapes which have so far proved most valuable are
those obtained by Rogers between the American
Vitis Labrusca and the European wine grape; and
that the attempts of Haskell and others to hybrid-

INSTABILITY OF HYBRIDS. 67

ize associated species of native grapes have given,
at best, only indifferent results. To these good
results from hybrids of fruit trees and vines I
shall revert presently.

Another theoretical point, which is borne out
by practice, is the conclusion that, because of the
ereat differences and lack of affinity between par-
ents, pronounced hybrid offsprings are unstable.
This is one of the greatest difficulties in the way
of the summary production of new varieties by
means of hybridization. It would appear, also,
that, because of the unlikeness of parents, hybrid
offspring must be exceedingly variable; but, as
a matter of fact, in many instances the parents
are so pronouncedly different that the hybrids
represent a distinct type by themselves, or else
they approach very nearly to the characters of
one of the parents. There are, to be sure, many
instances of exceedingly variable hybrid offspring,
but they are usually the offspring of variable par-
ents. In other words, variability in offspring
appears to follow rather as a result of variability
in parents than as a result of mere unlikeness
of characters. But the instability of hybrid off-
spring when propagated by seed is notorious.
Wallace writes that “the effect of occasional
crosses often results in a great amount of varia-
tion, but it also leads to instability of character,
and is therefore very little employed in the

68 PHILOSOPHY OF CROSSING PLANTS.

production of fixed and well-marked races.” I
may remark again that, because of the unequal
and unknown powers of the parents, we can never
predict what characters will appear in the hybrids.
This fact was well expressed by Lindley a half
century ago,in the phrase, “ Hybridizing is a
game of chance played between man and plants.”

V. CHARACTERISTICS OF CROSSES.

Bearing these fundamental propositions in mind,
let us pursue the subject somewhat in detail.
We shall find that the characters of hybrids,
as compared with the characters of simple crosses
between stocks of the same variety, are ambiguous,
negative, and often prejudicial. The fullest dis-
cussion of hybrids has been made by Focke (see
Lecture IV.), and he lays down the five following
propositions concerning the character of hybrid
offspring : —

1. “ All individuals which have come from the
crossing of two pure species or races, when pro-
duced and grown under like conditions, are
usually exactly like each other, or at least scarcely
more different from each other than plants of the
same species are.’ This proposition, although
perhaps true in the main, appears to be too
broadly and positively stated.

2. “The characters of hybrids may be different

HYBRIDS AND CROSS-BREEDS. 69

from the characters of the parents. The hybrids
differ most in size and vigor and in their sexual
powers.”

3. “Hybrids are distinguished from their par-
ents by their powers of vegetation or growth.
Hybrids between very different species are often
weak, especially when young, so that it is difficult
to raise them. On the other hand, cross-breeds
are, as a rule, uncommonly vigorous; they are dis-
tinguished mostly by size, rapidity of growth, early
flowering, productiveness, longer life, stronger
reproductive power, unusual size of some special
organs, and similar characteristics.”

4. “Hybrids produce a less amount of pollen
and fewer seeds than their parents, and they
often produce none. In cross-breeds this weaken-
ing of the reproductive powers does not occur.
The flowers of sterile or nearly sterile hybrids
usually remain fresh a long time.”

d. “Malformations and odd forms are apt to
appear in hybrids, especially in the flowers.”

Some of the relations between hybridization
and crossing within narrow limits are stated as
follows by Darwin: “It is an extraordinary fact
that with many species flowers fertilized with
their own pollen are either absolutely or in some
degree sterile; if fertilized with pollen from
another flower on the same plant, they are some-
times, though rarely, a little more fertile; if

TO PHILOSOPHY OF CROSSING PLANTS.

fertilized with pollen from another individual
or variety of the same species, they are fully
fertile; but if with pollen from a distinct species,
they are sterile in all possible degrees, until utter
sterility is reached. We thus have a long series
with absolute sterility at the two ends; at one
end due to the sexual elements not having been
sufficiently differentiated, and at the other end
to their having been differentiated in too great
a degree, or in some pecular manner.” |

The difficulties in the way of successful results
through hybridization are, therefore, these: the
difficulty of effecting the cross; infertility, in-
stability, variability, and often weakness and
monstrosity of the hybrids; and the absolute im-
possibility of predicting results. The advantage
to be derived from a successful hybridization is
the securing of a new variety which shall combine
in some measure the most desirable features of
both parents; and this advantage is often of so
ereat moment that it is worth while to make re-
peated efforts and to overlook numerous failures.
From these theoretical considerations it is apparent
that hybridization is essentially an empirical sub-
ject, and the results are such as fall under the
common denomination of chance. And, as it does
not rest upon any legitimate function in nature,
we can understand that it will always be difficult
to codify laws upon it.

_ HYBRIDS AND BUD-PROPAGATION. T1

Among the various characters of hybrid off-
spring, I presume that the most prejudicial one is
their instability, their tendency still to vary into
new forms or to return to one or the other parent
in succeeding generations. It is difficult to fix
any particular form which we may secure in the
first generation of hybrids. At the outset, we
notice that this discouraging feature is manifested
chiefly through the fact of seed-reproduction, and
we thereby come upon what is perhaps the most
important practical consideration in hybridization,
—the fact that the great majority of the best
hybrids in cultivation are increased by bud-propa-
gation, as cuttings, layers, suckers, buds, or grafts.
In fact, I recall very few instances in this country
of good undoubted hybrids which are propagated
with practical certainty by means of seeds. You
will recall that the genera in which hybrids are
most common are those in which bud-propagation
is the rule; as begonia, pelargonium, orchids,
gladiolus, rhododendron, roses, cannas, and the
fruits. This simply means that it is difficult to
fix hybrids so that they will come “true to seed,”
and makes apparent the fact that if we desire
hybrids we must expect to propagate them by
means of buds.

This is a point which appears to have been over-
looked by those who contend that hybridization
must necessarily swamp all results of natural se-

iz PHILOSOPHY OF CROSSING PLANTS.

lection; for, as comparatively few plants propagate
habitually by means of buds, whatever hybrids
might have appeared would have been speedily
lost, and all the more, also, because, by the terms
of their reasoning, the hybrids would cross with
other and dissimilar forms, and therefore lose their
identity as intermediates. Or, starting with the
assumption that hybrids are intermediates, and
would therefore obliterate specific types, we must
conclude that they should have some marked de-
eree of stability if they are to swamp or obliterate
the characters of species; but, as all hybrids tend
to break up when propagated by seeds, it must
follow that bud-propagation would become more
and more common, and this is associated in nature
with decreased seed-production. Now, seed-pro-
duction is the legitimate function of flowers; and
we must concede that, as seed-production de-
creased, floriferousness must have decreased; and
that, therefore, pronounced intercrossing would
have obliterated the very organs upon which it
depends, or have destroyed itself!

But I may be met by the objection that there
is no inherent reason why hybrids should not
become stable through seed-production by in-breed-
ing, and I might be cited to the opinion of Darwin
and others that in-breeding tends to fix any va-
riety, whether it originates by crossing or other
means. And it is a fact that in-breeding tends to

IN-BREEDING OF CROSSES. ia

fix varieties within certain limits, but those limits
are often overpassed in the case of very pro-
nounced crosses, whether cross-breeds or true
hybrids. And if it is true, as all observation and
experiments show, that sexual or reproductive
powers of crosses are weakened as the cross be-
comes more violent, we should expect less and less
possibility of successful in-breeding; for in-breed-
ing without disastrous results is possible only with
comparatively strong reproductive powers. As a
matter of fact, it is found in practice that it is
exceedingly difficult to fix pronounced hybrids by
means of in-breeding. It sometimes happens, too,
that the hybrid individual which we wish to per-
petuate may be infertile with itself, as I have often
found in the case of squashes. It is often advised
that we cross the hybrid individual which we wish
to fix with another like individual, or with one of
its parents. These results are often successful,
but oftener they are not. In the first place, it
often happens that the hybrid individuals may be
so diverse that no two of them are alike; this has
been my experience in many crosses. And, again,
crossing with a parent may draw the hybrid back
again to the parental form. So long ago as last
century Koélreuter proved this fact upon nicotiana
and dianthus. A hybrid between Nicotiana rus-
tica and N. paniculata was crossed with N. pani-
culata until it was indistinguishable from it; and

T4 PHILOSOPHY OF CROSSING PLANTS.

it was then crossed with N. rustica until it became
indistinguishable from that parent. Yet there is
no other way of fixing a hybrid to be propagated
by seeds than by in-breeding, and by constant at-
tention to selection. Fortunately, it occasionally
happens that a hybrid is stable, and therefore
needs no fixing.

In this connection I may cite some of my own
experience in crossing ege-plants and squashes ;
for, although the products were not true hybrids
in the strict interpretation of the word, many of
them were hybrids to all intents and purposes,
because made between very unlike varieties, and
they will serve to illustrate the difficulties of which
Ispeak. Offspring of egg-plant crosses were grown
in 1890, and upon some of the most promising
plants some flowers were self-pollinated. But
these self-pollinated seeds gave just as variable
offspring in 1891, as those selected almost at
random from the patch; and, what was worse,
none of them reproduced the parent, or “ came
true to seed,” and all further motive for in-breed-
ing was gone. My labor, therefore, amounted to
nothing more than my own edification. My
experience in crossing pumpkins and squashes has
now extended through many years; and, although
I have obtained about one thousand types not
named or described, I have not yet succeeded in
fixing one. The difficulty here is an aggravated

EXPERIMENTS IN IN-BREEDING. T5

one, however. The species are so exceedingly
variable that all the hybrid individuals may be
unlike, so that there can be no crossing between
identical stocks; and, if in-breeding is attempted,
it may be found that the flowers will not in-breed.
And the refusal to in-breed is all the more strange
because the sexes are separated in different flowers
upon the same plant. In other words, in my expe-
rience, it is very difficult to get good seeds from
squashes which are fertilized by a flower upon the
same vine. The squashes may grow normally to
full maturity, but be entirely hollow, or contain
only empty seeds. In some instances the seeds
may appear to be good, but may refuse to grow
under the best conditions. Finally, a small number
of flowers may give good seeds. I have many
times observed this refusal of squashes (Cucurbita
Pepo) to in-breed. It was first brought to my
attention through efforts to fix certain types into
varieties. The figures of one season’s tests will
sufficiently indicate the character of the problem.
In 1890, one hundred and eighty-five squash flowers
were carefully pollinated with staminate flowers
taken from the same vine which bore the pistillate
flowers. Only twenty-two of these produced fruit,
and of these only seven, or less than one-third, bore
good seeds, and in some of these the seeds were
few. Now, these twenty-two fruits represented as
many different varieties, so that the inability to set

76 PHILOSOPHY OF CROSSING PLANTS.

fruit with pollen from the same vine is not a
peculiarity of a particular variety. The records
of the seeds of the seven fruits in 1891 are as
follows : —

Fruit No. 1.— Four vines were obtained, with
four different types, two of them being white, one
yellow, and one black.

Fruit No. 2.— Twenty-three vines. Fifteen
types very unlike, twelve being white and three
yellow.

Fruit No. 3.—'Two vines. One type of fruit
which was almost like one of the original parents.

Fruit No. 4.— Thirty-two vines. Six types,
differing chiefly in size and shape.

Fruit No. 5. — Twenty vines. Nineteen types,
of which ten were white, eight orange, one striped,
and all very unlike.

Fruit No. 6.— Thirteen vines. Eleven types,
-— eight yellow, two black, one white.

Fruit No. T.— One vine.

These offspring were just as variable as those
from flowers not in-bred, and no more likely,
apparently, to reproduce the parent. ‘These tests
leave me without any method of fixing a pro-
nounced cross of squashes, and lead me to think
that the legitimate process of origination of new
kinds here, as, indeed, if not in general, is a more
gradual process of selection, coupled, perhaps, with
minor crossing.

ATTEMPT TO FIX A CROSS. Verg

I will relate a definite attempt towards the fixa-
tion of a squash which I had obtained from cross-
ing. The history of it runs back to 1887, when a
cross was effected between a summer yellow crook-
neck and a white bush scallop squash. In 1889
there appeared a squash of great excellence, com-
bining the merits of summer and winter squashes
with very attractive form, size, and color, and a
good habit of plant. I showed the fruit to one of
the most expert seedsmen of the country, and he
pronounced it one of the most promising types
which he had ever seen; and, as he informed me
that he had fixed squashes by breeding in and in,
I was all the more anxious to carry out my own
convictions in the same direction. It is needless
to say that I was very happy over what I regarded
as agreat triumph. Of course I must have a large
number of plants of my new variety, that I might
select the best, both for in-breeding and for cross-
ing similar types. So I selected the very finest
squash, having placed it where I could admire it
for some days, and saved every seed of it. These
seeds were planted upon the most conspicuous
knoll in my garden in 1890. It was soon evident
that something was wrong. I seemed to have
everything except my squash. One plant, how-
ever, bore fruits almost like the parent, and upon
this I began my attempts towards in-breeding.
But flower after flower failed, and I soon saw that

78 PHILOSOPHY OF CROSSING PLANTS.

the plant was infertile with itself. Careful search
revealed two or three other plants very like this
one, and I then proceeded to make crosses upon it.
I was equally confident that this method would
succeed. When I harvested my squashes in the
fall and took account of stock, I found that the
seeds of my one squash had given just as many
different types as there were plants, and I actually
counted one hundred and ten kinds distinct enough
to be named and recognized. Still confident, in
1891 I planted the seeds of my few crosses, and as
the summer days grew long and the crickets
chirped in the meadows, I watched the expanding
squash blossoms and wondered what they would
bring forth. But they brought only disappoint-
ment. Not one seed produced a squash like the
parent. My .squash had taken an _ unscientific
leave of absence, and I do not know its where-
abouts. And when the frost came and killed
every ambitious blossom, my hope went out and
has not yet returned!

Let us now recall how many undoubted hybrids
there are, named and known, among our fruits
and vegetables. In grapes there are the most.
There are Rogers’ hybrids, hke the Agawam,
Lindley, Wilder, Salem, and Barry; and there is
some reason for supposing that the Delaware,
Catawba, and other varieties are of hybrid origin.
And many hybrids have come to notice lately

LIST OF COMMON HYBRIDS. 79

through the work of Munson and others. But it
must be remembered that grapes are naturally ex-
ceedingly variable, and the specific limits are not
well known, and that hybridization among them
lacks much of that definiteness which ordinarily
attaches to the subject. In pears there is the
Kieffer class. In apples, peaches, plums, cher-
ries, gooseberries, and currants, there are no im-
portant commercial hybrids. In blackberries there
is the blackberry-dewberry class, represented by
the Wilson Early and others. Some of the rasp-
berries, like the Philadelphia and Shaffer, are hy-
brids between the red and black species. Hybrids
have been produced between the raspberry and
blackberry by two or three persons, but they pos-
sess no promise of economic results. Among all
the list of garden vegetables (plants which are
propagated by seeds) I do not know of a single
important hybrid ; and the same is true of wheat,
—unless the Carman wheat-rye varieties become
prominent, — oats, the grasses, and other farm
crops. But among ornamental plants there are
many; and it is a significant fact that the most
numerous, most marked, and most successful hy-
brids occur in the plants most carefully cultivated
and protected, those, in other words, which are
farthest removed from all untoward circumstances
and an independent position. ‘This is nowhere so
well illustrated as in the case of cultivated orchids,

80 PHILOSOPHY OF CROSSING PLANTS.

in which hybridization has played no end of freaks,
and in which, also, every individual plant is nursed
and coddled.!. With such plants the struggle for
existence is reduced to its lowest terms; for it
must be borne in mind that, even in the garden,
plants must fight severely for a chance to live, and
even then only the very best can persist, or are
even allowed to try.

I am sure that this lst of hybrids is much more
meagre than most catalogues and trade-lists would
have us believe, but I am sure that it is approxi-
mately near the truth. It is, of course, equivalent
to saying that most of the so-called hybrid fruits
and vegetables are myths. There is everywhere a
misconception of what a hybrid is, and. how it
comes to exist; and yet, perhaps because of this
indefinite knowledge, there is a wide-spread feel-
ing that a hybrid is necessarily good, while the
presumption is directly the opposite. The identity
of a hybrid in the popular mind rests entirely upon
some superficial character, and proceeds upon the
assumption that it is necessarily intermediate be-
tween the parents. Hence we find one of our
popular authors asserting that, because the kohl-
rabi bears its thickened portion midway of its stem,
it is evidently a hybrid between the cabbage and
turnip, which bear respectively the thickened parts

1 Consult E. Bohnhof, ‘‘ Dictionnaire des Orchidées Hy-
brides,’’ Paris, 1895.

INFLUENCE OF PARENTS. 81

at the opposite extremities of the stem! And then
there are those who confound the word hybrid
with high-bred, and who build attractive castles
upon the unconscious error. And thus is confu-
sion confounded!

But, before leaving this subject of hybridization,
I must speak of the old yet common notion that
there is some peculiar influence exerted by each
sex in the parentage of hybrids; for I shall thereby
not only call your attention to what I believe to
be an error, but shall also find the opportunity to
still further illustrate the entanglements of hybri-
dization. It was held by certain early observers, of
whom the great Linnzeus was one, that the female
parent determines the constitution of the hybrid,
while the male parent gives the external attributes,
as form, size, and color. The accumulated experi-
ence of nearly a century and a half appears to con-
tradict this proposition, and Focke, who has recently
gone over the whole ground, positively declares
that it is untrue. There are instances, to be sure,
in which this old idea is affirmed, but there are
others in which it is contradicted. The truth ap-
pears to be this, —that the parent of greater
strength or virility makes the stronger impression
upon the hybrids, whether it is the staminate or
pistillate parent; and it appears to be equally
true that it is usually impossible to determine be-
forehand which parent is the stronger. It is cer-

G

82 PHILOSOPHY OF CROSSING PLANTS.

tain that strength does not he in size, neither in
the high development of any character. It appears
to be more particularly associated with what we
call fixity or stability of character, or the tendency
towards invariability.

This has been well illustrated in my own experi-
ments with squashes, gourds, and pumpkins. The
common little pear-shaped gourd will impress itself
more strongly upon crosses than any of the edible
squashes and pumpkins with which it will effect a
cross, whether it is used as male or female parent.
Even the imposing and ubiquitous great field
pumpkin, which every New Englander associates
with pies, is overpowered by the little gourd.
Seeds from a large and sleek pumpkin which had
been fertilized by gourd pollen, produced gourds
and small hard-shelled globular fruits which were
entirely inedible. A more interesting experiment
was made between the handsome green-striped Ber-
gen fall squash and the little pear gourd. Several
flowers of the gourd were pollinated by the Bergen
in 1889. The fruits raised from these seeds in
1890 were remarkably gourd-like. Some of these
crosses were pollinated again in 1890 by the Ber-
gen, and the seeds were sown in 1891. Here,
then, were crosses into which the gourd had gone
once and the Bergen twice, and both the parents
are to all appearances equally fixed, the difference
in strength, if any, attaching rather to the Bergen.

POLLINATION IS UNCERTAIN. 83

Now, the crop of 1891 still carried pronounced
characters of the gourd. [Even in the fruits which
most resembled the Bergen, the shells were almost
flinty hard, and the flesh, even when thick and
tender, was bitter. Some of the fruits looked so
much like the Bergen that I was led to think that
the gourd had largely disappeared. The very hard
but thin paper-like shell which the gourd had laid
over the thick yellow flesh of the Bergen, I
thought might serve a useful purpose, and make
the squash a better keeper. And I found that it
was a great protection, for the squash could stand
any amount of rough handling, and was even not
injured by ten degrees of frost. All this was an
acquisition, and, as the squash was handsome and
exceedingly productive, nothing more seemed to
be desired. But it still remained to have a squash
for dinner. The cook complained of the hard
shell, but, once inside, the flesh was thick and
attractive, and it cooked nicely. But the flavor!
Dregs of quinine, gall, and boneset! The gourd
was still there !

VI. UNCERTAINTIES OF POLLINATION.

We have now seen that uncertainty follows
hybridization, and, in closing, I will say that
uncertainty also attaches to the mere act of
pollination. Between some species which are

84 PHILOSOPHY OF CROSSING PLANTS.

closely allied and which have large and strong
flowers, four-fifths of the attempts towards cross-
pollination may be successful; but such a large
proportion of successes is not common, and it
may be infrequent even in pollinations between
plants of the same species or variety. Some of
the failure is due in many cases to unskilful opera-
tion, but even the most expert operators fail as
often as they succeed in promiscuous pollinating.
There is good reason to believe, as Darwin has
shown, that the failure may be due to some selec-
tive power of individual plants, by which they re-
fuse pollen which is, in many instances, acceptable
to other plants even of the same variety or stock.
The lesson to be drawn from these facts is that
operations should be as many as possible, and that
discouragement should not come from failure.
In order to illustrate the varying fortunes of the
pollinator, I will transcribe some notes from my
field-book.

Two hundred and thirty-four pollinations of
gourds, pumpkins, and squashes, mostly between
varieties of one species (Cucurbita Pepo), and in-
cluding some individual pollinations, gave one
hundred and seventeen failures and one hundred
and seventeen successes. These crosses were made
in varying weather, from July 28 to August 30.
In some periods. nearly all the operations would
succeed, and at other times most of them would

RECORDS OF POLLINATIONS. 85

fail. I have always regarded these experiments as
among my most successful ones, and yet but half
of the pollinations “took.” But one must not
understand that I actually secured seeds from even
all these one hundred and seventeen fruits, for
some of them turned out to be seedless, and some
were destroyed by insects before they were ripe, or
they were lost by accidental means. A few more
than half of the successful pollinations —if by suc-
cess we mean the formation and growth of fruit —
really secured us seeds, or about one-fourth of the
whole number of efforts.

Twenty pollinations were made between potato
flowers, and they all failed; also, seven pollinations
of red peppers, four of husk-tomato, two of Nico-
tiana affinis upon petunia and two of the reciprocal
cross, twelve of radish, one of Mirabilis Jalapa upon
M. longiflora and two of the reciprocal cross, three
Convolvulus major upon C. minor and one of the
reciprocal, one muskmelon by squash, two musk-
melons by watermelon, and one muskmelon by
cucumber.

This is but one record. Let me give another: —

Cucumber, ninety-five efforts: fifty-two suc-
cesses, forty-three failures. Tomato, forty-three
efforts: nineteen successes, twenty-four failures.
Ege-plant, seven efforts: one success, six failures.
Pepper, fifteen efforts: one success, fourteen fail-
ures. Husk-tomato, forty-five efforts: forty-five

86 PHILOSOPHY OF CROSSING PLANTS.

failures. Pepino, twelve efforts: twelve failures.
Petunia by Nicotiana affinis, eleven efforts: eleven
failures. Nicotiana affinis by petunia, six efforts:
six failures. General Grant tobacco by Nicotiana
affinis, eleven efforts: eight successes, three fai!-
ures. Nicotiana affinis by General Grant tobacco,
fifteen efforts: fifteen failures. General Grant
tobacco by General Grant tobacco, one effort: one
success. Nicotiana affinis by Nicotiana affinis, three
efforts: two successes, one failure. Tuberous be-
gonia, five efforts: five successes.

Total, three hundred and twelve efforts: eighty-
nine successes, two hundred and twenty-three
failures.

CONCLUSION.

And now, the sum of it all is this: encourage in
every way crosses within the limits of the variety
and in connection with change of stock, expecting
increase in vigor and productiveness; hybridize
if you wish to experiment, but do it carefully,
systematically, thoroughly, and do not expect too
much. Extend Darwin’s famous proposition to
read: Nature abhors both perpetual self-fertiliza-
tion and hybridization.

LECTURE III.

HOW DOMESTIC VARIETIES ORIGINATE.

“THE key is man’s power of accumulative
selection: nature gives successive variations ;
man adds them up in certain directions useful
to him.” ‘This, in Darwin’s phrase, is the essence
of the cultivator’s skill in ameliorating the vege-
table kingdom. So far as man is concerned, the
origin of the initial variation is largely chance,
but this start or variation once given, he has the
power, in most cases, to perpetuate it and to
modify its characters. There are, then, two very
different factors or problems in the origination of
garden varieties, —the production of the first de-
parture or variation, and the subsequent breeding
of it. Persons who give little thought to the sub-
ject, look upon variation as the end of their endeav-
ors, thinking that a form comes into being with all
its characters well marked and fixed. In reality,
however, variation is but the beginning ; selection
is the end.

I. INDETERMINATE VARIETIES.

There are two general classes of garden varie-
ties as respects the method of their origin, —
87

88 HOW DOMESTIC VARIETIES ORIGINATE.

those which come into existence somewhat sud-
denly and which require little else of the hus-
bandman than the multiplication of them, and
those which are the result of a slow evolution or
direct breeding. ‘The former are indeterminate
or uncertain, and the latter are determinate or
definite. The greater part of those in the first
class are plants which are multiphed or divided
by bud-propagation. ‘They comprise nearly all
our fruits, the woody ornamental plants, and such
herbaceous genera as begonia, canna, gladiolus,
lily, dahlia, carnation, chrysanthemum, and the
like,—in fact, all those multiplied by grafting,
cuttings, bulbs, or other asexual parts. The
original plant may be either a seedling or a bud-
sport. The gardener, who is always on the look-
out for novelties, discovers its good qualities and
propagates it.

Varieties which are habitually multiphed by
buds, as in those plants which I have mentioned
in the last paragraph, vary widely when grown
from seeds, so that every seedling may be
markedly distinct. As soon, however, as varie-
ties are widely and exclusively propagated by
seeds, they develop a capacity of carrying the
greater part of the individual differences down
to the offspring. That is, seedlings from bud-
multiplied plants do not “come true,” as a rule,
whilst those from seed-propagated plants do

INDETERMINATE VARIETIES. 89

““come true.” The reason of this difference will
become apparent upon a moment’s reflection. In
the seed-propagated plants, like the kitchen-gar-
den vegetables and the annual flowers, we select
the seeds and thereby eliminate all those varia-
tions which would have arisen had the discarded
seeds been sown. In other words, we are con-
stantly fixing the tendency to “come true,” for
this feature of plants is as much a variation as
form or color or any other attribute is. Suppose,
for instance, that a certain variation were to re-
ceive two opposite treatments, the seeds from one-
half of the progeny being carefully selected year
by year, and all those from untypical plants dis-
earded, whilst in the other half all the seeds from
all the plants, whether good or bad, are saved
and sown. In the one case, it will be seen, we
are fixing the tendency to “come true,” for this
is all that constitutes a horticultural variety, —
a brood which is very much like all its parents.
In the other case, we are constantly eliminating
the tendency to “come true”’ by allowing every
modifying agency full sway. So the very act of
taking seeds only from plants which have “ come
true,” tends to still more strongly fix the heredi-
tary force within narrow hmits. Working against
this restrictive force, however, are all the agencies
of environment, so that, fortunately, now and
then a seed gives a “rogue,” or a plant widely

90 HOW DOMESTIC VARIETIES ORIGINATE.

unlike its parent, and this may be the start for a
new variety.

With bud-multiplied varieties, however, the
case is very different. Here every seed may be
sown, as in the illustrative case above, because
the seedlings are not wanted for themselves, but
simply as stocks upon which to bud or graft the
desired varieties. So there is no seed selection in
the ordinary propagation of apples, pears, peaches,
and the other orchard fruits. ‘The seeds are taken
indiscriminately from pomace or the refuse of can-
ning and evaporating factories. But every annual
garden vegetable is always grown from seeds more
or less carefully saved from plants which possess
some desired attribute. ‘There is no reason why
the tree fruits should not reproduce themselves
from seeds just as closely as the annual herbs do, if
they were to be as carefully propagated by selected
seeds through a long course of generations. There
is excellent proof of this in the well-marked races
or familes of Russian apples. In that country,
grafting has been little employed, and conse-
quently it has been necessary to select seeds
only from acceptable trees in order that the off-
spring might be more acceptable. So the Russian
apples have come to run in groups or families, each
family bearing the mark of some strong ancestor.
Most of the seedlings of the Duchess of Olden-
burg are recognizable because of their likeness to

PLANT-BREEDING. 91

the parent. We may thus trace an incipient ten-
dency in our own fruits towards racial characters.
The Fameuse type of apples, for example, tends to
perpetuate itself ; and a similar tendency is very
well marked in the Damson and Green Gage
plums, the Orange quince, Concord grape, and
Hill’s Chili and Crawford peaches. But inas-
much as bud-multiplication is so essential in
nursery practice, we can hardly hope for the
time when our trees and shrubs, or even our per-
ennial herbs, will ‘“‘come true” with much cer-
tainty. In them, therefore, we get new varieties
by simply sowing the seeds; but in seed-propa-
gated varieties we must depend either upon chance
variations or else we must resort to definite plant-
breeding.

Il. PLANT-BREEDING.

The breeding of domestic animals is attended,
for the most part, with such definite and often
precise results that there has come to be a gen-
eral desire to extend the same principles to
plants. It is not unusual to hear well-informed
people say that it is possible to breed plants with
as much certainty and exactness as it is to breed
animals. The fact is, however, that such exact-
ness will never be possible, because plants are
very unlike animals in organization, and because,

92 HOW DOMESTIC VARIETIES ORIGINATE.

also, the objects sought in the two cases are fun-
damentally unlike. Plants, as we have seen, are
made up of a colony of potential individuals, and
to breed between two plants by crossing means
that we must choose the sex-parents from amongst
as many individuals as there are flowers or branches
on the two plants, whilst in animals we choose two
definite personal parents. And these personal
parents are either male or female, and the union
is essential to the production of offspring, whilst
in plants each parent —that is, each flower — is
generally both male and female, and the union of
two is not essential to the production of offspring,
for the plant is capable of multiplying itself by
buds. The element of chance, therefore, is one
hundred, or more, to one in crossing plants as
compared with crossing animals. Then, again,
the plant-parents are modified profoundly by every
environmental condition of soil and temperature
and sunshine, or other external condition, since
they possess no bodily temperature, no choice of
conditions, and no volition to enable them to
overcome the circumstances in which they are
placed. Animals, on the contrary, have all these
elements of personality, and the breeder is also
able to control the conditions of their lives to a
nicety. In view of all these facts, it is not strange
that animals can be bred by crossing with more
confidence than plants can. But there is another

ANIMALS AND PLANTS. 93

and even more important difference between the
breeding of animals and the breeding of plants.
In animals, our sole object is to secure simply one
animal or one brood of offspring. In plants, our
object is, in general, to secure a race or genera-
tions of offspring, which may be disseminated
freely over the earth. In the bovine race, for
example, our object in breeding is to produce one
cow with given characters ; in turnips, our object
is to produce a new variety, the seed of which
will reproduce the variety, whether sown in Penn-
sylvania or Ceylon. It is apparent, therefore, that
any comparisons drawn between the breeding of
animals and plants are fundamentally fallacious.
Is there, then, any such thing as plant-breeding,
any possibility that the operator can proceed with
some confidence that he may obtain the ideal
which he has in mind? Yes, to a certain extent.
It is apparent that the very first effort on the
part of the plant-breeder must be to secure indi-
vidual differences; for so long as the plants which
he handles are very closely alike, so long there
will be little hope of obtaining new varieties. He
must, therefore, cause his plants to vary. In
plants which are comparatively unvariable, it is
frequently impossible to produce variations in the
desired direction at once, but it is more important
to “ break ” the type,— that is, to make it depart
markedly from its normal behavior in any or

94. HOW DOMESTIC VARIETIES ORIGINATE.

many directions (page 19). If the type once
begins to vary, to break up into different forms,
the operator may be sure that it will soon become
plastic enough to allow of modification in the
manner which he desires. But whilst it is im-
portant or even necessary to break a well-marked
type into many forms, it would no doubt be un-
wise to encourage this tendency after it once
appears, lest the plant acquire a too strong habit
of scattering. ‘This initial variation is induced
by changing the conditions in which the plant has
habitually grown, as a change of seed, change of
soil, tillage, varying the food supply, crossing and
the like.

As a matter of fact, however, nearly all plants
which have been long cultivated are already suf-
ficiently variable to afford a starting-point for
breeding. The operator should have a vivid
mental picture of the variety which he designs to
obtain ; then he should select that plant in his
plantation which is the nearest his ideal, and sow
the seeds of it. From the seedlings he should
again select the individuals which most nearly
approach his type, and so on, generation after
generation, until the desired object is attained.
It is important, if he is to make rapid. progress,
that he keep the same ideal in mind year by year,
otherwise there will be vacillation and the prog-
ress of one year may be undone by a counter

ANTAGONISTIC FEATURES. 95

movement the following year. In this way, it
will be found that almost any character of a plant
may be either intensified or lessened. ‘This is
man’s nearest approach to the Creator in his
dominion over the physical forms of life, and it
is great and potent in proportion as it sets for
itself correct ideals in the beginning and adheres
to them until the end.

When beginning this selection or breeding for
an ideal, it is important that impossible or contra-
dictory results should be avoided. Some of the
cautions and suggestions which need to be con-
sidered are these : —

1. Avoid striving after features which are antag-
onistic or foreign to the species or genus with which
you are working. Every group of plants has be-
come endowed with certain characters or lines
of development, and the cultivator will secure
quicker and surer results if he works along the
same lines, rather than to attempt to thwart them.
Nature gives the hint: let men follow it out,
rather than to endeavor to create new types of
characters. Let us take some of the solanaceous
plants as examples. ‘There are certain types of
the genus solanum which have a natural habit of
tuber-bearing, as the potato. Such species should
be bred for tubers and not for fruits. There are
other solanums, however, like the egg-plants and
the pepinoes, which naturally vary or develop in

96 HOW DOMESTIC VARIETIES ORIGINATE.

the direction of fruit-bearing, and these should be
bred for fruits and not for tubers; and the same
should be true in the related genera of tomatoes,
red peppers, and physals. Those ambitious per-
sons who are always looking for a tuber-bearing
tomato, therefore, might better concentrate their
energies on the potato, for the tomato is not devel-
oping in that direction; and even if the tomato
could be made to produce tubers, it would thereby
lessen its fruit production, for plants cannot main-
tain two diverse and profitable crops at the same
time. It is more reasonable, and certainly more
practicable, to grow potatoes on potato plants and
tomatoes on tomato plants.

2. The quickest and most marked results are to
be expected in those groups or species which are nor-
mally the most variable. ‘There are a greater num-
ber of variations or starting-points in such species ;
but it also follows that the forms are less stable
the more the species is variable. Yet the varia-
tions, being very plastic, yield themselves readily
to the wishes of the operator. Carriere puts the
thought in this form: “The stability of forms, in
any group of plants, is, in general, in inverse ratio
to the number of the species which it contains,
and also to the degree of its domestication.”

The most variable types are the most dominant
ones over the earth; that is, they occur in greater
numbers and under more diverse conditions than

VARIABLE TYPES. 97

the comparatively invariable types do. The com-
posite, or sunflower-like plants, comprise a ninth
or tenth of the total species of flowering plants,
and the larger part of the subordinate types or
genera contain many forms or species. Aster,
goldenrod, the hawkweeds, thistles, and other
groups, are representative of the cosmopolitan or
variable types of composites. Whenever, for any
reason, any type begins to decline in variability,
it also begins to perish; it is then tending towards
extinction. Monotypic genera— those which con-
tain but a single species — are generally of local
or disconnected distribution, and are, for the most
part, vanishing remnants of a once dominant or
important type. As a rule, most of our widely
variable and staple cultivated species are mem-
bers of large, or at least polytypic genera. Such,
for example, are the apples and pears, peaches
and plums, oranges and lemons, roses, bananas,
chrysanthemums, pinks, eucurbits, beans, potato,
orapes, barley, rice, cotton. A marked exception
to this statement is maize, which is immensely
variable and is generally held to have come from
a single species ; but the genesis of maize is un-
known, and it is possible, though scarcely proba-
ble, that more than one species 18 concerned in it.
Wheat is also a partial exception, although the
original specific type is not understood; and the
latest monographers admit three or four other spe-

H

98 HOW DOMESTIC VARIETIES ORIGINATE.

cies to the genus, aside from wheat. There are
other exceptions, but they are mostly unimportant,
and, in the main, it may be said that the domi-
nant domestic types of plants represent markedly
polytypic genera.

3. Breed for one thing at a time. The person
who strives at the same time for increase or modi-
fication in prolificacy and flavor will be likely to
fail in both. He should work for one object
alone, simply giving sufficient attention to sub-
sidiary objects to keep them up to normal stand-
ard. ‘This is really equivalent to saying that
there can be no such thing as the perfect all-
around variety which so many people covet. Va-
rieties must be adapted to specific uses, — one for
shipping, one for canning, one for dessert, one for
keeping qualities, and the like. The more good
varieties there are of any species, the more widely
and successfully that species can be cultivated.

4. Do not desire contradictory attributes in any
variety. A variety, for example, which bears the
maximum number of fruits or flowers cannot be
expected to greatly increase the size of those
organs without loss in numbers. This is well
shown in the tomato. The original tomato pro-
duced from six to ten fruits in a cluster, but as
the fruits increased in size the numbers in each
cluster fell to two or three. That is, increase in
size proceeded somewhat at the expense of numer-

SELECT FOR THE ENTIRE PLANT. 99

ical productivity ; yet the total weight of fruit
per plant has greatly increased. ‘The same is
true of apples and pears; for whilst these trees
bear flowers in clusters, they generally bear their
fruits singly. Originally, every flower normally
set fruit. The reason why blackberries, currants,
and grapes do not increase more markedly in size,
is probably because the size of cluster has been
given greater attention than the size of berry.
Plants which now bear a full crop of tubers
cannot be expected to increase greatly in fruit-
bearing, as I have already explained under
Rule 1. This fact is illustrated in the potato,
in which, as tuber production has increased, seed
production has decreased, so that potato growers
now complain that potatoes do not produce bolls
as freely as they did years ago.

5. When selecting seeds, remember that the char-
acter of the whole plant 1s more wmportant than the
character of any one branch or part of the plant ;
and the more uniform the plant in all its parts, the
greater is the likelihood that it will transmit tts
characters. If one is striving for larger flowers,
for instance, he will secure better results if he
choose seeds from plants which bear large flowers
throughout, than he will if he choose them from
some one large-flowering branch on a plant which
bears indifferent flowers on the remaining branches,
even though this given branch produce much larger

100 HOW DOMESTIC VARIETIES ORIGINATE.

flowers than those borne on the large-flowered .
plant. Small potatoes from productive hills give
a better product than large potatoes from unpro-
ductive hills. The practice of selecting large
ears from a bin of corn, or large melons from the
grocer’s wagon, is much less efficient in produc-
ing large products the following season than the
practice of going into the fields and selecting the
most uniformly large-fruited parents would be.
A very poor plant may occasionally produce one
or two very superior fruits, but the seeds are
more likely to perpetuate the characters of the
plant than of the fruits.

The following experiences detailed by Henri L.
de Vilmorin illustrate my proposition admirably:
“] tried an experiment with seeds of Chrysanthe-
mum carinatum gathered on double, single, and
semi-double heads, all growing on one plant,
and found no difference whatever in the propor-
tion of single and double-flowered plants. In
striped verbenas, an unequal distribution of the
color is often noticed; some heads are pure white,
some of a self color, and most are marked with
colored stripes on white ground. I had seeds
taken severally from all and tested alongside
one another. The result was the same. All the
seeds from one plant, whatever the color of the
flower that bore them, gave the same proportion
of plain and variegated flowers.”

UNIFORMNESS IN THE PARTS. 101

The second part of my proposition is equally as
important as the first,—the fact that a plant
which is uniform in all its branches or parts is
more likely to transmit its general features than
one which varies within itself. It is well known
that bean plants often produce beans with various
styles of markings on the same plant or even in
the same pod, yet these variations rarely ever
perpetuate themselves. The same remark may
be applied to variations in peas. ‘These illustra-
tions only add emphasis to the fact that intending
plant-breeders should give greater heed than they
usually do to the entire plant, rather than confine
their attention to the particular part or organ
which they desire to improve.

At first thought, it may look as if these facts
are directly opposed to the proposition which I
emphasized in my first lecture, that every branch
of a plant is a potential autonomy, but it is really
a confirmation of it. ‘The variation itself shows
that the branch is measurably independent, but it
is not until the conditions or causes of the vari-
ation are powerful enough to affect the entire
plant that they are sufficiently impressed upon
the organization of the plant to make their effects
hereditary.

There is an apparent exception to the law that
the character of the entire plant is more impor-
tant than any one organ or part of it, in the case

102 HOW DOMESTIC VARIETIES ORIGINATE.

of the seeds themselves. ‘That is, better results
usually follow the sowing of large and heavy
seeds than of small or unselected seeds from the
same plant. This, however, does not affect the
main proposition, for the seed is in a measure
independent of the plant-body, and is not so
directly influenced by environment as the other
organs are. And, again, the seed receives a part
of its elements from a second or male parent.
The good results which follow the use of large
seeds are, chiefly, greater uniformity of crop,
increased vigor, often a gain in earliness and
sometimes in bulk, and generally a greater ca-
pacity for the production of seeds. These results
are probably associated less with any innate he-
reditable tendencies than with the mere vegeta-
tive strength and uniformness of the large seeds.
The large seeds usually germinate more quickly
than the small ones, provided both are equally
mature, and they push the plantlet on more
vigorously. This initial gain, coming at the
most critical time in the life of the new indi-
vidual, is no doubt responsible for very much of
the result which follows. ‘The uniformity of crop
is the most important advantage which comes of
the use of large seeds, and this is obviously the
result of the elimination of all seeds of varying
degrees of maturity, of incomplete growth and
formation, and of low vitality.

PROGENY OF IMMATURE SEEDS 103

Another important consideration touching the
selection of seeds is the fact that very immature
seeds give a feeble but precocious progeny. ‘This
has long been observed by gardeners, but Sturte-
vant, Arthur, and Goff have recently made a
critical examination of the subject. “It is not
the shghtly unripe seeds that give a noticeable
increase in earliness,” according to Arthur, “ but
very unripe seeds, gathered from fruit [tomatoes]
scarcely of full size and still very green. Such
seeds do not weigh more than two-thirds as much
as those fully ripe. They germinate readily, but
the plantlets lack constitutional vigor and are
more easily affected by retarding or harmful
influences. If they can be brought through the
early period of growth and become well estab-
lished, and the foliage or fruit is not attacked by
rots or blights, the grower will usually be re-
warded by an earlier and more abundant crop of
slightly smaller and less firm fruit. These char-
acters will be more strongly emphasized in sub-
sequent years by continuous seed propagation.”
Goff remarks that the increase in earliness in
tomatoes, following the use of markedly immature
seeds, ‘is accompanied by a marked decrease in
the vigor of the plant, and in the size, firmness,
and keeping quality of the fruit.” These results
are probably closely associated with the chemical
constitution and content of the immature seeds.

104. HOW DOMESTIC VARIETIES ORIGINATE.

The organic compounds have probably not yet
reached a state of stability, and they therefore
respond quickly to external stimuli when placed
in conditions suitable to germination; and there
is little food for the nourishment of the plantlet.
The consequent weakness of the plantlet results
in a loss of vegetative vigor, which is earliness
(see Rule 11).

Still another feature connected with the choice
of seeds is the fact that in some plants, as in some
Ipomeeas, for example, the color of the seed is
more or less intimately associated with the color
of the flower which produced them and also with
the color of the flowers which they will produce.

6. Plants which have any desired characteristics
in common may differ widely in their ability to
transmit these characters. It is generally impos-
sible for the cultivator to determine, from the
appearance of any given number of similar plants,
which of them will give progeny the most unvari-
able and the most lke its parent; but it may be
said that those individuals which grow in the most
usual or normal environments are most likely to
perpetuate themselves. A very unusual condi-
tion, as of soil, moisture, or exposure, is not easily
imitated when providing for the succeeding gen-
eration, and a return to normal conditions of envi-
ronment may be expected to be followed by a more
or less complete return to normal attributes on the

SELECT SEVERAL STARTING-POINTS. 105

part of the plant. If the same variation, there-
fore, were to occur in plants growing under widely
different conditions, the operator who wishes to
preserve the new form should take particular care
to select his seeds from those individuals which
seem to have been least influenced by the imme-
diate conditions in which they have grown.

Again, if the same variation appears both in
uncrossed and crossed plants, the best results
should be expected in selecting seeds from the
former. We have already seen, in the second
lecture, how it is that crosses are unstable, and
how the instability is apt to be the greater the
more violent the cross. ‘ Cross-breeding greatly
increases the chance of wide variation,” writes
Henri L. de Vilmorin, “but it makes the task of
fixation more difficult.”

It is very important, therefore, when selecting
seeds from plants which seem to give promise of
a new variety, to sow the seeds of each plant
separately, and then make the subsequent selec-
tions from the most stable generation; and it is
equally important that the operator should not
trust to a single plant as a starting-point, when-
ever he has several promising plants from which
to choose.

17. The less marked the departure from the
genius of the normal type, the greater, in general,
as the likelihood that it will be perpetuated. ‘That

106 HOW DOMESTIC VARIETIES ORIGINATE.

is, widely aberrant forms are generally unstable.
This is admirably illustrated in crosses. The
seed-progeny of crosses between closely related
varieties, or between different plants of the same
variety, is more uniform and generally more easy
of improvement by selection than the progeny of
hybrids. In uncrossed plants, the general ten-
dency is to resemble their parents, and the greater
the number of like ancestors, the greater is the ten-
dency to “come true.” There is thought to be a
tendency, though necessarily a weak one, to return
to some particular ancestor, or to ‘“‘date back.”
This is known as atavism. The so-called atavistic
forms are likely to be unstable, to break up into nu-
merous forms, or to return more or less completely
to the type of the main line of the ancestry. The
following statements touching some of the rela-
tions of atavism to the amelioration of plants, are
the results of an excellent study of heredity in
lupines by Louis Leveque de Vilmorin : —

“1. The tendency to resemble its parents is
generally the strongest tendency in any plant;

«2. But it is notably impaired as it comes
into conflict with the tendency to resemble the
general line of its ancestry.

«3. This latter tendency, or atavism, is con-
stant, though not strong, and scarcely becomes
impaired by the intervention of a series of gen-
erations in which no reversion has taken place.

CROSSING NOT AN END. 107

“4, The tendency to resemble a near pro-
genitor (only two or three generations removed),
on the other hand, is very soon obliterated if the
given progenitor is different from the bulk of
its ancestors.”

8. The crossing of plants should be looked upon
as a means or starting-point, not as an end. We
cross two flowers and sow the seeds. The result-
ing seedlings may be unlike either parent. Here,
then, is variation. ‘The operator should select
that plant which most nearly satisfies his ideal,
and then, by selection from its progeny and the
progeny of succeeding generations, gradually ob-
tain the plant which he desires. It is only in
plants which are propagated by asexual parts—as
grafts, cuttings, layers, bulbs, and the like —that
hybrids or crosses are commonly immediately val-
uable; for in these plants we really cut up and
multiply the one individual plant which pleases us
in the first batch of seedlings, rather than to take
the offspring or seedlings of it. Thus, if any
particular plant in a lot of seedlings of crosses
of cannas, or plums, or hops, or strawberries, or
potatoes, is valuable, we multiply that one in-
dividual. There is no occasion for fixing the
variety. But any satisfactory plant in a lot of seed-
lings of crosses of pumpkins, or wheat, or beans,
must be made the parent of a new variety by sow-
ing the seeds of it and then by selecting for seed-

108 HOW DOMESTIC VARIETIES ORIGINATE.

parents, year by year, those plants which are best.
“The unsettled forms arising from crosses,” Focke
writes, “are the plastic material out of which
gardeners form their varieties.”

But even in the fruits, and other bud-propa-
gated plants, crossing may often be used to as
good advantage for the purpose of originating
variation as it can in peas or buckwheat. It only
requires a longer time to fix and select variations
because the plants mature so slowly. Ordinarily,
if the operator does not find satisfactory plants
amongst the seedlings of any cross of fruit trees,
he roots up the whole batch as profitless. But if
he were to allow the best plants to stand and
were to sow seeds from them, the second gen-
eration might produce something more to his
liking. But it is generally quicker to make
another cross and to try the experiment over
again, than to wait for unpromising seedlings to
bear. This repeated repetition of the experiment,
however, — continual crossing and sowing and
uprooting,—is gambling. Throwing dice to see
what will turn up is a comparable proceeding.
The sowing of uncrossed seed is little better.
Peter M. Gideon sowed over a bushel of apple
seed, and one seed produced the Wealthy apple.1

1'The facts in the origination of the Wealthy apple, as re-
lated to me by Mr. Gideon, are these: he first planted a bushel
of apple seeds, and then each year, for nine years, he planted

GUIDES TO CROSSING. 109

D. B. Wier raised a million seedlings of soft
maple, and one plant of the lot had finely divided
leaves, and is now Wier’s Cutleaved maple. Teas’
Weeping mulberry, which is now so deservedly
popular, was, as Mr. Teas tells me, “merely an
accidental seedling.” So this explains why the
production of new varieties of fruits is always
‘chance, whilst a skilled man can sit in his study
in the winter time and picture to himself a new
bean or muskmelon, and then go out in the next
three or four summers and produce it.

9. Lf wt ts desired to employ crossing as a direct
means of producing new varieties, each parent to the
proposed cross should be selected in agreement with
the rules already specified, and also because it pos-
sesses in an emphatic degree one or more of the
qualities which it is desired to combine; and the
more uniformly and persistently the parent pre-
sents a given character, the greater is the chance that
wt will transmit that character. It has already
been said that crossing for the instant production
of new varieties is most certain to give valuable

enough seed to give a thousand trees. At the end of ten years,
all the seedlings had perished (this was in Minnesota) except
one hardy seedling crab. Then a small lot of seeds of apples
and crab apples was obtained in Maine, and from these the
Wealthy came. There were only about fifty seeds in the
batch of crab seed which gave the Wealthy; but before this
variety was obtained, much over a bushel of seed had been
sown,

110 HOW DOMESTIC VARIETIES ORIGINATE.

results in those species which are propagated by
buds, because the initial individual differences
are not dissipated by seed-reproduction. This
is especially true of hybridization, or crossing
between distinct species; for in such violent cross-
ing as this the offspring is particularly likely to
be unstable when propagated by seeds. ‘The re-
sults of hybridization appear to be most certain
in those plants which are grown under glass, and
in which, therefore, the selection of the seed-
parents 1s most carefully made, and where the
conditions of existence are most uniform. ‘The
most remarkable results in hybridization which
have yet been attained are with the choicer glass-
house plants, such as orchids, begonias, anthu-
riums, and the hke. (Lecture IT.)

The more violent the cross, the less is the likeli-
hood that desirable offspring will follow. Species
which refuse to give satisfactory results when
hybridized directly or between the pure stocks,
may give good varieties when the “blood” has
become somewhat attenuated through previous
crossings. ‘The best results in hybridizing our
native grape with the European grape, for ex-
ample, have come from the use of one parent
which is already a hybrid. ‘Two notable examples
are the Brighton and Diamond grapes, raised by
Jacob Moore. The Brighton is a cross of Con-
cord (pure native) by Diana-Hamburg (hybrid of

IMPORTANT HYBRIDS OF FRUITS. tht

impure native and European). Diamond is a
cross of Concord by Iona, the latter parent un-
doubtedly of impure origin, containing a trace of
the European vine. T. V. Munson’s Brilliant is
a secondary hybrid, its parents, Lindley and Dela-
ware, both containing hybrid blood. Others of
his varieties have similar histories. Even when
the cross is much attenuated —or three or four
or even more times removed from a pure hybrid
origin by means of subsequent crossings — it may
still produce marked effects in a cross without
introducing such contradictory characters as to
jeopardize the value of the offspring.

Amongst American fruit plants there are com-
paratively few valuable hybrids. The most con-
spicuous ones are in the grapes, particularly the
various Rogers varieties, such as Agawam, Lind-
ley, Wilder, Barry, and others, which are hybrids
of the European grape and a native species.
Other hybrids are the Kieffer and allied pears
(between the common pear and the Oriental
pear), the Transcendent and a few other crabs
(between the common apple and the Siberian
crab), the Soulard and kindred crabs (between
the common apple and the native Western crab),
a few blackberries of the Wilson Early type
(between the blackberry and the dewberry), the
purple-cane raspberries (between the native red
and black raspberries, and possibly sometimes

112 HOW DOMESTIC VARIETIES ORIGINATE.

combined with the European raspberry), the
Utah Hybrid cherry (between the Western sand
cherry and the sand plum), and probably a few
of the native plums. There is undoubtedly a
fertile field for further work in hybridizing our
fruits, particularly those of native origin, and
also many of the ornamental plants; the danger
is that persons are apt to expect too much from
hybridization, and too little from the betterment
of all the other conditions which so profoundly
modify plants. Violent hybridizations generally
give unsatisfactory and unreliable results; but
subsequent crossings, when the “blood” of the
original species to the contract is considerably
attenuated, may be expected to correct or over-
come the first incompatibility, as explained above.

10. Establish the ideal of the desired variety
firmly in the mind before any attempt is made at
plant-breeding. If one is to make any progress
in securing new varieties, he must first be an
expert judge of the capabilities and merits of
the plants with which he is dealing, otherwise
he may attempt the impossible or he may obtain
a variety which has no merit. It is important,
too, that the person bear in mind the fact that
a variety which is simply as good as any other
in cultivation is not worth introducing. It
should be better in some particular than any
other in existence. ‘The operator must know the

PRODUCE AN INITIAL VARIATION. 113

points of his plant, as an expert stock-breeder
knows the points of an animal, and he must
possess the rare judgment to determipe which
characters are most likely to reappear in the
offspring. Inasmuch as a person can be an ex-
pert in only a few plants, it follows that he can-
not expect satisfactory results in breeding any
species which may chance to come before him.
Persistent and uniform effort, continued over a
series of years, is generally demanded for the
production of really valuable varieties. Thus it
often happens that one man excels all competitors
in breeding a particular class of plants. The hor-
ticulturist will recall, for instance, Lemoine in
the breeding of gladiolus, Eckford in peas, Crozy
in cannas, Bruant in pelargoniums, and others.
There are now and then varieties which arise
from no effort, but because of that very fact they
reflect no credit upon the so-called originator,
who is really only the lucky finder. So far as
the originator is concerned, such varieties are
merely chance. If, however, the operator — him-
self an expert judge of the plant with which he
deals — chooses his seeds with care and discrimi-
nation, and then proposes, if need be, to follow up
his work generation by generation by means of
selection, the work becomes plant-breeding of the
highest type.

First of all, therefore, the operator must know

a

114 HOW DOMESTIC VARIETIES ORIGINATE.

what he can likely get, and what will likely be
worth getting. Most persons, however, begin at
the other end of the problem,— they get what
they can, and then let the public judge if the
effort has been worth the while.

11. Having obtained a specific and correct ideal,
the operator must next seek to make his plant vary
in the desired direction. ‘This may be done by
crossing, or by modifying the conditions under
which the plant grows, as indicated in Lectures
I. and Il. If there are any two plants which
possess indications of the desired attributes, cross
them: amongst the seedlings there may be some
which may serve as starting-poimts for further
effort.

A change in the circumstances or environment
of the plant may start the desired attribute. If
the plant must be dwarfer, plant it on poorer or
drier soil, transfer it towards the poles, plant it
late in the season, or transplant it repeatedly (see
pages 25 and 143). Dwarf peas become climb-
ing peas on rich, moist soils. If the plant must
have large fruits, allow it more food and room,
and give attention to pruning and thinning. Cer-
tain geographical regions develop certain charac-
ters in plants, as we have seen (page 24); if,
therefore, the desired feature does not appear
spontaneously or as a result of any other treat-
ment, transfer the plant for a time to that region

PRODUCE AN INITIAL VARIATION. 115

which is characterized by such attributes, if there
is any such.

The importance of growing the plant under
conditions or environments in which the desired
type of characters is most frequently found, is
admirably emphasized in the evolution of varieties
which are adapted to forcing under glass. Within
a century,—and in many instances within a
decade or two,—species which were practically
unknown to glass-houses have produced varieties
which are perfectly adapted to them. This has
been accomplished by growing the most tractable
existing varieties under glass, and then carefully
and persistently selecting those which most com-
pletely adapt themselves to their environment and
to the ideals of the operator. One of the most
remarkable examples of this kind is afforded by
the carnation. In Europe it is chiefly a border
or out-door plant, but within a generation it has
produced hosts of excellent forcing varieties in
America, where it is grown almost exclusively as
a glass-house flower. So the carnation types of
Europe and America are widely unlike, and the
unlikeness becomes more emphatic year by year
because of the rapid aberrant evolution of the
American forms. :

Sowing the seeds of hardy annual plants in the
fall often generates a tendency to produce thick-
ened roots. The plant, finding itself unable to

116 HOW DOMESTIC VARIETIES ORIGINATE.

perfect seeds, stores its reserve in the root, and
it therefore tends to become biennial. In this
manner, with the aid of selection and the varia-
tion of the soil, Carri¢re was able to produce good
radishes from the wild slender-rooted charlock
(Raphanus Raphanistrum).

Lessened vigor, so long as the plant continues
to be healthy, nearly always results in a compara-
tive increase of fruits or reproductive organs.
It is an old horticultural maxim that checking
growth induces fruitfulness. It is largely in con-
sequence of this fact that plants bear heaviest
when they attain approximate maturity. Trees
are often thrown into bearing by girdling, heavy
pruning, the attacks of borers, and various acci-
dental injuries. The gardener knows that if he
keeps his plants in vigorous growth by con-
stantly potting them on into larger pots, he will
get little, or at least very late, bloom. The
plant-breeder, therefore, may be able to induce
the desired initial variation by attention to this
principle. (See discussion of variation in rela-
tion to food supply, page 16.) Arthur has re-
cently put the principle into this formula: “A
decrease in nutrition during the period of growth
of an organism, favors the development of the re-
productive parts at the expense of the vegetative
parts.”

A most important means of inducing variation

SIMULTANEITY OF VARIATION. ee

is the simple change of seed, the philosophical
reasons for which are explained on pages 59 and
28. A plant becomes closely fitted or accus-
tomed to one set of conditions, and when it is
placed in new conditions, it at once makes an
effort to adapt itself to them. ‘This adaptation
is variation. No doubt the free interchange of
seeds. between seed-merchants and customers is
one of the most fertile causes of the enormous
increase in varieties in recent times.

When once a novel variety appears, others of a
similar kind are likely soon to follow in other
places, and some persons have supposed that there
is a synchronistic variation in plants, or a tendency
for like variations to appear simultaneously in
widely separated localities. There is perhaps
some remote reason for this belief, because there
is, as Darwin expresses it, an accumulative effect
of domestication or cultivation, by virtue of which
plants which long remain comparatively invariable
may within a short time, when cultivation has
been continued long enough, vary widely and in
many directions; and it is to be expected that
even when plants have long since responded to
the wishes of the cultivator, an equal amount or
accumulation of the force of domestication would
tend to produce like effects in different places.
But it is probable that by far the greater part of
this synchronistic variation is simply an apparent

118 HOW DOMESTIC VARIETIES ORIGINATE.

one, for whenever any marked novelty appears the
attention of all interested persons is directed to
looking for similar variations amongst their own
plants.

12. The person who is wishing for new varieties
should look critically to all perennial plants, and
particularly to trees and shrubs, for bud-varieties
or sports. It has already been said (pages 28, 6)
that the branches of a tree may vary amongst
themselves in the same way in which seedlings
vary, and for the same reasons. As a rule, any
marked sport is capable of being perpetuated by
bud-propagation. The number of bud-varieties
now in cultivation is really very large. Many of
the cut-leaved and colored or variegated varieties
of ornamental plants were originally found upon
other trees as sports. The “ mixing in the hill”
of potatoes is bud-variation. Nectarines are
derived from the peach, some of them as sports
and some as seedlings. ‘The moss-rose was prob-
ably originally a sport from the Provence rose.
Greening apple trees often bear russet apples, and
russet trees sometimes bear greenings. So far as
I know, there are no varieties of annual plants
which have originated as sports. The probable
reason for this is the fact that the duration of the
plant is short and that its constitution is not pro-
foundly modified in a single generation by the
new circumstances in which it is placed every

BUD-VARIETIES. 119

year. The effects of the conditions in which it
lives are recorded in the seeds, and the plant dies
without allowing a second season of growth to
express the impressions which were received in
a former generation. The fact that every branch
of an annual plant—as of perennials —is unlike
every other branch, is evidence enough that the
annual is not unlike the perennial in fundamental
constitution ; and there is every reason to believe
that 1f any given annual were to become a peren-
nial, it would now and then develop differences
sufficiently pronounced to make them worthy the
name of sports, the same as hyacinths, bouvardias,
trees, and all other perennial plants are apt to do.

Bud-varieties may not only come true from buds
—as grafts, cuttings and layers, —but they occa-
sionally perpetuate themselves by seeds. Now,
these seedlings are amenable to selection, just the
same as any other seedlings are; the bud-variety,
therefore, may give the initial starting-point for
plant-breeding. But, more than this, it is some-
times possible to improve and fix the type by
bud-selection as well as by seed-selection. Dar-
win cites this interesting testimony: “ Mr. Salter
brings the principle of selection to bear on varie-
gated plants propagated by buds, and has thus
greatly improved and fixed several varieties. He
informs me that at first a branch often produces
variegated leaves on one side alone, and that the

120 HOW DOMESTIC VARIETIES ORIGINATE.

leaves are marked only with an irregular edging,
or with a few lines of white and yellow. To im-
prove and fix such varieties, he finds it necessary
to encourage the buds at the bases of the most
distinctly marked leaves and to propagate from
them alone. By following, with perseverance, this
plan during three or four successive seasons a dis-
tinct and fixed variety can generally be secured.”
Ernest Walker, a careful gardener at New Albany,
Indiana, is of the opinion that the abnormal char-
acter of sports often intensifies itself if the sport
is allowed to remain upon the parent plant for a
considerable time. He has observed this particu-
larly in coleus, where color sports are frequent.
“In these,” he says, “the sport begins with a
branch, which may be taken off and propagated as
anew variety. If left on the parent, other parts
of the plant are apt to show similar variations.
Indeed, I think it is not best to be in too great a
hurry to remove a sporting branch, for its char-
acter seems to tend to become more fixed if it
remains on the plant.”’

13. The starting-point once given, all permanent
progress lies in continued selection. ‘This, as I have
already pointed out, is really the key to the whole
matter. In the greater number of cases, the oper-
ator cannot produce the initial variation which he
desires, but, by looking carefully amongst many
plants, he may find one whith shows an indication

SELECTION THE KEY-NOTE. 121

of his ideal. This plant must be carefully saved,
and all the seeds sown in a place where crossing
with other types cannot take place. Of a hun-
dred seedlings from this plant, mayhap one or
two will still further emphasize the character
which is sought. ‘These, again, are saved and all
the seeds are sown. So the operation goes on,
patiently and persistently, and there is reward
at the end. ‘This is the one eternal and funda-
mental principle which underlies the amelioration
of plants under the touch of man; and because
we know, from experience, that it is so important,
we are sure, as Darwin was, that selection in
nature must be the most potent factor in the
progress of the vegetable world.

But suppose this suggestion of the new variety
does not appear amongst the batch of plants
which we raise? Then sow again; vary the con-
ditions; select the most widely variable types;
cross; at length —if the ideal is true —the sug-
gestion will come. “Cultivation, diversification
of the conditions of existence, and repeated sow-
ings” are the means which Verlot would employ
to induce variations. But the skill and the char-
acter of the final result he not so much in the
securing of the initial start, as in the subsequent
selection. Nature affords starting-points in end-
less number, but there are few men alert and skil-
ful enough to take the hint and improve it. If I

122 HOW DOMESTIC VARIETIES ORIGINATE.

want a new tomato, I first endeavor to discover
what I want. I decide that I must have one like
the Acme in color, but more spherical, with a
firmer flesh, and a little earlier. Then I shall
raise an acre of Acme tomatoes, and closely allied
varieties; or if I cannot do that, I make arrange-
ments to inspect my neighbor’s fields. I scruti-
nize every plant as the first fruits are ripening.
Finally, I find one plant — not one fruit — which
is something like the variety which I desire.
Very well! Wait two to five years, and you shall
see my new variety !

Some of these initial variations possess no ten-
dency to reproduce themselves. The seedlings of
them may break up into a great diversity of
forms, no form representing the parent closely.
In such cases, it is generally useless to proceed
further with this brood. Another start should be
made with another plant. So it is always impor-
tant, as we have already seen (Rule 6), to have as
many starting-points as possible, to lessen the risk
of failure. Whilst it requires nice judgment to se-
lect those plants which possess the most important
and the most transmissible combination of charac-
ters, the greatest skill is nevertheless required to
carry forward a correct system of selection.

14. Hven when the desired variety is obtained, tt
must be kept up to the standard by constant attention
to selection. ‘That is, there is no real stability in

SELECTION TO MAINTAIN PURITY. 5 83

the forms of plant life. So long as the conditions
of existence vary, so long will plants make the
effort to adapt themselves to the changes. No
two seasons are alike, and no two fields, or even
parts of fields, are alike; and there are no two
cultivators who give exactly the same and equal
attention to tillage, fertilizing, and the other
treatments of plants. All forms or varieties,
therefore, tend to’ “rum out” by variation or
gradual evolution into other forms ; but because
we keep the same name for all the succeeding
generations, we fancy that we still have the same
variety. In 1887 I found a single tomato plant
in my garden in Michigan, which had several
points of superiority over any other of the one
hundred and seventy varieties which I was then
growing. It came from a packet of German seed
of an inferior variety. The tomato was very
solid, an unusually long keeper, productive, and
attractive in size and appearance. The variation
was so promising that I named it in a sketch of
tomatoes which I published that year, calling it
the Ignotum (that 1s, unknown), to indicate that
the origin of it was no merit of my own. I sent
seeds to a few friends for testing. I sowed the
seeds for about five hundred plants in 1888 in
an isolated patch upon uniform soil. The larger
part of the plants were more OF less like the
parent. A few reverted. A few of the best

124 HOW DOMESTIC VARIETIES ORIGINATE.

plants were selected, and the seed saved. I
then moved to New York and took the seed with
me. This was sown in uniform soil in an iso-
lated position in 1889. ‘This crop, probably as
a result of the careful selection of the year before
and of the change of locality, was remarkably
uniform and handsome. Of the 442 plants which
I grew that year, none reverted to the little
Eiformige Dauer, the German variety from which
it had come, but there was some variation in them
due to different methods of treatment. I again
saved the seeds, and I was now ready to intro-
duce the variety. I therefore sold my seed, six
pounds, to V. H. Hallock & Son, Queens, New
York, who introduced it in 1890. The very next
year, 1891, I obtained the Ignotum from fifteen
dealers and grew the plants side by side. Of the
fifteen lots, eight bore small and poor fruits which
were not worth growing and which could not be rec-
ognized as Ionotum! Grown from our own seed,
it still held its characters well. Here, then, only
a year after its introduction, half the seedsmen
were selling a spurious stock. It is possible that
some of this variation arose from substitution of
other varieties by seedsmen, although I have yet
secured no evidence of any unfair dealing. It is
possible, also, that the product of some of the
samples which I early sent out for testing had
found their way into seedsmen’s hands. But I am

DURATION OF VARIETIES. 125

convinced that very much of this variation was
a legitimate result of the various conditions in
which the crops of 1890 had been grown, and the .
varying ideals of those who saved the seeds. I
am the more positive of this from the fact that
the Ignotum tomato, as I first knew it and bred
it, appears to be lost to cultivation, although the
name is still used for the legitimate family of
descendants from my original stock. All this
experience illustrates how quickly varieties pass
out by variation and by the unconscious and
unlike selection practised by different persons.
The duration of any variety is inversely propor-
tional to the frequency of its generations. Annual
plants, other conditions being the same, run out
sooner than perennials, because seed-reproduc-
tion—or the generations—intervenes more fre-
quently. ‘Trees, on the other hand, carry their
variations longer, because the seed-generations —
in which departures chiefly take place — are far-
ther apart. Of all the so-called fruit plants, the
strawberry runs out soonest and the varieties
change the oftenest, because a new generation can
be brought into fruit-bearing in two years, whilst
it may require a decade or more to bring a new
generation of apples or chestnuts into bearing.
Yet, my reader will remind me that the Wilson
strawberry has been and is the leading variety in
many places for nearly forty years, to which I

126 HOW DOMESTIC VARIETIES ORIGINATE.

reply that the Wilson of to-day is not necessarily
the same as that introduced by James Wilson,
simply because the name is the same. Every dif-
ferent soil or treatment tends to produce a different
strain or variation in the Wilson strawberry, as it
does in any other plant ; and every grower, when
setting a new plantation, selects his plants from
that part of his field which pleases him best,
rather than from those plants which most nearly
correspond to the original type of the Wilson.
That is, this unconscious selection on the part of
the grower takes no account of what the variety
was, but only of what it ought to be, and this
ideal differs with every person. It is not surpris-
ing, therefore, to find strains of Wilson strawberry
which are as unlike as many named varieties are ;
and it is to be expected that all of the strains now
in existence have departed considerably from the
original type.

This example borrowed from the strawberry is
a most important one, because it illustrates how a
variety may vary and pass out of existence even
though it is propagated wholly asexually, or by
buds. ‘There are to-day several different types of
Rhode Island Greening apple in cultivation, which
have originated from variations produced by enyi-
ronment and by the different models which propa-
gators have had in mind; and the same is true of
many other fruits.

AMELIORATION DUE TO SELECTION. 127

All the foregoing remarks demonstrate the
importance of constant attention to selection if
one desires to maintain the exact type of any
variety which he has produced. They explain
the value of the ‘“‘roguing’’—or systematic de-
struction of all “rogues” or non-typical plants —
which is invariably practised by all good seed-
erowers. But they still more emphatically show
that every variety is essentially unstable, and that
the only abiding result is constant evolution, the
old forms being left behind as the type expands
into new and better strains. Varieties to be valu-
able, therefore, ought not to be rigidly fixed, and,
fortunately, nature has prescribed that they can-
not be. Probably every decade sees a complete
change in every variety of any annual species
which is propagated exclusively from seeds, and
every century must see a like change in the tree
fruits. ‘These changes are so gradual, and the
original basis of comparison fades away so com-
pletely, that we generally fail to recognize the
evolution.

15. It is evident, therefore, that the most abiding
progress in the amelioration of plants must come as a
result of the very best cultivation and the most intel-
ligent selection and change of seed. Every reflec-
tive person must admit that the cultivation of
plants — which is the fundamental conception of
agriculture — has been and is crude and imperfect,

128 HOW DOMESTIC VARIETIES ORIGINATE.

and that there has been no conscious effort on the
part of the human race to produce any given final re-
sult upon the cultivated flora. Yet, this imperfect
cultivation has already modified plants so pro-
foundly that we cannot determine the originals of
many of them, and we can trace the evolution of
but few. The science of rural industry is now
fairly well understood in its essential fundamental
principles, and the intelligence of those classes of
persons who deal with plants is rapidly enlarging.
The opening of the twentieth century will virtu-
ally mark a new era for agriculture, and from
that time on the onward evolution of plants should
proceed confidently and unchecked. Our eyes are
too often dazzled by the novelties which suddenly
thrust themselves upon us, and we look for some
mystic power which shall enable us to produce
varieties forthwith at our will. We need not so
much varieties with new names as we do a general
increase in productiveness and efficiency of the
types which we already possess ; and this augmen-
tation must come chiefly in the form of a gradual
evolution under the stimulus of good care. The
man who will accomplish most for the amelioration
and unfolding of the forms of plants, is he who
fixes his eyes steadily upon the future, and with
the inspiration of along forecast, urges the better-
ment of all conditions in which plants grow.

DEWBERRY AND BLACKBERRY. 129

III. Sprctric EXAMPLES.

The foregoing principles and discussions will
become more concrete if a few actual examples of
the origination of varieties are given. In order
to begin with a very simple case, I will relate the
introduction of the varieties of dewberries, for
this fruit is yet little cultivated, the varieties are
few, and the domestication of it is not yet thirty
years old.

The Dewberry and Blackberry.

The dewberries are native fruits, and it is only
within the last ten years that they have become
prominent among fruit-growers. The most impor-
tant one is the Lucretia. This was found grow-
ing wild upon a plantation in West Virginia in
war time. In 1876, a few of the plants were sent
to Ohio, and from this start the present stock has
come. It is probable that similar wild varieties are
growing to-day in many parts of the country, but
they have not chanced to have been seen by per-
sons who are interested in cultivating them. It is
a form of the common wild dewberry, which grows
all over the northeastern states. Just why this
particular patch in West Virginia should have been
so much better than the general run of the species,
nobody knows, but it was undoubtedly the prod-
uct of some local environment of soil or position.

K

130 HOW DOMESTIC VARIETIES ORIGINATE.

Early in the seventies, IT’. C. Bartel, of Huey,
Clinton Co., Illinois, observed very excellent dew-
berries growing in rows between the lines of
stubble in an old cornfield, where the plant had
evidently been quick to avail itself of unoccupied
land. This was introduced as the Bartel dew-
berry, and is now the second in point of promi-
nence amongst the cultivated varieties. Other
varieties have appeared in much the same way.
A fruit-grower in Michigan found an extra good
dewberry in a neighboring wood-lot, and intro-
duced it under the name of Geer, in compliment
to the owner of the place. In Florida, an un-
usually good plant of the common wild dewberry
of that region was discovered, and introduced by
Reasoner Brothers, under the name of Manatee.
There are now about twenty named varieties of
dewberries in cultivation, as described in our
horticultural writings, all of which, so far as I
know, are chance plants from the wild.

As the dewberries become more widely grown,
good seedlings will now and then appear in cul-
tivated ground, and these will be named and
sold. After a time persons will begin to sow
seeds for the purpose of producing new varieties ;
and those seedlings which chance to possess un-
usual merit will be propagated, and in due time
introduced. This is the history of the cultivated
blackberries and raspberries which have come

EVOLUTION OF THE APPLE tat

from the wild plants in less than half a century.
These fruits are now so far developed that we no
longer think of looking to the woods and copses
for new varieties of promise, but the novelties are
mostly chance seedlings from cultivated varieties.
A few years ago a friend purchased plants of the
Snyder blackberry. When they came into bear-
ing he noticed that one plant was better than the
rest. It bore larger fruits, and the bearing season
was longer. He took suckers from this plant,
and from these others were taken, until he now has
a large plantation of the novelty, mostly selected
from plants which pleased him best. The variety
has such distinct merit that I have named it the
Mersereau, in honor of the man who recognized
and propagated it. He will continue selecting
from the best plants, as he propagates year by
year, and it may be that in a few years he will
have so much improved it that it will no longer
be the variety with which he started.

The Apple.

The original apple is not definitely known, but
it was certainly a very small and inferior, crabbed
fruit, borne mostly in clusters. When we first
find it described by historians, it was still of small
value. Pliny said that some kinds were so sour
as to take the edge off a knife. But better and

132. HOW DOMESTIC VARIETIES ORIGINATE.

better seedlings continued to come up about habi-
tations, until, when printed descriptions of fruits
began to be made, three or four hundred years
ago, there were many named kinds in existence.
The size had vastly improved, and with this in-
crease came the reduction of the number of fruits
in the cluster; so that, at the present time, whilst
apple flowers are borne in clusters, the fruits are
generally borne singly. That is, most of the
flowers fail to set fruit, and they complete their
mission when they have shed their pollen for the
benefit of the one which persists.

The American colonists brought with them the
staple varieties of the mother countries. But
the needs of the new country were unlike those
of the old, and the tastes and fashions of the
people were changing. So, as seedlings came up
about the buildings and along the fences, where
the seeds had been scattered, the ones which prom-
ised to satisfy the new needs best were saved, and
many of the old varieties were allowed to pass
away. In 1817, the date of the first American
fruit-book, over sixty per cent of the varieties
particularly recommended for cultivation in this
country were of American origin. In 18465,
nearly two hundred varieties of apples were de-
scribed as having been fruited in this country,
of which over half were of American origin. Be-
tween these two dates, introductions of foreign

EVOLUTION OF THE APPLE. 133

varieties had been freely made, so that the per-
centage of domestic varieties had fallen. But
the next thirty years saw a great change. Of
1823 varieties described in 1872, nearly or quite
seventy per cent were American, and a still greater
proportion of the most prized kinds were of
domestic origin. In the older states, the apple
had now become so thoroughly accustomed to its
environment, and the tastes of the people were
so well supplied, that there was no longer much
need for the introduction of foreign kinds. It
was not so in the Northwest. There the apples
of the eastern states did not thrive. The climate
was too cold and too dry. Attention was turned
to other countries with similar or rigorous cli-
mate. In 1870, the Department of Agriculture
at Washington imported cions of many varieties
of apples from Russia; but these did not satisfy
many fruit-growers of the northern states. It
was then conceived that the great interior plain
of Russia should yield apples adapted to the upper
Mississippi valley, whilst those already imported
had come from the seaboard territory. Accord-
ingly, early in the eighties, Charles Gibb, of the
province of Quebec, and Professor Budd, of Iowa,
went to Russia to introduce the promising fruits
of the central plain. ‘The result has been a most
interesting one to the pacific looker-on. There
are ardent advocates of the Russian varieties, and

134 HOW DOMESTIC VARIETIES ORIGINATE.

there are others who see nothing good in them.
There are those who believe that all progress
must come by securing seedlings from the hardi-
est varieties of the eastern states; there are others
who would derive everything from the Siberian
erabs, and still others who believe that the final
result lies in improving the native crabs. There
is no end of discussion and cross-purposes. In
the meantime, nature is quietly doing the work.
Here is a good seedhng of some old variety, there
a good one from some Russian, and now and then
one from the crab stocks. ‘The new varieties are
gradually supplanting the old, so quietly that few
people are aware of it; and by the time the con-
testants are done disputing, it will be found that
there are no Russians and no eastern apples, but
a brood of northwestern apples which have grown
out of the old confusion.

All these new apples are simply seedlings,
alinost all of them chance trees which come up
here and there wherever man has allowed nature
a bit of ground upon which to make garden as
she likes. In 1892, there were 878 varieties of
apples offered for sale by American nurserymen,
and it is doubtful if one in the whole lot was the
result of any attempt on the part of the originator
to produce a variety with definite qualities. And
what is true of the apple, is about equally true
of the other tree fruits. In the small fruits and

BEANS. 135

the grapes, where the generations are shorter and
the results quicker, more has been done in the
way of direct selection of seeds and the crossing
of chosen parents; but even. here, the methods
are mostly haphazard.

Beans.

Perhaps there are no plants more tractable in
the hands of the plant-breeder than the garden
beans. Some two or three years ago, a leading
eastern seedsman conceived of a new form of bean
pod which would at once commend itself to his
customers. He was so well convinced of the
merits of this prospective variety, that he made
a descriptive and “ taking” name for it. He then
wrote to a noted bean-raiser, describing the pro-
posed variety and giving the name. “Can you
make it for me?” he asked. ‘Yes, I will make
you the bean,” replied the grower. ‘The seeds-
man then announced in his catalogue that he
would soon introduce a new bean, and, in order
to hold the name, he published it, along with the
announcement. ‘Two years later, I visited the
bean-grower. ‘Did you get the bean?” I asked.
“Yes, here it is.” Sure enough, he had it, and it
answered the requirements very well. Another
seedsman would like a round-podded, stringless,
ereen-podded bean. This same man produced

136 HOW DOMESTIC VARIETIES ORIGINATE.

it, and I went into a field of fifteen acres of it,
where it was growing for seed, and the most fas-
tidious person could not have asked for a closer
approach to the ideal which the breeder had set
before him some four or five years before.

How is all this done? It looks simple enough.
The ideal is established first of all. The breeder
revolves it in his mind, and eliminates all the
impracticable and contradictory elements of it.
Then he goes carefully and critically through his
bean fields, particularly those varieties which are
most like the desired kind, and marks those plants
which most nearly approach his ideal The seeds
of these are carefully saved, and they are planted
in isolated positions. If he finds no promising
variations amongst his plantations, then he must
start off the variation in some other way. ‘This
is usually done by crossing those varieties which
are most like the proposed kind. He has got a
start; but now the science and skill begin. Year
by year he selects just those plants which please
him best and which he judges, from experience,
will most surely carry their features over to the
offspring. He starts with one plant; the next
year he may have only two. If he has ten or
twenty good ones, then the task is an easy one,
for the variety has elements of permanence — that
is, of hereditability —in it. But he may have no
plants the second year. In that case, he begins

BEANS. 137

again ; for if the ideal is true, it can be attained.
This bean-breeder to whom I have referred, and
upon whom many of our best seedsmen rely for
new varieties, tells me that he has discarded fully
three thousand varieties and forms as profitless.
This only means that he is a most astute judge
of beans, and that he knows when any type is
likely to prove to be a poor breeder.

The bean also affords an excellent example of
the care which it is generally necessary to exercise
to keep any variety true to the type. The person
of whom I have spoken, in common with all care-
ful seed-growers, searches his field with great
pains to discover the “rogues,” or those plants
which vary perceptibly from the type of the given
variety. The rogue may be a variation in size or
habit of plant, season of maturity, color or form
of pods, productiveness, susceptibility to rust, or
other aberrance. In the dwarf or bush beans,
which are now most exclusively grown, the most
frequent rogue is a climbing or half-climbing
plant. This is a reversion to the ancestral type
of the bean, which was no doubt a twining plant.
This rogue is always destroyed, even though it
may be, itself, a good bean. In some cases, the
men who perform the roguing are sent along
every row of a whole field on their hands and
knees, critically examining every plant. The ef-
fect of this continual selection is always to push

138 HOW DOMESTIC VARIETIES ORIGINATE.

on the variety to greater excellence. The vari-
ous “improved” strains of plants are obtained in
essentially this fashion. If the grower has been
painstaking with his roguing, he soon finds that
his seed gives better and more uniform crops than
the common stock of the variety. If the improve-
ment is marked, he may dignify his strain with
a distinct name, and it thereby becomes a new
variety. The improvement may be a very im-
portant one to a careful bean-grower, and at the
same time be so slight as to escape the attention
of the general farmer, or even of experimenters
who are not particularly skilled in judging the
merits of beans.

All these examples drawn from the bean are
excellent illustrations of the best and most scien-
tific plant- breeding, and the same methods — varied
to suit the different needs — apply to the ameliora-
tion of all other plants. The recent dwarf Lima
beans may be cited as examples of accidental or
fortuitous varieties, in which the preconstructed
ideal of the plant-breeder had no place. Four or
five of these beans have attained some prominence.
Henderson and Kumerle dwarf Limas were intro-
duced in 1889, Burpee in 1890, and Barteldes in
1892 or 1893. The variety which is now called
the Henderson was picked up twenty or more
years ago by a negro, who found it growing along
a roadside in Virginia. It was afterwards grown

BEANS. 139

in various gardens, and about 1885 it fell into the
hands of a seedsman in Richmond. Henderson
purchased the stock of it in 1887, grew it in
1888, and offered it to the general public in 1889.
The introduction of Henderson’s bean attracted
the attention of Asa Palmer, of Kennett Square,
Pennsylvania, who had also been growing a dwarf
Lima. He called upon Burpee, the well-known
seedsman of Philadelphia, described his variety,
and left four beans for trial. These were planted
in the test grounds and were found to be valuable.
Mr. Palmer’s entire stock was then purchased, —
comprising over an acre, which had been carefully
inspected during the season—and Burpee Bush
Lima was presented to the public in the spring of
1890. Mr. Palmer’s dwarf Lima originated in
1883, when his entire crop of Large White (Pole)
Limas was destroyed by cut-worms. He went
over his field to remove the poles before fitting
the land for other uses, but he found one little
plant, about ten inches high, which had been cut
off about an inch above the ground but which had
re-rooted. It bore three pods, each containing one
seed. These three seeds were planted in 1884,
and two of the plants were dwarf, like the parent.
By discarding all plants which had a tendency to
climb, in succeeding crops, the Burpee Bush Lima,
as we now have it, was developed. The Kumerle,
Thorburn, or Dreer, Dwarf Lima originated from

140 HOW DOMESTIC VARIETIES ORIGINATE.

occasional dwarf forms of the Challenger Pole
Lima, which J. W. Kumerle, of Newark, New
Jersey, found growing in his field. The stock
which came from these selected dwarf plants was
introduced by Thorburn and Dreer, under their
respective names. The singular Barteldes Bush
Lima came from Colorado, and is a similar dwarf
sport of the old White Spanish or Dutch Runner
bean. Barteldes received about a peck of the
seed and introduced it sparingly. It attracted
very little attention, and as the following season
was dry, Barteldes himself failed to get a crop,
and the variety was lost to the trade.

Cannas.

Few plants have shown more remarkable evolu-
tions in very recent years than the cannas. At
the present time, the Crozy cannas—so named
from Crozy, of Lyons, France, who has introduced
the greater number of them —are most popular.
This type is often called the French Dwarf, or
the Flowering Canna, and it is marked by a com-
paratively low stature, and very large and showy
spreading flowers in many colors, whereas the can-
nas of a few years ago were very tall plants, with
small and late dull red, narrow flowers, and they
were grown exclusively for their foliage effects.
How has this transformation come about ?

CANNAS. 141

In the first place, it should be said that there
are many species of canna, and about a half dozen
of these were well known to gardeners at the
opening of the century. About 1830, the cannas
began to attract much attention from cultivators,
and the original species were soon variously hybrid-
ized. Crossed seeds, and seeds from the succes-
sive generations of hybrids, introduced a host of
new and variable forms. ‘The first distinct fash-
ion in cannas seems to have been for tall, late-
flowering forms. In 1848, Année, a cultivator in
France, sowed seeds of Canna Nepalensis, a tall
oriental species, and there sprung up a race of
plants which has since been known as Canna
Annei. It is probable that this Canna Nepalensis
had become fertilized with other species growing
in Année’s collection, very likely with Canna
glauca. At any rate, this race of cannas became
popular, and was to its time what the French
dwarfs are to the present day. ‘The plants were
freely introduced into parks, beginning about
1856, but their use began to wane by 1870
or before. Descendants of this type, variously
crossed and modified, are now frequently seen
in parks and gardens.

The beginning of the modern race of dwarf,
large-flowered cannas was in 1863, when one of
the smaller-flowered Costa Rican species (Canna
Warscewiczii) was crossed upon a large-flowered

142 HOW DOMESTIC VARIETIES ORIGINATE.

Peruvian species (Canna iridiflora). The off-
spring of this union came to be called Canna
EKhemanni. This hybrid has been again variously
crossed with other species, and modified by culti-
vation and selection, until the present composite
type is the result. Seeds give new varieties; and
any seedling which is worth saving is thereafter
multiplied by divisions of the root, and the result-
ing plants are introduced to commerce.

These various examples are but types of what
has been and can be accomplished in a given group
of plants. There is nothing mysterious about the
subject, so far as the cultivator is concerned. He
simply sets his ideal, makes sure that it does not
contradict any of the fundamental laws of devel-
opment of the plant with which he is to work,
then patiently and persistently keeps at his task.
He must have good judgment, skill, and inspira-
tion, but he does not need genius.

“In the improvement of plants,” writes Henri
L. de Vilmorin, “the action of man, much like
influences which act on plants in the wild state,
only brings about slow and gradual changes, often
scarcely noticeable at first. But if the efforts
toward the desired end be kept on steadily, the
changes will soon become greater and greater, and
the last stages of the improvement will become
much more rapid than the first ones.”

LECTURE IV.

BORROWED OPINIONS; BEING EXTRACTS FROM
THE WRITINGS OF B. VERLOT, E. A. CARRIERE,
AND W. O. FOCKE.

I. VERLOT’S CLASSIFICATION OF VARIETIES OF
ORNAMENTAL PLANTS.

VerRLor (Sur la Production et la Fixation des
Variétés dans les Plantes d’ Ornement) distributes
the varieties of ornamental plants into sixteen
groups. I shall now transcribe these groups, and
under each shall give a very brief quotation or
abstract of some of his remarks concerning them.

1. Varieties distinguished by diminution of stat-
ure, or dwarfing. — Dwarfing is one of the most
frequent variations in the vegetable kingdom, but,
unlike many similar phenomena in the animal
kingdom, these dwarfs are nearly always very
fertile. If we question the cultivators upon the
subject, they respond that these variations are
purely accidental, and that whenever the varia-
tions offer any reward they are propagated and
distributed.

Dwarfing may be brought about by sowing the
seeds in the autumn (page 115), and at the same

145

144 BORROWED OPINIONS.

time successively transplanting the plants, as they
need. Suppose, for example, we sow seeds of Co-
reopsis tinctoria in August or September. When
the plants have developed leaves, they are trans-
planted, leaving sufficient space between them
to allow of liberal growth. When the plants
begin to touch each other, transplant again, per-
haps three or even four times. The plants
become strong, vigorous, and stocky ; we encour-
age the development of the lowest branches and
thereby tend to shorten the leading stem, thus
making the individual comparatively dwarf. The
seeds saved from plants thus treated during several
generations will be more apt to produce dwarf
varieties than seeds taken from other plants.
The greater part of dwarf varieties appear in
those plants which are sown in autumn, and in
those, if sown in spring, which are submitted to
successive transplantings. Thus, amongst the -
annual species which we habitually sow in July
and September, the following have produced dwarf
varieties : —

Calceolaria plantaginea.

Senecio cruentus.

Lychnis (or Agrostemma) Cceli-rosa.

Coreopsis (or Calliopsis) tinctoria.

Qinothera Drummondii.

Helichrysum bracteatum.

Leptosiphon densiflorus.

VERLOT’S CLASSIFICATION. 145

Dianthus Chinensis.
Scabiosa atropurpurea.
Schizanthus retusus.
Iberis umbellata.

Amongst those species which we sow in spring,
but frequently transplant, the following have
dwarf forms : —

Impatiens Balsamina.
Callistephus hortensis.
Tagetes patula.
Tagetes erecta.
Tagetes signata.

2. Varieties distinguished by augmentation of
stature, or giant forms. —'These varieties result
from various causes, amongst which are amount
and fertility of soil, the employment of newly
harvested seeds, and crossing.

3. Hardy varieties. —These are produced by
successive selections from the most hardy indi-
viduals. Hardy races are also obtained by cross-
ing with hardy species or types. ‘Thus, the forms
of Rhododendron arboreum are rendered hardier
when crossed with R. Catawbiense.

4. Large-flowered varieties. —These variations
are always due to a good soil which is rich in humus,
and above all by thorough and intelligent cultiva-
tion. ‘These are easily fixed, but they pass away

L

146 _ BORROWED OPINIONS.

insensibly when the conditions under which they
were produced are neglected. This return to
small flowers is well illustrated in the pansy. [All
plants which are grown for the beauty of their
flowers tend to increase the size of those organs,
because of the vigor which comes of good care,
and the selection which necessarily follows. The
frequency of the varietal name “ grandiflora” is
proof of this. As soon as the plant has made any
perceptible gain in the size of its flowers, some
nurseryman adds this adjective to its name, as in
Hydrangea paniculata grandiflora. No doubt the
name is sometimes bestowed without warrant, for
the purpose of selling the plant. Frequently the
catalogue-maker drops the proper specific name
and uses the varietal adjective as if it were the
legitimate name of the plant, as in Gazania grandi-
flora, which is properly Gazania rigens, variety
grandiflora. |

5, 6. Early and late varieties. — The chief
agency affecting the duration of plants is climate,
and desired variations in earliness or lateness are
obtained by transporting the plants to those cli-
mates which produce such effects as we seek, and
erowing them there for a time; or we can procure
seeds from such climates, if the given plants are
already grown there.

The age of the seed also has an influence upon
the resulting individual: the fresher the seed, the

VERLOT’S CLASSIFICATION. 147

more rapid its germination, and consequently the
more prompt the development of the plant. We
have reason to expect that fresh seeds will have a
tendency to produce early varieties, and that, on
the contrary, old seeds, by germinating more
slowly, will produce variations more or less late.
[Not only does the age of the seed seem to be
important in this connection, but recent experi-
ments seem to show that the degree of maturity
also modifies the offspring. Seeds which are
barely ripe enough to germinate have a tendency
to give earlier progeny than those which are fully
matured and ripened. See page 108. |

The first seeds to mature on any plant may be
expected to give early plants, and the subsequent
seeds give later plants. [Verlot states that a cold
region tends to make the plants later when trans-
ferred to it, but this seems to be an error. ‘The
fact is generally just the reverse. Plants taken
towards the poles or to higher altitudes, become
earlier in two ways, by shortening their period
of growth, and by vegetating at a less sum-
temperature in spring. See page 26. |

7. Odoriferous varieties. — Odor varies greatly,
even amongst varieties of one species. ‘The causes
of the differences in fragrance are not numerous
and they are little understood. Climate, exposure,
and the nature of the soil are leading factors.
The odor of plants which grow on dry and arid

148 BORROWED OPINIONS.

hills is much more penetrating than that of the
same species cultivated in humid and shady places.
It is possible, even, to entirely change the odor by
transporting the plant from one place to another.
For example, Satyrium hircinum exhales a most
pronounced goat-like odor in the vicinity of Paris
and northward, whilst in the east, and particularly
in the southern regions, its flowers have an odor
which is somewhat hke that of vanilla.

8. Varieties with colored parts.— Coloration may
be either complete or partial, and it may reside
in any or all parts of the plant, as follows: —

The stems.
The leaves.
variegated,

fl \
The flowers, coated

The fruits.
The seeds.

Variations in color are the most frequent of all
modifications in cultivated plants. These depart-
ures may be expected to arise under the influence
of continued cultivation and repeated sowings;
and the variations must then be selected until
they are fixed.

9. Varieties without color, or niin = Partial
albinism, or variegation, is as frequently observed
in spontaneous plants as it is in cultivated ones.
It usually occurs in the leaves only, but it is some-

VERLOT’S CLASSIFICATION. 149

times a feature of the entire plant. A variegated
plant does not exist of which we do not know the
non-variegated type. These variegated plants
appear both from seeds and from bud-variations,
and they are most surely propagated in the latter
case. It has been said that when the albinism
affects the margin of the leaf it is more likely to
be transmitted than when it occupies the central
part of the blade, but this generalization has many
exceptions. Page 157.

It is a curious fact that variegation and double-
ness of flowers are generally antagonistic, for they
do not appear in the same plant. One excludes
the other. It is supposed by Morren [and gen-
erally accepted] that doubling is the result of
excessive vigor and that partial albinism comes
of an enfeebling of the vital functions.

Variegations sometimes disappear entirely and
then, after two or three years, reappear in the
same individual. The first leaves of seedlings
from variegated plants may be perfectly green,
and the seedlings may afterwards take on the varie-
gated character. This behavior is well marked
in some ferns. .

Complete albinism, or chlorosis, indicates a pro-
found alteration in the tissues, and it is impossi-
ble of propagation. This decoloration is most
commonly a bud-variation.

10. Double varieties, or those distinguished by

150 BORROWED OPINIONS.

the transformation of the stamens and pistils into
petaloid organs —'There are various degrees of
doubling or duplication in flowers. ‘The calyx
and corolla alone may be duplicated, in which
case the flower is still fertile. Sometimes the sta-
mens only are transformed into petal-lke organs,
and the flower is then fertile if pollen is trans-
ferred from another flower. Sometimes all the
floral series — calyx, corolla, stamens, pistils —
may be duplicated or transformed ; then we have
a full (pleine) flower, which is incapable of pro-
ducing seeds or of fecundating another flower.
[Annual plants, and others not propagated by
buds and other asexual parts, which bear full
double flowers, must be propagated by seeds taken
from flowers which are nearly full double, but
which bear a few seeds; or, sometimes from a
nearly single flower which is fertilized by pollen
from a nearly full double flower. In these cases,
it is unusual for all the seedlings to produce full
double flowers.]

A rich soil, a cultivation which produces a luxu-
riant vegetation, are the conditions which gen-
erally produce doubling in flowers. But we can
repeat with De Candolle, “ That if we are gen-
erally ignorant of the causes of the doubling of
flowers, we also know that if we gather seeds
from an individual with semi-double flowers, the
plants which result have a greater tendency to pro-

VERLOT’S CLASSIFICATION. 161

duce double flowers than seeds taken from simple
flowers.”

Doubling may occur in all plants, whether an-
nuals, biennials, herbaceous or woody perennials,
and in all of them, when they are fertile, we can
finally make them reproduce the character iden-
tically.

We must always choose for seed-parents the
individuals of which the flowers are very double,
and exclude with the greatest care the single-
flowered plants, which are the most fertile and
the progeny of which quickly smother the progeny
of the double flowers.

11. Proliferous varieties. —['These are varia-
tions which are characterized by growths arising
from unusual places, as one flower springing out
of another flower, a branch or rosette growing out
of a flower, an unusual production of bulbs or
young plants from the root, bulbs from leaves or
the fronds of ferns, and the like.] These forms
are infrequent in cultivated plants and very rare
amongst wild plants. ‘They are generally asso-
ciated with the fertility of the soil. The pro-
liferous form of Papaver somniferum known as
Papaver monstruosum, perpetuates itself per-
fectly by seeds, but these variations are usually
unstable.

12. Varieties with conjoined parts (variétés par
soudures).— We know of a single example of this

152 BORROWED OPINIONS.

monstrosity in ornamental plants: it is that of
Papaver bracteatum, in which the corolla has
become monopetalous by the growing together of
the petals. ‘This monstrosity (described and fig-
ured in Revue Horticole) is cultivated by Vilmorin.
They can propagate it only by cuttings. They
have tried in vain to multiply it by seeds. [Simi-
lar forms of other species are known. |

12. Abortive varieties. —This type of monstros-
ity, which constitutes one of the most interesting
chapters in vegetable teratology, has been ob-
served in all parts of the flower. ‘They are mal-
formations which have no interest from the point
of view of ornament. [The petals, or other or-
gans, sometimes almost entirely disappear in this
type of variations. ]

14. Pelorie varieties. —[Peloria is a name ap-
plied by Linneus to a form of the toad-flax, Lina-
ria vulgaris, in which all of the five petals have
spurs, while the normal form has only one petal
spurred. ‘The term is now appled generically to
all similar regularity of structure in normally ir-
regular flowers.] The causes which produce this
transformation are not known, but aridity and dry-
ness of the soil, and new conditions of vegetation,
appear to favor its development. These mon-
strosities, at least in linaria, are propagated easily
by cuttings or buds, and Willdenow records an
experiment in which they came true from seeds.

VERLOT’S CLASSIFICATION. 153
15. Chloranthic varieties. — Here are included,
in a general way and in lieu of a better name, all
those transformations which render the flowers
absolutely sterile, and transform them more or
less completely into branches or leaf-lke organs.
They are purely bud-variations, and can be per-
petuated only by cuttings, buds, or other asexual
parts.
16. Various or polymorphous varieties, compris-
ing the following types : —

‘ thornless,
| spineless,
Stems { fastigiate,
filiform,
ae etc.
{ crisped,
areas fasciated,
bullate or blistered,

laciniate, ete.

The various modifications originate both by
seed-variation and bud-variation.

II. CARRIERE’s ACCOUNT OF BUD-VARIETIES.

The subject of bud-variations or sports never
fails to interest the student, and however familiar
he may be with these forms he never ceases to

154 BORROWED OPINIONS.

wonder at them. I have taken pains, therefore,
in addition to what I have already said upon the
subject (see pages 28, 117), to translate almost
bodily Carriére’s account of bud-varieties (an Pro-
duction et Fixation des Variétés dans les Végé-
taux), because, although written in 1865, it is
the most extended lst of bud-varieties which I
know. The catalogue might be greatly extended
by inserting the current varieties in commerce in
this country, but the original list is sufficiently
full for all purposes of illustration. Carriere’s
account now follows :—

1. General Remarks upon Bud- Variation.

Plants being composed of a certain number of
elements disposed in a certain order, and, more-
over, these elements, under the influence of organic
laws, being able to separate or group themselves
in different ways, it follows that the same plant
can, upon its different parts, present characters
and properties more or less different from those
which it normally presents. It is this fact which
constitutes that which in practice we call an acci-
dent [or bud variety], either of dimorphism? [of
form] or of dichroism [of color].

1 French writers use the word accident in the sense in which
we use bud-variation. The word dimorphism, used by Car-
riére for one of the features of bud-variation, is now applied to

CARRIERE’S REMARKS ON BUD-VARIATION. 155

We refer to bud-variation the phenomenon,
whose cause is unknown, which allows a bud on
any part of the plant to develop a member whose
form and appearance differ from those borne on
other parts of the plant. ‘Thus the common beech
producing a branch with laciniate leaves, Podocar-
pus Koraiana producing a branch whose ramifi-
cations are whorled and spreading instead of being
scattered, and whose leaves are distichous instead
of being alternately disposed about the branches
as they are normally, are examples of bud-varie-
ties.

Taken in its most absolute sense and considered
in the sum of all its characters, bud-variation, aside
from the details which it presents, can be divided
into two sections: one which includes all the phe-
nomena which are manifested suddenly, as in the
case of the fern-leaved beech, the hemp-leaved
rose, the English willow-leaved cherry, sour
grapes with long seeds, etc.; the other includes
all slower transformations, as in the case of Rosa
Eglanteria, tulips, Iris Xiphium, Viola Rotho-
magensis, var. pallida, etc. Strictly, we could
establish a third section to include all the trans-
formations resulting from the age of the individ-

different permanent and characteristic forms of individuals of
the same species. It is most commonly observed in the different
relative lengths of stamens and pistils. I have substituted other
words for it in most places in the text. —L. H. B.

156 BORROWED OPINIONS.

ual, which are the consequences of its adultness.
However, this last series of phenomena is seen
only in polymorphous species, which change in
appearance, form, and nature when they grow old
and especially when they bear fruit; such are the
ivies, Ficus stipulata or scandens, eucalyptus, ete.
Horticulture often profits by this peculiar prop-
erty of plants; multiplying separately the parts
with the exceptional characters, it obtains individ-
uals which present an appearance different from
the plants from which they arise. [This dissimi-
larity between young and mature individuals of
the same species is well marked in some of the
Conifers, as the cedars and retinosporas. ]

In a general way, then, dimorphism refers to
a different form on the same individual, whether
the change be complete or partial. Dichroism is
exactly analogous to it in essential points, only
that it refers to color instead of to form. Thus,
Flon’s Pink [Dianthus semperflorens of gardens,
introduced by M. Flon of Angers], which has red
flowers, developing a branch similar to the plant
in aspect and form, but bearing white flowers,
the ovate-leaved privet and the Japanese fusain
[EKuonymus Japonicus] producing buds giving
rise to variegated leaves, white kidney-beans pro-
ducing black ones, and vice versa, are examples of
dichroism.

Let us say that in bud-variation, less than any-

CARRIERE ON BUD-VARIATION. LSz

where else, we can do nothing towards obtaining
or producing the variations. Bud-varieties most
often spring up spontaneously, so to speak, and in
this respect our work is purely passive, consisting
in superintending these digressions or accidents
in the endeavor to take advantage of them when
they are presented. Let us state, also, that in
these series of varieties we find a considerable
diversity, either in the habit or aspect of the
plants or in their foliage or flowers, or sometimes
even in their fruits, and that we oftener find vari-
egations than among plants which come from seeds.

We ought to recall just here, —what we have
said on the subject of plants issuing from seeds, —
that variegations are the more constant the more
completely they circumscribe the organs upon
which they occur, whether upon the flowers or the
leaves; also, that when, on a plant whose variega-
tions are disposed in stripes or bands, we find a
part upon which they are disposed circularly, we
can be almost certain that, if we detach and graft
or make a cutting of this part, we shall preserve
its new character. This phenomenon is very fre-
quent in the camellias and especially in the azaleas.
The greater part of the varieties of azalea which
present these characters have had no other origin.*

1'To preserve variegations, it is best to resort to graftage,
generally speaking, as cuttings tend to produce individuals
more vigorous, and which therefore tend to return to the green

158 BORROWED OPINIONS.

Certain species are much more disposed than
others to produce these bud-varieties, either of
dimorphism or dichroism. We give an example
from the Chinese chrysanthemum. About 1836,
the horticultural establishment of Fromont re-
ceived from England three varieties of this
chrysanthemum; one had the flowers red, one
variegated, and one white flesh-colored. Planted
in the open air, the following year we saw the
three varieties on one plant, which seems to
show that these three varieties were only sports
from a common form. A phenomenon analogous
to the preceding, and which, like it, concerns the
Chinese chrysanthemum, was shown at the Mu-
seum! in 1856 upon a variety called Surprise.
This, which bore flowers scarcely rose flesh-col-
ored, produced, on one of its branches, flowers of
a deep rose-lilac. Cuttings having been made,
it has preserved all its characters, and to-day it
is still one of the most beautiful of the section.
We call it Gain du Muséum. In 1862, upon this
same Gain du Muséum, a branch developed which
bore flowers perfectly white, of almost the same
size and form as those of the type; then upon dif-

color, or even to the normal form, if the variety differs also in

form. We must select the parts, in perpetuating variegations,

in which the variegation is very pronounced, although we must

exercise care that the variegation be not too intense, else the

offspring will be weak and poor. —Carrikre. (See page 149.)
1 Muséum d’ Histoire Naturelle de Paris.

CARRIERE ON BUD-VARIATION. 159

ferent branches beside it were found others bear-
ing flowers half red, half white. In making cut-
tings from these two kinds of branches, we would
then obtain from the Surprise still other varieties.
Let us look at the variety Sophie. This, which
has dirty white flowers very slightly tinged with
red, with yellow centre, has produced, by bud-
variation, a plant known as Trophée. The latter,
which has flowers of a rose-lilac-violet, bears some
resemblance to the Gain du Museum. ‘There
were also upon the same branch, but on different
twigs, flowers similar to those borne by the varie-
ties Trophée and Sophie. These new flowers
were flat and had narrow and imbricated petals,
whilst the Trophée has convex flowers, large and
slightly serrate petals. The Madame Richard
chrysanthemum, of which the flowers are whitish
very lightly bordered with rose, has produced on
one of its branches violet flowers stronger than
those of the plant from which it sprung; the
petals are also larger and more imbricated. In
1863 we observed on certain varieties of chrysan-
themum the following sports: the variety called
Cedo nulli, with double white flowers very lightly
rose, produced a branch which bore flowers much
larger and much more spreading than those of
Cedo nulli. The Argentine, with small white
flowers, pompon-form, gave a branch more vigor-
ous than itself, whose spreading, very large flow-

160 BORROWED OPINIONS.

ers, of a beautiful yellow, resembled to a certain
extent those of the large-flowered chrysanthemum,
a fact which tends to show that from the pompons
to the large-flowered sorts there is but a step. In
1864 we saw upon a stem of the Vesta (a pompon
chrysanthemum which has white flowers) several
branches which bore flowers entirely deep yellow.
The dimensions, as well as the form of the
flowers, were the same.

Varieties obtained by bud-variation are very
numerous. ‘There is not a genus among those
which comprise a number of species which has
not produced them. Although we shall mention,
farther on, a certain number of these bud-varie-
ties, adding some observations, there are some
which, in our opinion, are so interesting that, by
anticipation, we ought to speak of them here.
One of them relates to a kind of pink which is
known in commerce as Flon’s pink. This Flon’s
pink, which is closely related to those which we
call Spanish pink, Badin pink, etc., has flowers
very deep red, almost double, so that it does not
produce seeds, and we are obliged to multiply it
by cuttings. Nevertheless, it has already given,
by bud-variation, several varieties, of which the
most remarkable, a very beautiful white, was de-
veloped in 1858. Since that time this variety
has been maintained with all its characters. Ob-
tained by M. Pare, horticulturist at Paris, this

CARRIERE ON ROSE SPORTS. Toi

variety has been called Marie Paré, for one of
the children of the originator. Other varieties,
presenting colors different from that of which we
have spoken, have been developed from Flon’s
pink by M. Pare. [The pinks are fertile in bud-
varieties, particularly the carnation. Many of
the carnations which are now well known to com-
mercial growers first appeared as sports. ‘The
Portia, which is a deep self-red, frequently sports,
sometimes into almost pure white. |

The genus which, probably, has produced the
most examples of this nature is the Rose. The
examples are so very interesting that we cannot
resist the temptation to say something in detail
concerning them. We will cite several remark-
able examples, commencing with those which
have sprung from the Hundred-leaved Rose [ Pro-
vence rose, Rosa centifolia]. The bud-varieties
which have issued from this rose can be arranged
in two series: one which includes all individuals
which are but little removed from the type, which
differ from it only in color or form, either of the
flowers or sometimes of the leaves, and comprise
the ordinary Hundred-leaved roses; the other
series includes individuals possessing the charac-
ters of the first series, but which, in addition, are
provided with small bracts or glandular hairs
which give the name “ Moss-rose.” Bud-varieties
produced by Rosa centifolia : — :

M

162 BORROWED OPINIONS.

A. Ordinary Hundred-leaved roses.
I. Flowers more or less large.
Cabbage-leaved or lettuce-leaved R. cen-
tifolia.
Celery-leaved.
Anemone.
Nancy.
Peintres.
Flore magno, or Foliaceous.
Apetalous.
Unique white.
Unique variegated.
II. Flowers small.— Pompons.
Burgogne pompon.
White pompon.
Bordeaux pompon.
Kingston pompon.

B. Moss-roses.
I, Flowers more or less large.
Ordinary.
Cristata.
White-flowered
Variegated.
Sage-leaved.
Unique Provence.
Zoé, or Mousseuse partout.
II. Flowers small. — Pompons.

Pompon.

CARRIERE ON ROSE SPORTS. 163

One must not suppose that all the moss-roses
which he meets with to-day in commerce are the
result of bud-variation. ‘The larger part, on the
contrary, come from seeds. The moss-rose is
nearly arace. From seeds taken from the moss-
rose, we have obtained a certain number of
individuals which have preserved the general
characters of the plants from which they came ;
they are more or less “mossy.” Let us state,
however, that this “‘mossy ” character is not pecu-
liar to any section of roses, but that we find it
in most garden species, as the hybrid remontants,
rose-of-four-seasons, etc. The fact of the repro-
duction of the “mossiness” of roses by seeds,
proves again, what we have asserted several times,
that everything in a plant tends to reproduce
itself, that the peculiarities, properties, monstrosi-
ties even, may become hereditary.

The Zoé moss-rose is one of the most remark-
able bud-varieties which has been produced by
Rosa centifolia. This variety, instead of being
“mossy” only upon the peduncle or calyx, as
most of the other varieties of this group are,
is “mossy” on all its parts, whence the name
Mousseuse partout, [“* mossy everywhere” ]. This
variety was produced again in 1864, at M. Ja-
main’s, horticulturist, Paris, where we followed
the development of it. We also learned that at
this place, in two beds planted with ordinary

164 BORROWED OPINIONS.

moss-roses, beside the Zoé, there were several
stems which tended likewise to modify them-
selves, some in their leaves, others in their
flowers.

We must remark that it often happens that
certain individuals of bud-varieties return, on .
some one of their parts, to the type from which
they came. Thus, on a moss-rose from R. cen-
tifolia, we have seen a branch of the ordinary
Hundred-leaved rose. We should observe, how-
ever, that most usually the parts which seem to
return to the type present, notwithstanding, differ-
ences from it. There has been a step in advance,
and it is contrary to nature to retrace completely.

The Rose du Roi, known by nearly every one,
has produced the following six bud-varieties: —

1. Bernard Perpetual. This rose has the
branches more slender than those of the parent;
its flowers and leaves are also smaller. Its pom-
pon flowers are very pretty, with a rose-color very
much brighter than that of the Rose du Roi.

2. Long-peduncled Rose du Roi. This has
branches much longer than those of the type;
the internodes are more distant, and the peduncles
are also longer. It is only a sort of degeneration.

3. Madame Tellier. Very similar to the last,
being distinguished only by its flowers, which are
less colored, possessing a very bright rose flesh-
color.

CARRIERE ON ROSE SPORTS. 165

4. Mogador. This rose differs from Rose du
Roi by its stronger flowers, of a more vivid, deeper
red; its branches more colored, permitting it to
be distinguished even in winter. Horticulturists
do not like this variety, because it is hard to
‘force, and because it passes very quickly to a
dirty violet.

5. Capitaine Renard, or Variegated Rose du
Roi. This variety differs from Madame ‘Tellier
by its flowers being variegated or ribboned with
white. It was found at Orleans by M. Desfosseé-
Thuillier.

6. Celina Dubos. Found by M. Dubos, hor-—
ticulturist at Pierrefitte, near Saint-Denis, upon
Rose du Roi. It has the branches more slender
and the leaves a little smaller than the parent ;
its flowers, very similar in form to those of the
type, are white, slightly flesh-colored.

The Rose de la Reine has produced two sports:
one, Belle Normande, whose flowers, rose flesh-
colored, recall those of Souvenir de la Malmaison;
the other, Madame Cambel of Isly, or Triomphe
de Valenciennes, which differs from the parent
only in its marbled-variegated flowers.

The Duchesse de Cambacérés rose, which has
uni-colored, deep rose flowers, has produced by
bud-variation Belle de Printemps, which has rose
flowers marbled with brown.

The Baronne Prévost has produced, to our

166 BORROWED OPINIONS.

knowledge, five varieties, two of which have
variegated flowers and one marbled. One of the
two variegated varieties, Madame Désiree Giraud,
was found at the place of M. Desiré Giraud at
Marly, near Valenciennes. It is not vigorous.
The second variety, Panachée d’Orléans, which »
was observed for the first time at Orléans, is
very vigorous. Its branches are more slender
than those of Baronne Prevost, and the very
smooth and shiny bark has few prickles. In
short, its branches recall those of Cuisse de
Nymphe. It sometimes happens that this variety
produces large branches, vigorous and very thorny,
but less so than those of Baronne Prevost; its
flowers also resemble the type more closely. It is
an intermediate produced by the single matter of
vegetation. The Baronne Prévost marbre dif-
fers from the type only in its flowers, which are
marbled with brown. Another variety, placed
in the trade by M. Pierre Oger, horticulturist
at Caen, differs from the type only in the color
of the flowers, which are very much paler. The
fifth sport produced by the Baronne Prevost is
more recent. We observed it first in 1864, at
Vitry-sur-Seine, in a garden under the care of
M. Lachaume. We called it Madame Lachaume.
It differs from the type by its branches being a
little less thorny, but especially by its inflores-
cence, which, long-paniculate, very much branched,

CARRIERE ON ROSE SPORTS. 167

recalls that of certain Noisettes. The flower, also,
is a little weaker than that of the type. Buta
very remarkable fact is that the hip, instead of
being very regularly attenuated at its base and
becoming confounded with the peduncle, as in
Baronne Prévost, is abruptly and slightly in-
flated, then contracted, and inflated again near
the summit. ‘The peduncles are also much more
slender and longer than those of the parent.

The Duchesse d’Orléans, whose flowers are
violet-rose, produced by bud-variation, in 1858, a
variety known as Sceur des Anges. ‘This variety
differs from its parent particularly in the color of
the flowers, which is pale flesh-rose, like that of
the flowers of Souvenir de la Malmaison.

The rose called Quatre-Saisons has produced
the following sports : —

1. White Moss, or de Thionville. This was
first observed at Thionville about 1855. It dif-
fers from the type by its branches being more
slender and supplied with hispid, glandular hairs.
Its light green leaves are also softer to the touch
and slightly tomentose. Its flowers are pure
white. Sometimes it produces strong branches
which bear rose-colored flowers. In this latter
condition it is the ordinary Quatre-Saisons, a fact
observed by M. Duval of Montmorency, later by
M. Victor Verdier, Paris, and recently (1864) at
the Museum.

168 BORROWED OPINIONS.

2. Quatre-Saisons pompon.

3. White.

The Provence roses have likewise produced a
number of bud-varieties. Among the best known

are :—
Pompon Saint-Frangois.

Pompon Saint-Jacques.
Camaieu.

Panaché semi-double.
Tricolore de Flandre.

The last variety, which appeared in Belgium
some years ago, is remarkable for its variegated
flowers; it is a slender grower, although it comes
from a very vigorous variety. It sometimes re-
turns to the type. The variety Camaieu is remark-
able for its striped flowers, very pretty, and al-
most unique in the genus. Its wood is meagre
and its leaflets are toothed.

In the Damask roses, which are sorts of Quatre-
Saisons roses, not remontants, we consider as bud-
varieties the three following :—

Damask York and Lancaster.
Damask with blistered leaves.

The ordinary Bengal rose has sported into the
Bengale a bois strié [striped-stemmed Bengal].
The branches are often almost completely yellow.

A very curious sport of the rose is the plant
which we have called Rosier a feuilles de Chanvre

BUD-VARIATION IN THE ROSE. 169

[hemp-leaved rose]. By its flowers and especially
by its leaves, this variety differs considerably from
Rosa alba, from which it comes. Its leaflets are
hooded, long and narrow, and very coarsely den-
tate-serrate, sometimes as if gnawed on the edges,
strongly nerved, of a dark green, rugose-scabrous.
It happens sometimes, also, that its leaves are
opposite upon certain branches. The flowers of
this variety are smaller than those of Rosa alba,
often irregular, and somewhat monstrous, and
always sterile. [Probably no plants are so prolific
of bud-varieties as the roses. Every gardener of
“experience has observed the fact. The follow-
ing experiences of a single horticulturist (Ernest
Walker, New Albany, Indiana), with one rose,
illustrate this fact admirably. “I have had a
number of sports of the Perle des Jardins rose,”
he writes me, “in our greenhouses. The first one
was a double silvery pink with a short bud, and a
very double, somewhat quartered flower. The
stock of this I sold, as a new variety, for fifty
dollars. The next sport wasa white Perle. [The
Perle is a golden-yellow rose.] I sold a plant of
Perle to a local customer, who afterwards com-
plained that it was not true to name, because the
flower was white. She took it to be Cornelia
Cook. I went to see the rose, and found a Perle
rose in everything but color. I secured the plant,
and was intending to introduce it, when, within

170 BORROWED OPINIONS.

a few months, I heard that Nanz & Neuner, of
Louisville, Kentucky, had one, and that a London
firm had another; and later I found that one had
originated in Germany. Another sport of Perle
was a single rose, like Isabella Sprunt. Another
was like a Madame Falcot. At another time a
whole branch sported into a form with a long,
slender bud (about two inches long and _five-
eighths inch in diameter), with only two calyx
lobes, and only two petals,— which were very
broad, —in each cycle or series. This sport was
really a monstrosity, and I could not propagate
tis.

The so-called ornamental plants are not the
only ones which present these examples of hetero-
morphism. Fruit trees furnish very remarkable
examples. We will cite some cases, beginning
with those furnished by the cherry called Anglais
hatit [Early English]. The most curious sport
given us by this cherry is that which we call
Cerisier Anglais heterophylle or a feuilles de
saule [heterophyllous or willow-leaved English
cherry]. This is the history of the sport:
Upon a young tree whose parts are normal, we
see, sometimes suddenly and without apparent
cause, a vigorous bud develop, which bud, instead
of producing leaves of the ordinary form, bears
those which are very long and narrow, often
somewhat falcate, and often irregularly erose.

CARRIERE ON CHERRY SPORTS. Liz

Grafted, this variety presents very singular pecu-
liarities, as follows: so long as it preserves its
exceptional characters the plant does not flourish,
but as it constantly tends to lose them we observe
that when the leaves have almost returned to the
normal form the trees flourish and bear. Never-
theless, this variety never resumes identically the
characters of the type from which it came. Its
aspect is always distinct. The tree is never
fertile, and its fruit also differs from that of
the Early English. The young shoots preserve
their accidental character, and each year the leaves
which it develops are nearly identical to those
which the variety produced when it was first
developed.

This variety is not the only one which is pre-
sented by the Early English. Thus, when the
trees are old, it frequently happens that we find
on the same individual three kinds of fruits, dis-
tinct in their times of maturing. ‘There is, first,
the Early English, whose fruits become black ;
the Late English, whose fruits, of a beautiful
deep red, shining as if varnished, ripen later.
Finally, we nearly always find another variety,
very late, whose fruits, a little smaller, are still
entirely green when the other two have been
gathered a long time. In these three sports, the
differences are shown only on the fruits. The
Indule cherry is also only a sport from the Early

Li? BORROWED OPINIONS.

English. It is distinguished by its foliage and
earliness. The Early English cherry is not the
only one which furnishes bud-varieties pertaining
to the fruits. We find analogous examples in
the May Duke, Cherry Duke, and Reine Hor-
tense. These varieties, indeed, have produced
on different branches of the same individual sub-
varieties whose fruits ripened a fortnight later
than normally. Grafted, each of these sub-
varieties preserves its accidental character.

A phenomenon analogous to the preceding ones
is shown each year at the Museum upon an or-
dinary double-flowered cherry. The tree upon
which this anomaly was developed is nearly four-
teen inches in diameter, is grafted on the Sainte-
Lucie about twenty-seven inches above the ground.
Above the junction the stem is naked for about six
and a half feet. At this height is a large branch,
which every year is covered with extremely double
flowers, whilst the flowers of other branches, ex-
panding very much later, are scarcely half double,
and yield fruits.

The Coé violette plum is an example of dichro-
ism. It is a bud-variety which was produced on
the White-fruited Coé, and which, grafted, is
maintained with all the characters which it pre-
sented at the time of its appearing. We have
very often observed upon the Damas de Tours
plum an instance almost the same as the preced-

CARRIERE ON SPORTS OF FRUITS. 173

ing. On the same tree there were branches
which bore fruits different in form and color, and
differing a fortnight in time of maturing. Thus,
while the fruits of the type are very large, length-
ened, of a deep red color which recalls the Pond
Seedling, marked only on one side by a very
slight furrow, the fruits of the later sub-variety
are a little smaller, and their form is that of the
ordinary Reine Claude; they are of an _ herba-
ceous green, which passes more or less into a very
clear red ; the stem, arched, swollen at the base,
is inserted in a cavity quite large by the widening
of the furrow, whilst the stem of the typical fruits
is erect, little or not at all swollen, inserted in a
very small cavity placed almost on the surface of
the fruit.

Another plum, the Prunier Puget, presents the
following peculiarities: Upon the same branch it
very frequently happens that there are fruits of
a violet-red, dotted or striped with red-green.
We find some, also, which present all the inter-
mediate tints and others which are almost uni-
colored. By multiplying them separately, there
may be a chance to establish these varieties and
to obtain several from one tree.

We have seen on a red-fruited currant bush a
branch which bore fruits as white as those of the
Hollande a fruits blancs [White Dutch].

The fact of the nectarine coming suddenly from

174 BORROWED OPINIONS.

a peach can no longer be doubted. Recent ex-
amples have come to support the experiments of
certain authors, notably Sieulle.

Two other similar examples, of which we ought
to speak, are furnished by two varieties of Chasse-
las grapes, one known as Chasselas panache [ Va-
riegated Chasselas] and Chasselas Suisse [Swiss
Chasselas]. Both appear to have come from a va-
riety with black fruits, the color which predomi-
nates in them. ‘These are the peculiarities which
they present: almost all the bunches bear some
fruits more or less variegated or striped, white
in Chasselas Suisse, red in Chasselas panache.
But it happens frequently that the elements are
separated and that we have then, upon different
shoots, sometimes upon the same shoot, bunches
of grapes of different color, almost entirely white
if they belong to the Chasselas Suisse, and red if
they belong to the Chasselas panaché. One of
the varieties is only a modification of the other,
which is itself only a modification of some other.

The pear Saint-Germain gris, whose deep gray
fruits are very different in appearance from those
of the ordinary Saint-Germain, is a bud-variety
which was produced upon a branch of the latter,
and which, multiplied by grafting, is maintained
in all its characters. A similar variety was pro-
duced on the Messire-Jean, so that at present we
possess in the gardens a Messire-Jean gris, and

CARRIERE ON SPORTS OF FRUITS. 175

a Messire-Jean jaune [gray and yellow Messire-
Jean]. ‘To these examples we will add two other
analogous ones, which were recorded in the Bul-
letin de Académie des Sciences, xxxiv., meeting
of May 17th. One, given by M. Dureau de la
Malle, refers to the Bon Chrétien pear, which
produced sometimes typical fruits and at others
“of a form entirely different and unknown.” The
other example, cited by M. Mourriere, professor
at Bernay, has reference to an apple which, on the
same branches, produced fruits which had the
appearance of a Reinette rousse and others which
resembled a kind of Reinette du Canada. The
latter is smooth, punctated, and often of a bright
red upon one side. [The recent experiments
of Waite, in this country, respecting the imme-
diate influence of pollen, raise the question if
some of these minor variations in form of the
pear fruit may not have arisen from vagaries of
pollination. |

The various examples which we have cited are
common to a very large number of plants, among
which we will cite the banana and sugar-cane.
Indeed, although these plants do not produce
seeds, we find in each species a large number of
varieties which are very distinct in vigor, aspect,
habit, and in the banana in form, size, and quality
of fruit. All these varieties are produced by
bud-variation. These remarks can be applied to

176 BORROWED OPINIONS.

other monocotyledonous plants, as Arundo, Pha-
laris, Bamboos, Draczena, Yucea, ete.

[Carriere cites the different shapes and colors
of beans in the same pod as examples of bud-
variation, but it is a question if these differences
are not determined in the seed of the previous
year. At all events, since there is only a single
year in the life of the bean, we prefer to ascribe
variations in it to the generation of the parents
from which it has just sprung. There is no pre-
vious year’s growth of the same individual with
which to compare variations and to ascertain if
they are bud-departures from the type. Page 118. ]

2. List of Bud-varieties.

After having sought to present certain examples
of bud-variation which, by their importance, seem
to be sufficient to fix the attention, we will con-
tinue by the enumeration of a certain number of
others, without, however, entering into details for
each one of them. Sometimes we shall give only
the name of the variety. If, however, they pre-
sent particular interest, either in a practical or
scientific point of view, we shall dwell upon them
more at length, considering either their origins or
peculiarities. [The garden names of the plants

1 The student should also consult Darwin’s ‘‘ Animals and
Plants under Domestication.’’

CARRIERE’S LIST OF BUD-VARIETIES. 177

are given essentially as they stand in the original,
for, as the purpose of this list is to acquaint hor-
ticulturists with the nature and frequency of bud-
variations, I have considered it unnecessary to
make any particular attempt to revise the nomen-
clature. ‘The names are familiar, and, therefore,
useful as they stand. |

It would have been easy to extend this enumera-
tion of examples of bud-variation. We have not
thought it necessary because, aside from leading
us too far, the real interest of the subject would
gain nothing by it. We have, then, thought it
our duty to put limits upon a subject which has
no limits.

Acer erlocarpum, fasciatum.

Very remarkable for its much fasciated branches.
This variety showed itself at the Museum in 1857
upon a seedling which, during the first two years,
presented nothing abnormal. In the third year,
when the tree had been cut back, the sport
appeared, since which time it has maintained itself
with all its characters. This variety is to A.
eriocarpum what the variety montrosa is to Sam-
bucus nigra.

Acorus gramineus, variegata.!

1 When a name is not followed by remarks, the reader is to
understand that it represents a known variety and that the
name of the species indicates the origin of the variety. — L. H. B.

N

178 BORROWED OPINIONS.

Aisculus rubicunda, variegata.

[Ausculus Hippocastanum, double - flowered.
Upon a well-known Horse Chestnut tree in the
environs of Geneva the owner, in 1822 or 1823,
detected a single branch bearing double flowers.
This still continues to bear double flowers and
grafts from it do the same. It is thought to be
the original of all the doubled-flowered Horse
Chestnuts in the world. — A. De Candolle in Acad.
Sct., Paris, 1875, quoted by Asa Gray, Stlliman’s
Journ. 3d ser. x. 238].

Agatha amelloides, variegata.
Ageratum Mexicanum, nanum.

This plant, which is now used to so much ad-
vantage for borders, is the product of a branch
which developed accidentally from A. Mexicanum.
Its heads are almost sessile and a little irregular,
borne so close to the leaves as to make the plant
undesirable from some points of view. The type
plants, on the contrary, which are very much
larger, have the heads large and regular and raised
on long peduncles.

Ageratum Mexicanum, intermedium.

This variety, which is a bud-variety of sec-
ond degree, that is, a sport from a sport (from
the variety nanum), is intermediate. The plants
are very floriferous. ‘Their heads are also better

CARRIERE’S LIST OF BUD-VARIETIES. 179

than those of the type, and as they are borne
upon longer peduncles, the plants are not only
suitable for garden ornament but for cut flowers.
The dimensions of this variety are intermediate
between the last variety and the specific type.

Ageratum Mexicanum, variegatum.

This differs little from the type except by its
leaves being variegated with yellowish-white on
the margins. Its inflorescence is, however, a little
more slender and its heads are smaller. In gen-
eral, the plant is “leggy,” weak.

Almond with variegated leaves.

Leaves bordered and made satin-like with white ;
vegetation delicate. It sometimes returns to the

type.

Anemone Japonica, Honorine Jobert.

Very vigorous and very beautiful. This vari-
ety, of which the flower is white, is a bud-variety
from the so-called A. hybrida or A. elegans,
which was obtained in England by M. Gordon by
crossing A. Japonica with A. vitifolia. The va-
riety was produced some years ago at M. Jobert’s,
amateur at Verdun.

Apricot with variegated leaves.

180 BORROWED OPINIONS.

Aralia trifoliata, Cookii.

This plant has its leaves, in general, simple,
long, and narrow.

Arundo Donax, variegata argentea, and A. Donax,
variegata aurea.

These varieties differ from the type by the
leaves being bordered with white in the first,
and with yellow in the second. They are much
more delicate than the type.

Aspidistra elatior, variegata.

Aster bicolor.

This plant, whieh we believe not to be a dis-
tinct species but simply a dwarf form, very prob-
ably a bud-variety of A. versicolor, produced at
the Museum in 1856, upon one of its stems, a
vigorous bud which presents all the characters of
A. versicolor except that it is a little smaller.
This variety, to which we have given the vari-
etal name Major, has preserved all its characters
when multiplied by root-cuttings, and to-day is
still one of the most beautiful perennial plants.

Azalea Indica, Dieudonné Spae.

Flowers salmon, margined with white. It is
a sport from A. formosa, Ivery, which has rose
flowers.

CARRIERE’S LIST OF BUD-VARIETIES. 181

Azalea Indica, Beauté de Europe.

This variety has flowers white at the base,
variegated with red. It is a sport from A. deli-
cata, which has deep salmon flowers.

Azalea Indica, Criterion.

Flowers deep rose bordered with white. This
is a sport from A. Iveriana, which has flowers
white, striped with rose.

Azalea Indica, alba rosea.

Flowers rose, slightly bordered with white. <A
bud-variety from A. Iveriana.

Azalea Indica, exquisita grandiflora.

Flowers deep rose bordered with white. It 1s
a bud-variety from A. alba perfecta, which has
flowers white, very lightly striped with rose.

Buxus Balearica, cucullata.

This bud-variety of B. Balearica differs from
its parent by its smaller leaves, which are very
strongly convex, and rounded in the middle.

Buxus sempervirens, argentea.
Buxus sempervirens, aurea.
Buxus sempervirens, marginata.
All these varieties are distinguished from the

type by their leaves being variegated or bordered
with either white or yellow.

182 BORROWED OPINIONS.

Camellia Japonica, Comte de Paris.

This variety, which has strongly striped rose-
flesh-colored flowers, is a sport from the Duchesse
d@’Orleans, which bears white striped flowers. This
variety is not only much more vigorous than its
parent, but it has the merit of fully expanding its
flowers, while the buds of the Duchesse d’Orléans
almost always fall before opening.

Camellia Japonica, Montironi rosea.

This plant, whose flowers are entirely rose, 1s
a sport from the Montironi, which has white, very
lightly striped flowers.

Camellia Japonica, Giardino Franchetti.

Flowers deep rose, bordered with white. It is
an offshoot from C. Targioni, which has white
flowers lightly striped with rose.

Camellia Japonica, Comtesse W oronzoff.

This variety, which has delicate rose flowers, is
a sport from C. centifolia alba, whose flowers
are pure white.

Camellia Japonica, Giardino Schmitz.

Flowers delicate rose-color. It is a bud-variety
from the Elisa Centurion, which bears very lightly
rose-striped white flowers.

CARRIERE’S LIST OF BUD-VARIETIES. 183

Camellia Japonica, Impératrice Eugenie.

Flowers rose-flesh-colored. A bud-variety from
Montironi, whose flowers are very striped with
rose.

Camellia Japonica, Paolina Armari.

Flowers deep rose. Bud-variety from Miss
Abby Wilder, which has white, lightly rose-striped
flowers.

Camellia Japonica, Princesse Aldrovandi.

Flowers rose, bordered with white. Sport from
Teutonia, which bears flowers white, rose-striped.

Camellia Japonica, Bicolor de la Reine.

Flowers rose, bordered with white. It is a
sport from de la Reine, whose flowers are white,
lightly striped with rose.

Last year we saw upon a camellia with rose
flowers, some branches bearing flowers completely
white.

Cephalotaxus pedunculata, fastigiata.

This variety, which has been described and
figured as being a species of Podocarpus (P.
Koraiana), is an example of bud-variation. We
had proof of this statement at the Museum in
1863. Having made cuttings from a certain num-
ber of branches of the so-called Podocarpus, one

184 BORROWED OPINIONS.

of them, instead of producing simple and _ scat-
tered, strictly erect branches bearing scattered
leaves, produced whorled, horizontal branches
bearing distichous leaves. ‘The variety fastigiata
is to C. pedunculata what Taxus baccata, fastigi-
ata, is to T. baccata.

Cereus Peruvianus, monstrosus.

Sometimes returns to the type.

Clematis bicolor or Sieboldii.

This plant, of which the flowers, violet inside,
are almost double from the transformations of the
stamens, is a sport from C. florida, which has
single greenish-white flowers. We have several
times had occasion to ascertain that such is the
origin of this clematis.

The variety known as C. bicolor, flore pleno,
which we sometimes call Atragene Americana,
so remarkable for its enormous greenish-white
flowers, is a direct sport from C. bicolor, conse-
quently a bud-variety of the second degree from
C. florida, a fact which we have been able to
verify again this year. On a plant of C. bicolor,
planted in the open air, there is developed, almost
from the base, a branch which bore flowers en-
tirely full, monstrous, yellowish-green, so that the
two bud-varieties — C. bicolor and its variety flore
pleno — were united upon the same individual.

CARRIERE’S LIST OF BUD-VARIETIES. 185

Clematis, Helena monstrosa.

This plant is none other than C. Helena which,
by bud-variation, is transformed and has become
double flowered. This example is analogous to
that which is produced by C. bicolor.

Cheiranthus Cheiri, variegata flore pleno.

Sport from the double-flowered yellow gilly-
flower of the walls.

Cherry. See page 170.
Cornus sanguinea, variegata.
Cornus Mas, variegata.

Cytisus Adami.

Whatever may be the origin of this plant,
whether a hybrid, as is generally believed, or a
peculiar form, we propose here to say nothing
concerning it beyond a verification of its pecular-
ities. It develops very frequently and normally,
so to speak, some branches of C. Laburnum
and others belonging to C. purpureus. When
we graft separately these two kinds of branches,
these species remain invariable, although the
grafts were taken from C. Adami.

Dactylis glomerata, variegata.

186 BORROWED OPINIONS.

Echinocactus multiplex, cristata.

This variety, instead of having a _ regular,
lengthened, melon-like stem, forms a thick mass
which extends itself into little fan-shaped
bunches, and instead of longitudinal furrows,
large and deep, and separated by protuberances
upon which are borne long, very rigid spines
(about three-fourths to one and_ two-fifths
inches); the variety has only very slight fur-
rows or kind of folds disposed transversely to
the direction of the fasciation, consequently in
a contrary direction to those which are presented
by the type, and upon the borders are spurs
(about two-fifths inch long) disposed in stars.
In a word, the variety is entirely different from
its parent.

Eleagnus reflexa, variegata argentea and varie-
gata aurea.

These two varieties differ from the type in
having their leaves bordered, respectively, with
white or yellow.

Eleagnus pungens, variegata.

Euonymus Japonica, argentea and aurea.

These, especially aurea, return sometimes to the
type.

CARRIERE’S LIST OF BUD-VARIETIES. 187

Euonymus Japonica, flavida.

This plant, which developed upon a type plant
with green leaves in 1862, is distinguished by its
leaves being bordered with yellowish-green, some-
times with nearly white. It is vigorous.

Euonymus Japonica, fasciata.

Very remarkable for its much fasciated branches.
This variety appeared at the Museum in 1864, upon
a typical E. Japonica.

Euonymus Japonica, calamistrata.

This sprung from the variety argentea, from
which it differs in its more slender parts. Its
leaves are smaller and crisped as if erose.

EK. Japonica has also produced many other bud-
varieties, which differ in variegation, or sometimes
even by the form of the leaves. It is more than
probable that the various varieties which have been
introduced recently from Japan are bud-varieties.

Fagus sylvatica, fern-leaved.

This variety once presented the following
peculiarity: Having grafted it upon the com-
mon beech, the branches developed from each
side of the stem almost distichously. All those
upon one side bore leaves similar to those of the
common beech, whilst those upon the other side
bore only laciniated leaves.

188 BORROWED OPINIONS.

Ficus scandens, microphylla.

This variety, which we sometimes meet in com-
merce under the name of F. buxifolia, is a bud-
variety which appeared in 1856 at the Botanic
Garden of Orléans upon a plant of F. scandens
grown in a greenhouse. Its leaves are very small,
somewhat suborbicular and marked with brown.
This variety is preserved in all its characters,
both upon the original plant and in all the multi-
plications which have been made of it.

Fontanesia phyllireoides, variegata.

This very pretty variety appeared at the
Museum in 1854. Since its appearing, this vari-
ety has not varied. Its branches, of a yellowish-
green, are slender, and the leaves are deeply
bordered with yellowish-white.

Fraxinus Americana, variegata.

Fraxinus excelsior, jaspidea.
This variety is distinguished by its bark being
striped or slightly ribboned with yellow.

Fraxinus excelsior, variegata.

The common ash has produced several sports
which are marked by the variegation of their
leaves. This one has yellow and white disposed
in bands and bordering the leaves, or sometimes

CARRIERE’S LIST OF BUD-VARIETIES. 189

in spots upon all parts of the blade, as upon the
leaves of Aucuba, for example ; hence the various
names, argentea, aurea, striata, maculata, aucube-
folia, ete.

Gardenia radicans, variegata.

In this instance the variation is two-fold. The
leaves are bordered, and are also much narrower
than in the type.

Gillyflower, called Savoyarde, with variegated
leaves.

This is a sport from the double-flowered brown
gillyflower.

Grape. See Vine.

Hedera. Variegated tree-ivy.

This sub-variety is a bud-variety from the so-
called tree-ivy (lierre en arbre,) from which it
varies only in the yellowish-white variegations of
the leaves.

What we call tree-ivy is a common ivy, or one
of its varieties, arrived at the full-grown state and
which then fruits. The branches are large, short,
cylindrical, and destitute of climbing roots. The
leaves, instead of being lobed, are heart-shaped,
more or less lengthened, sometimes very obtusely
rounded. As there are several forms of creeping
ivy, so there are several sub-varieties of the tree-

190 BORROWED OPINIONS.

ivy. They partake of the character of the varieties
from which they come and are distinguished by the
form and dimensions of the leaves, by the size of
the branches, these characters all depending upon
the vigor and appearance of the mother varieties.
We obtain the tree-ivy either by cuttings or by
grafting from adult branches, that is, branches
which have been modified by fructification. They
then branch and form very pretty bushes. Some-
times, especially near the ground or in badly aired
places, branches arise supplied with climbing roots,
bearing leaves more or less lobed, and which creep
and take root as soon as they touch the ground.
Here, in the case of the tree-ivy, is an instance of
bud-variation due to the maturity of the individual.

Hibiscus Syriacus, flore pleno variegata.

This variety, whose leaves are variegated with
yellowish-white, appeared in 1858 upon a plant
with entirely green leaves.

Hibiscus Syriacus, variegata.

Remarkable in the variegation of its leaves. Its
flowers are similar to those of the last. It is a va-
riation directly from the type. It is not vigorous.

Hyacinth.
The double blue or Globe terrestre is a bud-
variety from the double white or Sultan Achmet.

CARRIERE’S LIST OF BUD-VARIETIES. 191

The double white with blue eye, or Spheera Mundi,
is a sport from the double white. ‘The single red,
called Acteur, cultivated for a very long time
without varying, has produced by bud-variation
at Hemstede, near Haarlem, a variety with double,
imbricated red flowers. The hyacinth Ami du
Coeur, with single blue flowers, also long culti-
vated without varying, has produced, trom the
same bulb, two flower stalks, one of which bore
flowers dregs-of-wine color, while the other bore
flowers of a delicate flesh-colored rose.

Hydrangea Hortensia.

This sterile plant is a sport from the form called
H. Japonica, analogous to those which are pro-
duced upon Viburnum Keteleerii and V. Opulus.

Hydrangea Japonica, variegata.

Differs from the type only by its leaves being
bordered with white.

Nex Aquifolium, calamistrata variegata.

This variety is a sport from the I. calamis-
trata, which is a sport from the common holly.

ex Aquifolium, ferox aurea, and ferox argentea.

Bud-varieties from the variety ferox, from
which they differ in the variegation of the leaves,
yellow in the first, white in the second.

192 BORROWED OPINIONS.

The very numerous varieties of common holly
in cultivation are for the most part fixed bud-
varieties.

Iris spectabilis.

This plant, so remarkable for its color, is a bud-
variety from Iris Xiphium, from which, however,
it is very different.

Juniperus communis, variegata.
Juniperus excelsa, variegata.
Juniperus Virginiana, variegata.

Juniperus Virginiana, monstrosa.

This variety, which arises from knaurs or burrs,
is shown frequently upon the Virginian juniper
(ved cedar).

Lamium album, variegatum.

Laurocerasus vulgaris, angustifolia.

This plant, which for a long time has gone under
the name of Hartogia Capensis, is a bud-variety, a
fact which we have been able to ascertain several
times. Its leaves are very straight, long, of a
clear green, and more strongly toothed than those
of the plant from which it comes. It is very con-
stant. We have a record of its variation.

CARRIERE’S LIST OF BUD-VARIETIES. 193

Laurocerasus vulgaris, variegata.
Laurocerasus Lusitanica, variegata.

Ligustrum Japonicum, variegatum.

The L. Japonicum appears to be subject to bud-
variations, especially in the direction of dichroism.
We have already produced from it several varie-
ties distinct in the color or disposition of the
variegations, the varieties receiving names accord-
ing to their character. There is one which differs
somewhat in the form of its leaves.

Ligustrum ovalifolium, aureum.

This variety, which is distinguished by having
its leaves bordered or ribboned with yellow, was
produced at the Museum in 1861. It comes from
a branch which was developed spontaneously upon
a type plant. It is unstable.

Ligustrum vulgare, variegatum.

This variety has leaves variegated with yellow.
It occurs quite frequently in the wild state. We
have found it several times in the woods. It is
not stable.

Lilac, common, variegated.

Lilac, Persian, laciniate-leaved and white-flowered.

Although we are not able to state precisely when
these two sports appeared, we have no doubt that
oO

194 BORROWED OPINIONS.

they are bud-varieties, as the Persian lilac never
gives seeds. The origin of the Persian lilac itself,
even, 1s in much doubt.

Mamillaria nivea, dedalea.

This variety forms a compact mass whose folds
and circumvolutions are disposed in a sort of laby-
rinth (whence dedalea), giving it a little the ap-
pearance of calves’ pluck. ‘The type from which
this variety comes forms a melon-formed cylinder
which is slightly swollen at the summit. It bears
spines disposed in bundles and from about four-
fifths inch to one inch and a fifth in length, rigid,
very sharp, surrounded at the base by a series of
smaller ones disposed in the form of a star. The
variety, on the other hand, aside from its peculiar
form, has no spines. It is invested upon all its
parts with silky hairs, silvery and as soft as felt
to the touch. The parent and offspring have
nothing in common in their general form.

Mentha rotundifolia, variegata.
Molinia ccerulea. variegata.

Musa paradisiaca, vittata.

This is distinguished from the type by the white
bands upon its leaves. The variegation is con-
spicuous upon the yellow plants. It often disap-
pears with time, so that in the old plants we do
not often find any trace of it.

CARRIERE’S LIST OF BUD-VARIETIES. 195

Myrtle, common, variegated-leaved.
Frequently returns to the type.

Opuntia cylindrica, cristata.

In exterior characters this plant has nothing in
common with the type, which forms a regular,
cylindrical column. ‘The variety, on the contrary,
is made up of enlarged pieces placed against each
other in different ways, much the same as those
presented by the various species of opuntia which
we call “Semelles du Pape.”

Orange tree, ‘Turkish.

This variety, which is a sport from a kind of
Seville orange (probably from the Horned Seville
orange), bears at times upon its various branches
leaves narrow and irregular (as erose), variegated
or rather satin-bordered, white, and, upon other
branches, green leaves, large and strongly eared,
as well as fruits which recall those of the Horned
Seville orange.

Orontium Japonicum, variegatum.

Osmanthus Fortunei,! ovata.

This variety is unstable. After having pre-
served it for more than a year without varying, it
has resumed in large part its primitive character,
which is to have leaves long, exceedingly thorny

1 The Olea ilicifolia of commerce. — CARRIERE.

196 BORROWED OPINIONS.

and strongly nerved. Sometimes we find branches
bearing leaves of different forms.

Osmanthus Aquifolius, variegatus.

Ditters from the type by the yellowish-white
variegations of the leaves.

O. Aquifolius, which we can consider as the
representative in Japan of our common holly,
appears, like the holly, to be very subject to bud-
variation. We have no doubt that the varieties
recently introduced from Japan originated in this
manner.

Peach, carnation-flowered (Persica dianthiflora),
and many-colored (P. versicolor).

These two varieties are sportive forms of P.
roseflora, of which the flowers are very deep
red. Like this, the two varieties have double
flowers, but of very different colors from those of
their parent. The carnation-flowered has flowers
of a flesh-colored rose. The many-colored, on
the contrary, has white flowers striped or rib-
boned with brilliant rose. This last is very much
more delicate than P. roszflora or the carnation-
flowered.

Peach, willow-leaved red Madeleine.

This variety, remarkable for the form of its
leaves, which are very long and narrow, plane,

CARRIERE’S LIST OF BUD-VARIETIES. 197

glossy, very shortly toothed, is the result of a
bud-variation from the variety designated by cer-
tain horticulturists as Madeleine de Courson (red
Madeleine). It appears to us to have great re-
semblance to that very anciently known as the
willow-leaved. |

Peach, laciniate-leaved red Madeleine.

Leaves very strongly and coarsely toothed or
laciniated.

Pears, variegated fruited.

The following pears have given by bud-varia-
tion variegated varieties : Duchesse d’Angouléme,
Amanlis, Guenette or Madeleine, Saint-Germain,
Bergamotte d’automne, Culotte de Suisse, ete.
These bud-varieties are further remarkable in that
the variegations extend to the branches and fruit,
but not to the leaves, a character which distin-
guishes them from the next variety, which is like-
wise a bud-variety.

Pear, Amanlis, with yellow bark and leaves.

A very remarkable variety. We could almost
say that it is pretty. It was developed upon a stem
of Amanlis which presented nothing abnormal.
It is very vigorous, and produces a most singular
effect, with its parts all yellow except the bark,
which is grayish-white. It has not yet fruited.

198 BORROWED OPINIONS.

[One of the most marked cases of bud-variation
which ever came under my notice was observed
a few years ago upon a tree of Onondaga pear.
One branch, so placed as to remove all possibility
of its being a root-sprout or a graft, bore about a
dozen pears which were intensely and uniformly
russeted. They were so different in appearance
from the pears upon the remainder of the tree that
no one would suppose for a moment that they were
the same variety. Even the Sheldon does not
differ more widely from the Onondaga in appear-
ance than did this singular sport. |

See page 174 for further notes on the pear.

Pelargonium zonale, Manglesii.

Distinguished from the species by its white
variegated leaves, which are more deeply lobed,
and by the weaker branches. It has, in its turn,
produced several varieties by bud-variation.

The bud-varieties produced by P. zonale and P.
inquinans (which are in reality only one) are very
numerous. There are among them some varieties
of such pronounced characters that, if we ignored
their origin, we might consider them species.

Pelargonium hederefolium, variegatum.

Phalaris arundinacea, picta and aurea.

These two varieties differ from the species in
the variegation, which is produced by white in

CARRIERE’S LIST OF BUD-VARIETIES. 199

the first case and yellow in the second. They
are exactly representative of the phenomenon
which is observed in Arundo Donax, as well as
in the sugar-cane.

Phlox decussata, white Croix de Saint-Louis.

This variety, of which the flowers are entirely
white, appeared upon the variety Croix de Saint-
Louis in 1863. The parent variety has white-
striped rose flowers, and cross-form, whence its
name.

Phragmites vulgaris, variegata.
Leaves bordered or margined with white.

Picea excelsa, tabuleeformis.

This variety, which attains a height of scarcely
more than a foot or two, and which spreads out
horizontally so as to form a sort of carpet, is a
most remarkable variation which resulted from
a knaur or burr upon the stem of a very large
spruce. It was produced in the park of Trianon
at Versailles.

Pinus sylvestris, nana monstrosa.

Produced from a knaur from the stem of a
large pine. It is dwarf and monstrous, remarka-
ble for its long, unequal, crowded leaves, and
by its slender, sometimes almost filiform, irreg-
ular branches, which are produced in such quan-

200 BORROWED OPINIONS.

tity that they sometimes completely conceal the
branches and even the trunk.

Pinus sylvestris, nana compacta.

This variety is also the result of a knaur from
a large pine. It attains but a foot or two in
height. Its very short and numerous branches
have already borne two crops of cones, some
nearly ripe, small, though well formed, others
much younger, still herbaceous.

Pittosporum Tobira, variegatum.

Populus Greeca,! pendula.

We cannot say whence this variety came nor
how it was obtained. It has been in cultivation
very long. We have a singular experience to
record concerning it. In 1858 we grafted fifteen
plants of P. nivea with P. Greca, and, seven of
the grafts growing, there was one which produced
slender and drooping branches just like the variety
pendula of P. Greca, of commerce. This phe-
nomenon is one of the most curious which we
know. The tree is planted in the nursery of the
Museum at the side of one of its brothers, to
which, physically, it bears almost no resemblance,
although coming from the same parent plant.

1 Undoubtedly our native large-toothed aspen, Populus
grandidentata.— L. H. B.

CARRIERE’S LIST OF BUD-VARIETIES. 201

Both are pistillate and are covered each year
with catkins.

Potato.

Potatoes furnish many examples of bud-va-
riation [mixing in the hill”]. Many of our
cultivated varieties are bud-varieties from the
subterranean parts. Every year at digging time,
if we wish to keep the varieties true, we are
obliged to throw out those which, we say, are
“degenerated.” This so-called degeneracy con-
stantly tends to remove the products from the
starting-point, and has, then, the result of pro-
ducing new varieties.

Modifications in potatoes may also occur in
the manner of vegetation or growth of the under-
ground parts. Such is the case in the variety
called Pouse-debout [* tubers standing on end”’].
This name was given the variety because the
tubers, instead of lying horizontally, or nearly so,
are placed upright, one against the other, much as
small pieces of wood are arranged for the making
of charcoal.

The Marjolin we consider nothing else than a
peculiarity of vegetation. This is proved by the
fact that its characters — not blossoming and
maturing very early — are not constant. It has
produced two other varieties by modifications of
its underground parts. One variety is the Mar-

202 BORROWED OPINIONS.

jolin tardive [Late Marjolin], called also Marjolin
de deuxieme Saison, which is sometimes sold in
the Paris markets for the Hollande jaune [Yellow
Holland]. It is remarkable for the period of its
growth, which is more prolonged than that of the
type, and it is also covered each year with flowers,
while its parent scarcely ever blossoms. The
other variety has no resemblance to the Marjolin
in form. It is round, and its sunken eyes give
it exactly the appearance of the ordinary yellow
potato. When we cultivated the Marjolin there
was not a year when we did not obtain round
ones, although we had planted long ones very true
in appearance.

A very remarkable example of the modifications
furnished by the ordinary yellow potato is the
following: In a square planted exclusively with
this variety, very true in appearance, we gathered
a certain number of which the skin was more or
less dark; some had yellow flesh, others white.
Planted separately, these bud-varieties have given
us potatoes round in form like the parent type,
but among which there were found some entirely
violet in both exterior and interior, and some had
black flesh shghtly marbled with white. This
modification of color was not the only change.
In some cases the quality was very much modified.
Thus, instead of being nearly like the yellow
potato, the flesh of these varieties was compact,
neither good nor bad.

CARRIERE’S LIST OF BUD-VARIETIES. 208

We give two other examples of bud-variation
in potatoes, observed by us at the Museum in
1864: Half of a plat was planted with the smooth
long yellow called Hollande, and half with the
regular long red commonly called Vitelotte lisse.
The first half yielded tubers similar to those
which we had planted. The second half, on the
contrary, produced tubers differing from the pa-
rent in color, being of a reddish-yellow, although
the form remained about the same. The quality,
also, did not vary, so that while we confounded
them sometimes with the Hollande, we were able
to distinguish. them readily when cooked, as
they remained whole, while the others fell to
pieces.

On the end of a plat where we had planted fifty
of the ordinary round yellow potato, one plant
grew until late in the season and gave round
potatoes of a deep red.

In this same year, 1864, in a square planted
entirely to Chardon potato, we observed some
plants exactly similar to the others in growth and
appearance, but which differed entirely in the
color of the flowers, being dull white, a little
sulphur-colored, while those of Chardon are violet-
rose or rose-violet. The tubers from these white-
flowered plants differed from those of the type in
being more round and regular and having less
pronounced eyes. Aside from these variations,

204 BORROWED OPINIONS.

we have found among the Chardon both earlier
and later varieties, and this in spite of the fact
that we had planted only such tubers as appeared
to be entirely true and which for a long time had
produced no variations whatever. Here, as in
the preceding cases, the modifications were from
the tubers, seed not having been sown.

An instance similar to the above is reported by
M. Joigneaux in the Journal de la Ferme et des
Maisons de Campagne: Nine or ten years ago six
beautiful tubers of a long, pale yellow potato were
given us. In order to increase the number of
hills we divided each tuber into three pieces. We
planted them ourselves. ‘The cultivating was also
done by us. Some of the potatoes, a very small
number, resembled the type, but the larger num-
ber were spherical, some yellow like the parent,
others deep red...

All cultivators know that the smooth or even
Vitelottes, whose eyes are few in number and
scarcely perceptible, often produce tubers of vari-
ous forms and with eyes so much sunken that it
is almost impossible to peel them. Once we ob-
tained a variety which, besides the many and deep
eyes, produced, in considerable quantity, agglom-
erations which gave to the whole a monstrous
form. They were veritable hydras. Although
coming from the Vitelotte, which is a good potato,
this variety was very acrid and bad.

CARRIERE’S LIST OF BUD-VARIETIES. 205

All these examples show without doubt how
a part of the varieties of potatoes are pro-
duced, and proves that they do not all come
from seeds. We may convince ourselves of it
when, having observed the growth of the plants,
we mark the peculiar plants and gather their
tubers separately. See, also, page 209.

The phenomena presented by potatoes prove
that the cause of the appearing of new varieties
is not always due, as we generally suppose, to
crossing, as fecundation can act only upon the
seeds. It is also very rarely that we practise
crossing in potatoes, but we can number the varie-
ties by the hundred. But it often happens that
cuttings made from portions of the top of the plant
produce varieties different from the parent. More-
over, the existence of numerous varieties of certain
plants which we cultivate and which never pro-
duce seeds, proves beyond a doubt that there are
causes aside from crossing which tend to the pro-
duction of new varieties.

Prunus Mahaleb, variegata.

Aside from this variety, which is very pretty
with its long and very slender branches and
white-variegated leaves, P. Mahaleb has by bud-
variation given several sub-varieties which are
distinguished by the form of the leaves, and
especially by the color of their variegations.

206 BORROWED OPINIONS.

Almost all these varieties are more delicate than
the type.

Rheum australe, variegatum.

Remarkable for the beautiful white variegation
of the leaves.

Ribes nigrum, variegatum.
Leaves variegated with yellowish-white.

Ribes rubrum, variegatum.

Robinia hispida, arborea and macrophylla.

R. hispida, var. arborea of the gardens (R.
macrophylla DC.), differs from the type by its
greater vigor, its branches very much larger,
the bark dark, glossy, and smooth, and by the
thicker coriaceous leaves, which are glossy as if
varnished. R. hispida, var. macrophylla of the
gardens, is nearer the type than the last; it differs
from it, however, by its greater vigor, and espe-
cially in its flowers, which, less abundant and a
little more developed, are paler in color. Like
the type, of which even the origin is doubtful,
these varieties do not produce seeds.

The fact of the sportive production of the
variety inermis upon R. hispida is wholly be-
yond doubt. Several times we have found the
two sorts of twigs growing side by side upon the
same branch. It is only necessary to multiply

CARRIERE’S LIST OF BUD-VARIETIES. 207

them separately in order to obtain distinct varie-
ties.1

Robinia Pseudacacia, umbraculifera.

This plant, now so commonly used either for
ornament, under the common name of Acacia
boule, or as a dwarf shrub and considered as a
forage plant, and called in consequence Acacia A
faucher, comes, according to Turpin, from a knaur
which appeared on the stem of a R. Pseudacacia.
This fact does not surprise us. It shows us how
important these singular exerescences may become.

Rose Eglanteria, punicea.

This differs from the ordinary yellow-flowered
capucine rose (R. Eglanteria) in the color of the
flowers, which is an orange-red. In many soils
this variety returns more or less quickly to the
type. It frequently happens that we may see
even upon the same stem a red and a yellow
flower. Sometimes we find a flower which pre-
sents these colors separately on opposite sides and
of nearly equal extent, or some petals may be half
red and half yellow. In general, the sport is less
vigorous than the type, so that under the influ-

1 Plants of normally thorny species are often found which
bear no thorns. There is a so-called variety inermis of the
honey-locust, Gleditschia triacanthos. Wild blackberries with
smooth canes are occasionally found. In fact, most prickly or
thorn-bearing plants vary much in these characters. —L. H. B.

208 BORROWED OPINIONS.

ence of a slow modification we sometimes see the
color gradually disappear, and at the end of a
certain time we have a rose with completely yel-
low flowers in the place where we planted one
with orange-red flowers.

For further notes on the rose, see pages 161 to
170.

Salix Babylonica, annularis.

Very remarkable for the form of its leaves.
For a long time we have noticed it showing itself
each year upon an old tree. The parts upon
which it appeared, being generally weak, pro-
duced, instead of long, linear, plane leaves, those
which were rolled up on the edges and distorted
into rings. It is very constant. We have no
examples of its reverting. It is much less vigor-
ous than the type.

.

Sambucus nigra, variegata aurea and variegata
argentea.

These two varieties differ from the species in
the variegation, which is made by yellow in the
first, white in the second. ‘The second is much
less vigorous than the type.

Sambucus nigra, monstrosa.

Analogous to Euonymus Japonica, fasciata. The
flowers are also monstrous, and, up to this time,
the seeds which they have produced have always

been poor.

CARRIERE’S LIST OF BUD-VARIETIES. 209

Solanum Dulcamara, variegatum.

Solanum tuberosum, variegatum.

Remarkable for its yellow-variegated stems and
leaves. During the year previous to the appear-
ing of this variety, its parent type presented no
unusual characters. See Potato, page 201.

Spireea Ulmaria, variegata.
Symphoricarpus vulgaris, variegata.
Symphytum officinale, variegatum.

Thuyopsis dolibrata, variegata.

This variety, of which the leaves are variegated
with white, is remarkable for its vigor and its
facility for forming heads when multiplied by
cuttings.

Ulmus campestris, variegata, argentea, aurea picta,
etc.

Variegated varieties of the common elm are
numerous. They are distinguished both by the
color and form of the variegations.

Viburnum Opulus, sterilis or Boule de neige.
Viburnum Opulus, sterilis variegatum.

Viburnum Keteleerii, macrocephalum.

Analogous to the sterile varieties of V. Opulus.
:

210 BORROWED OPINIONS.
Viburnum Tinus, variegatum.

Viola Rothomagensis, pallida.

This variety, of which the two superior petals
are pale lilac and spotted, while the three others
are yellowish-white and lightly striped, was pro-
duced by bud-variation at the Museum.

We have said elsewhere that the phenomena of
bud-variation could be divided into two classes,
one group including those variations which appear
abruptly, the other those which take place slowly.
This violet falls under the latter group. In
1863 we received from the hills of Vernon a
certain number of plants of V. Rothomagensis.
Planted at the Museum, they preserved nearly all
their characters except the villosity, which in large
part disappeared the first year. During this year
1863 and the entire year 1864, they produced blue
flowers abundantly. In the winter of 1864-665 all
the plants perished except one. The remaining
plant, instead of being covered with beautiful
blue flowers as it had been the two preceding
years, produced flowers almost white.!

Vine (Grape).
Bud-variation is comparatively common in the

vine. [Frequent cases occur in the American

1 This variation appears, after all, to have been an abrupt
one. —I.. H. B.

CARRIERE ON GRAPE SPORTS. 911

grapes.] It is well understood in this case, as
the vine is one of the oldest of cultivated plants
and it is multiplied almost always by cuttings;
and as cuttings are made by millions each year
a bud-variety soon becomes widely disseminated.
It frequently happens that a shoot will produce
grapes differing in form or color from those which
are borne upon other shoots of the same vine. We
may add that these variations nearly always pre-
sent peculiar qualities also. We will cite ex-
amples.

Upon a plant of black-fruited Muscat grape we
have observed for several years that some shoots
produce white grapes.

The white seedless Corinth is a bud-variety
from a variety which has much larger fruits, with
seeds. ‘This is a fact which we have several times
observed upon bunches where some fruits were
unusually developed and which contained seeds.
The white Cornith is analogous to the Chasselas
de Demoiselles.

A proprietor of large vineyards in the middle
of France, the late Cazalis Allut, wrote some years
ago as follows : —

«A stock of Teret produced with me, for sev-
eral years, black grapes upon shoots of two of
its arms, and gray grapes upon shoots of the other
arms. A stock of Epiran gris, trained in cordon,
is now about forty feet long. The first twenty

O12 BORROWED OPINIONS.

feet produces constantly gray grapes and the re-
mainder produces white ones. I have in an
enclosure a stock of Epiran noir having several
arms. ‘The shoots of one of the arms give grapes
almost twice as large as those on other parts of
the vine.”

Another viticulturist, M. Henri Bouschet of
Montpellier, wrote very recently: —

‘“‘T had occasion for several years to see in my
collection at Lot-et-Garonne, a plant of Prunella
eris, which, sometimes upon one stem, sometimes
upon two, bore black grapes, while the remainder
of the vine bore gray ones. I have noticed for
two years in my collection at Calmelte a most
curious fact upon three grafts of a Spanish va-
riety which came to me from the collection of
Luxembourg, where it is called Parrel del Reyno
de Lorea and which I have recognized as our
Morastel noir. One of these three stocks has
borne on one side, to my great surprise, black
grapes similar to those of the Morastel, and upon
the other side, constantly, white bunches having
an appearance very different from an ordinary
white Morastel, and presenting a folhage very
different in size and form. ‘This odd foliage ap-
pears to me to be identical with that of the Oyo
de Rey de Morada, of which the bright yellow-
ish-eteen leaves present very shallow rounded
dentate lobes, while the leaves of the Morastel are

CARRIERE ON GRAPE SPORTS. 213

deep green with deep divisions, the lobes acute,
with teeth detached and terminating in a point.”

A passage which we find in the Parfait
Vigneron (edition of 1811) seems to confirm the
opinions which we give concerning bud-variations
in grapes: —

« A citizen of Vilmorin has observed a stock of
Meunier to bear, upon some shoots, leaves and
fruits of Maurillon précoce. A citizen of Ju-
milhac has seen likewise the Meunier become
Maurillon.”

Therefore the grape called Madeleine Juillet,
Maurillon hatif, etc., is only a sport from Meunier,
a fact which shows, as we have said before, that
the varieties produced by bud-variation may pre-
sent qualities different from those presented by
the plants from which they come.

Upon a plant of Pinot gris there appeared at
the Museum in 1863 a shoot whose leaves were
much variegated or striped with yellow. It
produced a grape very similar to the variety from
which it came. It appeared to be much less fer-
tile, however.

In 1863 we observed two other very remarkable
examples. One example concerns the Précoce
Malingre, the other the variety designated by
the name of Verjus. ‘These examples present
contrary results. Thus, while Précoce Malingre
has long, oval, scattered fruits, and the bud-variety

214 BORROWED OPINIONS.

which appeared upon it had round fruits borne
close together and larger than those of the type,
the Verjus has shghtly oblong or nearly spherical
fruits and the bud-variety which was developed
upon it had fruits long-oval and attenuated at
both ends, and somewhat later than those of the
type.

The Chasselas gros Coulard is a bud-variety
which appears frequently upon the ordinary
Chasselas. Its fruits are large and spherical.
They often drop. It differs especially from the
Chasselas by its stronger shoots with joints
much closer together, and by its leaves being
less lobed, a little longer and thicker, of a
olossy green as if varnished. It differs also
from the ordinary Chasselas in its temperament.
It needs much heat and also shelter from the
influence of the air. It generally succeeds well
in forced culture.

The Chasselas de Demoiselles, remarkable for
its fruits, which are scarcely larger than shot, is a
bud-variety from the ordinary Chasselas. This
phenomenon appears to be due to the partial abor-
tion of the sexual organs and particularly of the
anthers, whence results the lack of impregnation
of the flowers and the consequent abortion of seeds.
Propagated by cuttings, it preserves its characters.
A variety with variegated leaves has appeared
from the ordinary Chasselas.

FOCKE ON THE PRIMARY CROSS. O15

Wigandia Caracassana, variegata.

Distinct by its leaves and even its branches
being variegated with white. The variety ap-
peared in 1862 upon a plant which, placed in the
open air at the beginning of the season, presented
no unusual characters.

IlJ.. Focksr’s Discussion oF THE CHARACTER-
ISTICS OF CRossES. (Translation of Chapter
IV. of Die Pflanzen-Mischlinge.*)

There is no absolute distinction between plants
of a pure and those of a hybrid or mixed descent;
there are therefore no signs by which one can,
under all circumstances and with certainty, rec-
ognize the mongrel nature of a certain plant.
Hybrids, nevertheless, often show a _ series of
characteristics which indicate with greater or
less accuracy their mixed descent. Certain rules
can be made in regard to them, of which none,
of course, is without exception.

I. THE SIMPLE PRIMARY CROSS (A x B).2
1. All individuals formed by the crossing of

two pure species or races are, if they have been

1 Focke uses the word mischling (derived from mischen, ‘to
mix’’) in about the same sense in which we use the noun cross ;
i.e. it is a generic term for both cross-breeds and hybrids.

2In these formulas, the letters are used to designate the
parents of any cross. In common with usage amongst botanists,

216 BORROWED OPINIONS.

produced and grown under the same conditions,
exactly like each other, as a rule, or they differ
hardly more than specimens of one and the same
species are apt to do.

This carefully stated proposition, founded upon
experience, appears to be sufficiently justified by
numerous experiments, but it is, nevertheless,
subject to many exceptions. Some students of
crosses have so limited its application that they
dared only to assert the similarity of the speci-
mens obtained from a capsule fertilized by the
same plant. At all events, the rule proves itself
to a certain degree trustworthy only in those
cases in which the similarity of origin and con-
dition of growth demanded by the terms of the
rule are really present.

The question easiest to answer is Just the one
about which there has been the most violent dis-
cussion, namely, that relating to the stronger
influence of the one or the other sex upon the
offspring. The crosses of the two species or races
A and B resemble each other whether A was male
or female in the crossing. Experimenters, es-
pecially Kélreuter, Gartner, Naudin, and Wichura,
have not, on the whole, been able to find any

arbitrary signs are used in the text to designate the sex of each
parent. The sign ¢ represents the male, or the parent which
furnished the pollen; Q stands for the female or seed-bearing
parent. —L. H. B.

FOCKE ON THE PRIMARY CROSS. 217

difference between AQ? xBs, and BE xAe.
More than one hundred years had passed since
Kolreuter had proved the conformity of Nicotiana
rustica 2 x paniculata ¢, and N. paniculata ? x
rustica ¢, when one of the acutest florists of our
time, Timbal-Lagrave, was also astonished in the
highest degree by a similar experience. All the
rules and supposed experiences according to which
the florists were to know, from the morphological
characteristics of a hybrid, which of the progeni-
tors had furnished the pollen for its formation,
and which had borne the seed, are entirely ground-
less and foolish. Besides, it has been proved by
many experiments, that in the vegetable kingdom,
with regard to species, as a rule, the form-deter-
mining power of the male and female elements in
the progeny is entirely equal.

As with all other rules touching the crossing of
plants, this one of the similarity of the products
of reciprocal crossings is not without its excep-
tions. It is a matter of course that an observed
difference can be ascribed, with any probability,
only to a stronger influence of the male or female
elements, provided that the experiments are car-
ried on in exactly the same manner, and if they
always lead to the same result after repeated
trials. So far, almost all the recorded experi-
ments could be improved in this respect, for they
are open to justifiable doubt. The following

218 BORROWED OPINIONS.

declarations of the differences in the products of
reciprocal crossing seem worthy of notice.

a. The influence of the female element in Pe-
largonium fulgidum x grandiflorum, P. peltatum
x zonale, Epilobium hirsutum x Tournefortii,! ex-
ceeds that of the male as regards general form.
This influence is also shown more strongly in the
colors of the flowers of several hybrids of Dig-
italis, in some also in the form of the corolla. In
Nymphiea rubra x dentata, the seed-leaves resemble
those of the female progenitors.

6b. The female element shows, apparently, an
overpowering influence in the power of with-
standing cold in the Rhododendron (hybrids of
R. arboreum), Lycium, and perhaps also in Cri-
num (hybrids of C. Capense).

ce. The male element influences the general
form especially in Papaver Caucasicum x somni-
ferum, and Cypripedium barbatum x villosum (aif
constant ?); it shows a stronger influence upon
the colors of the flower in Petunia.

d. Giirtner says that he has sometimes observed
differences in the fruitfulness and progeny of re-
ciprocal hybrid forms, e.g. in Dianthus barbatus
x superbus. The experiences of Gartner should,
however, hardly suffice to establish the uniformity
of this occurrence.

The principal differences between A? and Bé

1 The various examples cited in the text are fully explained
in the body of Focke’s work, Die Pflanzen-Mischlinge.

FOCKE ON THE PRIMARY CROSS. 219

and B? and A ¢, have been observed by Koélreuter
and Girtner in some Digitalis hybrids. ‘That
these differences really show themselves each time
and in the same manner, is by no means proved.

Much oftener, the departures from the regular
uniformity of single specimens, observed among
hybrids, are entirely independent of the influence
of the species in the hybrid formation. Often
important differences appear among the seedlings
of the same cross which have been treated exactly
alike. These differences show themselves in dif-
ferent ways : —

1. Single specimens of the cross may show but
shght differences among themselves, especially in
the color of the blossoms and similar character-
istics, which are easily changed; compare, for
example, the hybrids of Verbascum phoeniceum,
Salix Caprea x daphnoides.

2. ‘The cross may appear in two different types,
each of which represents a different combination
of the characteristics of the progenitors. As a
rule, one type more closely resembles the one, and
the second the other progenitor; the numbers of
the two types are often very unequal. Giartner
designated the rarer forms “exceptional types”
(Ausnahmetypus). For examples, study Cistus,
Dianthus, Geum, Cinothera, Lobelia, Verbascum
Thapsus x nigrum, Nicotiana quadrivalis x Taba-
cum, var. macrophylla.

220 BORROWED OPINIONS.

9

3. The cross may appear in several different
types. Giartner gives some examples of these,
although in his cases it is probably a question of
three known forms of one polymorphous union.

4. The cross may appear in a typical interme-
diate form, and a number of vacillating ones
approaching one or the other parent, and among
which no distinct type can be distinguished.
Such is Medicago falcata x sativa, usually also
Melandrium album x rubrum.

5. The cross may have from the beginning
very many forms. ‘The experiments which have
been made leave it doubtful if in these cases one
or more constant types, with a similar combina-
tion of characteristics, can be distinguished among
the vacillating forms. Study Abutilon, hybrids
of Pelargonium glaucum, P. radula x myrrhi-
folhum, Passiflora, Hieracium, Nepenthes, Narcis-
sus. Gartner made the assertion that hybrids
between two species are of similar form, whilst
crosses of varieties are polymorphous. If, by va-
rieties, is understood the unsettled garden forms
or garden crosses, then the remark is justifiable ;
but if fixed races of pure descent are also included,
then it is decidedly wrong. See ‘“ Cross-breeds
and Hybrids” (section III., page 247).

Entirely different results are obtained by com-
paring hybrids which, although springing from
the same species, were produced and grown in

FOCKE ON THE PRIMARY CROSS. oot

different places. Spontaneous hybrids are, as a
rule, much more variable than those produced
artificially, e.g. Verbascum Lychnitis x Thapsus
and V. Lychnitis x nigrum. My hybrids between
Digitalis purpurea and D. lutea resembled each
other much when I had sown the seeds; on the
other hand, very different forms arose when the
seeds had accidentally sown themselves. It may
be that in this case there was no real causative
connection between the many forms and the man-
ner of sowing; nevertheless it is certain that
several growers have very often obtained different
results in their crossings from the same species.
The uniformity of all the products of the same
crossing, which undoubtedly is the rule with the
experiments of growers, appears in nature rather
to be the exception than the rule. It remains to
be ascertained what influence the unequal nour-
ishment of the parent species, or of the hybrid
seedlings, has upon the variability of the hybrids.

2. The characteristics of the crosses may be differ-
ent from the characteristics of the parent species.
It is in size and luxuriance (see proposition 3), as
well as in their sexual ability (see proposition 4),
that they differ most from both parent species.

The manner differs in which the characteristics
of the parent species are united in the cross.
Generally, a blending or mutual union takes
place, but often in such a manner that in one

999 BORROWED OPINIONS.

respect one, and in another respect the other
parent seems to appear. Sometimes, for example,
the cross resembles one parent in the leaves, and
the other parent in the flower. Sometimes there
appears a variety of the cross, in which the char-
acteristics are distributed in the reverse order.
Some crosses resemble at first one species, and
later the other parent species; as their leaves
show in the spring the one and in the fall the
other type (Cistus, Populus); or the colors of
the flowers change during the blossoming (Melan-
drium album x rubrum, Epijlobium roseum x
montanum ; compare also Lantana), or in the fall
(Nicotiana rustica x Tabacum, ‘Tropzeolum, Lobe-
lia, etc. ), also sometimes in different years (Bletia
crispa X cinnabarina, Galium cinereum x verum).
By the union of races (rarely in hybrids in the
more restricted sense of the term), one finds,
among other things, the characteristics of the par-
ent species unmixed beside each other (compare
Cucumis Melo, the thorniness of the Datura
fruits, color of the flower in Rhododendron Rho-
dora xX calendulaceum, R. Ponticum x flavum,
Anagallis, Linaria vulgaris x purpurea, Calceo-
laria, Mimulus, Mirabilis). The colors of the
flower often appear in an unexpected and unex-
plainable manner: the hybrids of Verbascum
phceniceum are very variable in the colors of
the flowers, and in other respects quite uniform;

FOCKE ON THE PRIMARY CROSS. 293

on the hybrids of Helianthemum have sometimes
been found differently colored flowers on the
same stalk, at the same time.

From the crossing of closely related races,
especially varieties in color, there often appear
plants which exactly or very closely resemble the
parent types; compare Brassica Rapa, var., Linum,
Pisum, Phaseolus; Anagallis, Atropa, Datura
Stramonium, Salvia Horminum, etc. Usually
the influence of the second parent race does not
show itself till the second generation; and, in
fact, in such a manner that a part of the seed-
lings return to that form, either entirely or only
in certain respects. Only in Atropa, no return
to the (slightly fixed) yellow form has _ been
observed.

In some eases, the cross resembles one of the
parent forms so much that it could be considered
as only a slight variation of the same. Even in
crosses between two considerably different species,
the predominating influence of one parent species
is sometimes shown in a striking degree. The
hybrid of Dianthus Armeria x deltoides resem-
bles D. deltoides much more than the other
parent species; of D. Caryophyllus x Chinensis
resembles D. Caryophyllus ; of Melandrium ru-
brum x noctiflorum resembles M. rubrum; of Ver-
bascum Blattaria x nigrum resembles V. nigrum ;
of Digitalis lutea x purpurea resembles D. lutea.

994 BORROWED OPINIONS.

Sometimes even the primary hybrids show
characteristics which are entirely different from
either parent species ; especially is this the case,
amongst other things, in the colors of the flowers.
The most curious is the regularly blue-flowering
hybrids of the white Datura ferox with the
equally white D. levis and D. Stramonium,
var. Bertolonii. There are many examples of
unexpected coloring of the blossoms of hybrids
from species of colored flowers, while the crosses
by no means always show the shades of color
which would be obtained by a mixture of the
parental pigments. Noticeable instances are
shown, e.g., by Clematis recta x integrifola, Aqui-
legia atropurpurea x Canadensis (and others),
Nicotiana suaveolens x glutinosa, Verbascum
pulverulentum x thapsiforme, hybrids of V.
pheeniceum, Anemone patens x vernalis, Be-
gonia Dregei x Sutherlandi (and others). In
the crosses of races, e.g., of Papaver somniferum
and Datura Stramonium, many characteristics
appear which belong not to the parent forms, but
to other races of the same species. Nicotiana
rustica x paniculata sometimes shows the colors
of the blossoms of N. Texana, a foreign sub-
species of N. rustica. Other characteristics
which the hybrids show in a greater degree than
the parent forms, are, e.g., the greater stickiness
of some hybrids of Nicotiana (rustica x paniculata),

LL

ee ee eee ee

FOCKE ON THE PRIMARY CROSS. 225
the apparently greater wealth of honey in N.
rustica x paniculata, the stronger nauseous odor
of the hybrids of Melandrium viscosum, the
supposed increase in the quantity of quinine (?)
in hybrids of Cinchona (according to Kuntze).

In later generations of hybrid growth, devia-
tions from the characteristics of the parent species
are much oftener observed.

3. Crosses of different races and species are dis-
tinguished from plants of a pure race, as a rule,
by the power of vegetation. Hybrids between very
different species are often very weak, especially
when young, so that tt is difficult to successfully
raise the seedlings. On the other hand, crosses of
more closely related species and races are, as a
rule, uncommonly luxuriant and strong; they are
distinguished mostly by size, rapidity of growth,
early flowering, abundance of flowers, longer life,
stronger reproductive power, unusual size of some
special organs, and like characteristics.

For a closer confirmation of this proposition, it
will be to the purpose to refer to some examples.
Delicate seedlings are mentioned, e.g., in Nym-
pheea alba crossed with foreign species, Hibiscus,
Rhododendron Rhodora with other species, R.
Sinense with Eurhododendron, Convolvulus, poly-
phyllous Salix hybrids, Crinum, Narcissus. The
experience that seedlings of hybrid-fertilized seeds
are delicate, is frequent. A dwarfish growth has

Q

226 BORROWED OPINIONS.

rarely been observed in hybrids, although there
are some instances of it. Large growth is, on the
other hand, much more common, e.g., Lycium
Datura, Isoloma, Mirabilis. Usually the hybrids
exceed in height both parent species, or at least
their medium height ; compare, e.g., many hybrids
of Nicotiana, Verbascum, Digitalis. The vege-
tation sometimes takes place exceedingly fast.
Klotzsch emphasizes the rapid growth of his
hybrids of Ulmus, Alnus, Quercus, and Pinus.
The flowering time often comes earlier than in
the parent species, e.g., in Papaver dubium x
somniferum, some Dianthus hybrids, Rhododen-
dron arboreum x Catawbiense, Lycium, Nicotiana
rustica x paniculata, Digitalis, Wichura’s six-fold
Salix hybrid, Gladiolas, Hippeastrum vittatum x
Regine, etc., but especially in many hybrids of
Verbascum. On the other hand, there are, no
doubt, also single hybrids which blossom only
after a long period, or not at all, e.g., in the genera
Cereus and Rhododendron. Only one example
of earlier maturity of seeds, independent of an
earlier maturing of the blossoms, is known to me,
namely, in Nuphar. Very often and very com-
monly an extraordinary abundance of flowers has
been observed in hybrids ; compare, e.g., Capsella,
Helianthemum, Tropeolum, Passiflora, Begonia,
thododendron, Nicotiana (rustica x paniculata,
elutinosa x Tabacum, and others), Verbascum,

FOCKE ON THE PRIMARY CROSS. OL

Digitalis, many Gesneracewe, Mirabilis, Cypripe-
dium. The size of the blossoms is often increased
in hybrids; in a cross of two species with different
sized flowers it is not rare that the flowers of the hy-
brid attain the size of those of the larger-flowered
parent, or almost that size. Dianthus arenarius x
superbus, Rubus cesius x Bellardii, hybrids of Rosa
Gallica, Begonia Boliviensis, Isoloma Tydeum,
give examples of uncommonly large flowers.

A strong vegetative reproductive power is very
common among hybrids ; compare, e.g., Nymphea,
hybrids of Rubus cesius, Nicotiana suaveolens x
Tabacum, var. latissima, Linaria striata x vulgaris,
Potamogeton. <A longer life is especially notice-
able in some hybrids of Nicotiana and Digitalis.
A greater power of resistance against cold is also
noticeable in Nicotiana suaveolens x Tabacum, var.
latissima, while Salix viminalis x purpurea is said
to be more sensitive to frost than either of the
parent species.

These facts indicate partly a certain loss of
vigor which is inherent to the hybrids on account
of their abnormal descent, and partly, on the con-
trary, an exceptional vegetative power. ‘This
latter fact, which occurs much oftener than the
former, has only lately been elucidated to any
extent. The important experiments of Knight,
Lecoq, and others, had been known for some time,
but only through the careful researches of Charles

228 BORROWED OPINIONS.

Darwin has the favorable influence of a cross
between different individuals and races of one
and the same species been clearly shown. The
strengthening of the vegetative power in some
hybrids is evidently a universal experience which
needs no special explanation, on account of the
peculiar circumstances of the formation of the
hybrids. It was formerly thought that the di-
minished sexual fruitfulness is compensated by a
greater vegetative luxuriance, a statement the
untenableness of which, as Girtner showed, is
most plainly shown by the fact that many of the
most fruitful crosses (Datura, Mirabilis) are also
distinguished by an enormous growth.

4. Hybrids of different species often form in their
anthers a smaller number of normal pollen grains,
and in their fruits a smaller number of normal seeds,
than plants of a pure descent; sometimes they pro-
duce neither pollen nor seeds. In crosses of nearly
related races, this weakening of the sexual organs
does not oceur as a rule. The blossoms of sterile or
slightly fruitful hybrids usually remain fresh for
a long time.

No characteristic of hybrids has received more
attention than the diminution of the sexual power
which has been observed in them. Even Ko6lreuter
believed that this characteristic permitted the
drawing of a sharp line between species and vari-
eties. The same thought has prevailed amongst

ae Sai |

FOCKE ON THE PRIMARY CROSS. 229

botanists to a great extent since then, and even
in very recent times Naudin, Decaisne, and Cas-
pary have defended the ideas of Kolreuter in a
more or less modified form. Knight, Klotzsch,
and formerly also Gordon, considered the pollen
of hybrids to be entirely impotent, a view which
was even then contradicted by the accurate ex-
periments of Koélreuter. It has often been wrongly
declared that K6olreuter had himself spread the
doctrine of the total sterility of hybrids; this
assertion 1s to be explained only by ignorance or
by a misunderstanding of the Latin text. K6l-
reuter, in fact, does not speak of sterility, but of
lessened fruitfulness, as being a common charac-
teristic of hybrids.

The fruitfulness of hybrids is considerably dif-
ferent, according to the individual genera. For
instance, the hybrids of Papaver, Viola, Verbas-
cum, and Digitalis show but little fruitfulness ;
the hybrids of Anemone, Nicotiana, Mentha, Cri-
num, the Cucurbitacee, and Passiflora are much
oftener fruitful, while in Aquilegia, Dianthus,
Pelargonium, Geum, Epilobium, Fuchsia, Coty-
ledon, Begonia, Cirsium, Erica, Rhododendron,
Calceolaria, Quercus, Salix, Gladiolus, Cypripe-
dium, Hippeastrum, the Gesneracee, and Orchide,
the fruitful hybrids are commoner than the bar-
ren ones. In the genera Vitis, Prunus, Fragaria,
and Pyrus, we use the crosses of closely related

230 BORROWED OPINIONS.

species as fruit plants; in Cereus, hybrids of even
widely different species show undiminished fruit-
fulness.

The sterility of hybrids is sometimes shown by
their exhibiting no inclination to blossom, a char-
acteristic which has been observed particularly in
some hybrids of Rhododendron, Epilobium, Ce-
reus, and Hymenocallis. These are, however,
are exceptions, for as a rule hybrids bloom earlier
and more profusely than the pure species (see
page 226).

In hybrids which have flowers of only one sex,
the staminate blossoms often fall off while they
are yet buds, as in Cucurbitacee and Begonias
(hybrids of B. Froebeli). Sometimes the anthers
are arrested in their growth and form hermaphro-
dite flowers, as has been observed in some hybrids
of Pelargonium and Digitalis (D. lutea x pur-
purea form tubiflora Lind1.). The common result
of the production of hybrids is a different forma-
tion in the pollen grains in the hybrids from that
of the parents. Often the anthers of the hybrids
are dead and contain no pollen, or they are small
and do not open at all. Such deficiency of pollen
can be observed, for example, in Rubus Ideus x
odoratus, Ribes aureum x sanguineum, Alopecurus
geniculatus x pratensis. In other cases, the pollen
dust consists of small powdery grains of irregular
form and size, which do not swell when moistened,

FOCKE ON THE PRIMARY OROSS. ee |

and among which are usually found a few well-
formed pollen cells which are capable of germina-
tion. Often the number of normal pollen grains
is greater, and comprises ten or twenty per cent,
or more, of the whole number. There are often
found, in a greater or less degree, large angular
grains capable of swelling, as well as well-formed
ones, by the side of those dwarfed or stunted.
Among crosses of closely related species, as Melan-
drium album x rubrum, there are usually found
only few irregularities in the form of the pollen
grains. In a hybrid Sinningia, the pollen of the
second year of flowering was better than that of
the first.

In hybrids of undoubted different species, a
normal formation of the pollen is seldom seen.
The statements on this subject require, for the
most part, corroboration, nevertheless I refer to
Nymphea Lotus x rubra, Begonia rubrovenia x
xanthina, Isoloma Tydeum x sciadocalyx, Salix
purpurea X repens; almost perfectly formed pollen
grains were found in Salix aurita x Caprea, and 8.
viminalis x repens.

On the other hand, it is still more rare that a
deficiency of pollen is found in an evident cross
of races. Perhaps it could be found oftener if
one sought for it. The only certain example of
which I know is an Anagallis cross by myself.
Whether Raphanus sativus and R. Raphanistrum

ee BORROWED OPINIONS.

are to be considered as species or races is doubtful.
Nevertheless there are some crosses of very closely
related species which appear to be entirely sterile,
as, for example, Capsella rubella x Bursa-pas-
toris, Viola alba x scotophylla, Papaver dubium
x Rheeas.

The sexual capabilities of the female organs
are, as a rule, not so much weakened in hybrids
as the male. Nevertheless they are greatly
diminished. Many hybrids never produce fruit.
Even after many experiments, one must not make
definite assertions about the absolute sterility of
a hybrid: in Rubus cesius x Ideeus, for example,
one can see several thousand blossoms remain
sterile, and nevertheless, now and then, fruit is
produced. Compare, further, Digitalis lutea x
purpurea, Lobelia fulgens x syphilitica, Crinum Ca-
pense x scabrum. <A morphologically distinguish-
able defect of the ovule of hybrids has seldom been
referred to so far, although it has been observed in
Cistus, by Bornet. If one wishes to obtain a defi-
nite judgment on the female creative power in
hybrids, he must fertilize the ovules with pollen of
the parent species, which, as a rule, develops more
perfect fruit than the pollen of hybrids which has
been weakened in its generative power. In some
cases, hybrids whose pollen shows but slight
potency produce normal fruit with pollen from
the parent, as in Luffa.

FOCKE ON THE PRIMARY CROSS. 238

Some hybrids drop their entire blossoms un-
wilted, with the calyx and flower stalk intact,
e.g., Ribes, Nicotiana rustica x paniculata, and
other hybrids of Nicotiana.

As a rule, the corolla wilts, after a longer
period, in a normal manner, or is thrown off as
in the parent species; but no setting of fruit thus
takes place, or only a little of very poor character.
Sometimes the fruit is well formed externally,
but contains no seed. In many cases hybrids set
fruit, but to a less extent and with fewer seeds
than the parent species. Even in crosses of
closely related species, the number of seeds seems
to be smaller than in the parent species; so, for
example, according to Girtner, in Melandrium
album x rubrum, Lobelia cardinalis x fulgens, and
even in undoubted crosses of races of Verbascum.

Hybrids of essentially different races rarely
show an undiminished fruitfulness, although no
noticeable diminution has been proved in Brassica
Napus x oleracea, Dianthus Chinensis x pluma-
rius, var. Sibiricus, Pelargonium pinnatum x hir-
sutum, Abutilon, Medicago, a few Cereuses and
Begonias, Hieracium aurantiacum x echioides, Ni-
cotiana alata x Langsdorffi, a few hybrids of
Erica, Calceolaria, Isoloma, Veronica, and several
Orchide. Fruits and seeds in abundance are also
found in many other garden hybrids, and in many
wild ones, as in Roses, Epilobiums, Fuchsias, Cir-

934 BORROWED OPINIONS.

siums, Hieraciums, Willows, of Lobelia Lowi,
etc. In these cases, however, one cannot exactly
ascertain if the plants are primary hybrids, or,
which is much more probable, if they belong to
later generations or arose through derivative
hybridization (Riickkreuzungen).

In order to set seeds, or at least to produce a
vigorous progeny, some hybrids require to be fer-
tilized by other individuals, even though they are
themselves hybrids; compare, for example, hybrids
of Cistus, Begonia, Gladiolus, and Hippeastrum.,
In some hybrids, only the first flowers produce
seeds, as Aquilegia, Dianthus, Silene, Lavatera
Thuringiaca x Pseudolbia, Rubus foliosus x Spren-
gelii; in other cases, the first flowers are regu-
larly sterile, while the later ones are often fruitful,
as in Datura, Nicotiana rustica x paniculata, N.
rustica x quadrivalvis, Mirabilis. In longer-lived
plants, often all the flowers of the first years are
sterile, while later, when the plant has reached a
certain age, a few fruits are formed; this has
been noticed, for example, in Rubus Ideeus x
cesius, R. Bellardii x ceesius, Calceolaria integri-
folia x plantaginea, Crinum Capense x scabrum.

Although, as a rule, the female creative power
in hybrids is less weakened than the male,
still there are some cases in which the reverse
is true ; compare Nymphea Lotus x rubra, Cico-
nium x Dibrachya in the genus Pelargonium,

_——S lL

FOCKE ON THE PRIMARY CROSS. B85

Lobelia fulgens x syphilitica, Verbascum thapsi-
forme x nigrum, Narcissus montanus, etc; these
are probably, for the most part, only accidental
occurrences.

The longer duration of the flowers (especially
the pollinized ones) on many sterile hybrids, is
an occurrence which is analogous to the longer
duration of unfertilized or incompletely fertilized
flowers. Often in sterile hybrids, especially after
dusting with the pollen of the parent or a related
species, the fruits swell more or less without per-
fect seeds’ being formed in them. Externally
well-formed but seedless fruits are found in Cacti,
Passifloras, Cucurbits, and Orchids. Gartner has
studied this characteristic, which is of no special
use or value in the culture of hybrids, very care-
fully ; however, it offers an important proof of
the truth of the statement that the development of
the outside coats of the fruits takes place in a
normal manner on account of the irritation which
the germinating pollen produces, but that it is
nevertheless independent of the fertilization of
the ovule, and the development of the embryo and
of the seed.

One can, in general, make the statement that
hybrids of closely related races are more fruitful
on the average than those of considerably differ-
ent species. One can also consider, as a rule, as
has been shown above, that closely related species

236 BORROWED OPINIONS.

form hybrids among themselves more easily than
those considerably different. Both rules are true
only to a certain extent. If one should infer from
these that hybrids are more fruitful the easier they
are formed, then he would go far wrong. ‘There
is no definite relation between ease of formation
and fruitfulness of crosses.

From a teleological point of view, we formerly
saw in the sterility of hybrids a means of keeping
“species? separate. What purpose this separa-
tion was to serve was not explained, unless it was
for the convenience of the systematist. Now, on
the contrary, we ask if the formation and sepa-
ration of species does not really depend upon
the fruitfulness of the crosses between the dis-
tinctly marked races of the parent types. The
noticeable resemblance between illegitimate and
hybrid progeny gives us no standpoint for further
investigation of the cause of unfruitfulness. More
light may be given by the fact that in the hybrid
pteridophytes and mosses the formation of sexless
spores is just as much wanting as is the formation
of pollen grains in hybrid phanerogams. The
obstacle to the regular propagation of hybrids
seems to lie in the development of certain cells
which have the power to maintain the type of the
parent form, it being immaterial if these cells have
sexual functions or not. In any case, more facts
must be gathered to justify the adoption or pres-

FOCKE ON THE PROGENY OF CROSSES. 237

entation of a principle of such scope. This view
of the situation can even now be considered as a
hypothesis which certainly as yet offers no expla-
nation of the limitations of species, but it leads
the way to a final solution, because it brings a
long list of different but plainly analogous phe-
nomena as facts in the animal and vegetable king-
doms under one common point of view.

5. Malformations and curious forms are much
more common, especially in the flower parts of
hybrids, than in individuals of a pure descent.

Compare Papaver, Dianthus, Pelargonium, Ni-
cotiana, Digitalis. Double flowers appear to be
formed especially easily in hybrids.

Il. THE PROGENY OF CROSSES.

Hybrids are more easily and completely fertil-
ized by the pollen of the parent forms than by
their own. Exceptions to this rule are hardly
known (although compare Hieracium echioides x
aurantiacum), although no large number of ex-
periments have been made in this direction. By
their own pollen is to be understood the pol-
len of hybrids of the same cross, as well as that
of the same individual. When hybrids grow in
the vicinity of parent forms, they must naturally
often be fertilized by them. In the progeny there
will be, therefore, a number of forms between the

238 BORROWED OPINIONS.

primary hybrids and the parent species. In sow-
ing the seeds of hybrids, one cannot always deter-
mine if fertilization from the parent species could
have taken place or not. The general statement
that the progeny of a hybrid has shown itself to
be very variable is therefore of little weight or
worth. Sometimes a hybrid is more easily fertil-
ized by a third species than by its own pollen.
Compare, for example, Nicotiana rustica x panicu-
lata, and Linaria purpurea x genistefolia.

1. Progeny of Crosses with their own Pollen.
(A ane x Ga Be

a. If one protects the fruitful hybrids from the
influence of the parent forms or other related spe-
cies, then he obtains hybrids of the second genera-
tion. It seems to me that the progeny of hybrids
appears or acts very differently according to the
leneth of their life. In long-lived plants, the
mingling and mutual blending of the two types
which have been joined in the hybrid often ap-
pears to be more complete, so that the progeny also
inherits the characteristic of the new intermediate
form or type in an equal or uniform manner.

The progenies of one- or two-year hybrids are,
as a rule, very different and varied in form;
compare Pisum, Phaseolus, Lactuca, Tragopogon,

|
|

FOCKE ON THE PROGENY OF CROSSES. 239

Datura, Nicotiana alata x Langsdorffii, etc. Ex-
ceptions are found in Brassica, Gtnothera, Nico-
tiana rustica x paniculata, Verbascum Austriacum
x nigrum.

The progenies of hybrids of several generations
(mehryahriger) are generally similar, although the
cases in which the intermediate type shows itself
to be constant appear to be much more frequent.
Some hybrids of Aquilegia, Dianthus, Lavatera,
Geum, Cereus, Begonia, Cirsium, Hieracium, Pri-
mula, Linaria, Veronica, Lamium, and Hippeastrum
appear to come very true to seed.

The progenies of shrubs and trees are, in the
majority of cases, very constant or invariable ;
compare Atsculus, Amygdalus, Prunus, Erica,
Quercus, Salix. Also many hybrids of Fuchsia
and Calceolaria are said to be constant. The
Rhododendron hybrids are partly true to seed
and partly variable. The progeny of the crosses
of Vitis, Pyrus, and Cratzegus, on the other hand,
seem to be very variable.

6. The several forms in which some primary
hybrids appear seem to be unsettled in the prog-
eny. In Dianthus, according to Girtner, the
“exception types,” when sown, mostly return to
the normal mongrel form. The various primary
forms of the hybrids of Hieracium, Mendel found
to be true to seed.

e. C.F. von Gartner and other botanists have

240 BORROWED OPINIONS.

made the assertion that the progeny of hybrids
becomes, from generation to generation, weaker
and less fruitful. It is certain that their growth,
which was at first increased, gradually decreases
when they fertilize themselves. Gartner’s experi-
ments, however, were made on a very small scale,
so that his hybrids were influenced not only by
close in-breeding, but also by the various circum-
stances which so often result in the loss of garden
plants cultivated only in small numbers. Even
Girtner noticed exceptions, as in Aquilegia, Dian-
thus barbatus x Chinensis, D. Armeria x del-
toides. Crosses of closely related species evidently
can be continually propagated or kept up with
ease ; compare Brassica, Melandrium, Medicago,
Petunia. Many gardeners assert with much con-
fidence that many hybrids can be propagated very
well for several generations by means of seeds ;
compare Erica, Lychnis, Primula Auricula x hir-
suta, Datura. Many observations of wild plants
seem to confirm this view. ‘The principle has also
been laid down that the fruitfulness of hybrids
increases again in later generations. It does not,
however, appear that this rule can have a very
broad application. It is much more probable that
single fruitful specimens arise among hybrids,
which can easily reproduce themselves under favor-
able external conditions by inheriting this pe-
culiarity. Fruitful descendants of hybrids are

FOCKE ON THE PROGENY OF CROSSES. 241

probably sometimes the product of derivative
hybridization.

d. Complete reversions to the parent forms
without the aid of pollen from parent forms arise
only in crosses of closely related races. Even in
such crosses, true reversions only seldom take
place, e.g., Phaseolus.

e. From the variable progeny of fruitful crosses
a few principal types often arise after a few — per-
haps three or four — generations. If one protects
these new types from crossing, they tend to be-
come constant or fixed. Scientific trials or exper-
iments which confirm this statement have been
made only to a small extent, especially by Lecoqg
in Mirabilis, by Gordon in Linaria and especially
in Datura. ‘These gardeners have produced many
new races by the crossing of related species and
well-fixed races. Also many wild, fixed, inter-
mediate forms may have arisen in this manner ;
compare Brassica, Lychnis, Zinnia, Primula, Petu-
nia, Nicotiana commutata, Pentstemon, Mentha,
Lamium. The new types of the descendants of
crosses frequently differ in some characteristic
from both parent forms. My Nicotiana rustica
x paniculata had in the second and third genera-
tions, on the whole, much smaller leaves than
either of the parent forms.

J. The sterility and inconstancy of the progeny
of hybrids have often led botanists to conclusions

R

QA?, BORROWED OPINIONS.

which are not confirmed by experience. It is
entirely wrong, as can be seen by the facts which
have been stated, to assert that all crosses would
necessarily soon be lost on account of these char-
acteristics which have been promiscuously ascribed
tothem. The unsettled forms arising from crosses
are the plastic material out of which not only
gardeners form their new varieties, but which
material is biologically the more valuable as it
furnishes new species in the household of nature.

2. Derivative Hybridization of Crosses with the
Parent Forms.

(AQOxBS)OxASG, APxBS)Sx BS, AUX oe

As long as one laid much stress upon the male
or female influence which one or the other parent
species may have had upon the hybrid, a differ-
ence was carefully made between the advancing
hybrid forms, or those which more closely resem-
bled the male parent, and the degenerating
hybrid forms, or those which more closely resem-
bled the female parent. But these differences
are, according to experiments that have been
made, of very subordinate importance, or, per-
haps, of none at all.

By treating hybrids with parental pollen, one
obtains, as a rule, a rather varied progeny. The

FOCKE ON DERIVATIVE HYBRIDS. 243

intermediate form between the hybrid and respec-
tive parent is apt to be more numerous and more
fruitful. Besides this, there are formed a less
number of individuals, some of which resemble
the hybrid, and some the parent species. Both
are apt to be but slightly fruitful.

The three-fourths hybrids, (A x B) 2? x Aé,
are often quite fruitful with their own pollen and
appear to give races true to seed easier than the
original hybrid; compare A¢gilops spelteformis.
Gartner observed frequently that in later genera-
tions the pollen became more regular and the
fruitfulness greater, as in Dianthus (Chinensis
barbatus) x barbatus, and also in other three-
fourths hybrids of Dianthus, Lavatera, and
Nicotiana.

If one treats the three-fourths hybrid (A x B) 9
x AS again with pollen from A, then he
obtains a seven-eighths hybrid, or the third
hybridized generation, which, as a rule, is very
much like the parent which furnished the seven-
eighths part, but it is still apt to show marked
differences in form and fruitfulness in some indi-
viduals. The last traces of the one original
parent species disappears mostly in the fourth,
fifth, or even sixth hybridized generation.

Kolreuter and Gartner have completed the
transition of one parent species into the other in
many cases. ‘They found that for a complete

244 BORROWED OPINIONS.

change, three to six generations were necessary —
as a rule, four to five. HEvidently the greater or
less duration of the change depends more or less
upon surrounding circumstances. Gordon found
that Melandrium album x rubrum was like the
parent species in the second generation, when —
fertihzed with its own pollen, while Gartner
found three to four generations necessary to
bring the one into the other by means of parental
pollen.

As arule, the products of the fertilization of a
parent species with pollen from a hybrid, as
A? x (A x B)¢, are similar to those of the
opposite cross; nevertheless the statements of
observers agree that the variety of forms is apt
to be greater when one uses the hybrid as the
male element ; compare Dianthus and Salix.

There appears, in the products of derivative
hybrids of crosses, as among the direct progeny,
new characteristics, which are lacking in the
parent forms, but are for the most part found in
related races or species.

3. Hybrids of Several Species.
a. Triple Hybrids.
In the first years of his experience, Koélreuter

succeeded in uniting three entirely different Nico-
tiana species into one hybrid form. The simplest

FOCKE ON MULTIPLE HYBRIDS. 245

formulas according to which such a union could
take place are: (A xB)? x CS,C? x (AxB)é,
and, (A.x B) 9ox (A x.C)¢é-> In. the genera
Dianthus, Pelargonium, Begonia, Rhododendron,
Nicotiana, Achimenes, Calceolaria, Salix, Hippeas-
trum, Gladiolus, and a few others, there has been
made a number of such unions without any par-
ticular difficulty. One must nevertheless deter-
mine if he unites three essentially different species,
or if two of the factors, or even all three are only
closely related to each other. ‘There are similar
but evidently different species which, in crossings
among themselves, behave almost like races of the
same species, as for example: —

Melandrium album and rubrum.

Vitis vinifera, cordifolia, zestivalis and Labruscea.

Lobelia fulgens, splendens, and cardinalis.

Rhododendren Ponticum, arboreum and Cataw-
biense.

R. flavum, viscosum, nudiflorum, and calendu-
laceum.

Berberis Aquifolium, and the most closely
related species.

Hybrids between the crosses of two species of
these groups with the third species of the same
genus, can no more be called true triple hybrids
than crosses of species belonging to some smaller
or narrower group of Vitis, Lobelia, and Rhodo-
dendron. True triple hybrids which have been

246 BORROWED OPINIONS.

formed from three essentially different species are
apt to be much varied in form, especially if the
male parent species was a hybrid. On the other
hand, in those unions which are most easily formed
and are made by the formula (A xB) 2 x Cé,
the type of C is apt to predominate strongly,

as, for instance, Nicotiana (rustica x paniculata) 9-

x Longsdorffii ¢, Achimenes (grandiflora x can-
dida) 2 x longiflora ¢, and other Gesneracee.
The hybrids of Erica are said to produce just as
uniform a progeny as the pure species. Several
Salix hybrids have acted in the same manner.

For gardeners, therefore, the triple hybrids in

some genera (as in Pelargonium, Begonia, Rhodo-
dendron, Achimenes, Isoloma, Cypripedium, Gladi-
olus) are very valuable. If they produce seeds,
their progeny is very variable.

b. Hybrids of Four to Six Species.

If one does not count the crossings of very
nearly related species (as Vitis, Rhododendron,
etc. ), these hybrids of four or more parent forms
are somewhat rare. We know them especially in
the genera Dianthus, Pelargonium, Begonia, Rho-
dodendron, Nicotiana, Salix, Hippeastrum, Gladi-
olus. The artificial union of different species in
a single hybrid form has been carried farthest by
Wichura, who united six Salix species.

. ee hee A ee

—— SO ee

FOCKE ON GROSS-BREEDS AND HYBRIDS. 247

c. Crosses of Plants Grown Together.

In some genera, as Pelargonium, Fuchsia, Be-
gonia, Rosa, Erica, Rhododendron, Achimenes, Cal-
ceolaria, Gladiolus, Hippeastrum, gardeners have
crossed species and hybrids in the most manifold
manner, intentionally and unintentionally, and
have used the most promising forms obtained for
further propagation. The progeny of this com-
plex crossing is naturally almost always very vari-
able. There appear, however, to be exceptions to
_ this rule; Sweet plainly asserts that one always
obtains the same cross from the crossing of some
complex Pelargonium hybrids. Such constant
complicated hybrids are, according to him, P.
(hyb.) involucratum x (hyb.) ignescens and P.
(hyb.) Mostyne x (hyb.) ignescens. That the
Erica and some Salix hybrids produce a uniform
progeny has already been mentioned.

II. CROSS-BREEDS AND HYBRIDS.

According to usage, we designate unions of two
different varieties of one species as cross-breeds,
unions of two different species as hybrids. It is
necessary, on account of the indefiniteness of the
term “variety,” to remember that only varieties
true to seed, or races and sub-species, can bequeath
their characteristics with any degree of certainty ;

248 BORROWED OPINIONS.

inconstant species, which are so often designated
as varieties, are not considered in the theory of
hybridization. : 3
Many writers have taken great pains to find a
difference between cross-breeds and hybrids; they
held firmly to the hope that by means of trials in
crossing a boundary between species and _ sub-
species could be formed. Gartner, who expresses
himself plainly in several parts of his work that the
appearance of crosses clearly proves the specific

differences of relationships of the parent forms,

becomes very reticent as soon as he attempts, on
pages 574-582, connectedly to unfold the princi-
ples of ‘‘ variety hybrids.” Herbert and Naudin
have formed the opinion, after their many experi-
ments, that it is impossible to draw the line be-
tween cross-breeds and hybrids ; but, nevertheless,
later botanists have again tried to find precise dif-
ferences between them.

The following propositions have been made : —

1. The pollen of cross-breeds is normal: hy-
brids have a greater or less number of imperfectly
formed grains in their pollen.

2. The fruitfulness of cross-breeds is normal:
that of hybrids plainly diminished.

3. Hybrids of two species with differently col-
ored blossoms produce flowers of mixed or uni-
formly modified colors: plants with irregular,
mottled flowers have always been produced by

FOCKE ON CROSS-BREEDS AND HYBRIDS. 249

the crossing of varieties. It is the same with the
coloring, marking, covering of the fruits, and
other characteristics.

4. Cross-breeds have a strong inclination to

return to the parent form in later generations.
_ These four propositions are in the main correct,
but they offer little help, in a case of doubt, to a
right decision as to specific merits. The cross of
the red and white Anagallis arvensis would have
to be considered as a hybrid on account of its pol-
len, and as a cross-breed on account of the appear-
ance of flowers of two colors. In Datura, crosses,
which in other respects are plainly characterized
as hybrids, easily show complete returns to the
parent forms. Hybrids whose fruitfulness appears
to be in no way diminished have already been men-
tioned (page 229). One can, consequently, make
the rule, that crosses of closely related races are
apt to show the characteristics ascribed to cross-
breeds, but it is impossible by that means to es-
tablish any sharp line between race crosses and
species hybrids.

Usually a few other characteristics are ascribed
to cross-breeds by which they are distinguished
from the hybrids of species. Giirtner has asserted
that cross-breeds of like descent are even in the
first generation very dissimilar, while hybrids of
the first generation are always very uniform.
This assertion, which is also repeated by others,

250 BORROWED OPINIONS.

is entirely wrong. ‘The polymorphism of the hy-
brids of the species of Abutilon, Passiflora, Hiera-
cium, ete., has already been shown, while, on the
other hand, the crosses of races, in the first gener-
ation, are usually just as uniform as the real hy-
brids. Again, it has sometimes been asserted that
the “varieties” of one and the same species, when
crossed with another species, always produce the
same hybrid forms. Giirtner, especially, has laid
particular stress upon this supposed behavior of
varieties, although he must have known that IXol-
reuter had already observed the inheritance of
color of the blossom in the races of Mirabilis,
Dianthus, and Verbascum, the doubling of flowers
in Aquilegia and Dianthus, the carriage and form
of the leaf in the races of Nicotiana Tabacum and
Hibiscus. The white-blossoming Datura ferox
gives with D. Stramonium a white-blooming cross,
and with the smooth-fruited race (var. Bertolonil)
of the same species, a blue-blossoming cross.
Nymphea Lotus x rubra is different from N.
Lotus x dentata. It cannot be in the least doubt-
ful that the inheritable characteristics of races and
so-called varieties are also bequeathed to their
progeny.

One will hardly go wrong if he assumes that
Girtner came to make this rule about the be-
havior of varieties through the behavior of unfixed
garden crosses and garden sorts. It is a matter

FOCKE ON CROSS-BREEDS AND HYBRIDS. 251

of course that forms which show themselves un-
fixed in their normal progeny should produce
polymorphous hybrids, and that unfixed variety-
signs are apt to disappear entirely in the products
of crossing with pure species.

The true situation is, in short, as follows : —
The nearer the morphological and systematic rela-
tionship of the parent forms is, the less the sexual
capacity of reproduction in the cross is apt to
depart from the normal direction ; the greater the
difference between the parent forms, the more, on
the average, is the fruitfulness of the cross weak-
ened. Exceptions are not rare.

The nearer the parent forms are related to each
other, the oftener the progeny of crosses show
complete returns to the parent forms.

Crosses from nearly related parent forms some-
times show in their blossoms and fruits the pe-
culiar characteristics of the parent forms unmixed
beside each other ; this rarely takes place in crosses
whose parent forms were considerably different.
Most unsymmetrically colored flowers (Mirabilis,
Camellia, Mimulus, Petunia, etc.) first originated
in the progeny of crosses.

LECTURE V.

POLLINATION; OR HOW TO CROSS PLANTS.

1. THE STRUCTURE OF THE FLOWER.

POLLINATION is the act of conveying pollen from
the anther to the stigma. It is the manual part
of the crossing of plants. The word fertilization
is often used in a like sense, although erroneously;
for it is the office of the pollen, not of the opera-
tor, to fertilize or fecundate that part of the flower
which is to develop into a seed.

The chief requirement in pollinating flowers is
to know the parts of the flower itself. The con-

Fic. 1. — Bell-flower.

spicuous or showy part of the flower is the envelope,
which is endlessly modified in size, form, and color.
252

STRUCTURE OF THE FLOWER. PASS}

This envelope protects the inner or essential organs,
and it also attracts insects, which often perform the
labor of pollination. This floral envelope is usu-
ally of two series or parts, —an outer and commonly
green series known as the calyx, and an inner
and generally more showy series known as the
corolla. ‘These two se-
ries are well shown in
the bell-flower, Fig. 1.
The calyx, with its re-
flexed lobes, is at C,
and the large bell-form
portion is the corolla.
When the calyx is com-
posed of separate parts
or leaves, each part is
called a sepal; in like
manner each separate
part of the corolla is
a petal. In the lily,
Fig. 2, there is no dis-
tinction between calyx
and corolla; or, it may
be said, the calyx is wanting. These envelopes
of the flower are often much disguised. This is
particularly true in the orchids, one of which, a
lady-slipper, is illustrated in Fig. 8. The sepals
are seen at DD. They are apparently only two,
but there is reason to believe that the lower sepal

Fic. 2.— Flower of white lily.

954 POLLINATION.

is really made up of a union of two. The three
inner leaves are the petals, the lower one, H,
being enlarged into the sac or slipper.

The most important organs of the flower, how-
ever, to one who wishes to make crosses, are. the
so-called sexual organs, the stamens and _ pistils.
They can be readily
distinguished in the
lily, Fig. . 2.) Bhe
six bodies shown at
S are the ends of
the stamens, or so-
called male organs.
These stamens gen-
erally have a stalk
or stem, known as a
filament, and the en-
larged tip as the
anther. It is in this
anther that the pol-
lenis. borne: 9 The
pollen is generally
made up of very mi-

Fic. 3.— Flower of greenhouse
cypripedium. nute yellow or brown-

ish grains, although
it is sometimes in the form of a more or less glu-
tinous or adhesive mass, as in the milk-weeds and
orchids. The irritating dust which falls from the
corn tassels at the later cultivatings is the pollen.

THE ESSENTIAL ORGANS. 255

The pistil, or so-called female organ, is shown at
OP, Fig. 2. The enlarged portion at O is the
ovary, Which will develop into the seed-pod. The
stigma, or the enlarged and roughened part which
receives the pollen, is at P. Between these two
parts is the slender style, a portion which is absent
in many flowers.

The stamens and pistils are known as the essen-
tial organs of the flower, for, whilst the calyx and
corolla may be entirely absent, either one or both
of these organs is present; and these are the parts
which are directly concerned in the reproduction
of the species. Like the floral envelopes, these
essential organs are often greatly modified, so
much so that botanists are sometimes perplexed
to distinguish them from each other or from mod-
ified forms of the petals or sepals. ‘The particu-
lar features of these organs which the plant-breeder
must be able to distinguish are the anther and the
stigma; for the anther bears the pollen, and the
stigma must receive it. In Fig. 1, the stamens
are shown at E. In the flower A, which has just
expanded, these stamens are rigid and in condition
to shed the pollen, but in the flower B, they have
shed the pollen and have collapsed. ‘The stigma
in this case is divided into three parts, but when
the flower first opens, these parts are closed to-
gether, H in flower A, so that it is impossible
that they receive any pollen from the same flower ;

256 POLLINATION.

when the stamens have withered, however, as in
B, the stigma, H, spreads open and is ready to

Fic. 4. — Flower of night-blooming cereus.

receive any pollen which may be brought to it by
insects or other agencies. In this case, the ovary

THE ESSENTIAL ORGANS. IST

or young seed-pod, which is in the bottom of the
flower, is not shown in the engraving.

Some of the particular forms of essential organs
are well illustrated in the accompanying photo-
eraphs. In the night-blooming cereus, Fig. 4,
the many-rayed stigma is shown just below the

Fic. 5. — Flower of the shrubby hibiscus (Hibiscus Syriacus).

centre of the mouth of the flower, and the nu-

merous stamens are arranged in a circular manner

outside of it. The many petals and numerous

spreading sepals are also well shown. ‘The hibis-

cus, Fig. 5, has a central column with the anthers

hanging upon it, and a large stigma raised beyond
Ss

258 POLLINATION.

Fic. 6.— Bugbane (Cimicifuga,
racemosa).

them. The wild bugbane,
or cimicifuga, is seen in
Fig. 6, natural size. Here
is a long spike or cluster
of flowers. At the top
are the unopened buds, in
the centre the expanded
flowers with the floral
envelopes fallen away, —
the fringe-like stamens
very prominent, — and
below are seen the pis-
tils, the stamens having
fallen. These pistils will
now ripen into pods, but
the tip-like stigma may
still be -seen on them.
The stamens and the long
protruding style, tipped
with its stigma, are also
shown in the fuchsia,
Fig. 15. The essential
organs of orchids are cu-
riously disguised. They
are combined into a sin-
gle body. In the lady-
slipper, Fig. 3, the lip-like
stigma is shown at P.
Upon either side, at its

STAMINATE AND PISTILLATE FLOWERS. 259

base, is an anther $. Projecting over the stigma
is a greenish ladle-like body, T, which is a trans-
formed and sterile anther. In all lady-slippers,
these organs are essentially the same as in the
drawing, although they vary much in size and
shape; but in most other orchids, the two side
anthers, S, are wholly wanting, and the terminal
organ, IT, is a pollen-bearing anther. In numer-
ous plants, there are many distinct pistils in each
flower. Such is the case in the strawberry, where
each little yellow ‘‘seed” on the ripened berry
represents a pistil; and the blackberry and the
raspberry, where each little grain or drupelet of
the fruit stands for the same organ. <A flowering
raspberry is illustrated natural size in Fig. 7, for
the purpose of showing the ring of many anthers
near the centre_of the flower, inside of which, in
the very centre, is a little head of pistils.

It frequently occurs that the stamens and pistils
are borne in different flowers, rather than together
in the same flower as they are in the examples
which we have studied. In these cases the flower
is said to be staminate, or male or sterile, in one
case, and pistillate, female or fertile, in the other
case. If these two kinds of flowers are borne
together upon the same plant, as in pumpkins,
melons, cucumbers, chestnuts, oaks, and begonias,
the plant is said to be monecious; but if the stami-
nate and pistillate flowers are on entirely different

260 POLLINATION.

plants, as in willows and poplars, the plant is dea-
cious. The two kinds of squash flowers are
shown in Fig. 8. The pistillate flower is on
the left, and it is at once distinguished by the
ovary or little squash below the colored portion,

Fic. 7. — Blossom of flowering raspberry (Rubus odoratus).

or corolla of the flower. The lobed stigma is
seen in the centre. The staminate flower is on
the right. It has a longer stem, no ovary, and the
anthers are united into a conspicuous cone in the
centre. The flowers expand early in the morning.
Insects carry pollen to the pistillate flower, which

STAMINATE AND PISTILLATE FLOWERS. 261

then begins to set its fruit, whilst the staminate
flower dies. The flowers of the common wild
clematis are shown in Fig. 9. Upon the right

Fic. 8.— Squash flowers of each sex.

are the sterile flowers, which are wholly stami-
nate. On the left, the flowers with larger sepals
—the petals are absent — have a cone of pistils in

262 POLLINATION.

the centre, and a few short and sterile stamens
spreading from the base of the cone. These dif-
ferent flowers are borne on different plants in this
species of clematis, and the plants are therefore
practically dicecious, because the stamens of the
pistillate flowers generally bear no pollen. A sim-
ilar mixed arrangement occurs in some strawber-

Fic. 9. — Flowers of clematis (Clematis Virginiana).

ries, except that there are no purely staminate
flowers. There are purely pistillate varieties,
others, like the Crescent, with a few nearly or
quite abortive stamens at the base of the cone of
pistils, and others in which the flowers are per-
fect or hermaphrodite, that is, containing the two
sexes.

Ee

COMPOSITOUS FLOWERS. 2638

The compositous flowers — like the asters, daisies,
goldenrods, sunflowers, dahlias, zinnias, chrysan-
themums, and their kin—need to be considered
in still a different category. In these plants, the
head, or so-called flower, is an aggregation of sev-
eral or many small flowers or florets. Each seed
in a sunflower head, for example, represents a dis-
tinct flower. Sometimes all of these flowers are
perfect, — contain the two sexes, — and sometimes
they are pistillate or staminate in different parts
of the head; and in some cases the plants are
dicecious. In many plants of the composite fam-
ily, the flowers near the border of the head are
unlike those of the centre or disc, in having a
long ray-lke corolla; and these ray-flowers are
frequently of different form from the others in the
character of the essential organs. Very frequently
the ray-flowers are pistillate, whilst the disc-flow-
ers are generally hermaphrodite. The anthers, in
these plants, are united in a ring closely about the
style and below the stigma.

The ovary, as we have seen, ripens into the
pod, berry, or other fruit; but it is not able to
bear seeds until it is assisted by the pollen. The
pollen falls upon the roughish or sticky surface
of the stigma, and there germinates or sends a
minute tube downwards through the style and
finally reaches the ovule, which, when fertilized,
rapidly ripens into the seed. ‘The nature of this

964 POLLINATION.

fecundation is not germane to the present subject;
but it may be said that only one pollen grain is
necessary to the fertilization of a single ovule, but
the addition of a superabundance of pollen greatly
stimulates the growth of the fleshy or enveloping
parts of the fruit. It is important that the person
who desires to cross plants should become familar
with the stigma when it is “ripe,” receptive, or
ready to receive the pollen. This condition is gen-
erally indicated by the glutinous or sticky or moist
condition of the stigma, or in those stigmas which
are not glutinous it is told by the appearing of-a
distinctly roughened or papillose condition. ‘This
receptive condition generally occurs about as soon
as the flower opens. If pollen is withheld, the
stigma will remain receptive much longer than
when fertilization has taken place, —in some flow-
ers for two or three days.

The pollen is discharged from the anther in
various ways, but it most commonly escapes
through a chink or crack in the side of the
anther. Sometimes it escapes through pores at
one end of the anther; and in other cases there
are more elaborate mechanisms to admit of its dis-
charge. In most plants, the anthers and stigma
in the same flower mature at different times, so
that close-fertilization or in-breeding is avoided.
This is well illustrated in the bell-flower, Fig. 1.
Here the anthers wither and die before the stig-

PREPARING THE FLOWERS. 265

matic lobes open. In other cases, the stigma
matures first, although this is not the usual con-
dition.

IJ. MANIPULATING THE FLOWERS.

We are now familiar with the essential principles
in the pollination of flowers. Before a person pro-
ceeds to operate upon a flower with which he is
unfamiliar, he should carefully study its structure,
so as to be able to locate the different organs, and
to discover when the pollen and the stigma are
ready for the work.

The first and last rule in the pollinating of plants
is this: Hvzercise every precaution to prevent any
other pollination than that which you design to give.
The anthers, therefore, must be removed from the
flower before it opens. ‘This removal of the anthers
is known as emasculation. Just as soon as this
is done, tie up the flower securely in a bag to
protect it from foreign pollen which may be
brought by wind or insects. As soon as the
stigma is ripe, remove the bag and apply the de-
sired pollen, placing the bag on the flower again,
where it must remain until the seeds begin to
form. The stigma may be receptive the day fol-
lowing emasculation, or, perhaps, not until a week
afterwards. Much depends upon the age of the
bud when emasculation takes place. It is gener-

266 POLLINATION,

ally best to delay emasculation as long as possible
and not have the flower open; but the operator
must be sure that the anthers do not discharge or
that insects do not get into the flower before he
has emasculated it. The bud at B, in Fig. 3, 1s

Fic. 10.— Tobacco flowers, showing the parts of the flower, a bud
ready to be emasculated, and an emasculated subject.

nearly ready to emasculate. The older buds on
the top of the spike of bugbane, Fig. 6, are ready
to operate upon; and so is the bud seen at the
left-in Fig. 7.

The manner of emasculating the flower varies

EMASCULATING. 267

with the operator. It is a common practice to
clip off the anthers with a pair of small scissors,
or to hook them out with a bent pin or a crochet
hook. Others use tweezers. For myself, how-
ever, I do not lke any of these methods, because
the anthers are apt to drop into the bottom of the
corolla, where it is sometimes difficult to rescue
them; and if one uses tweezers, there is always
danger that the anthers may be crushed and that
some of the pollen may adhere to the instrument
and contaminate future crosses. I therefore usu-
ally cut the corolla completely off just above the
ovary, with a pair of small, long-handled surgeon’s
scissors (see Fig. 12), removing everything but
the pistil. The operation is explained in Fig. 10,
which shows the tobacco flower. The flower at
the left shows the pin-head stigma in the centre of
the throat, and the five anthers surrounding it.
The second flower is spread open for the purpose
of showing these organs. The third figure is a
bud in the right condition for operation. The
right-hand figure shows this bud cut around with
the points of the scissors, leaving only the pistil.
The line at W, in Fig. 2, shows where the flower
of the lily might be cut off. The manner of oper-
ating upon a compositous flower is shown in the
picture of the zinnia, Fig. 11. In this plant the
outer florets of the head are pistillate, whilst those
of the disc are perfect. It is only necessary,

268 POLLINATION.

flowers before any of them open, and to cover the

E
therefore, to remove the central stamen-bearing
flower up before any of the pistils near the border

’

Fia. 11.— Zinnia tlowers; the upper head ready for emasculation,
the lower one showing the operation performed.

have protruded themselves. The upper head in
Fig: 11 shows the untreated sample, whilst the
lower one shows the same with the cone of central

EMASCULATING. 269

flowers pulled out. This treated head should now
be covered, to await the maturing of the stigmas.
In many compositous plants, however, the case is
not so simple as this, because all the flowers are
perfect. In such cases, nearly all the florets should
be removed from the head, and a few remaining
ones emasculated in essentially the same manner
as described for the tobacco, Fig. 10. Whenever
flowers are borne in clusters, nearly all of them
should be removed and the attention confined to
only two or three of them. One is then more cer-
tain of getting seeds to set. In some cases, like
the apple cluster, only one or two flowers of any
cluster ever set fruit, and the operator should then
choose the two or three strongest and most prom-
ising buds, and cut all the others off.

Flowers which bear no stamens, as the pistillate
flowers of squashes, strawberries, and many other
plants, of course do not require emasculating.
They should be tied up while in bud, however, to
prevent the access of any foreign pollen. Indian
corn is a case in point. ‘The pistillate flowers are
on the ear, each kernel of corn representing a single
flower. The silks are the stigmas. If it is desired
to cross corn, therefore, the ear should be covered
before any silks are protruded, and the pollen should
be applied some days later, when the silks are full
grown. ‘The staminate or male flowers are in the
tassel.

270 POLLINATION.

The pollen should be derived from a flower

which has also been protected fr

Fic. 12.—Instruments used in pollinating flow

Pin scalpel, scissors, lens,

om wind and in-

ers, natural size,

= eee

at a ne i fo 5 to. 9

a iy *

APPLYING THE POLLEN. EL

sects, because foreign pollen may have been
dropped upon an anther by an insect visitor and it
may be unknowingly transferred by the operator.
The pollen-bearing parent needs no operation, of
course, but the flower should have been tied up
in a bag when it was in bud. ‘The pollen is best
obtained by picking off a ripe anther and crush-
ing it upon the thumb-nail. ‘Then it is trans-
ferred to the stigma by a tiny scalpel made by
hammering out the small end of a pin, as shown,
full size, at the left in Fig. 12. The stigma
should be entirely covered with the pollen, if pos-
sible. It is often advised to use a camel’s hair

Fic. 13. — Ladle for pollinating house tomatoes.

brush to transfer the pollen, but much of the
pollen sticks amongst the hairs of the brush and
is ready to contaminate a future cross; and where
the pollen is scarce it cannot be conserved to
advantage by a brush. In some cases the pollen
is discharged so freely that the anther may be
rubbed upon the stigma, or even shaken over it,
but in most instances it will be necessary to actu-
ally place the pollen upon the stigma with some
hard instrument. When pollinating house-grown
melons and cucumbers, the staminate flower is
broken off, the corolla stripped back, and the

Te, POLLINATION.

anther-cone inserted into the pistillate flower,
where it is allowed to remain until it dries and
falls away. In pollinating house tomatoes, an
implement shown in Fig. 13, one-third size, is
used. This is simply a watch-glass, T, secured
to a handle. When the house
is dry, at midday, the watch-
glass is held under the flowers,
which are tapped, and the pol-
len falls into the glass. The
glass is then held up under
another flower until the stigma
rests in the pollen. It should
be said, however, that this pol-
lination of tomatoes is for the
purpose of making the fruit
set in the absence of insects,
not to effect a cross: It the
latter purpose were the object
sought, the flowers which are
to bear the seeds would need
to be emasculated.

Sometimes it is impossible to secure the pollen
at the time the stigma is ready. In some cases
of this kind, the intended parents can be grown
under glass so as to bring them into bloom at the
same time. In other cases, it is necessary to keep
the pollen for some time. The length of time
that pollen will keep varies with the species and

Fic. 14. — Bag for cov-
ering the flowers.

KEEPING THE POLLEN. Bie

probably also with the strength and vigor of the
plant which bears it. As a rule, it will not keep
more than a week or two, and, in general, it may

perme et —

Fic. 15.— Fuchsias, showing the stamens and pistils, and a bud
ready to be emasculated.

be said that the fresher it is the better it may be
expected to act. It is best kept in dry and tight

T

274 POLLINATION.

paper bags, such as are used for covering the
flowers. .

Something more should be said about the bags
which are used for covering the flowers. After
having tried every kind which is recommended,
I tind grocer’s manilla bags much the most satis-
factory. For most flowers the four-ounce size is
the handiest. When the bags are still flat, as

Fie. 16.— Fuchsia flower emasculated.

they come from the packages, a hole is made
through the two overlapping folds near the open-
ing, and a string is passed through it and then
tied at one of the folds, as shown in Fig. 14.
The bag is then ready for use. Before it is put
on the flower, the lower end of it is dipped in
water to soften it so that it can be puckered
tightly about the stem and thereby prevent the

ee ee

BAGGING THE FLOWER. 975

entrance of any in-
sect. A bag is put
upon the seed-bear-
ing flower when
emasculation is per-
formed, and upon
the intended pol-
len parent when
the flower is still
in bud. The bag
may be removed
from the emascu-
lated flower from
time to time to ex-
amine the stigma,
and again when
the pollen is ap-
pled; butit should
not be taken off
permanently until
the pod or fruit
begins to grow.
By way of re-
capitulation, let us
consider the cross- ae
ing of a fuchsia Fre. 17.—Fuchsia flower tied up aiter
flower. In Fig. 15 emasculation.
two flowers are shown in full bloom, with the long
style and the eight shorter stamens. ‘The single

276 POLLINATION.

bud is just the right age to emasculate. We
therefore cut off the two flowers and emasculate
the bud, as in Fig. 16. The pollen of another
flower is applied and the bag is tied on, as seen in
Fig. 17. The best label is a small merchandise
tag, and this records the staminate parent and
the date.

It will be seen that in the operation of emascu-
lating the fuchsia flower we cut off the sepals as

Fic. 18. — Tomato and quince, showing how the sepals were cut off
in emasculating.

well as the petals. In some plants the calyx
adheres to the full-grown fruit, as on the apple,
pear, quince, gooseberry, or persists at the base
of the fruit, as in the tomato, pea, raspberry. In
these fruits, therefore, the cutting away of the
calyx leaves an indelible mark which at once dis-
tinguishes the fruits which have been crossed,

CROSSING IN FLOWERLESS PLANTS. et

even if the labels are lost. In Fig. 18 a tomato
and quince are shown which are thus marked.

All the foregoing remarks do not apply to the
crossing of ferns, lycopods, and the like, because
these plants have no flowers; yet cross-fertiliza-
tion may take place in them. When the spores

U it
\
«

)

4

i LM
Hie
a

LLL. AISA

PNA

Fic. 19. — Pollinating kit.

of these flowerless plants are sown, a thin green
tissue, or prothallus, appears and spreads over the
ground. In this tissue the separate sex-organs
appear, and after fecundation takes place, the
fern, aS we commonly understand it, springs forth.
Thereafter, this fern lives an asexual life and

278 POLLINATION.

produces spores year after year; but it is only in
this primitive prothallic stage that fertilization
takes place, once in the lifetime of the plant. If
these plants are to be crossed, the only procedure
open to the gardener is to sow the spores of the
intended parents together in the hope that a nat-
ural mixing may take place. There are various
well-authenticated fern hybrids.

The pollination of flowers is such a simple work
that few implements are required for its easy
performance. Great care is more important than

= SSS

TERN,

Fig. 20. — Pollinating kit.

any number of tools. Every one who expects to —
cross plants should provide himself with the three
instruments shown in Fig. 12,—a pin scalpel,
sharp-pointed scissors, and a large hand-lens. If
one contemplates much experimenting in this
direction, however, it is economy of time to have
some sort of a box in which there are compart-
ments for the various necessities. ‘These various
compartments suggest at once whatever accesso-
ries are wanting, and they hold a sufficient supply

IMPLEMENTS USED IN CROSSING. 279

for several hundred operations. There should
be a compartment for bags, string, lens, scissors,
and pencils, tags, note-book, and the like. Figs.
19 and 20 show a convenient case for an experi-
menter, and one which I have used with satisfac-
tion for several years. This kit is twelve inches
long, nine inches wide, and three inches deep.

The chances of success in pollinating are dis-
cussed in Lecture II. (page 83).

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GLOSSARY.

1. THe FLOWER.

Anther.— That portion of the stamen which bears the
pollen. It is the uppermost portion of the stamen.

Calyx.— The outer series of floral envelopes, usually
green. The various separate parts of the calyx are
sepals.

Corolla. — The inner series of floral envelopes, usually
colored and forming the showy part of the flower. If
it is divided into separate parts, these are called petals.

Essential organs. — The stamens and pistils.

Female. — Said of flowers which have only pistils or the
seed-bearing part, or of plants which bear only such
flowers ; applied also to the pistils in any flower.

Filament. — The stalk or stem of a stamen, bearing the
anther.

Floral envelopes. —The calyx and corolla.

Male. — Said of flowers which bear only stamens, or of
plants which have only staminate flowers; also applied
to the stamens or pollen-bearing organs of flowers.

Ovary. — The lowest part of the pistil, containing the
ovules. It is the most thickened portion of the pistil,
and it may stand either below or above the petals.
The ovary ripens into the fruit.

Ovule. — A body in the ovary which ripens into a seed.

Pet'-al.— The separate portions or leaves of the corolla.

Pistil. — The seed-bearing organ of the flower. It always
comprises two parts, the ovary —which becomes the
pod or fruit—and the stigma. Usually there is a

281

282 GLOSSARY.

style connecting the two. Often called the fertile or
female organ.

Pistillate. — Said of a plant or flower which has only pis-
tils or female organs.

Pollen. — The contents of the anther, capable of fertil-
izing the ovules. It is usually composed of minute
yellow or brown grains.

Se'-pal. — The separate portions or leaves of the calyx.

Spore. — The reproductive organ of flowerless plants, by
means of which they propagate, as other plants propa-
gate by means of the seed. The spore is asexual.

Stamen. — The pollen-bearing organ of the flower. Often
called the male or sterile organ. Its essential part is
the anther. The stalk, when present, is called the
filament.

Staminate. — Said of a flower or plant which bears only
stamens or male organs.

Stigma.— The top end of the pistil, where the pollen
lodges and germinates. It is usually a somewhat ex-
panded surface, and is roughened, or sticky, or moist
when ready to receive the pollen.

Style. —The more or less slender portion of the pistil
which lies between the stigma and ovary. The pol-
len-tubes pass through it in reaching the ovary.

2. CROSSING.

Bigener; bigeneric-hybrid. — A hybrid between species of
different genera.

Bigeneric half-breed.— The product of a cross between
varieties of species of different genera.

Close-fertilization ; self-fertilization. — The action of pollen
upon the pistil of the same flower.

Close-pollination ; self-pollination. — The transfer of pollen
to a pistil of the same flower.

Cross. — The offspring of any two flowers which have
been cross-fertilized.

GLOSSARY. 283

Cross-breed ; half-breed; mongrel ; variety-hybrid. — A cross
between varieties of the same species.

Cross-fertilization. — The action of pollen upon the pistil
of another flower of the same species.

Crossing. — The operation or practice of cross-pollinating.

Cross-pollination. —The conveyance of pollen to the
stigma of another flower.

Derivative- or derivation-hybrid; secondary-hybrid. — A hy-
brid between hybrids, or between a hybrid and one of
its parents.

Fertilization; fecundation ; impregnation. — The action of
the pollen upon the ovules.

Half-hybrid. — The product of a cross between a species
aud a variety of another species.

Hybrid.— The offspring of crossed plants of different
species.

Hybridism; hybridity. — The state, quality, or condition
of being a hybrid.

Hybridization. —The state or condition of being hybrid-
ized, or the process or act of hybridizing.

Hybridizing. — The operation or practice of crossing be-
tween species.

Individual cross. — The offspring of two crossed flowers
on the same plant.

Individual fertilization. — Fertilization between flowers
upon the same plant.

Mongrel. — A cross.

Mule. — A sterile (seedless) hybrid.

Pollination. — The conveyance of pollen from the anther
to the stigma (page 252).

The term cross is used to denote the offspring of any
sexual union between plants, whether of different
species or varieties, or even different flowers upon the
same plant. It is a general term. And the word is

284 GLOSSARY.

also sometimes used to denote the operation of per-
forming or bringing about the sexual union. There
are different kinds of crosses. One of these is the
hybrid. A hybrid is a cross between two species, as a
plum and a peach, or a raspberry and a blackberry.
There has lately been some objection urged against
this term, because it is often impossible to define the
limitations of species,— to tell where one species ends
and another begins. And it is a fact that this diffi-
culty exists, for plants which some botanists regard as
mere varieties others regard as distinct species. But
the term hybrid is no more inaccurate than the term
species, upon which it rests; and, so long as men talk
about species, so long have we an equal right to talk
about hybrids. Here, as everywhere, terms are mere
conveniences, and they seldom express the whole truth.
In common speech the word hybrid is much misused.
Crosses between varieties of one species are termed
half-breeds or cross-breeds, and those between different
flowers upon the same plant are called individual
Crosses.
3. CLASSIFICATION.

Break. — A radical departure from the type. Ordinarily
used in the sense of sport, but in its larger meaning it
refers to the permanent appearance of apparently new
or very pronounced characters in a species.

Bud-variation. — Variation or departure from a type
through the agency of buds (pages 28, 153).

Bud-variety. — A variety resulting from bud-variation.
Bud-sport.

Family (Order in botany.)— A group of genera and
species; as Cupulifere, the Oak Family, Rosacee, the

- Rose Family.

Form. — A minor variety, usually transient, produced by

some local environment.

GLOSSARY. 285

Genus (plural, genera). — A group or kind comprising a
greater or less number of closely related species; as
Acer, the maples, Fragaria, the strawberries.

Race.— A fixed cultural variety; that is, a cultural va-

riety which reproduces itself more or less uniformly
from seeds.

Seedling. — A plant growing directly from seed, without
the intervention of grafts, layers, or cuttings.

Seed-variation. — Variation or departure from a type
through the agency of seeds.

Seed-variety. — A variety resulting from seed-variation.

Species (plural, species).— An indefinite term applied to
all individuals of a certain kind which come or are
supposed to come from a common parentage. A per-
ennial succession of normal or natural similar indi-
viduals perpetuated by means of seedage. “All the
descendants from the same stock.” — Gray.

Sport.— A variety or variation which appears suddenly
and unaccountably, either from seeds or buds; more
properly restricted to varieties originating from buds,
and so used in this book.

Stock. — The parentage of a particular strain or variety.

Strain. — A sub-variety, or individuals of a variety, which
has been improved and bred under known conditions.

Variation.—1. The act or condition of varying or be-
coming modified. 2. A transient variety, more or less
incapable of being fixed or rendered permanent.

Variety. — A form or series of forms of a species marked
by characters of less permanence or less importance
than are the species themselves.

Wilding. — A wild individual from a cultivated species.

INDEX.

—_>e—_—_-
Abortive varieties, 152. Aralia, bud-variety, 180.
Abutilon, crosses, 220, 288, 250. Arthur, 103, 116.
Accident, 154. Arundo, variation, 176, 180.
Acclimatization, 24, 26. Asexual propagation, 7.
Acer, bud-variety, 177. Aspidistra, sport, 180.
Achimenes, crosses, 245, 246, 247. Aster, varieties, 180.
Acorns, bud-variety, 177. ‘ Atavism, 106.
Acquired characters, 14. Atragene, 184.
Adult forms, 156, Atropa, crosses, 223.
Aigilops, crosses, 248. Azalea, bud-varieties, 180.
fsculus, bud-variety, 178.
Aisculus, crosses, 239. Bag for covering flowers, 272.
Agatha, bud-variety, 178. Bamboos, variation, 176.
Ageratum, bud-varieties, 178. Banana, varieties, 175.
Agrostemma Ceeli-rosa, dwarfs, 144. Bartel, T. C., 130.
Albinos, 148. Barteldes, 140.
Allut, Cazalis, 211. Bean, bud-variation, 176.
Almond, bud-variety, 179. . Beans, types of, 135.
Alnus, crosses, 226..- Beet, crosses, 56.
Alopecurus, crosses, 230. Begonia, crosses, 224, 226, 227, 229,
Altitude and plants, 25. 230, 281, 288, -284, 239, 245, 246,
Amelioration, gradual, 50. 247.
Amygdalus, crosses, 239. Begonia pollinations, 86.
Anagallis, crosses, 222, 223, 231. Bell-flower, 252.
Anemone, crosses, 224, 229. Berberis, crosses, 245.
Anemone, varieties, 179. Bigness, variation in, 18.
Animal and plant contrasted, 5, 91. Blackberry, crosses, 79, 111.
Année, 141. Blackberry, introduction of, 129.
Antagonistic features, 95. Bletia, crosses, 222.
Anther, 254. Bohnhof, 80.
Apple, Wealthy, 108. Bornet, 232.
Apples, bud-variation in, 118, 175. Bouschet, Henri, 212.
Apples, hybrid, 66, 79, 111. Brassica, crosses, 223, 239, 240, 241.
Apples, races of, 90. Braun, Alexander, 1T.
Apples, variations in, 3, 27, 37, 99, 131. | Breaking the type, 19, 28, 93.
Apricot, bud-variety, 179. Bruant, 113.

Aquilegia, crosses, 224, 229, 234, 239, | Buckwheat, crosses, 56.
240, 250. Budd, Professor, 133.

287

288

Bud-variation, 6, 21, 28, 37, 101, 118,

126, 153.
Bugbane, 258.
Burpee, 139.
Buxus, bud-varieties, 181.

Cabbage, crosses, 56.
Cacti, crosses, 235.

Calceolaria, crosses, 222, 229, 283, 234,

239, 245, 247.
Calceolaria plantaginea, dwarfs, 144.
Calliopsis tinctoria, dwarfs, 144.
Callistephus hortensis, dwarfs, 145.
Calyx, 253.
Camellia, bud-varieties, 182.
Camellia, crosses, 251.
Canary-grass, crosses, 57.
Cannas, 140.
Capsella, crosses, 226, 231.
Carman, 79.
Carnation, 115.
Carriére, 96, 116, 153.
Caspary, 229.
Cedars, 156.
Cephalotaxus, sports, 183.
Cereus, bud-variety, 184.
Cereus, crosses, 226, 230, 238, 289.
Cereus, night-blooming, 256.
Change of seed, 28, 59, 116.
Checking growth, 116.
Cheiranthus, sport of, 185.
Cherry, hybrid, 112.
Cherry, sports of, 171.
Chloranthic varieties, 153.
Chlorosis, 149.
Choice of variations, 31.
Chrysanthemum carinatum, 100.
Chrysanthemum, sports of, 158.
Cimicifuga racemosa, 258.
Cinchona, crosses, 225.
Cirsium, crosses, 229, 288, 239.
Cistus, crosses, 219, 222, 231, 234.
Clematis, crosses, 224.
Clematis, flowers, 262.
Clematis, varieties, 184,
Climate and variation, 24, 114, 146.
Coleus, sports in, 120.
Coloration, 148.
Colors, modified by climate, 25.
Conifers, 156.

INDEX.

Contradictory attributes, 98.

Convolvulus pollinations, 85.

Coreopsis tinctoria, dwarfs, 144.

Cornus, bud-varieties, 185.

Corolla, 253.

Cotyledon, crosses, 229.

Crabs, hybrid, 66, 111.

Cratzegus, crosses, 239.

Crinum, crosses, 218, 225, 229, 232, 234.

Cross-breeds, Focke on, 247.

Cross, function of, 50.

Cross, primary, 215.

Crosses, characteristics of, 68.

Crosses, Focke on, 215.

Crossing a means, 107.

Crossing and change of seed, 59.

Crossing, limits of, 44.

Crossing, philosophy of, 39.

Crossing, rule for, 109.

Crozy, 113, 140.

Cucumber pollinations, 85.

Cucumis, crosses, 222.

Cucurbita Pepo, 75, 84.

Cucurbitacee, crosses, 46, 58, 74, 82
229, 230, 235.

Cultivation, philosophy of, 22.

Currant, sports of, 1738. (See Ribes.)

Cypripedium, 254.

Cypripedium, crosses, 218, 227, 229,
246.

Cytisus Adami, 185.

Dactylis, bud-variety, 185.

Darwin, 17, 23, 32, 42, 47, 51, 54, 56, 60,
68, 69, 72, 84, 87, 117, 119, 121, 176,
228.

Dating back, 106.

Datura, crosses, 222, 223, 224, 226, 228,
234, 239, 240, 249, 250.

Decaisne, 229.

De Candolle, 150, 178.

Derivative crosses, 238.

Dewberry crosses, 79, 111.

Dewberry, introduction of, 129.

Dianthus Chinensis, dwarfs, 145.

Dianthus, crosses, 218, 219, 228, 226,
227, 299, 238, 234, 237, 239, 240, 2438,
244, 245, 246, 250.

Dianthus semperflorens, 156.

Dichroism, 154.

INDEX.

289

Digitalis, crosses, 218, 219, 221, 223, | Fuchsia flowers, 273, 274, 275.

226, 227, 229, 280, 232, 237.
Dimorphism, 154.
Diccious plants, 260.
Divergence of character, 28.
Division of labor, 42.
Doubleness in hybrids, 237.
Doubleness of flowers, 149.
Dracana, variation, 176.
Duval, M., 167.
Dwarfing, 25, 114, 148.

Early varieties, 146.

Echinocactus, sports, 186. -

Eckford, 118.

Egg-plant, crosses, 57, 74.

Egg-plant pollinations, 85.

Egg-plants, variation in, 95.

Egypt, plagues of, 40.

Eleagnus, bud-varieties, 186.

Emasculation, 265.

Envelopes, floral, 252.

Environment and variation, 12.

Epilobium, crosses, 218, 222, 229, 230,
233.

Equilibrium of organisms, 20, 61.

Erica, crosses, 229, 233, 239, 240, 246,
247,

Essential organs, 255.

Euonymus Japonicus, 156, 186.

Fagus, fern-leaved, 187.

Fall sowing, 115, 148.

Ferns, crossing, 277.

Fertility of soil, 18, 22.

Ficus, forms of, 188.

Filament, 254.

Fittest, survival of, 32, 39.
Fixation of plants, 31.

Flavor, modified by climate, 25.
Flon, M., 156.

Flowerless plants, crossing, 277.
Focke, 68, 81, 108, 215.
Fontanesia, sport, 188,

Food supply, 16, 116.
Fortuitous variation, 9.
Fragaria, crosses, 229,
Fraxinus, bud-varieties, 188.
Fromont, 158.

Fuchsia, crosses, 229, 238, 239, 247,

U

Function of the Cross, 50.
Fusain, 156.

Galium, crosses, 222.

Gardenia, bud-variety, 189.

Gartner, 216, 218, 219, 220, 228, 2383,
235, 239, 240, 243, 244, 248, 249, 250.

Gazania rigens, 146.

Genera, monotypic, 97.

Gesneracee, crosses, 227, 229, 246.

Geum, crosses, 219, 229, 239.

Giant forms, 145.

Gibb, Charles, 133.

Gideon, Peter M., 108.

Gillyflower, bud-variety, 189.

Giraud, Désiré, 166.

Gladiolus, crosses, 226, 229, 234, 245,
246.

Gleditschia triacanthos, 207.

Glossary, 282.

Goff, 103.

Gordon, 229, 241, 244,

Gourd, crosses, 58, 74, 82.

Grape, bud-varieties, 174, 210.

Grapes, hybrid, 66, 78, 110, 111.

Gray, Asa, 33, 178.

Greenhouses, produce variation, 115.

Hallock, V. H., & Son, 124.

Hardy varieties, 145.

Hartogia Capensis, 192.

Hedera, forms of, 189.

Helianthemum, crosses, 223, 226.

Helichrysum bracteatum, dwarfs, 144.

Henderson, 138.

Herbert, 248.

Hibiscus, bud-varieties, 190.

Hibiscus, crosses, 225, 250,

Hibiscus Syriacus, 257.

Hieracium, crosses, 220, 238, 234, 237,
239, 250.

Hippeastrum, crosses, 226, 229, 234,
239, 245, 246, 247.

Holly, sports, 191.

Horse-chestnut, bud-variety, 178.

Husk-tomato, 60, 85.

Hyacinth, forms, 190.

Hybrids, characters of, 68, 215.

Hybrids, Focke on, 215.

290

Hybrids, multiple, 246, 247.
Hybrids, rarity of, 53.
Hybrids, seven-eighths, 243,
Hybrids, three-fourths, 243.
Hybrids, triple, 244.
Hydrangea, 146, 191.
Hymenocallis, crosses, 280.

Iberis umbellata, dwarfs, 145.
Ignotum tomato, 123.

Ilex, bud-varieties, 191.
Impatiens Balsamina, dwarfs, 145.
In-breeding, 72.

Indeterminate varieties, 87.
Individuality, causes of, 8.
Individuality, fact of, 2.
Instruments for pollination, 270.
Ipomeeas, colors of seeds, 104,
Iris, bud-variety, 192.

Isolation of the plant, 22.

INDEX.

Leafiness, 25,

Lecogq, 227.

Lemoine, 113.

Lens for pollinating, 270.

Leptosiphon densiflorus, dwarfs, 144.

Lettuce, crosses, 56.

Ligustrum, sports, 193.

Lilac, bud-varieties, 193.

Lily, white, 253.

Lima beans, 188.

Limits of crossing, 44.

Linaria, crosses, 222, 227, 238, 289,
241,

Lindley, 68.

Links, missing, 41, 48.

Linneus, 81, 152.

Linum, crosses, 223.

Lobelia, crosses, 219, 222, 282, 238,
234, 235, 245,

Luffa, crosses, 232,

Isoloma, crosses, 226, 227, 231, 233, | Lupines, heredity in, 106.

246.
Ivy, forms of, 189.

Jamain, M., 163.

Jobert, M., 179.

Joigneaux, M., 204.
Juniperus, bud-varieties, 192,

Klotzsch, 226, 229.

Knight, Thomas Andrew, 17, 54, 227,
229.

Kohl-rabi, 80.

Kdlreuter, 54, 73, 216, 217, 219, 228,
229, 248, 244, 250.

Kumerle, W. J., 140.

Kuntze, 225.

Labor, division of, 42, 48.
Lachaume, M.,-166.

Lactuca, crosses, 238.

Ladle for pollinating, 271.
Lamium, bud-variety, 192.
Lamium, crosses, 289, 241.
Lantana, crosses, 222.
Large-flowered varieties, 145.
Late varieties, 146.

Latitude and plants, 25.
Laurocerasus, sports, 192.
Layatera, crosses, 234, 239, 243.

Lychnis Ceeli-rosa, dwarfs, 144,
Lychnis, crosses, 240, 241.
Lycium, crosses, 218, 226,
Lycopods, crossing, 277.

Maize, crosses, 56.

Malle, Dureau de la, 175.

Mamillaria, sports, 194,

Maple, Wier’s, 109.

Meadow, plants in, 28.

Medicago, crosses, 220, 233, 240.
Melandrium, crosses, 220, 222, 228,
225, 231, 238, 240, 244, 245.

Mendel, 239.

Mentha, bud-variety, 194.

Mentha, crosscs, 229, 241,

Mersereau, 131.

Mimulus, crosses, 222, 251.

Mirabilis, crosses, 222, 226, 227, 228,
234, 241, 250, 251.

| Mirabilis pollinations, 85,

Missing links, 41.

Mixing in the hill, 118, 201.
Molinia, bud-variety, 194.
Moncecious plants, 259.
Monotypie genera, 97.
Moore, Jacob, 110.
Morning-glory, 54.

Morong, Dr. Thomas, 60.

INDEX.

Morren, 149.

Mourriére, M., 175.
Mulberry, Teas’, 109.
Multiple hybrids, 246, 247.
Munson, Professor, 58.
Monson, TV, (9, 2i1,
Musa, bud-variety, 194,
Muskmelon pollination, 85,
Myrtle, bud-variety, 195,

Nanz & Neuner, 170.

Narcissus, crosses, 220, 225, 236.

Natal variations, 15.

Natural selection, 32, 51.

Naudin, 216, 229, 248.

Nectarine, origin of, 118, 178.

Nepenthes, crosses, 220.

Nicotiana, crosses, 72, 217, 219, 222,
224, 225, 226, 227, 229, 233, 234, 237,
238, 239, 241, 248, 244, 245, 246, 250.

Nicotiana pollinations, 85, 86.

Nuphar, crosses, 226,

Nymphea, crosses, 218, 225, 227, 231,
234, 250. ;

Odoriferous varieties, 147.
(nothera, crosses, 219, 239,
(nothera Drummondii, dwarfs, 144.
Oger, Pierre, 166.

Olea ilicifolia, 195.

Opuntia, bud-variety, 195,
Orange, bud-variety, 195.
Orchide, crosses, 229, 233, 235.
Orchids, hybrids, 79.
Orontium, sport, 195.
Osmanthus, sport, 195,

Ovary, 255, 263.

Palmer, Asa, 139.

Pansy, 146.

Papaver, crosses, 218, 224, 226, 229,
231, 237.

Papayer, forms of, 151.

Paré, M., 160.

Parents, influence of, 81, 217.

Passiflora, crosses, 220, 226, 229, 235,
250.

Peach, bud-variation in, 118, 178, 196.

Peach, hybrids, 47.

Peaches, races of, 91.

291

Pear, bud-varieties, 174, 197,

Pears, hybrid, 66, 79, 111.

Pears, variation in, 99.

Peas, viney, 16.

Pelargonium, crosses, 218, 220, 229,
280, 233, 234, 237, 245, 246, 247.

Pelargonium, sports in, 198.

Peloric varieties, 152.

Pentstemon, crosses, 241,

Pepino pollinations, 86,

Pepino, variation in, 95.

Pepper, red, pollination, 85.

Peppers, variation in, 96,

Persica, 196,

Petal, 253.

Petunia, crosses, 218, 240, 241, 251.

Petunia pollinations, 85, 86.

Phalaris, sports, 198.

Phaseolus, crosses, 228, 288, 241,

Phlox, bud-varieties, 199.

Phragmites, bud-varieties, 199.

Physalis, 60,

Physalis, variation in, 96.

Picea, bud-variety, 199.

Pink, 156, 160.

Pinus, bud-varieties, 199.

Pinus, crosses, 226,

Pistil, 255.

Pisum, crosses, 223, 238.

Pittosporum, sport, 200.

Plant-breeding, 91.

Pliny, 131,

Plum, hybrids, 47, 112.

Plum, sports of, 172.

Plums, Japanese, 27.

Podocarpus, 155, 183.

Pollen, 254, 264.

Pollinating kit, 277, 278.

Pollination, 252.

Pollination, uncertainties of, 83.

Polymorphous varieties, 153.

Polytypic genera, 97.

Populus, bud-variety, 200.

Populus, crosses, 222.

Position, advantage of, 22.

Post-natal variations, 15.

Potamogeton, crosses, 227.

Potato, 37, 117.

Potato and tomato, 95.

Potato, bud-varieties, 201, 209.

292

Potato, seedlessness, 99.
Precocious varieties, 146.
Primula, crosses, 2389, 240, 241.
Progeny of crosses, 237.
Proliferous varieties, 150.
Propagation, asexual, 7.
Pruning, 23.

Prunus, bud-variety, 205.
Prunus, crosses, 229, 239.
Pumpkin, crossing, 46, 58, 74, 82.
Pyrus, crosses, 229, 239.

Quercus, crosses, 226, 229, 289.
Quince, pollinated, 276.

Races in fruits, 90.

Radish pollinations, 85.

Raphanus Raphanistrum, 116, 231.

Raphanus sativus, 231.

Raspberry, flowering, 260.

Raspberry, hybrids, 79, 111.

Representative species, 66.

Retinosporas, 156.

Rheum, bud-variety, 206.

Rhododendron, crosses, 145, 218, 222,
225, 226, 229, 230, 239, 245, 246, 247,

Ribes, bud-varieties, 206.

Ribes, crosses, 230, 233.

Robinia, bud-varieties, 206.

Rogue, 89, 127.

Rosa, 161.

Rose, bud-varieties in, 118, 161, 207.

Roses, crosses. 233, 247.

Rubus, crosses, 227, 230, 232, 234.

Rubus odoratus, 260.

Running out of varieties, 36, 125.

Russia, fruits from, 27, 90, 133.

Rye, hybrids, 79.

Salix, bud-variety, 208.

Salix, crosses, 219, 225, 226, 227, 229,
231, 234, 239, 244, 245, 246, 247.

Salter, 119.

Salvia, crosses, 223.

Sambucus, sports, 208.

Satyrium hircinum, 148.

Scabiosa atropurpurea, dwarfs, 145.

Scalpel for pollinating, 270.

Schizanthus retusus, dwarfs, 145.

Scissors for pollinating, 270.

INDEX.

Secondary crosses, 288.

Seed, change of, 28, 59.

Seeds, colors of, 104.

Seeds, early, 147.

Seeds, immature, 103, 147.
Seeds, large and small, 101.
Selection and progress, 120, 122, 127.
Selection, natural, 32, 51.
Self-fertilization, effects of, 54.
Senecio cruentus, dwarfs, 144.
Sepal, 253.

Seven-eighths hybrids, 248.
Sex and variation, 11, 48.
Silene, crosses, 234.

Sinningia, crosses, 231.
Solanum, bud-varieties, 209.
Solanum, variations in, 95.
Spencer, 61.

Spirea, bud-varieties, 209.
Sports, 22, 28, 37, 153.
Sprengel, 54.

Squash, crosses, 58, 74, 82.
Squash flowers, 261.

Squash, Hubbard, 46.
Stamens, 254.

Stigma, 255.

Strawberry, Wilson, 125.
Struggle for life, 20, 29, 39.
Sturtevant, 103.

Style, 255.

Sugar-cane, varieties, 175.
Survival of the fittest, 32, 39.
Swamping effects of inter-crossing, 46.
Sweet, 247.

Symphoricarpus, sport. 209.
Symphytum, bud-varieties, 209.
Synchronistic variations, 117.

Tagetes, dwarfs, 145.

Teas, 109.

Teleology of hybrids, 236.
Thinning, 28.
Three-fourths hybrids, 243.
Thuyopsis, sport, 209.
Tillage and food supply, 17, 22.
Timbal-Lagrave, 217.
Toad-flax, 152.

Tobacco flowers, 266.
Tobacco pollinations, 86.
Tomato and potato, 95.

INDEX.

Tomato, crosses, 58.

Tomato, Ignotum, 123.
Tomato, pollinated, 276.
Tomato pollinations, 85.
Tomato, Trophy, 37.
Tomato, variation in, 98.
Tomatoes, breeding, 103.
Tragopogon, crosses, 288.
Triple hybrids, 244.
Tropxolum, crosses, 222, 226.

Ulmus, bud-variety, 209.
Ulmus, crosses, 226.

Variability, variation in, 25.

Variation and environment, 12.

Variation caused by sex, 11, 48.

Variation, fortuitous, 9.

Variation, philosophy of, 1.

Variations, choice of, 31.

Variations, fixation of, 31.

Variations, natal and post-natal, 15.

Variations, origin of, 8, 41. [157.

Variegation, perpetuating, 120, 149,

Varieties, running out, 36, 125.

Variety, what is a, 35. -

Verbascum, crosses, 219, 221, 222, 228,
224, 226, 229, 285, 239, 250.

293

Verdier, Victor, 167.

Verlot, 121, 148.

Veronica, crosses, 233, 239.

Viburnum, sports, 209.

Vilmorin, 152.

Vilmorin, Henri
142.

Vilmorin, Louis Levyéque de, 106.

Vine, bud-varieties, 174, 210.

Viola, bud-variety, 210.

Viola, crosses, 229, 231.

Vitis, crosses, 229, 239, 245, 246.

L. de, 100, 108,

Walker, Ernest, 120, 169.
Wallace, 47, 60, 67.
Watermelon pollination, 85.
Weismann, 13, 14.

Wheat, hybrids, 79.
Wichura, 216, 246.

Wier, D. B., 109.
Wigandia, sport, 215.
Willdenow, 152.

Yucca, variation, 176.

Zinnia, crosses, 241.
Zinnia, flowers, 268,

NEW EDITION.

The Horticulturist’s Rule-Book.

A COMPENDIUM OF USEFUL INFORMATION FOR FRUIT-
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