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U. S. DEPARTMENT OF AGRICULTURE.
BUREAU OF PLANT INDUSTRY— BULLETIN NO. 243.
B. T. GALLOWAY, Chief of Bureau.
HETEROZYGOSIS IN EVOLUTION AND
IN PLANT BREEDING.
E. M. EAST,
Assistant Professsor of Experimental Plant Morphology, Harvard University,
and Collaborator of the Bureau of Plant Industry,
ASSISTED BY
H. K. HAYES,
Plant Breeder of the Connecticut Agricultural Experiment Station.
[In Cooperation with the Connecticut Agricultural Experiment Station and Harvard University.]
Issued June 5, 1912.
WASHINGTON:
GOVERNMENT PRINTING OFFICE.
1912.
BUREAU OF PLANT INDUSTRY.
Chief of Bureau, Beverly T. Galloway.
Assistant Chief of Bureau, William A. Taylor.
Editor, J. E. Rockwell.
Chief Cleric, James E. Jones.
Tobacco Investigations.
scientific staff.
W. W. Garner, Physiologist in Charge.
E. H. Mathewson and G. W. Harris, Crop Technologists.
H. A. Allard,.C. W. Bacon, E. G. Beinhart, JJ. E. Brown, C. L. Foubert, W. M. Lunn, E. G. Moss, and
Otto Olson, Assistants.
J. S. Cuningham and B. F. Seherffius, Experts.
J. E= Blohm, Special Agent
B. G. Anderson, R P. Cocke, EL M. East, W. W. Green, E. K. Hibshman, and True Houser, Collaborators.
243
2
LETTER OF TRANSMITTAL.
U. S. Department of Agriculture,
Bureau of Plant Industry,
Office of the Chief,
Washington, D. C, January 20, 1912.
Sir: I have the honor to transmit herewith and to recommend for
publication as Bulletin No. 243 of the series of this Bureau a manu-
script entitled " Heterozygosis in Evolution and in Plant Breeding,"
by Dr. E. M. East, Assistant Professor of Experimental Plant Mor-
phology, Harvard University, and Collaborator of this Bureau, and
Mr. H. K. Hayes, Plant Breeder of the Connecticut Agricultural
Experiment Station. This paper reports results from experiments
that have at different times received aid from this Bureau, the Con-
necticut Agricultural Experiment Station, and the Bussey Institu-
tion of Harvard University and should be considered the product of
their joint collaboration.
Respectfully, B. T. Galloway,
Chief of Bureau.
Hon. James Wilson,
Secretary of Agriculture.
243
CONTENTS
Page.
Introduction . 7
The problem 8
Early investigations 8
The work of Darwin 13
Recent investigations 17
Experiments on a normally cross-fertilized species, Zea mays 19
Effects of inbreeding 19
Crossing inbred types 24
Experiments on species generally self-fertilized 26
The characters affected by heterozygosis 31
Theoretical interpretation of results. 32
Extension of the conclusions to the animal kingdom 39
Value of heterozygosis in evolution 43
Value of heterozygosis in plant breeding 46
Maize. 46
Truck crops 47
Plants reproduced asexually 48
Forestry 48
Bibliography 49
Index 53
243 5
LLUSTRATIONS.
Page.
Plate I. Tassels and ears of an almost sterile strain of corn isolated by inbreed-
ing 24
II. Watson's flint and Longfellow flint corn inbred two years with Fj
hybrid 24
III. Learning dent strains of corn, Xo. 9 and Xo. 12, after four years' in-
breeding, compared with Fx hybrid 26
IV. Inbred strains of Learning dent corn compared with F: and F2 genera-
tions 26
V. Strains 6 and 7 of Learning pure lines of corn and Fx generation of
crosses 26
VI. Fig. 1. — Xicotiana tabacum variety. Fig. 2. — Xicotiana tabacum
variety X X. silvestris, F1 generation. Fig. 3. — Xicotiana sil-
vestris 26
VII. Fig. 1. — Xicotiana rustica texana.- Fig. 2. — Xicotiana rustica tex-
ana X X. tabacum variety, Fx generation. Fig. 3. — Xicotiana
tabacum variety 28
VIII. Fig. 1. — Xicotiana alata grandiflora. Fig. 2. — Xicotiana tabacum
variety. Fig. 3. — Xicotiana alata grandiflora X X. tabacum 28
243
6
B. P. I.— 724.
HETEROZYGOSIS IN EVOLUTION AND IN
PLANT BREEDING.'
INTRODUCTION.
When a biologist begins any line of genetic work with either
plants or animals he generally has occasion to differentiate his stock
into more or less pure types by in-and-in breeding. Frequently in
the case of animals, and nearly always in the case of plants that are
naturally cross-fertilized, he finds there is a loss of vigor, usuaUy
unaccompanied by pathological symptoms. This loss of vigor is
generally expressed by a decrease in the size of the individual, but it
may be shown by a slight decrease in fertility. The phenomenon,
although it probably occurs in all great groups reproducing sexually,
is not general, however, for in many animals and in plants that are
normally self-fertilized it is unnoticeable.
If after obtaining his "pure" stocks the experimenter has occasion
to cross strains that differ in character, he often finds that the reverse
phenomenon occurs. The vigor of the hybrid is greater than that of
either parent.
These manifestations have been noticed for over a century by
plant breeders and for probably two thousand years or more by
animal hybridizers. Until the end of the nineteenth century the
interpretation of the phenomena, if, indeed, that which is only a
paraphrased statement of the facts can be called an interpreta-
tion, was that deterioration both morphological and physiological
is the direct result of inbreeding, and therefore occasional crossing
of genetically distinct blood lines is a necessary requisite to vigor in
every sexually propagated species.
Seven years ago an extended series of investigations was started
at the Connecticut Agricultural Experiment Station having as
their primary object an interpretation of these facts in keeping with
the more extended knowledge comprised in modern biology. This
paper presents a full account of the views that the writers have
come to hold through the data gathered in these experiments,
although it has not been thought necessary or advisable to confuse
the arguments by overloading it with all of the data in their posses-
1 Published also as a contribution from the Laboratory of Genetics, Bussey Institution of Harvard Uni-
versity.
28748°— BuL 243—12 2 7
8 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
sion. It is hoped that an adequate number of facts are cited to sup-
port the thesis, and it is sufficient on this occasion to say that not
a single fact has been discovered that is irreconcilable with it.
THE PROBLEM.
The experimental data upon which the defense of our thesis
is based have been obtained entirely from plants, but observations
of animal hybrids and published records lead us to believe that the
facts are the same among animals. We believe, therefore, that
our conclusions apply alike to the animal and the vegetable kingdoms,
for we believe the propositions upon which the arguments are based
are applicable to all organisms reproducing sexually. These propo-
sitions are:
(1) Mendel's law — that is, the segregation of character factors in
the germ cells of hybrids and their chance recombination in sexual
fusions— is a general law.
(2) Stimulus to development is greater when certain, or possibly
all, characters are in the heterozygous condition than when they
are in a homozygous condition.
(3) This stimulus to development is cumulative up to a limiting
point and varies directly with the number of heterozygous factors
in the organism, although it is recognized that some of the factors
may have a more powerful action than others.
We later in this bulletin take up briefly some of the specific reasons
for extending these theories to the animal kingdom, but at present
we shall confine ourselves to developing the botanical proof.
EARLY INVESTIGATIONS.
The number of cases in which hybridizers have noticed an increase
in vigor in crosses between subvarieties, between varieties, and between
species is so great that an extended citation of the facts is superfluous.
Without exception the horticultural writers of the nineteenth century
noted the phenomenon and many of them described it at great
length. We have taken some trouble to find out its generality, and
have found records of its occurrence in the gymnosperms (Darwin,1
1876; Focke, 1881) and pteridophytes (Focke, 1881) as well as
throughout the angiosperms. In fact, out of 85 families of angio-
sperms in which artificial hybrids have been made, instances of
hybrid vigor exceeding that of the parent species have been noted
in 59.
Kolreuter (1763), the earliest botanist to study artificial plant
hybrids — as Darwin notes — gives many exact measurements of his
hybrids and speaks with astonishment of their "statura portentosa"
1 Citations to literature throughout this bulletin refer to the " Bibliography " on pages 49-51.
243
EAELY INVESTIGATIONS. 9
and " ambitus vastissimus ac altitudo valde conspicua." Later,
after having been struck with certain natural adaptations for cross-
fertilization, he made a passing remark which plainly showed that
he thought nature had intended plants to be cross-fertilized and
that benefit resulted therefrom. The hybridists that followed
Kolreuter were all interested in the phenomenon, but up to the
time of Darwin only Knight and Gartner attempted to generalize
from their observations. Perhaps this was because each one noted
the fact that some species hybrids were small and weak. Knight
(1799), however, made the somewhat generalized statement that
nature had something more in view than self-fertilization and in-
tended that sexual intercourse should take place between neigh-
boring plants of the same species. On the whole, however, Gartner
has given the best expression of the views of the botanical experi-
menters down to 1849, and for this reason we have translated in
full his section on "Wachstum, Luxuriation und Sprossungsver-
mogen der Bastarde" (Gartner, 1849, p. 526). He writes as follows:
One of the most conspicuous and common characteristics of plant hybrids is the
luxuriance of all their parts, a luxuriance that is shown in the rankness of their growth
and a prodigal development of root shoots, branches, leaves, and blossoms that could
not be induced in the parent stocks by the most careful cultivation. The hybrids
usually reach the full development of their parts only when planted in the open, as
Kolreuter (1763) has already remarked; when grown in pots and thus limited in food
supply their tendency is toward fruit development and seed production.
Concerning the great vigor of hybrids all observers are agreed; on this point may
be cited Kolreuter (1763), Sageret (1826), Sabine Berthollet (1827), W. Herbert (1837),
Mauz (1825), and Lecoq (1845). The vigor of a plant can even serve to indicate its
hybrid nature in a doubtful case, as Kolreuter has done with Mirabilis jalapo-
dichotoma.
Besides possessing general vegetative vigor, hybrids are often noticeable for the
extraordinary length of their stems. In various hybrids of the genus Verbascum, for
example lychnitis-thapsus, the stem shoots up 12 to 15 feet high, with a panicle 7 to 9
feet, the six highest side branches 2 to 3 feet, and the stem 1£ inches in diameter at the
base; in Althaea cannabino-officinalis the stem is 10 to 12 feet; in Malva mauritano-
sylvestris 9 to 11 feet; in Digitalis purpureo-ochroleuca 8 to 10 feet, with panicles 4 to 5
feet; and in Petunia nyctaginifloro-phoenicea and Lobelia cardinali-syphilitica 3 to 4 feet
each. Prof. Wiegmann also corroborates these observations.
Hybrids in the genera Mirabilis and Datura are especially conspicuous for their
enormous size, as Kolreuter has already stated. The different hybrids of Datura —
Stramonio-tatula, quercifolia-ferox, laevi-tatula, and laevi-ferox — grew so large as to
be almost treelike, with branches and leaves that nearly weighed down the stems,
evejn before the time for developing their numerous blossoms. Likewise such species
hybrids as Nicotiana suavolenti-macrophylla, Nicotiana rustica-marylandica, and Trop-
aeolum majus-minus reach a noteworthy height and circumference.
The root system and the power of germination of hybrids are highly correlated with
their great vegetative vigor. Many hybrids, therefore, which are not so luxuriant
in growth as those just described, for example, Dianthus, Lavatera, Lycium, Lych-
nis, Lobelia, Geum, and Pentstemon hybrids, put forth stalks easily and therefore are
readily propagated by layers, stolons, or cuttings. The observations of Kolreuter
243
10 HETEKOZYGOSIS IN EVOLUTION AND PLANT BREEDING.
{1763), Sageret (1826), and Wlegmann (1828) agree with ours in this respect. This
extraordinary side branching and tillering, as well as the growth of the main stem, in
most hybrids continues until late in the fall and in many until frost, as we have ob-
served in Lobelia syphilitico-cardinalis, Petunia nyctaginifloro-phoenicea, Nicotiana
suaveolenti-macrophylla, Pentstemon gentianoideo-angustifolius, Digitalis purpureo-
ochroleuca, Malva mauritiano-sylvestris, Althaea cannabino-officinalis, etc. Sageret
(1826) makes the same statement about Nicotiana tabaco-undulata. There are other
hybrids, however, that are without this ability to form sprouts, such as Matthiola
annuo-glabra and those between several Nicotiana species.
Luxuriation expresses itself at times as proliferation; for instance, in Lychnis diurno-
flos cuculi the receptaculum is changed to a bud that puts forth branches and leaves.
If, moreover, the vigor of the hybrids especially affects the stem and the branches,
particularly their length, nevertheless the leaves take part in it by becoming larger.
Hybrids in the genera Datura, Nicotiana, Tropaeolum, Yerbascum, and Pentstemon
are examples.
Kolreuter (1763) expresses the opinion that the strength and luxuriance of hybrids
continued long after blooming rests upon the fact that the plants are not exhausted
and worn out by the production of seed. Similarly, Edw. Blyth (1837) sees in the
impotence or sterility of animal hybrids the explanation of their great muscular devel-
opment, while the considerable size which these hybrids reach in comparison with
their parents may be interpreted in the same manner, since capons are able to make a
like growth.
But if we take into consideration that: (1) Such a sex condition may exist in
dioecious plants without resulting in the luxuriance shown by hybrids, then the reason
given above may be no adequate explanation of that phenomenon. (2) The luxu-
riance of the hybrid plants is already present and visible before the development of the
flowers, although one may not doubt that the derangement of the sexual activities
and of the development of those organs is not without consequences to the inner life
of these plants and that there may obtain essential difference between the weakening
or the entire suppression of one or the other of the sexual activities of the hybrids and
of the normal separation of the sexes. (3) Not all partially fertile and sterile hybrids
are gifted with an increased vegetative power, since we have observed several abso-
lutely sterile hybrids with weakened and limited vegetative vigor; for example,
Nicotiana grandifloro-glutinosa, N. glutinosa-quadrivalvis, N. rustico-suavolens, N.
suaveolenti-quadrivalvis, Dianthus barbato-deltoides, D. caucasico-arenarius, Verbascum
blattaria-lychnitis, etc.; at the same time many other hybrids keep the growth rela-
tionships of the parent plants unchanged. (4) Among all the hybrids that we have
observed, those which show the greatest luxuriance in all their parts are precisely
those which show the greatest fertility, for example, Datura stramonio-tatula, Datura
' quercifolio-ferox, Tropaeolum majus-minus, Lavatera pseudolbio-thuringwca, Lycium
barbaro-afrum, and Mirahilis jalapo-dichotoma. (5) Planting partially fertile hybrids,
such as Nicotiana rustico-paniculata and Dianthus barbato-chinensis, etc., in pots makes
the production of fruit and seed easier through limiting the vegetative growth, but a
sterile plant is never made fertile by this method. Luxuriance is therefore a peculiar
quality of several hybrids, although it is not possessed by all in the same degree.
Although the early hybridizers paid more attention to crosses
between distinct species than they did to crosses between races that
differed by only a few relatively unimportant characters, there is no
question but that the}" noted a very great number of cases where
crosses of the latter character gave plants that were remarkable for
their vigor. In fact, we have found no record of intervarietal crosses
243
EAKLY INVESTIGATIONS. 11
where delicate or weak hybrids resulted. On the other hand, species
crosses sometimes result in hybrids constitutionally feeble. It is
obvious, therefore, that a reasonable interpretation of the facts must
include an explanation of each category. This matter must be left
until later, however, for the work of the early investigators is cited
only to show the prevalence of the phenomena under discussion.
Gartner's researches were followed by but little systematic study
of cross and self fertilization in plants until the time of Darwin, and
even Darwin's earlier work was confined to the natural means of plant
pollination. This early work, mainly a study of pollination in
orchids, was summed up in 1862 by the saying " Nature abhors per-
petual self-fertilization," a dictum that has become known as the
Knight-Darwin law. This important conclusion gave a great
impetus to the study of the means of flower pollination throughout
the angiosperms. A huge literature of several thousand titles was
built up, from which at times important compilations, such as those
of Muller (1873) and Knuth (1898), have been made. Every possible
variation in flowering habit was argued into an adaptation for cross-
fertilization with an ingenuity and zeal similar to that shown by
zoologists in their work upon protective coloration and mimicry,
and often with as enthusiastic prodigality of extravagant logic. The
earnestness of these observers extended our knowledge of the me-
chanics of pollination in the angiosperms beyond that of any one
phase of general botany, yet in the last half of the nineteenth cen-
tury Darwin was the only scientist who made a systematic experi-
mental inquiry into the physiological effect of cross-pollination com-
pared with self-pollination. The net result of the work of the other
observers was simply to show the widespread occurrence of means by
which cross-pollination might take place. This fact may be taken
to indicate that cross-fertilization is an advantage to a species, but
it certainly does not prove that cross-fertilization is indispensable.
The many plants naturally self-fertilized preclude it.
Darwin's later experimental work on this subject was so important,
both from the standpoint of completeness and brilliancy of analysis,
that it must be considered by itself. For this reason we will dis-
regard chronology and conclude this part of our historical summary
with the observations of the greatest hybridizer contemporary
with Darwin, W. O. Focke. In Focke's fine work "Die Pflanzen-
Mischlinge" he gives a chapter on the properties of hybrids, from
which the following extract is taken:
Crosses between different races and different varieties are distinguished from individ-
uals of the pure type, as a rule, by their vegetative vigor. Hybrids between mark-
edly different species are frequently quite delicate, especially when young, so that
the seedlings are difficult to raise. Hybrids between species or between races that
243
12 HETEROZYGOSIS IX EVOLUTION AND PLANT BREEDING.
are more nearly related are, as a rule, uncorrrmonly tall and robust, as is shown by
their size, rapidity of growth, earliness of flowering, abundance of blossoms, long
duration of life, ease of asexual propagation, increased size of individual organs,
and similar characters.
To undertake a closer examination of the above propositions, it will be necessary
to cite a few examples. The following hybrids are abnormally weak: Nymphaea alba
when crossed with foreign species, Hibiscus, Rhododendron rhodora with other species,
R. sinense with Eurhododendron, Convolvulus, the polyhybrids of Salix, Crinum,
and Narcissus. Moreover, it has often been noticed that other hybrid seedlings are
somewhat delicate and are brought to maturity with difficulty. Really dwarf growths
have been but seldom observed in hybrids; compare, however, certain hybrids of
Nicotiana. (Page 2S5 above, and especially N. quadrivalvis X tabacum macro-
phylla. p. 292.) Giant growths are more frequent; note for example Lycium, Datura,
Isoloma, and Mirabilis. In size the hybrids generally surpass both the parental
species, or at the least they surpass the average height of the two; compare many
hybrids of Nicotiana, Verbascum, and Digitalis. Development often goes on with
great rapidity, as Klotzsch has emphasized in his hybrids of Tllmus, Alnus, Quercus,
and Pinus. Further, the blossoms of hybrids often appear earlier than do those
of the parent species, for example, Papaver dubium X somniferum, many Dianthus
hybrids, Rhododendron arboreum X cataicbiense, Lycium, Nicotiana rustica X panicu-
lata, Digitalis, Wichura's six-fold Salix hybrids, Gladiolus, Hippeastrum vittatum X
reginae, etc., and especially many hybrids of Verbascum. On the contrary, it must
be admitted, there are several hybrids that blossom only after a long growth period
or not at all, examples of which may be found in the genera Cereus and Rhododen-
dron. Of earlier ripening of the seed independent of earlier blossoming only one
example has come down to me, namely Xuphar. Very frequently, one might say
very generally, an extraordinary numerical production of flowers has been observed,
for example, Capsella, Helianthemum, Tropaeolum, Passiflora, Begonia, Rhododen-
dron, Nicotiana (rustica X paniculata, glutinosa X tabacum, and others), Verbascum,
Digitalis, many of the Gesneracese, Mirabilis, and Cypripedium. The size of the
blossoms is often increased in hybrids. By crossing two species with flowers of dif-
ferent size, those of the hybrids very nearly reach (not seldom entirely reach) the size
of the larger variety. Examples of hybrids with unusually large blossoms are Dian-
thus arenarius X superbus, Rubus caesius X bellardii, and hybrids of Rosa gallica, Be-
gonia boliviensis, and Isoloma tydaeum.
A great capacity for vegetative propagation is very general in hybrids; among the
good examples of such a phenomenon may be mentioned Nymphaea, hybrids of
Rubus caesius, Nicotiana suaveolens X latissima, Linaria striata X vulgaris, and Pota-
mogeton. Great longevity may be mentioned as a characteristic of a few hybrids
of Nicotiana and Digitalis, ability to withstand cold is especially noticeable in Nico-
tiana suaveolens X tabacum latissima, while Salix viminalis X purpurea is more
sensitive to frost than either of the parent species.
These facts point in part to a certain weakness of constitution which is a peculiarity
of the hybrid as a result of its abnormal origin and in part to an extraordinary vegeta-
tive vigor. An explanation of the last phenomenon, which has been observed much
more frequently than the weakness, has only recently been found. The noteworthy
experiments of Knight, Lecoq, and others have been familiar for some time, but
only through the painstaking experiments of Charles Darwin has the benefit of a
cross between individuals and races of one and the same species been clearly demon-
strated. The intensification of vegetative vigor in species hybrids is obviously a cor-
responding experience which requires no especial explanation on the basis of peculiar
conditions in hybrids. It was formerly believed that the decreased sexual fertility of
243
THE WORK OF DAE WIN. 13
hybrids was compensated by a greater vegetative luxuriance, a conception the untena-
bility of which, as Gartner showed, is refuted in the simplest manner by the experience
that many of the most fertile crosses (Datura, Mirabilis) are at the same time character-
ized by the most excessive stature.
THE WORK OF DARWIN.
Through Darwin's work we get a very different insight into the
meaning of cross and self fertilization. At the beginning of his
work the knowledge on the subject gained from the experiments and
observations of the older hybridists might be summed up in one
sentence: Crosses between varieties or between species often give
hybrids with a greater vegetative vigor than is possessed by either
parent. To be sure there was also a belief that ill effects result
from inbreeding, but this belief was generally confined to the animal
kingdom. At the end of Darwin's brilliant experiments, or, rather,
brilliant analyses of simple but great experiments, not a single point
of the many ramifications into which the problem may be divided
but had been fully covered. Unfortunately Mendel's experiments
were unknown, and the master key of the situation was not available
to him. Had it been we can not doubt that he would have made
good use of it.
Darwin's interest in the subject arose of course from its connection
with the problem of evolution. If the offspring from a cross-fertiliza-
tion has an advantage over the offspring of a self-fertilization in the
struggle for existence, one can hardly doubt the power of natural
selection in fixing the structures of flowers. And this being granted,
it is obvious that in many flowers mechanical devices to procure
cross-fertilization would have been developed. Having found this
to be the case in several plants, he bent all his energies to interpreting
all flower structures in the same manner. As stated before, the
fascination of the work thus initiated has brought us a huge litera-
ture on the subject, some of the arguments of which are fantastic to
say the least. Darwin himself never allowed his conclusions to get
ahead of his facts, a trait that his followers did not always copy.
He firmly believed that self-fertilization was so injurious that plants
dependent upon it must ultimately perish, but he frankly admitted
the obstacles which self-fertilized families like Leguminosse placed
in the way of his conclusions. If he had known of the vigorous
plants that reproduce apogamously no doubt he would have
regarded the obstacles more seriously than he did. Nevertheless
one must admit that at that time, considering the importance of
placing evolution on an impregnable foundation, Darwin did not
overstate his conclusions. He proved conclusively the advantage
of cross-fertilization and the numerous means by which it is obtained.
243
14 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
If he did not distinguish between the advantage a process may hold
forth and the necessity of that process, it was because he was not
in possession of all the facts. One does not criticize Darwin, there-
fore, if in a careful examination of his data in the light of modern
knowledge many facts are found that may reasonably have some-
what different interpretations than those originally given.
The first point we will consider is the benefit arising from cross-
fertilization. It must be granted from the data already presented
that an increase in vigor generally results when different species
or markedly different varieties are crossed. It is also perfectly
obvious that many plants are naturally designed for cross-fertili-
zation. It can hardly be argued, however, that specific crosses
could have had a widespread value in the course of evolution. It must
be shown, therefore, that in plants not widely different in character
cross-fertilization shows an advantage over self-fertilization. In
Table A ("Cross and Self Fertilisation," p. 240) Darwin's results
on this subject are given. To be fair, 15 of these experiments
should be discarded, because the number of plants measured in the
comparison between those crossed and those selfed is less than five.
There are 37 experiments left. Of these, the crossed plants were
higher in 24 cases, provided an error of 5 per cent is allowed. In
13 cases, then, cross-fertilization showed no definite advantage.
In Table B, where the weights of entire plants are considered,
cross-fertilization showed to advantage in 5 experiments out of 8.
From these data it seems logical to argue that cross-fertilization
between nearly related plants is often a benefit, yet since types that
are self-pollinated in nature — legumes, wheat, tobacco, etc. — are
among the most vigorous of living plants, it can not be said to be
indispensable. Furthermore, about 25 of our most vigorous species
of angiosperms have given up sexual reproduction either partially
or entirely and have become apogamous.
Did the simple act of crossing produce these beneficial results?
If so, why was the advantage due to cross-fertilization not general
and without exception? Darwin himself answered these questions.
He says (loc. cit., p. 269):
A cross between plants that have been self -fertilized during several successive gen-
erations and kept all the time under nearly uniform conditions does not benefit the
offspring in the least, or only in a very slight degree. Mimulus and the descendants
of Ipomoea named Hero offer instances of this rule. Again, plants self -fertilized
during several generations profit only to a small extent by a cross with intercrossed
plants of the same stock (as in* the case of Dianthus) in comparison with the effects
of a cross by a fresh stock. Plants of the same stock intercrossed during several gen-
erations (as with Petunia) were inferior in a marked manner in fertility to those derived
from the corresponding self-fertilized plants crossed by a fresh stock. Lastly, certain
plants which are regularly intercrossed by insects in a state of nature and which were
artificially crossed in each succeeding generation in the course of my experiments, so
243
THE WORK OF DARWIN. 15
that they can never or most rarely have suffered any evil from self-fertilization (as
with Eschscholtzia and Ipomoea), nevertheless profited greatly by a cross with a fresh
stock. These several cases taken together show us in the clearest manner that it is
not the mere crossing of any two individuals which is beneficial to the offspring. The
benefit thus derived depends on the plants which are united differing in some manner,
and there can hardly be a doubt that it is in the constitution or nature of the sexual
elements. Anyhow, it is certain that the differences are not of an external nature,
for two plants which resemble each other as closely as individuals of the same species
ever do profit in the plainest manner when intercrossed if their progenitors have been
exposed during several generations to different conditions.
In other experiments that Darwin performed it was shown conclu-
sively that crosses between individual flowers borne on the same
plant conferred no benefit whatever on the progeny. It is evident,
therefore, since plants may differ in nonvisible transmissible charac-
ters, that differences in transmissible factors alone account for the
benefit produced by crossing and are indispensable to its occurrence.
This is clearly shown by the fact that even types naturally self-
fertilized, such as the garden pea (Pisum sativum), showed a remark-
able increase in vigor when entirely different strains were crossed.
We may well believe, then, that if Darwin's plants used in his Table
A had all been heterozygous at the start they would all have showed
a considerable difference in favor of the progeny of those continually
cross-fertilized. Furthermore, leaving out of consideration our own
beliefs, a study of his own experiments (Ipomoea) shows that if his
comparisons had been kept up for a considerable number of genera-
tions the cross-fertilized stocks would have become so nearly like the
self-fertilized stocks in constitution that the advantage due to cross-
fertilization would have been small. But to this point we shall
again recur.
Let us now consider whether the known effects of inbreeding and
crossbreeding are manifestations of the same phenomenon. In
" Animals and Plants Under Domestication" he says (vol. 2, p. 89):
The gain in constitutional vigor derived from an occasional cross between indi-
viduals of the same variety but belonging to different families, or between distinct
varieties, has not been so largely or so frequently discussed as have the evil effects
of too close interbreeding. But the former point is the more important of the two,
inasmuch as the evidence is more decisive. The evil results from close interbreeding
are difficult to detect, for they accumulate slowly and differ much in degree with
different species, whilst the good effects which almost invariably follow a cross are
from the first manifest. It should, however, be clearly understood that the advantage
of close interbreeding, as far as the retention of character is concerned, is indisputable
and often outweighs the evil of a slight loss of constitutional vigor.
It is obvious that Darwin believed in a definite accumulation of
evil effects from self-fertilization, but his experiments do not justify
this view. He is perfectly correct in saying that the good effects
of crossing are immediately evident. This is clear when it is remem-
bered that if two plants differ in several transmissible allelomorphs
28748°— Bui. 243—12 3
16 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
the first hybrid generation is heterozygous in all these characters,
while future generations as a whole are heterozygous in only part
of these characters. Furthermore, one may cross two plants differing
but slightly and obtain only a small increase in size; he may then
recross with a third plant of widely different nature and obtain a
great increase. When one inbreeds, however, he relies on chance
combinations to eliminate heterozygosis. This occurs through the
action of the laws governing probabilities. Many heterozygous
combinations are eliminated at once. This lowers the number of
such combinations, and, while the percentage of elimination is the
same, the effect of the inbreeding decreases. Complete homozygosis
is approached as a variable approaching a limit. It may be illus-
trated by the old story of the dog decreasing the distance from the
hare by half at each jump. The effects of inbreeding, therefore,
appear to accumulate, while the effects of crossbreeding are imme-
diately manifest. But is the apparent accumulation of evil effects
real? And are the effects evil? Our interpretation is that the
effects of inbreeding are not to accumulate ill effects, but to isolate
homozygous strains. One does away with a stimulus due to hetero-
zygosis, and one sometimes isolates strains with poor transmissible
qualities. But one also isolates good strains; strains that remain
good in spite of continued self-fertilization. In other words, the
apparent evil effects of self-fertilization decrease directly with the
number of' generations it is practiced, due to the increase in homo-
zygosis. On the theory entertained by us it should come to an end
with complete homozygosis; practically, complete homozygosis is
difficult to obtain.
Did such a decrease in deterioration actually occur in Darwin's
experiments as they were increased in duration? They did. Dar-
win himself noted the point. He says ("Cross and Self Fertilisa-
tion," p. 55):
As the plants which were self-fertilized in each succeeding generation necessarily
became much more closely interbred in the later than in the earlier generations, it
might have been expected that the difference in height between them and the crossed
plants would have gone on increasing; but so far was this from being the case that the
difference between the two sets of plants in the seventh, eighth, and ninth genera-
tions taken together is less than the first and second (and third) taken together.
This statement was made concerning his experiments with Ipo-
moea purpurea, which were continued for 10 generations. The ratio
of heights of crossed to heights of selfed plants varied from 100 to 68
in the third generation to 100 to 86 in the fourth generation, but in
the ninth generation the ratio was 100 to 79, which is higher than
that of the first generation. The tenth generation was indeed. low,
but it may with all fairness be rejected, for Darwin states that the
plants were diseased.
243
RECENT INVESTIGATIONS. 17
We know, further, that Darwin was not dealing with the same
strain at the end of his experiments that he was at the beginning.
This change was due, as we now know, to the elimination of Mende-
lian segregates. The plants in the beginning varied greatly in the
color of their flowers. Indeed, they varied during the whole time
of experimentation; but the color of the later generations was much
more uniform than that of the earlier generations. The selfed gen-
erations were so uniform, in fact, that his gardener said "they did
not need to be labeled."
In this experiment as well as in those with other species, such as
Mimulus luteus and Nicotiana tdbacum, remarkably vigorous self-
fertilized types appeared. It may be that new transmissible varia-
tions arose, but it is unnecessary to assume it. One may account
for every result obtained by Darwin by granting the isolation of
homozygous Mendelian segregates, accompanied by loss of the vigor
due to heterozygosis through self-fertilization.
RECENT INVESTIGATIONS.
Since the time of Darwin, several writers, whose results will be
discussed later, have investigated the effect of inbreeding on animals.
Botanists, however, have in general been interested only in the super-
ficial results of inbreeding and crossbreeding and have made no
attempts until recently to bring together and to correlate our knowl-
edge regarding them.
In 1905, Shull and the senior writer each started independent inves-
tigations concerning the effects of inbreeding in maize, which may be
regarded as an ideal cross-fertilized species. To supplement these
experiments we have made a large series of crosses with species of
the genus Nicotiana which are generally self-fertilized, as well as
minor observations on other plants. We will not discuss our previ-
ous papers (East, 1907, 1908, 1909, 1910; Hayes and East, 1911) as
the present paper gives a resume of those experiments. Concerning
Shull's work (1908, 1909, 1910, 1911), we wish to quote his own con-
clusions for they are stated very concisely. Furthermore, Shull's
data and our own, independently obtained, are corroborative in every
detail and therefore have greater weight than either alone. Even
the additional conclusions drawn from the data presented in this
paper are largely an application of the earlier analysis to the broader
problems that are legitimately concerned.
Shull's conclusions up to the year 1910 are summarized by him
as follows (Shull, 1910):
(1) The progeny of every self-fertilized corn plant is of inferior size, vigor, and pro-
ductiveness as compared with the progeny of a normally crossbred plant derived from
243
18 HETEROZYGOSIS IX EVOLUTION AND PLANT BREEDING.
the same source. This is true when the chosen parent is above the average condi-
tion as well as when below it.
(2) The decrease in size and vigor which accompanies self-fertilization is the great-
est in the first generation and becomes less and less in each succeeding generation
until a condition is reached in which there is (presumably) no more loss of vigor.
(3) Self-fertilized families from a common origin differ from one another in definite
hereditary morphological characters.
(4) Regression of fluctuating characters has been observed to take place away from
the common mean or average of the several families instead of toward it.
(5) A cross between sibs (sister and brother) within a self-fertilized family shows
little or no improvement over self-fertilization in the same family.
(6) A cross between plants belonging to two self-fertilized families results in a
progeny of as great vigor, size, and productiveness as are possessed by families which
had never been self-fertilized.
(7) The reciprocal crosses between two distinct self -fertilized families are equal
and possess1 the characters of the original corn with which the experiments were
started.
(8) The Fx generation from a combination of plants belonging to certain self-fertilized
families produces a yield superior to that of the original crossbred stock.
(9) The yield and quality of the crop produced are functions of the particular com-
bination of self-fertilized parental types and these qualities remain the same whenever
the cross is repeated.
(10) The Fj hybrids are no more variable than the pure strains which enter into
them.
(11) The F2 shows much greater variability than the Fv
(12) The yield per acre of the F2 is less than that of the F:.
TVe should also like to quote Shull (1911) upon one important
point upon which we have but few data:
Necessary corollaries of the view that the degree of vigor is dependent on the degree
of hybridity or, in other words, that it is dependent roughly upon the number, of
heterozygous elements present and not upon any injurious effect of inbreeding per se
are (a) that when two plants in the same self-fertilized family, or within the same
genotype, however distantly the chosen individuals may be related, are bred together
there shall be no increase of vigor over that shown by self-fertilized plants in the same
genotype, since no new hereditary element is introduced by such a cross; (b) that first-
generation hybrids produced by crossing individuals belonging to two self-fertilized
lines or pure genotypes will show the highest degree of vigor possible in progenies
representing combinations of those two genotypes, because in the first generation
every individual will be heterozygous with respect to all of the characters which dif-
ferentiate the two genotypes to which the chosen parents belong, while in subsequent
generations recombinations of these characters will increase the average number of
heterozygous genes present in each individual; (c) that crosses between sibs (sister
and brother) among the first-generation hybrids between two genotypes will yield
progenies having the same characteristics, the same vigor, and the same degree of
heterogeneity as will be shown by the progenies of self-fertilized plants belonging to
the same first-generation family.
All of these propositions have now been tested in a limited way. In 1910, nine
different self-fertilized families were compared with nine crosses between sibs within
the same self-fertilized family; ten crosses between sibs in F1 families were compared
i They are usually as vigorous or more vigorous than the original strains, but may or may not have the
original characters. Some characters may have been entirely eliminated. — E. M. E.
243
EXPERIMENTS ON ZEA MAYS.
19
with ten self-fertilizations in the same Fx families; seven families were raised as first-
generation hybrids between individuals belonging to different self-fertilized families;
and ten families were grown in which self-fertilization had been entirely precluded
during the past five years. The average height of plants in decimeters, the average
number of rows per ear, and the average yield in bushels per acre in these 55 families
are given in the following table:
Average height
Average rows. .
Average yield..
Selfed X
self.
19.28
12.28
29.04
Selfed X
sibs.
20.00
13. 26
30.17
Pi.
25.00
14.41
08. 07
F2.
23.42
13. 07
44.62
Fi X self.
23.55
13. 62
41.77
FiX
sibs.
23.30
13.73
47.77
Cross-
breds.
22.95
15.13
61.52
An examination of this table indicates to me that on the whole my self -fertilized
families are not yet quite pure bred ; for the sib crosses give on the average a slightly
greater height, number of rows per ear, and yield per acre than the corresponding
self -fertilized families as shown by a comparison of the first two columns of the table.
The same fact is apparent from a comparison of the UF1X self" and UF1 X sibs"
columns, except that in this case the heights and number of rows per ear are essentially
equal while the yield per acre is significantly higher in the sib crosses than in the
self -fertilized families.
These statements should be sufficient to indicate Shull's work
and point of view. Other writers have proposed methods designed
to utilize commercially the increase in vigor shown by first-generation
hybrids, and at least two other theoretical interpretations of this
increase have been submitted (Jost, 1907; and Keeble and Pellew,
1910). These papers will be considered later. We will now take up
the data obtained in our own experiments.
EXPERIMENTS ON A NORMALLY CROSS-FERTILIZED SPECIES,
ZEA MAYS.
EFFECTS OF INBEEEDING.
In these experiments over 30 varieties of maize, including all the
varieties widely differentiated from each other, have been artificially
self -fertilized for from one to seven generations. In every case a
loss of vegetative vigor has followed. At least, following the earlier
usage, one may say the result is a loss of vigor if it is kept clearly in
mind that pathological degeneration is not what is meant. The
universal decline in vigor consists simply in a somewhat less rapid
cell division or slower growth and a smaller total amount of cell
division resulting in smaller plants and plant organs.
Besides this phenomenon, to which there has been no exception,
the progeny always become more or less differentiated in normal
morphological characters, although this is less marked in some varie-
ties than in others. For example, from the yellow dent variety known
243
20 HETEROZYGOSIS IK EVOLUTION AND PLANT BREEDING.
as Learning various strains differing in the following characters have
been isolated during the several generations that they have been inbred :
Red pericarp and colorless pericarp
Red cob and colorless cob.
Red silks and colorless silks.
Red glumes and colorless glumes.
Profusely branched tassels and scantily branched tassels.
Long ears and short ears.
Ears with various numbers of rows.
Ears with large seeds and ears with small seeds.
Ears with straight rows and ears with crooked rows
Ears high on the stalk and ears low on the stalk.
Stalks with many tillers and stalks with few tillers.
Other minor differences have been observed, but these will serve
to show just what is meant by "normal differences." There were
also differences in yield of seed — described later in this bulletin —
some of which may not seem to be normal in character at first thought,
but which we hope to show are not less normal than those given
above.
Besides tnese variations, aberrant individuals appeared in a few
strains with characters which might well be called abnormal; that is,
they are monstrous characters. But this does not mean that they
might not have originated in the same manner as normal characters,
for they are transmitted as such. Two of these characters, fasciated
ears and bifurcated cobs, show a simple Mendelian segregation with
incomplete dominance; two others, striped leaves and dwarf plants,
are probably recessives. It is possible, however, that one form of
striped leaf is the heterozygote between pure white and normal
green. It may be that the first two of these abnormalities are not
simply isolated as Mendelian segregates. They have also appeared
in commercial varieties grown on very fertile soil, a fact that suggests
their origin through interference with normal processes of cell divi-
sion, acceleration in one case and retardation in the other.
The variability of the strains in the above characters decreased as
inbreeding was continued, until after four generations they were
practically constant for all grosser characters. This does not mean
that physiological fluctuation was not as great as in the original
strain. It was not reduced in the least degree. Nor can it be said that
no new heritable variations arose. Certain variations did appear
which may have been new to the strain — witness the fasciated ears —
but of this one could not be certain. Furthermore, it is not meant that
after four or five generations of inbreeding a type is homozygous in all
of its characters. Such a gametic condition is theoretical and could
never be recognized in a pedigree culture. But near homozygotes or
243
Experiments on zea mays. 21
near homozygous genotypes are obtained without selection simply
by inbreeding. The reason for this is simple.
Mendel in his original paper showed that if equal fertility of all
plants in all generations is assumed and, furthermore, if every plant
is always self-fertilized then in the nth generation the ratio of any par-
ticular allelomorphic pair (A, a) would be 2n- 1 AA: 2 Aa:2n-laa.
If we consider only homozygotes and heterozygotes, the ratio is
2n— 1:1. Of course the matter is not quite so simple when several
allelomorphs are concerned, but in the end the result is similar.
Heterozygotes are eliminated and homozygotes remain. The prob-
able number of homozygotes and any particular class of hetero-
zygotes in any generation r is found by expanding the binomial
[l + (2r— l)]n where n represents the number of character pairs
involved. The exponent of the first term gives the number of hetero-
zygous and the exponent of the second term the number of homo-
zygous characters. As an example, suppose we desire to know the
probable character of the fifth segregating generation (Fe) when
inbred, if three character pairs are concerned. Expanded we get
13 + 3[12(31)] + 3[1(31)^] + (31)3.
Reducing, we have a probable fifth-generation population consisting
of 1 heterozygous for three pairs; 93 heterozygous for two pairs;
2,883 heterozygous for one pair; 29,791 homozygous in all three
character combinations.
From this illustration we think it is fairly easy to see that no
matter in how many characters a plant is heterozygous, continued
inbreeding will sooner or later eliminate them. Close selection, of
course, tends toward the same eud, but not with the rapidity or cer-
tainty of self-fertilization.
Inbreeding a naturally crossbred plant, then, has these results:
(1) There is partial loss of power of development, causing a
reduction in the rapidity and amount of cell division. This phe-
nomenon is universal and therefore can not be related to inheritance.
Further, it continues only to a certain point and is in no sense an
actual degeneration.
(2) There is an isolation of subvarieties differing in morphological
characters accompanying the loss of vigor.
(3) There is often regression away from instead of toward the mean
of the general population.
(4) As these subvarieties become more constant in their characters
the loss of vigor ceases to be noticeable.
(5) Normal strains with such hereditary characters that they may
be called degenerate strains are sometimes, though rarely, isolated.
243
22
HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
(6) It is possible that pure strains may be isolated that are so
lacking in vigor that the mechanism of cell division does not properly
perform its function, and abnormalities are thereby produced.
The maize families shown hi Table I illustrate some of these facts,
if the yield of shelled corn per acre is taken as a basis of comparison
of vigor. These families are not selected to fit a theory, but include
representatives of four of the great subdivisions of the species out of
those grown in sufficient quantity to give considerable confidence in
the determination of yield. Many other types have been inbred for
from one to four years, but neither land nor time was available to
grow them in large quantities. Their behavior, however, was the
same. Inbreeding always reduced the yield of seed and the height
and delayed the time of flowering. In general, the decrease in vigor
lessened with the inbreeding. Further, both good and bad strains
were isolated.
Table I. — Effect of inbreeding on the yield of maize.
Variety.
Year
grown.
Num-
ber of
years
inbred.
Yield in
bushels
per acre.
Variety.
Year
grown.
Num-
ber of
years
inbred.
Yield in
bushels
per acre.
Watson's flint No. 5
No. 5-8
No. 5-8-3
Starchy No. 10 l
No. 10-3
No. 10-3-7
No. 10-3-7-3
No. 10-4
No. 10-4-8
1908
1909
1910
1908
1909
1910
1911
1909
1910
i'
2
r
2
3
1
2
75.7
47.5
36.1
70.5
56.0
67.0
39.1
43.0
48.7
29.3
93.2
58.7
51.2
53.6
42.1
88.0
59.1
95.2
57.9
80.0
27.7
88.0
60.9
59.3
Learning dent— Contd.
No. 1-7-1-1
No. 1-7-1-1-1
No. 1-7-1-1-1-4
No. 1-7-1-2
No. 1-7-1-2-2
No. 1-7-1-2-2-9
Learning dent No. 1
No. 1-9
1908
1910
1911
1909
1910
1911
1905
1906
1908
1909
1910
1911
1905
1906
3
4
5
3
4
5
i"
2
3
4
5
.....
46.0
63.2
25.4
59.7
68.1
41.3
88.0
42.3
No. 10-4-8-3 1911
No. 1-9-1
51.7
Stowell's sweet No. 19. . i 1909
No. 19-4 | 1910
No. 19-4-7 2 j 1911
No. 19-8 ! 1910
No. 19-8-2 2 < i9ii
1
2
1
2
i"
2
3
4
0
1
No. 1-9-1-2
No. 1-9-1-2-4
No. 1-9-1-2-4-6
Learning dent No. 1
No. 1-12
35.4
47.7
26.0
88.0
38.1
Learning dent No. 1 1905
No. 1-12-1
1907 2
32.8
No. 1-6
No. 1-6-1
No. 1-6-1-3
No. 1-6-1-3-4
No. 1-6-1-3-4-4
Learning dent No. 1
No. 1-7
1906
1908
1909
1910
1911
1905
1906
No. 1-12-1-1 1
No. 1-12-1-1-2
No. 1-12-1-1-2-4....
No. 1-12-1-1-2-4-11.
No. 1-12-1-1-4
No. 1-12-1-1-4-14...
No. 1-12-1-1-4-14-3.
1908
1909
1910
1911
1909
1910
1911
3
4
5
6
4
5
6
46.2
23.3
16.5
2.0
28.7
9.5
2.0
No. l-7-li
1907
2
Two selections from the progeny of this ear grown.
Probably a normal yield. Grown on a more fertile soil than the rest in 1911.
The different families were all planted on the same plat under uni-
form conditions each season, but, unfortunately, circumstances made
it necessary to grow them upon different fields each season. It is
therefore necessary to take into consideration the differences in soil
fertility and meteorological conditions each year to see the truth of
our conclusions, namely, that continued inbreeding caused only
isolation of strains of varying potency. The greatest differences in
the environmental conditions were in the years 1908, 1909, and 1911.
243
EXPERIMENTS ON ZEA MAYS.
23
In 1908 the land used was highly fertile and the general environmental
conditions much above the normal. Four stalks per hill were grown
this season, but as only three were grown in other years the actual
yields have been reduced one-fourth. Even at this disadvantage
the yields in 1908 are probably somewhat high. For opposite rea-
sons, poor soil and badly distributed rainfall, the yields of 1909 are
somewhat too low and the yields of 1911 are very much too low.
This will be appreciated if the low yields for 1911 are examined in
Table III.
Since the data on the Learning dent variety are the most interesting
they are repeated in a somewhat different form in Table II. There
they are shown in a regular line of descent.
Table II. — Effect of inbreeding on a variety of Learning dent maize.
(Yield, in bushels, of shelled corn per acre.)
Parent variety.
Generations inbred and years in which grown.
1
2
3
4
5
6
88.0(1905)....
f 59.1
(1906)
95.2
(1908)
57.9
(1909)
80.0
(1910)
27.7
(1911)
60.9
(1906)
59.3
(1907)
f 46.0
1 (1908)
63.2
(1910)
25.4
(1911)
59. 7
I (1909)
68.1
(1910)
41.3
(1911)
42.3
(1906)
51.7
(1908)
35.4
(1909)
47.7
(1910)
26.0
(1911)
38.1
1 (1906)
32.8
(1907)
46.2
(1908)
f 23.3
(1909)
16.5
(1910)
2.0
(1911)
28. 7
I (1909)
9.5
(1910)
2.0
(1911)
The Learning, a well-known commercial dent variety, yielded 88
bushels per acre the year before it was first inbred. The season was
normal, and this yield may be considered fairly typical of what the
variety will do on a moderately good soil. Four ears were inbred
and were grown in 1906. This was again an average year. The four
strains showed marked decreases in yield and notable differences in
their characters. The year 1907 was again an average year, and the
second inbred generations are normal. Two strains were not grown
as second inbred generations until 1908, however, and they are there-
fore too high. In 1909 the yields are too low; in 1910 normal, and in
1911 much too low. With these facts in mind, an examination of the
tables shows how the strains became more and more differentiated.
The first strain, No. 6, is a remarkably good variety of corn even after
five generations of inbreeding. It yielded 80 bushels per acre in 1910.
The yield was low in 1911, but since all yields were low that year it can
28748°— Bui. 243—12 4
24
HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
hardly be doubted that this strain will continue to produce good nor-
mal yields of grain. In the field, even in 1911, the plants were
uniformly vigorous and healthy and were especially remarkable for
their low variability. The poorest strain, No. 12, is partially sterile,
never fills out at the tip of the ear and can hardly exist alone. In
1911 it yielded scarcely any corn but will no doubt continue its exist-
ence as a partly sterile variety. Plate I shows ears and tassels of an
almost sterile stiain isolated by inbreeding.
CROSSING INBRED TYPES.
When two of these inbred strains are again crossed, the ¥t generation
shows an immediate return to normal vigor. The plants are earlier
and taller, and there is a greater total amount of dry matter per
plant. For example, in 1911 the average height of all the strains of
inbred Learning dent was 84 inches while the average height of the
16 hybrid combinations was 111 inches and the height of the shortest
hybrid combination was 1 foot greater than that of the tallest inbred
strain.
Table III gives the yields of shelled corn per acre of several inbred
types, together with the yields of many first-generation crosses.
Many interesting points may be learned from this table, provided it is
remembered that maize is greatly influenced by environmental con-
ditions and therefore only populations grown in the same season
should be compared with each other. For this reason the compari-
sons between first-generation hybrids and the unselected commercial
types from which the inbred strains came are not to be given too great
weight. On the other hand, there is such an enormous difference
between many of the first-generation hybrids and the normal com-
mercial varieties that the conclusion that the former are often better
is perfectly just.
Table III. — Comparative yields of inbred types of maize and their first-generation crosses.
Variety.
Year
grown.
Num-
ber of
years
inbred.
Yield in
bushels
per acre.
Comparison
between
Fi and
unselected ,
commercial
strains.
White dent No. 8
Learning dent No. 1-7
No. (8X1-7), Fi
Flint No. 5
Flint No. 11
No. (5X11), Fi
Flint No. 5
Learning dent No. 1-6
No. (5X1-6), F!
No. (5X1-6), Fi
No. (5Xl-6)-l, F2....
No. (5Xl-6)-2, F2....
Starchy No. 10
Learning dent No. 1-6
No. (10X1-6), Fi
243
1908
1908
1909
1909
1909
1909
1909
1909
1910
1910
1910
1910
1910
1910
121.0
62.0
142.0
47.5
44.2
76.3
47.5
57.9
88.9
105.5
54.1
48.9
48.7
80.4
139.0
121.0
88.0
142.0
48.' 0
76.3
75.7
88.0
88.9
105.5
54.1
48.9
70.5
88.0
139.0
Bui. 243, Bureau of Plant Industry, U. S. Dept. of Agriculture.
Plate I.
Tassels and Ears of an Almost Sterile Strain of Corn Isolated by Inbreeding.
(Photographed by Emerson.)
Bui. 243, Bureau of Plant Industry, U. S. Dept. of Agriculture.
Plate II.
Watson's Flint and Longfellow Flint Corn Inbred Two Years With Fi
Hybrid.
(All ears hand-pollinated.)
EXPERIMENTS ON ZEA MAYS.
25
Table III. — Comparative yields of inbred types of maize and their first-generation
crosses — Continued .
Variety.
Learning dent IN o. 1-7
Sweet No. 19 '
No. ( 1-7X 19) , Fi
Learning dent No. 1-9
Learning dent No. 1-12
No. (1-12X1-9), Fi
No. (1-12X 1-9), Fi
No. (l-12Xl-9)-l, F2
No. (1-12X l-9)-4, F2
No. (1-12X l-9)-12, F2
Learning dent 1-6
Learning dent 1-7-1
Learning dent 1-7-2
Learning dent 1-9-2
Learning dent 1-12-2
Learning dent 1-12-4
No. (1-6X1-7-1), Fi
No. ( 1-8X1-7-2) , Fi |
No. (1-6X1-9-2), Fi
No. (1-6X 1-12-2), Fi
No. (1-7-1X1-6), Fi
No. (1-7-1X1-7-2), Fi
No. (1-7-1X1-9-2), Fi
No. (1-7-1X1-12-2), Fi
No. (1-7-1X1-12-4), F,
No. (1-7-2X1-6) , Fi
No. (1-7-2X1-12-2), Fi
No. (1-9-2X1-6), Fi
No. (1-9-2X1-7-1), Fi
No. ( 1-9-2 X 1-12-2) , Fi
No. (1-12-2X1-7-2), Fi
No. (1-12-2X1-12-4), Fi
Year
grown.
1910
1910
1910
1909
1909
1909
1910
1910
1910
1910
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
1911
Num-
ber of
years
inbred.
Yield in
bushels
per acre.
65.5
53.6
142.7
23.3
35.4
110.2
117.5
102.2
91.5
91.5
27.7
25.4
41.3
26.0
2.0
2.0
75.6
58.3
31.6
10.2
58.8
41.3
51.5
16.9
60.2
57.7
63.5
37.3
46.2
3.6
76.9
24.5
Comparison
between
Fi and
unselected
commercial
strains.
88.0
93.2
142.7
88.0
88.0
110.2
117.5
102.2
91.5
91.5
Attention is called first to the fact that in combinations (5 X 1-6)
and (1-12 X 1-9) both the first and second hybrid generations are
grown in the same year. The first hybrid generation gives an enor-
mous increase over the inbred types. The second hybrid generation
is also much greater than the inbred strains, but recombination with
the production of homozygotes has taken place, and this generation
gives much lower yields than when the greatest possible heterozygosity
existed as in the first hybrid generation.
Attention should next be directed to the results of 1911, when
nearly all the possible combinations of the inbred Learning strains
were made. The yields of the inbred types given are those with one
more year of inbreeding than the real parents of the first-generation
hybrids. But considering the amount of previous inbreeding to
which they had been subjected this probably makes but little differ-
ence. As stated before, the yields in 1911 were very much reduced
by the unfavorable season, and this too must be given due weight in
studying the yields. As a whole the combinations into which
No. 1-7 was introduced were the best while those into which the poor
type No. 12 was introduced are the poorest. The combination
(1-7-1 X 1-12-4) was, however, a very good cross.
243
26 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
Possibly a question may arise as to whether the fine yields of the
combination (1-12x1-9) in 1909 and 1910 and the poor yields of
combination (1-9-2x1-12-2) in 1911 are not due to a difference
in the behavior of a reciprocal cross. This is probably not the correct
reason, for in general there is no difference in reciprocals. No. 1-12
was further inbred when the combinations grown in 1911 were made
and this is probably the cause of their poor showing. In the earlier
combination, No. 1-12 undoubtedly had a somewhat different
gametic constitution than when the later crosses were made. Some
essential factor may have been eliminated, therefore, during the
further inbreeding. On the other hand, the whole explanation may
lie in the poor season of 1911.
The marked increase in productiveness of the Ft hybrid over the
parent inbred types of maize is well shown in Plates II and III, while
Plate IV illustrates the falling off in productiveness of the F2 genera--
tion as compared with the Fj generation from inbred types. Plate V
serves to show the striking increase in vigor of the ¥t generation from
a cross of pure lines.
The logical conclusion from the facts brought out above is appar-
ently that good inbred strains are better than poor ones in combina-
tion, but that good and poor strains crossed together may give very
strong plants.
EXPERIMENTS ON SPECIES GENERALLY SELF-FERTILIZED.
As experimental material that contrasts well with maize, the
genus Nicotiana was selected. This genus contains a large number
of species and varieties, most of which have flowers adapted to self-
fertilization. No doubt cross-fertilization sometimes occurs in most
of them, but it is not the rule.
Seeds of several species and many varieties were obtained from
various parts of the world through the kindness of a number of
friends. The same species did not always arrive with the same
name, and we have not been fortunate enough to have the aid of
a Nicotiana specialist in their identification. "We have, however,
studied them in pure-line cultures during the past four years and
have compared them with specimens in the Gray Herbarium of
Harvard University. This gives us some confidence that the names
used are in accord with the species as accepted and described by
Comes in his "Monographic du Genre Nicotiana," Naples, 1899.
Many crosses have been made between different varieties within
the two species Nicotiana tabacum, L., and N. rustica, L. Some of the
varieties of N. tabacum have been practically identical as far as
external appearance is concerned, although received under different
names. When this has been the case, the results have been varied.
243
Bui. 243, Bureau of Plant Industry, U. S. Dept. of Agncultur
Plate III.
Leaming Dent Strains of Corn, No. 9 (at Left) and No. 12 (at
Right), after Four Years' Inbreeding, Compared with Fi Hybrid
(in Center).
(All ears hand-pollinated.)
Bui. 243, Bureau of Plant Industry, U. S. Dept. of Agriculture.
Plate IV.
CD
Bui. 243, Bureau of Plant Industry, U. S. Dept. of Agriculture.
Plate V.
"- — d
Z7
rx K
Sx
U. CD
O
co
LU
z
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LU
0C
=>
a.
Bui. 243, Bureau of Plant Industry, U. S. Dept. of Agriculture.
Plate VI.
2 —
EXPERIMENTS ON SPECIES GENERALLY SELF-FERTILIZED.
27
For example, two exceedingly similar varieties may give hybrids
with no greater luxuriance of growth than the pure parent strains;
other varieties as similar in appearance may give hybrids with as
much as 25 per cent greater vigor than the average of the two par-
ents. In this case the criterion of greater vigor is height of plant.
If one accepts the old view that nonrelationship between the indi-
viduals used as parents is the reason for the increased vigor of the
hybrids, there would be no logical reason why all such crosses should
not show the same condition. If, on the other hand, the correct
explanation is to be sought in the similarity or dissimilarity of the
gametic constitution of the parents, it is quite evident that different
crosses among varieties similar in external characters may behave in
a different manner. Plants having a close genetic relationship with
each other — that is, descendants of a previous cross — may be quite
different in gametic constitution and therefore show an increased
vigor in the ¥1 hybrid; but genetically unrelated plants of practi-
cally the same gametic constitution may be obtained from different
parts of the world under different names and not be expected to
show an increased vigor in the hybrid.
An example of the amount of increase in height in crosses between
Nicotiana rustica brazilia Comes and N. rustica scabra Comes, both
obtained from Italy, is shown in Table IV.
Table IV .—Height
of
crosses
between Nicotiana
hrazilia (349)
rustica
scabra
(352) and N. rustica
Variety or cross.
Class centers in inches.
24
27
30
33
36
39
42
45
48
51
54
57
60
63
66
69
72
75
78
349
4
10
22
14
7
352
2
1
5
n
l
16
3
17
0
3
6
5
5
352 X 349 Fi
5
2
5
4
6
6
1
5
1
1
349 X 352 Fi
The reciprocal crosses both showed a marked tendency to advance
the mode until in each case it is higher than the highest plant of the
taller parent. Different strains of N. tabacum var. "Sumatra/' of
N. tabacum var. "Havana," and of N. rustica var. brazilia, identical
in external appearance, obtained both from the same locality and
from opposite parts of the world, have also shown increased height
when crossed. On the other hand, strains of N. tabacum varieties
"Sumatra" and "Havana," from seed of plants grown in Connecti-
cut, when crossed with like varieties from seed of plants grown in
Italy have shown no increase in vigor. Accounts of other similar
crosses could be given, but it seems unnecessary to multiply exam-
ples. We will therefore pass to a consideration of the specific crosses
shown in Table V.
243
28 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
Table V. — Condition of hybrids in crosses between species of Nicotiana.
Cross.
Germina-
tion.
Fertility.
Condition of hybrid.
N. alata Lk. and Otto, yar. grandi-
flora Comes :
X N. forgetiana Hort. (Sand.).
X N. langsdorffii Weinm
X N. longiflora Cav
Per cent .
100
100
100
2
3
0
0
100
0
100
100
100
0
60
(?)
100
100
100
0
100
25
- 5
0
100
100
100
5
80
100
5
0
100
2
100
60
5
10
0
1
0
100
Fertile..
...do
Sterile...
Slightly
fertile .
SterileC?)
25 per cent in height; very vigorous and pro-
fuse in flowers.
105 per cent in height; vigorous and profuse in
flowers.
100 per cent in height; 100 per cent in vigor.
80 per cent in height; 80 per cent in general vigor.
Very weak; seedlings died.
X N. tabacum L
N. bigelovii Wats. :
X N. alata grandiflora Comes. .
X N. longiflora Cay
X N. quadriyalyis Pursh
X X.silyestrisSpeg. and Comes
Fertile..
125 per cent in height; 100 per cent in general
vigor.
Sterile...
Fertile..
...do
Sterile/.!
120 per cent in height; 120 per cent in vigor; pro-
fuse in flowers.
125 per cent in height; 130 per cent in general
vigor; profuse in flowers.
160 per cent in height; 125 per cent in general
vigor; profuse in flowers.
N. forgetiana Hort. (Sand.):
X N. alata grandiflora Comes. .
X N. langsdorffii Weinm
X N.tabacumL ,
N. glauca Gran, x N. tabacum L. .
N. glutinosa L. x N. tabacum L. . .
N. langsdorffii Weinm. :
X N. alata Lk. Otto, yar.
grandiflora Comes.
X N. bigeloyii Wats
80 per cent in height; less vigorous.
Gartner obtained plants higher and more vigor-
Fertile..
Sterile...
Fertile . .
ous than parents.
105 per. cent in height; 100 per cent in vigor.
110 per cent in height; very vigorous.
X N. forgetiana Hort. (Sand.) .
X N. paniculata L
110 per cent in height; 100 per cent in vigor; pro-
fuse in flowers.
N. longiflora Cay. X N. alata Lk.
and Otto, yar. grandiflora Comes.
N. paniculata L.:
X alata Lk. and Otto, yar.
grandiflora Comes.
X N. bigeloyii Wats
Sterile...
Slightly
fertile.
SterUe...
...do
100 per cent in height and general vigor.
95 per cent in height; rather weak.
100 per cent in height; 95 per cent in vigor.
X N. langsdorffii Weinm
15 per cent in height; very weak and stunted.
Partially
fertile.
SterileC?)
Fertile . .
...do.....
125 per cent in height; very vigorous and pro-
X N. tabacum L
fuse in flowers.
Plants very weak and small.
N.plumbaginifolia Viy. X N.
longiflora Cay.
N. quadriyalyis Pursh. X N. bige-
loyii Wats.
N. rustica L.:
X N. alata Lk. and Otto, yar.
125 per cent in height; 110 per cent in general
vigor.
110 per cent in height; 100 per cent in general
vigor; profuse in flowers.
So weak that plants lived only about two weeks.
grandiflora Comes.
X N. langsdorffii Weinm
Sterile(?)
Partially
feitile.
Sterile...
110 per cent in height; 110 per cent in vigor; very
profuse in flowers.
125 per cent in height; very vigorous; profuse in
flowers.
180 per cent in height; extremely vigorous; pro-
fuse in flowers.
N. silvestris Speg. and Comes:
X N. tabacum L
Sterile...
...do
Almost
sterile.
Sterile . .
140 per cent in height; 120 per cent in vigor; pro-
N. tabacum L.:
X N. alata Lk. and Otto, var.
grandiflora Comes.
fuse in flowers.
10 per cent of average of parents in height and in
general vigor.
120 per cent of average of parents in height and in
general vigor.
85 per cent of average of parents in height and 80
per cent in general vigor.
25 per cent of average of parents in height: Gart-
ner obtained plants more vigorous than parents.
60 per cent of average of parents in height; 75 per
cent in general vigor.
X N. langsdorffii Weinm
X N. longiflora Cay
...do....
Very small and weak; died before flowering.
X N. plumbaginifolia Viy
X N. silyestris Speg. and Comes
Sterile...
135 per cent of average of parents in height; 120
per cent in vigor*, profuse in flowers.
243
Bui. 243, Bureau of Plant Industry, U. S. Dept. of Agriculture.
Plate VII.
il. 243, Bureau of Piant Industry, U. S. Dept. of Agriculture.
Plate VI
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I
EXPERIMENTS ON SPECIES GENERALLY SELF-FERTILIZED. 29
The voluminous data that have been collected on these hybrids
have been condensed and approximated so that they include only
facts germane to the matter in hand. Those crosses designated as
not having germinated are crosses in which seed was obtained, but
from which no plant was obtained from a planting of at least one
hundred seeds. In some of these crosses the seed was poorly formed
(without embryo) and one may say conclusively that they would
never produce plants. Other crosses gave fully mature, perfect seed
which did not germinate. Possibly the proper conditions for their
germination were not obtained. At least it would be rash to conclude
that all of the crosses of which the seed did not germinate would
never produce plants under any conditions. But it is proper to say
that some crosses are possible in which the hybrid plant reaches no
further than the seed stage. A few hybrids, viz, Nicotiana tabacum, X
N. paniculata, N. rusticaX N. alata grandiflora, etc., germinated and
produced a few weak plants that died before flowering. There were
still others that produced mature plants, but plants shorter than
either parent and weak in character. By far the majority of the
hybrids, however, were taller than the average of the parents and
many were taller than either parent. The luxuriance of their growth
was also such that they may be said to be more vigorous than either
parent. Plate VI shows the result of a cross between Nicotiana
tabacum, var., and Nicotiana silvestris.
One gets the idea from a survey of the crosses in this genus that
there are (a) plants so different that they will not cross; (b) crosses
that produce seed that contain no proper embryo; (c) crosses that
produce seed with embryo, but which go no further than the resting
stage of the seed; id) crosses less vigorous than either parent;
(<?) crosses more vigorous than the average of the parents; and (/)
crosses" more vigorous than either parent. It seems probable, then,
that actual fusion may take place between gametes either so differ-
ent in character that the zygote can not develop or in which the
male cell does not bring in the proper substance to stimulate develop-
ment. On the other hand, when development does take place in a
normal manner the great majority of cases show a stimulus greater
in the hybrids than in the pure species. Compare Plate VII.
It might be supposed that the luxuriant development of many of
these hybrids is due to their sterility, that is, due to the fact that no
energy is used in seed formation. Such an idea was held by some
of the earlier hybridizers, but was disproved by Gartner. Nor is it
justified by our own experience. Fertile crosses between plants
differing in character either equal or exceed the parental vigor;
sterile crosses may show a great increase in vigor or they may show
a great diminution in vigor. Plate VIII represents a sterile hybrid
243
30 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
showing decided diminution in vigor. But there need be no con-
fusion in the interpretation of these facts. It is known that some
plants are so unlike that there is mechanical or chemical obstruction
to fertilization. In one case the stigmatic fluid may be poisonous
to certain foreign pollen; in another case the pollen tubes can not
penetrate the micropyle ; sometimes nuclei do not enter the micropyle ;
frequently the two nuclei will not fuse. Such conditions absolutely
prevent a cross. On the other hand, where crossing is possible, all
of the physiological processes normal to the plant may not be carried
out. The difficulty often lies in the maturation of the sex cells, the
reduction of the chromatin, and the preparation for a new sexual
act. In the proposed parent plants tins has already taken place
naturally. The male and female gametes are ready for fusion, and
if nothing interferes this fusion takes place. But this does not mean
that normal development can take place. Cell division may be so
difficult that no embryo is formed, there being simply a pericarp
formed by the reaction of maternal tissue to stimulation. Again,
development of the embryo may take place, primarily because the
difficulty of development is decreased through the nutrition furnished
by the mother plant. But it may stop at this point. Thus it is
obvious that where the parent plants are so different that normal
somatic cell division can not take place, weak plants result even
though they are heterozygous for many characters. If, however,
cell division is normal we may believe that the vigor of the hybrid
increases directly with the amount or the kind of heterozygosis
present, without regard to whether the plant is sterile or fertile.
Sterility, therefore, is often simply an inability to mature the sex
elements properly, possibly because of mechanical obstruction to
normal reduction of chromosomes differing widely in their character,
and sometimes it is correlated with abnormal ontogeny.
We make the statement that hybrid vigor increases with the
amount or with the kind of heterozygosis advisedly. The increased
vigor may vary roughly with the number of heterozygous characters
present, up to that limiting case where the action of other forces pre-
vents or obscures their influence, or it may depend largely upon the
quality of the characters that are heterozygous. This matter has
not been determined; in reality it makes no difference with the
thesis under discussion. It is an interesting problem, but can hardly
be tested experimentally by crossing owing to the number of unknown
characters that may be present in either a heterozygous or homozy-
gous condition. The proof submitted here rests entirely upon the
effects obtained by continued inbreeding as explained by the mathe-
matical expectancy of homozygotes and heterozygotes under con-
tinued inbreeding.
243
THE CHARACTERS AFFECTED BY HETEROZYGOSIS. 31
One further point ought to be noted here. It has been shown that
weak types are sometimes isolated from maize by inbreeding, their
delicate constitution being due, it is assumed, to homozygosis of
heritable characters that produce weakness and not to the mere fact
of inbreeding. Does one obtain weak types in self -fertilized species ?
Undoubtedly such strains arise, but it is difficult to obtain examples
because the weakness of individual plants is usually a physiological
fluctuation due to external conditions and is not transmitted. This
has been found to be true by growing seedlings from weak plants
that have been self -fertilized. They usually give normal plants.
Weak strains have been isolated, however, from Nicotiana tabacum,
from N. paniculata, and from N. attenuata that continued to transmit
their poor constitution. We may conclude, therefore, that weak
strains arise in self-fertilized species, but are eliminated by natural
selection.
THE CHARACTERS AFFECTED BY HETEROZYGOSIS
The term vigor has hitherto been used with the general meaning
which the biologist readily understands. We will now endeavor to
show in what plant characters this vigor finds expression. It is not
an easy task because of the possibility of confusing the phenomenon
of Mendelian dominance with the physiological effect due to hetero-
zygosis. The confusion is due to a superficial resemblance only.
Dominance is the expressed potency of a character in a cross and
affects the character as a whole. A morphological character like
the pods of individual maize seeds, or the product of some physio-
logical reaction like the red color of the seed pericarp in maize may
be perfectly dominant, that is, it may be developed completely when
obtained from only one parent. Size characters on the other hand
usually lack dominance or at best show incomplete dominance.
The vigor of the first hybrid generation theoretically has nothing to
do with these facts. This is easily demonstrated if one remembers
that the increased vigor manifested as height in the Ft generation
can not be obtained as a pure homozygous Mendelian segregate,
which would be possible if due to dominance. Furthermore, the
universality with which vigor of heterozygosis is expressed as height
shows the distinction between the two phenomena. If the greater
height were the expression of the meeting of two factors {T1t2xtlT2)
both of which were necessary to produce the character, one could not
account for the frequency of the occurrence. Nevertheless, in prac-
tice the confusion exists, and while we have considerable confidence
in the conclusions drawn from our experiments, we have no intention
of expressing them dogmatically.
It has been stated that the vigor due to heterozygosis is primarily
an increase and an acceleration of cell division; in other words, an
243
32 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
increased power of assimilation. This is first of all expressed by the
increased size of the root system, a fact noticed by Kolreuter and
Gartner as quoted on page 9. This is the first noticeable difference,
for the size of the cotyledons of the hybrid is largely influenced by
the size of the maternal pericarp, yet there is a slight increase in the
cotyledon size, as we have found in experiments with species of the
genus Imp aliens and with the tomato, Lycopersicum esculentum.
Hybrid seedlings next show the increased vigor by their rapidity of
growth tending toward an earlier maturity. This feature is the accel-
eration of cell division referred to above. Ultimately, however, there
is not only acceleration but increased cell division, resulting in taller
plants. Data supporting this fact have already been shown in
papers on maize (East, 1911, 1911a). The increased size is entirely
internodal. Neither in crosses between maize varieties nor between
varieties of Nicotiana tabacum is there any tendency to increase the
number of nodes. This stem growth is comparatively much greater
than is increased leaf surface in the plants investigated (N. tabacum),
although the latter can be definitely traced.
The size of the flower is not affected, at least not certainly. The
fruit also does not seem to be affected where there is a small natural
amount of cell division, as in the capsule of tobacco. In fleshy fruits
like the tomato or eggplant there is a marked increase. This is prob-
ably true also of the large pomes and pepos, but this is only a surmise
by analogy.
The increased vigor of the whole plant makes it possible for more
flowers and fruit to be produced, as we have determined in tobacco
and tomato. A more or less indeterminate inflorescence is always
prolonged, which probably accounts for the increased size that is
found in the ears of maize hybrids.
There are many less important plant characters upon which no
data have been gathered, but the action of heterozygosis is known well
enough to justify the former statement that it affects the amount and
rapidity of assimilation as expressed by cell division.
THEORETICAL INTERPRETATION OF RESULTS.
At this point it may be well to stop, collect our facts, and discuss
their theoretical interpretation, notwithstanding a certain repetition
it will involve. We believe it to be established that —
(1) The decrease in vigor due to inbreeding naturally cross-fertilized
species and the increase in vigor due to crossing naturally self-
fertilized species are manifestations of one phenomenon. This phe-
nomenon is heterozygosis. Crossing produces heterozygosis in all
characters by which the parent plants differ. Inbreeding tends to
produce homozygosis automatically.
243
THEORETICAL INTERPRETATION OF RESULTS. 33
(2) The phenomenon exists and is in fact widespread in the vege-
table kingdom.
(3) Inbreeding is not injurious in itself, but weak types kept in
existence in a cross-fertilized species through heterozygosis may be
isolated by its means. Weak types appear in self-fertilized species,
but are eliminated because they must stand or fall by their own
merits.
The logical interpretation of all of these facts rests, we believe, on
the acceptance of Johannsen's (1903, 1909) " genotype conception
of heredity." This conception in turn is an extension of Weismann-
ism1 without Weismann's mechanistic speculations, supported by
Mendelism. Johannsen (1911) gives the essential points of this con-
ception in these paragraphs:
The personal qualities of any individual organism do not at all cause the qualities
of its offspring, but the qualities of both ancestor and descendant are in quite the
same manner determined by the nature of the '"sexual substances" — i. e., the
gametes — from which they have developed. Personal qualities are then the reac-
tions of the gametes joining to form a zygote; but the nature of the gametes is not
determined by the personal qualities of the parents or ancestors in question. This is
the modern view of heredity.
The main result of all true analytical experiments in questions concerning genetics
is the upsetting of the transmission conception of heredity, and the two different ways
of genetic research, pure-line breeding as well as hybridization after Mendel's model,
have in that respect led to the same point of view, the "genotype conception''' as we
may call the conception of heredity just now sketched.
A simple illustration of what is meant by the above statement is
as follows: Suppose a maize with red pericarp (RR) be crossed with
one with a colorless pericarp (rr). In the hybrid the gametes R and
r are formed in equal quantities. By chance mating lRR:2Rr : Irr
are obtained. Now the homozygous dominant RR is exactly like the
heterozygote Rr in appearance, but the one breeds true to red pericarp
and the other again throws about 25 per cent white progeny. In
other words, the gametic composition of the z}^gotes determines
whether the resulting plants shall have ears with red or with colorless
pericarps, but the fact that a plant has an ear with a red pericarp
does not show what kind of gametes it will form.
The genotype conception of heredity, as stated before, rests on the
noninheritance of somatic modifications and the general truth of
Mendelism. The first part of the proposition now has almost univer-
sal support. All data point to a germ-cell-to-germ-cell hereditary
transmission. In certain animals it has been demonstrated that
there is an early segregation or setting apart of the material designed
i One need become a Weismannian only so far as to agree with the observed facts which have shown
that the transmission of acquired characters must be so relatively infrequent as to make the possibility
negligible in experimental genetics and plant breeding.
243
34 HETEROZYGOSIS IX EVOLUTION AND PLAXT BREEDING.
to become the germ cells. This fact naturally has been proved in but
few animals, but from it one must infer that in all metazoa there is
a relative independence of soma and germ plasm undreamed of a
few decades ago. In the higher plants no visible difference between
germ plasm and soma plasm has been proved, yet the recent experi-
ments of Baur and of Winkler on periclinal chimeras or false-graft
hybrids have shown that one of the subepidermal layers is probably
alone responsible for the sexual cells. In recent years few biologists
have believed that surrounding conditions did not occasionally
modify gametic structures. On the other hand, fewer and fewer
investigators have maintained that any sort of somatic adaptation
would impress the germ plasm with the ability to transmit the same
modification.
The experimental work on the genotype conception of heredity has
been largely a demonstration of the last statement. It has shown
that in general fluctuations caused by ordinary environmental
changes are not inherited. The idea involved is comparatively old.
Vilmorin's promulgation of his " isolation principle" in plant breed-
ing hi the middle of the nineteenth century might be called its start-
ing point. Vilmorin used the average character of a plant's progeny
as the index of that particular plant's breeding capacity. This is the
genotype conception, pure and simple. Since that time all plant
breeding by selection which has been at all profitable has been done
in this way, although the theoretical interpretation of the results
obtained was unknown. This was given by Johamisen through his
work upon barley and beans.
Since then corroborative results have been obtained by Jennings
(1908, 1910) on Paramaecium, Hanel (1907) upon Hydra, Pearl
(1909, 1911) upon fowls, Barber (1907) upon yeasts, TToltereck
(1909) upon Daphnia, Jensen (1907) upon bacteria, East (1910a)
upon potatoes, Love (1910) upon peas, and Shull (1911) and East
(1911) upon maize. And no one to my knowledge lias made a
successful attack upon the position taken. It is true that, attacks
have been made by Pearson (1910) and Harris (1911), but their main
argument is that the genotype theory is wrong, because it antago-
nizes the utterly erroneous biometric idea that heredity is measured
only by the correlation between parents and progeny in somatic
characters.
To be sure a caveat has been filed by Castle ( " Heredity ",
New York, 1911) to the effect that unit characters so called can
sometimes be modified by selection. This is no real criticism of the
genotype conception of heredity, however, for Castle firmly believes
in the generality of Mendelism and the general noninheritance of
somatic modifications. It must simply be understood that, like
243
THEORETICAL INTERPRETATION OF RESULTS. 35
most chemical compounds, characters are generally stable under ordi-
nary conditions, but also like chemical compounds they may some-
times be modified. This modification then becomes a new character
or is the old character in a slightly different form, depending on the
point of view.
The second part of the proposition rests upon the law of segrega-
tion and recombination of gametic factors, which is the essence of
Mendelism. Every day the generality of this law becomes more
probable. Leaving out of consideration experiments on apogamous
and parthenogenetic species almost every paper published since 1900
dealing with crosses between varieties fertile inter se in which quali-
tative differences have been studied has shown that factors repre-
senting these characters segregate in the germ cells of the hybrid
and recombine in the next generation. The few exceptions have
been papers dealing with characters evidently quantitative, treated
from a biometrical standpoint and not proving or disproving any-
thing.
Recently there have also been investigations (Emerson, 1910;
East, 1910, 1911; East and Hayes, 1911; Lang, 1911, Tammes,
1911) showing that size or quantitative characters also segregate.
Of course all selection experiments on cross-fertilized species using
Vilmorin's isolation principle and the investigations just cited in
support of Johannsen have really proved segregation and recombi-
nation of size characters, else strains differing in such characters
could not be isolated from complex hybrids. The senior writer
(1910), however, has shown how such segregation can be given a
strict Mendelian interpretation by postulating absence of dominance
and multiplicity of determinants affecting the same general charac-
ters. The experimental basis upon which it rests is the investiga-
tions of Nillson-Ehle (1909) upon oats and wheat and his own upon
maize.
It is possible that there are many apparent exceptions to the law
of segregation; it is even possible that practically there are real
exceptions, but these exceptions are likely to be in the nature of
changed conditions which modify the action of Mendel's law through
new sets of conditions. Our meaning is shown by parallels in the
domain of physics and chemistry, where certain laws act perfectly
only under ideal conditions which are very often not fulfilled in
nature. For example, De Vries (1907) states that Burbank's and
Janczewski's bramble hybrids have bred true. Without any data
upon which to base a critical judgment one does not know what to
say, but taking the statement at full value, any number of conditions
may cause this hybrid constancy without invalidating the law of seg-
regation. There may be apogamy, all zygotes may not develop,
243
36 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
selective fertilization may occur, or the action of the law may be
opposed or suspended by other conditions of which we know nothing.
Personally we consider the genotype conception not as a theory
but as a fact. Considering it as a fact, how does it aid the interpre-
tation of the results obtained by inbreeding and by crossing inbred
types of maize ? Maize as a cross-fertilized species of great variability
is in a constant state of hybridization. It is a collection of complex
hybrids. Its usual mode of pollination through the agency of the
wind keeps up this state of hybridization. Inbreeding, however,
tends to produce homozygous types. As already shown, if one
assumes equal fertility for all plants and that each plant lives and
produces offspring in the nth generation there is a ratio 2n— 1 pure
dominants, 2 heterozygotes and 2n — 1 pure recessives for each allelo-
morphic pair.
This theoretical state of affairs may not occur for other reasons
(as unpaired chromosomes) and the large number of allelomorphic
pairs in a complex hybrid may prolong the time required for isola-
tion of strains that are completely homozygous, but final isolation
of strains completely homozygous is the goal toward which inbreed-
ing tends. These completely homozygous strains are Johannsen's
homozygous genotypes. Perhaps no one has ever isolated a real
homozygous genotype, but strains homozygous for many characters
are constantly being separated. This, indeed, is the sole function
of selection.
Weismann assigned two purposes to the gametic fusion termed
sexual reproduction; one being to mingle the hereditary characters
carried by the two germ cells, the other to stimulate development
of the zygote. This general statement was so obviously a fact that
biologists were unanimous in its acceptance and two distinct lines of
investigation have developed from it. Research concerning trans-
mission phenomena has been almost divorced from the study of the
physiology of development in its intimate connection with sexual
reproduction. This separation, in view of the subject of this bulletin,
seems unnecessary and unwise, for it may permit only a partial and
distorted view of the results of reproduction. At any rate the data
given here are of interest from both view points, since they deal with a
purely physiological result brought about by a strictly morphological
transmission phenomenon.
The hypotheses in regard to the way by which the act of fertiliza-
tion initiates development are numerous, but since they are entirely
speculative it is not necessary to discuss them here. The only conclu-
sion that seems justified is that they are not immediately psychological
or vitalistic in nature. Loeb's remarkable researches prove this. But
whatever may be the explanation of the means by which the process
243
THEORETICAL INTERPRETATION OF RESULTS. 37
is carried out, the statement can be made unreservedly that the
heterozygous condition carries with it the function of increasing this
stimulus to development. It may be mechanical, chemical, or elec-
trical. One can say that greater developmental energy is evolved
when the mate to an allelomorphic pair is lacking than when both
are present in the zygote. In other words, developmental stimulus
is less when like genes are received from both parents. But it is
clearly recognized that this is a statement and not an explanation.
The explanation is awaited.
The developmental stimulus is to a certain degree cumulative.
In other words, the expression "the greater the degree of heterozy-
gous condition the greater is the vigor of the resulting plant" roughly
expresses the facts. This does not mean that the possession of cer-
tain allelomorphic pairs in a heterozygous condition is not more
necessary than others of normal development. Castle and Little
(1910), for example, have shown the probability that zygotes which
are potentially homozygous yellow mice are formed but do not
develop. Baur (1909) has shown that homozygous recessives of
pelargoniums that lack the necessary mechanism for chlorophyll
formation are formed but can live only a few days. Of course in
the latter case there is actual absence of a physiological mechanism
that is absolutely essential to development. Whether the condition
is similar in the yellow mice is unknown. It is quite possible that
lack of normal or presence of abnormal factors will account for many
cases of improper development, but these facts must not be con-
fused with the phenomenon under consideration. What we are con-
cerned with here is that developmental stimulus due to heterozygosity
increases roughly with the number of heterozygous allelomorphic
pairs, even though some of these pairs may produce a much greater
stimulus than others.
Inbreeding, then, tends to isolate homozygous strains which lack
the physiological vigor due to heterozygosity. Decrease in vigor
due to inbreeding lessens with decrease in heterozygosity and van-
ishes with the isolation of a completely homozygous strain. More-
over, these homozygous strains can be quite different from each
other in natural inherent vigor. From a single strain of Learning
dent maize one isolated type is a good profitable corn after four
generations of inbreeding, having yielded at the rate of 80 bushels
per acre in 1910; another type is partially sterile and can hardly
develop to maturity after five generations of inbreeding, and yielded
in 1910 only 9.5 bushels per acre. Thus we see the true explanation
of the apparent degeneration that so many observers have attributed
to inbreeding per se.
243
38 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
Wheii species that are naturally close fertilized produce variations
that are weak and degenerate, they perish in the natural struggle
for existence or are not allowed to propagate by man. Since only
the experimental breeder sees the origin of degenerate strains of
close-fertilized species (as we have done in the genus Nicotiana),
biologists have left them out of their consideration and have con-
cluded that some exception to the natural laws of physiology has
been made in their favor so that they could stand the inbreeding for
which they are naturally fitted. Nothing could be further from the
facts. Species which through their flower structure must be self-
fertilized produce as many degenerate strains as any species. They
are produced, but they do not survive; they are lost and forgotten.
Species which through their flower structure are naturally cross-
fertilized, on the other hand, produce strains poor in natural vigor,
degenerate strains, and they are kept from sight. They survive
the scythe of natural selection; they are selected for propagation by
man because they are crossed with other strains and are vigorous
through heterozygosity. Inbreeding tears aside their mask. They
must then stand or fall on their own merits. Those strains with a
high amount of inherent natural vigor, due to gametic constitution,
lose the added vigor due to a heterozygous condition, but are still
good strains, ready to stand up forever under constant inbreeding.
The poor strains that have had the help of hybridization with good
strains, combined with the added vigor due to heterozygosity, are
stripped of all pretense, shown in all their weakness, and inbreeding
is given as the cause for their degeneracy. At least inbreeding has
until recently been given as the cause, but it is hoped that the newer
interpretation will now be accepted as logically interpreting all the
facts.
Although the increased power of growth of hybrids and the de-
creased vigor attending inbreeding have not been recognized as the
same phenomenon until the work of Shull and the senior writer,
nevertheless there has been a so-called interpretation of the increased
vigor of hybrids current among plant physiologists. It is the theory
of rejuvenescence or renewal of youth in the protoplasm. Continued
self-fertilization is thought to be comparable to vegetative repro-
duction and continued vegetative reproduction is supposed to bring
about a senile condition in the protoplasm. This theory was borrowed
from zoology, having long since been proposed by Butschli to account
for conjugation in protozoa. It can not be considered a theory that
helps in interpreting the vigor of hybrids, for it tells us nothing.
Moreover, it may be based upon wrong premises. It is not at all
certain that conjugation is an absolutely necessary phenomenon.
Woodruff (1911) has demonstrated that protozoa can be kept in
243
EXTENSION OF CONCLUSIONS TO ANIMAL KINGDOM. 39
healthy condition without conjugation for at least 2,300 generations.
Jennings has been unable to make certain genotypes of Paramaecium
conjugate. Nuclear fusions sometimes occur in some of the ascomy-
cetes and basidiomycetes, but in general these fungi reproduce
asexually and possibly have produced hundreds of species in this
manner. In the higher plants there are many species in which
either no seed is produced or sexual propagation is seldom resorted
to, and yet they seem to be in no danger of degeneration. Among
them may be mentioned the banana, hop, strawberry, sugar cane,
and many of the grasses. There are also certain apogamous genera,
such as Taraxacum and Hieracium, that are exceedingly vigorous.
From these facts it is reasonably conclusive that sexual reproduction
may be a benefit, but is not a necessity.
Keeble and Pellew (1910) have recently suggested that "the greater
height and vigor which the Ft generation of hybrids commonly
exhibit may be due to the meeting in the zygote of dominant growth
factors of more than one allelomorphic pair, one (or more) provided
by the gametes of one parent, the other (or others) by the gametes
of the other parent." We do not believe this theory is correct. The
"tallness" and " dwarf ness" in peas which Keeble was investigat-
ing is a phenomenon apparently quite different from the ordinary
transmissible size differences among plant varieties. Dwarf vari-
eties exist among many cultivated plants, and in many known cases
dwarf ness is recessive to tallness. It acts as a monohybrid or possibly
a dihybrid in inheritance, and tallness is fully dominant. Varietal
size differences generally show no dominance, however, and are
caused by several factors. Transmissible size differences are un-
doubtedly caused by certain gametic combinations (East, 1911), but
this has nothing to do with the increase of vigor which we are dis-
cussing. The latter is too universal a phenomenon among crosses
to have any such explanation. Furthermore, such interpretation
would not fitly explain the fact that all maize varieties lose vigor
when inbred.
EXTENSION OF THE CONCLUSIONS TO THE ANIMAL KINGDOM.
Can the conclusions in regard to heterozygosis be extended to
animals? The answer is affirmative as far as an interpretation of
the known facts is concerned. No experimental attack from the
standpoint taken in this paper has been made, but the older work
furnishes many data that readily fit this view. As a matter of fact,
however, it is questionable whether it is necessary to make formal
proof in the matter. Sexual reproduction has undoubtedly arisen
several times in the vegetable kingdom and at least once independ-
ently in the animal kingdom. Why or how it arose, one need not
243
40 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
inquire; having arisen, the purposes served are essentially the same
if the similarity of the methods is an argument. The duplex nature
of organisms, the halving of the chromatin and the production of
simplex cells at the maturation of the sex cells, the fusion of two
simplex cells as the starting point of a new organism, the general
result of this fusion in the matter of development, and the trans-
mission of heritable characters, are so similar in their main points
that it would be quite wonderful if the process both in plants and
animals did not now fulfill like requirements.
Since our conclusions are based upon the generality of Mendehsm,
which has been rendered highly probable by the multiplicity of zoolog-
ical researches, it seems only necessary to show that heterozygosis in
annuals does cause (or accompany) an increase in vigor. It is easier
to do this than to attack the still widespread belief that inbreeding is
injurious per se. We have seen fertile crosses between different
varieties of cattle, of swine, of sheep, and of domestic birds that 'were
more vigorous than either parent. There are several swine raisers in
the Middle West who make a practice of selling only first-generation
crosses on account of their size. A number of very vigorous sterile
hybrids of both domestic and void animals might also be cited, but
with these crosses a complication is encountered. In plants we found
that the presence or absence of normal sexual organs made little if any
difference in the amount of vigor induced by heterozygosis. In ani-
mals the case is undoubtedly different. From their very mode of
development — annuals being closed forms and plants open forms —
internal secretions play a great rdle. And it is a matter of common
knowledge that castration, in vertebrates at least, causes an extra-
ordinary development of the body. In the human race this develop-
ment is especially noticeable in the femur bones, so that Havelock
Ellis states that the eunuchs of Cairo can be readily picked out of a
crowd by their great stature. It is obvious, therefore, that there are
two causes of vigorous somatic development, elimination of sexual
organs and heterozygosis. In sterile hybrids, therefore, one can not
say how much of the induced stimulation is due to each cause, but in
fertile crosses there is no question about its source.
It is much more difficult to argue against the supposed injurious
effects of inbreeding. Abhorrence of incest, which probably had a
religious origin among our ancestors, is so difficult to eradicate from
our minds that judgment is made before the facts are heard. This
belief is not universal in the human race if Westermarck, the greatest
authority on the history of marriage, *is to be trusted, but the retort
is made that the races that approve incestuous unions are low in intel-
ligence. The answer does not prove anything, however, as low races
with both beliefs are found, and, furthermore, as disapproval of inces-
243
EXTENSION OF CONCLUSIONS TO ANIMAL KINGDOM. 41
tuous relations is both religious and esthetic, it would only be expected
in races of some intelligence. Nor is the answer germane, for it is not
shown that incestuous tribes are less well developed physically than
related tribes with different customs, which is the real matter under
examination.
But let us confine the discussion to the lower animals. If this is
done there are two things to consider, the closeness of matings and
their result. The statement is often made that self-fertilization in
plants is a much closer sexual relationship than can obtain in bisexual
animals. With a germ-to-germ transmission conception of heredity
it is doubtful if this is true. After a wide cross, a self-fertilized plant
of the Fx generation produces markedly different progeny, due to
recombinations of gametic factors. After continuous self-fertiliza-
tion for many generations, the gametic factors tend to become homo-
zygous and their matings are close in relationship. Thus it is per-
fectly clear that it is not kinship of the two organisms furnishing the
sex cells that determines the closeness of the mating, but the simi-
larity of the constitution of the cells themselves. There is no a priori
reason, therefore, why bisexual animals may not be bred as thor-
oughly in-and-in as plants.
On this account the statement must be made very emphatic that
investigations such as studies of cousin marriages in the human race
amount to nothing. A cousin marriage may be a wide cross, it may
be very narrow.
There is a possibility that has not been mentioned, however, that
may prove to be an essential difference between the reproduction of
bisexual animals and hermaphroditic plants. There is no question
but that sex in the higher animals is essentially Mendelian in its
behavior. There is no necessity of tying its interpretation to the
chromosomes or to the accessory chromosome in particular. Castle's
(1909) simple explanation that the female is gametically x the male
plus a theoretical X factor has interpre'tated so many facts that its
correctness — possibly somewhat modified — is highly probable. Under
this interpretation one sex is always heterozygous. No similar expla-
nation has been advanced to account for hermaphroditism. Possibly
the same thing accounts for the differentiation into microgamete and
macrogamete in plants, although not accompanied there by somatic
changes. Since we are ignorant of the facts in plants, we can not say
that sex furnishes a real reason for believing bisexual animal matings
1 Note the words "gametically the male. " This is not at all the same thing as saying the male plus some-
thing else. The X may produce many important changes during ontogeny.
There are two classes of facts; in one the male is homozygous, having no X factors, while the female has one.
In the other the male is heterozygous, having one X factor, while the female is homozygous, with two X fac-
tors The human race probably belongs to the second type.
243
42 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
less incestous than plants. The facts are simply given for what they
are worth.
We are now ready to take up the actual effect of inbreeding in ani-
mals. In the statements of Darwin's correspondents we find through-
out the tendency to mix esthetic feelings and facts. But here and
there an independent observer maintained that breeding good stocks
in-and-in had no evil effect. Undoubtedly there is sometimes a
slight loss in vigor (we should say vegetative vigor as we have done in
plants, because constitutional vigor is not lost), but there is no degen-
eration. On the other hand, there is segregation toward homozygous
strains, and these strains differ in constitutional vigor. The greatest
breeds of horses, cattle, swine, and sheep have been developed by
in-and-in breeding. Breeders have worked for homozygous strains,
for they desired strains that bred true. Inbreeding has been accused
of producing everything undesirable in many of these strains, but the
accusations are extremely illogical. Consider one or two examples.
The race horse has undoubtedly been inbred more than the draft horse.
Did inbreeding produce the nervousness and delicate constitution of
the former ? Certainly not. It is absolutely essential that the race
horse be nervous. It has been thus selected for generations. Again,
the delicate constitution of the Boston terrier or even the toy terrier
is pointed out as the effect of inbreeding. We doubt very much if
there has been any more inbreeding in the case of the Boston terrier
than with the St. Bernard, but the selective ideals have been quite
different.
The necessity for heterozygosis may be very different in various
species of animals. In some the stimulus to zygotic development may
be insufficient when like germ cells conjugate; in others, it may pro-
duce normal development. Weismann has made much of the fact
that hermaphroditic animals are always cross-fertilized at times. It
may be necessary in these species or it may be coincidence. Possibly
hermaphroditic species will be found that are always self-fertilized yet
retain their vigor even as in plants. At any rate Weismann's argu-
ments seem to have little force, considering the widespread preva-
lence of parthenogenesis in the animal kingdom. It seems reasonable
to conclude that in animals as in plants cross-fertilization may be
advantageous but is not a necessity.
The actual experiments of Crampe (1883), Bitzema Bos (1894),
and Von Guaita (1898) on mammals, of Fabre-Domengue (1898) on
birds, and of Castle et al (1906) on the fly DrosopJiila ampelopMla
Low may all be interpreted in this way. Fertility was decreased in
some strains. Those strains needed the stimulus due to a certain
amount of heterozygosis for their proper development. Other strains
243
VALUE OF HETEROZYGOSIS IN EVOLUTION. 43
were perfectly fertile in spite of inbreeding. Sometimes combina-
tions of hereditary characters resulted in relatively weak strains;
other combinations of characters gave strong strains. In no case
was there absolute and universal degeneration due directly to
inbreeding.
As a final example of the simple way in which these experiments
on animals fit the heterozygosis theory, we will take a case that
puzzled Weismann (1904). Nathusius allowed the progeny of a
Yorkshire sow to inbreed for three generations. Weismann says:
"The result was unfavorable, for the young were weakly in consti-
tution and were not prolific. One of the last female animals, for
instance, when paired with its own uncle, Jcnown to be fertile with
sows of a different breed, produced a litter of 6 and a second lit-
ter of 5 weakly piglings; but when Nathusius paired the same
sow with a boar of a small black breed, which boar had begotten
7 to 9 young when paired with sows of his own breed (the black
breed evidently near homozygous through close breeding), the sow
of the large Yorkshire breed produced in the first litter 21 and in
the second 18 piglings."
VALUE OF HETEROZYGOSIS IN EVOLUTION.
Before undertaking to discuss the part that heterozygosis may have
played in evolution, emphasis must be laid upon one point of criti-
cism directed against almost all speculative evolutionary philosophy.
Unconsciously, perhaps, many of the conditions that are widespread
among living forms have been spoken of as having been selected to
continue their existence in nature because they are indispensable to
the organism. This is certainly untrue. One has only to recall
other epochs of geology to appreciate the fact. The huge reptiles of
the Cretaceous period were long in developing their peculiar speciali-
zations, yet they were swept away. In a present-day post-mortem
we can assign many reasons why they were eliminated from the
organic worlds but if their characters were so unfit for their environ-
ment, how did they come to be developed ? It is said the environ-
ment changed and left them too specialized for adaptive response.
This is plausible enough, but, nevertheless, possibly untrue.
Must we not be just as skeptical about the question of sexual dif-
ferentiation ? It has arisen several times; it has persisted. Having
arisen, it undoubtedly has a function. Perhaps it was necessary;
perhaps it was a fundamental blunder, as was once humorously
stated. Speculation is, of course, futile. We merely wish to point
out that in discussing a function intimately connected with sexual
reproduction it is absolutely unnecessary to suppose that sex fulfills
243
44 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
a protoplasmic necessity or demand.1 We do not say that the belief
is untrue, but that it is not known to be true and therefore should not
be treated as a fact.
In other words, electric drills and hammers are very useful in build-
ing a bridge, but good bridges have been built without them. Sexual
reproduction serves a purpose, but several of the most vigorous genera
of our higher plants have given it up. It is evidently unnecessary
to them. We must cast a vote, therefore, against the belief in the
rejuvenescence theory of sexual reproduction. Furthermore, we
believe that any hypothesis in which an endeavor is made to twist
the phenomena attending sexual reproduction into requisites indis-
pensable to the evolution of all species possessing it is without basis.
All one can do is to suggest how it may have been beneficial at times
to some species.
Transmissible variations are produced in great numbers by apoga-
mous genera such as Taraxacum and Hieracium, so that sexual reproduc-
tion is not the cause of variation. Johannsen's (1906) and many other
pedigree-culture studies have shown that it presumably never increases
variation. But it does permit recombination of the gametic factors of
the parents, and this has no doubt been of great service in evolution.
Galton and Quetelet (Weismann, 1904) have argued that the intercross-
ing thus allowed is a means of keeping the species constant, but even
with the old idea of blended inheritance this seems to us to be an
exaggeration. Greatest constancy in the actual descendants, if new
heritable variations are disregarded, would come from asexual repro-
duction. If the species group is considered as a whole and compara-
tively free from competition, a great amount of intercrossing — as in
a naturally cross-fertilized strain — would help toward a general fixa-
tion of type, even though it did not contribute toward the produc-
tion of homozygous factors; but if a rigid competition is allowed,
new and better combinations of characters would replace the old.
Perhaps this matter may be made clearer by an illustration drawn
from our maize studies. Height is a complex due to many contribut-
ing factors. Some of them are probably correlated in inheritance,
but a sufficient number are transmitted independently to give the
1 Vitalistic interpretations of the origin of characters, though largely confessions of ignorance of ulti-
mate causes, deserve consideration for calling attention to that fact; yet one must admit that if every-
thing is accounted for by some "perfecting principle" this creative force has made many trials and errors.
Of course things do not just happen. The chemist sees certain series of compounds give similar reac-
tions under like conditions, while other series give other reactions under those conditions. More complex
chemicals under the general term protoplasm probably act in the same manner and produce variations
through their reactions. Some of these variations are widespread— that is, they are general reactions;
others are less general— that is, they are specific reactions. Personally this analogy helps in the conception
of certain orthogenetic phenomena, but the conception leads back to the same blank wall of ignorance.
The vitalist and the believer in mechanico-chemical theories reach the same point, but the latter is pleased
if he is able to reduce a series of facts to the shorthand of a formula; the former is worried because knowledge
stops at the most interesting place.
243
VALUE OF HETEROZYGOSIS IN EVOLUTION. 45
example validity. There is no dominance, and when two individuals
differing in stature are crossed there is a blend in the first hybrid
generation. There is a real segregation, however, resulting in an
increased variability in the F2 generation. In the Fx generation
there is a normal frequency deviation due to noninherited fluctua-
tions. In the F2 generation there is a similar curve, but with greater
variability, due to fluctuating variability plus the variability due to
the recombination of gametic factors. This condition of affairs
tends toward the maintenance of a general mean in height, but this
mean is false. It is false because the modal class which Galton and
Quetelet took to be the type toward which the species is tending
actually contains more heterozygous individuals and individuals
heterozygous for more factors than any other. An individual
selected from this class is less likely to breed true than one selected
from the extremes. Cross-fertilization, therefore, may tend toward
the production of a mean that gives falsely an appearance of fixity
of type.
This preliminary discussion has necessarily been rather long in
order to have a basis for considering the part that heterozygosis
may have played in evolution. We shall confine ourselves to the
higher plants, although we think a portion of the statements made
are equally true when applied to animals. It can hardly be doubted
that heterozygosis did aid in the development of the mechanisms
whereby flowers are cross-fertilized. Variations must have appeared
that favored cross-fertilization. These plants producing a cross-
fertilized progeny would have had more vigor than the self-fertilized
relatives. The crossing mechanism could then have become homo-
zygous and fixed, while the advantage due to cross-fertilization
continued. But was this new mechanism an advantage? It must
have been often an advantage to the species as a whole. In compe-
tition with other species, the general vigor of those which were
cross-fertilized would aid in their survival. But the mechanism
may not have been useful in evolving real vigor in the species,
because of the survival of weak strains in combination. In self-
fertilized species, new characters that weakened the individual
would have been immediately eliminated. Only strains that stood
by themselves, that survived on their own merits, would have been
retained. On the other hand, weak genotypes in cross-fertilized
species were retained through the vigor that they exhibited when
crossed with other genotypes. The result is, therefore, that self-
fertilized strains that have survived competition are inherently
stronger than cross-fertilized strains. On this account weak geno-
types may often be isolated from a cross-fertilized species that as a
whole is strong and hardy.
243
46 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
VALUE OF HETEROZYGOSIS IN PLANT BREEDING.
First-generation hybrids of many economic plants give a yield
sufficiently greater than pure strains to pay for their production
and leave a profit. This is true only of crops where crossing is easy
and where profit is made from accelerated and increased cell divi-
sion or number of fruits. In general, it is not true where the selling
price is greatly increased by the possession of some special quality.
As Collins has remarked, value may at times accrue also from the
fact that a plant breeder who has found a great increase in yield
from growing the first hybrid generation of a particular cross may
keep the parents a secret and maintain a justly remunerative busi-
ness by selling hybridized seed or seedlings. A few suggestions as
to the crops to which this method may be applied are given below.
MAIZE.
Maize is our most important field crop, and an increase of one
bushel per acre to the average yield would add many millions of
dollars annually to the nation's resources. The methods now in
general use for its improvement all follow Vilmorin's isolation
principle. Progeny-row tests are grown from individual ears. This
means that good strains are isolated, but it also means that the
longer selection is carried on the nearer is a homozygous condition
approached. Thus the increased stimulus due to heterozygosis is
lost. Since from both Shull's tests and our own, strains made
almost homozygous by artificial inbreeding have yielded as high as
250 per cent increase over the average of the parents, this stimulus
is not to be lightly disregarded. Of course these tests were made
with strains so nearly homozygous that they gave very low yields.
But we have obtained yields of ear corn very much higher than are
ever given on land of like fertility by commercial types. Shull
(1909) has therefore suggested that near-homozygous strains be pro-
duced by self-fertilization, the best combination- determined by ex-
periment, and hybridized seed of this combination sold. This pro-
cedure is undoubtedly the best in theory, because the greatest degree
of heterozygosis is thereby obtained. Perhaps it can be made prac-
tical, but we are afraid very few commercial men would undertake it.
As a method whose practicability outweighs its theoretical disad-
vantage, the senior writer (East, 1909) has suggested that combina-
tions of commercial varieties be made, testing them until the most
profitable combination is found. Since maize is monoecious, this
method can be used on a large scale at a small cost. It is only neces-
sary to take two varieties, A and B, plant them in alternate rows,
and detassel all of the plants of one variety. The seed gathered
243
VALUE OF HETEROZYGOSIS IN PLANT BREEDING. 47
from this detasseled variety is all crossed seed and will give, in gen-
eral, a greater yield than the average of the two parents. Crossed
seed can be produced in this manner at an additional cost over
natural seed of not over 9 cents per bushel. If it averages two
bushels per acre increase in yield, the producer can sell it at one
dollar advance over natural seed and still allow the buyer a good
profit. The method is given in greater detail in another paper
(Hayes and East, 1911).
This plan we thought original, but Collins (1910) has shown that
it is comparatively old. It has been suggested time and again with-
out gaining a foothold in agricultural practice. Let us hope that
the time is now ripe for a scientific method to be understood, appre-
ciated, and used.
It is fortunate that we have at hand data from many agriculturists
showing the value of using first-generation hybrids in maize. They
are very convincing. We will not discuss them in detail, but refer
the reader to Collins's paper (1910). We may say, however, that the
following researches have shown that a commercial use of the method
is possible: Beal at the Michigan Experiment Station in 1880, Inger-
soll at the Indiana Experiment Station in 1881, Sanborn at the
Maine Experiment Station in 1889, Morrow and Gardner at the
Illinois Experiment Station in 1892, Shull of the Carnegie Institution
Station for Experimental Evolution in 1909, East at the Connecticut
Experiment Station in 1909, Collins and his assistants of the United.
States Department of Agriculture in 1910, Hayes and East at the
Connecticut Experiment Station in 1911, and Hartley and his assist-
ants of the United States Department of Agriculture in 1912.
TRUCK CROPS.
In some truck and garden crops, such as beans and peas, the diffi-
culty of making artificial crosses absolutely precludes a commercial
use of the stimulus due to heterozygosis. Other crops, such as
pumpkins and squashes, are too plentiful and cheap to be worth the
trouble. Besides, these crops are so often crossed naturally that
they are always more or less heterozygous. On the other hand,
there are garden crops that are in demand at all seasons of the year
and are grown under glass during the winter with profit. Some of
them are easily crossed and will pay for their crossing. As examples,
tomatoes and eggplants may be cited. Both are easily crossed and
are worth crossing. We grew a cross between Golden Queen and
Sutton's Best of All tomatoes in 1909. It outyielded both parents.
Further, we are informed that several unpublished experiments at
the New York Experiment Station by Wellington have shown that
crossed seed is worth its production.
243
48 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
Eggplants have another advantage that should be mentioned.
Varieties exist whose fruits are so large that the buyer does not care
for them, the seller makes no profit, and the plant produces a very
limited number. Other varieties have very small fruit. Now fruit
size is intermediate in the first hybrid generation, while the number
produced is increased and the time of ripening advanced.
PLANTS REPRODUCED ASEXUALLY.
The one type of plants where heterozygosis has been utilized,
though not purposely, is that class which is reproduced asexually by
cuttings, grafts, etc. Potatoes and grapes are good examples. Com-
mercial varieties are always hybrids, and the reason, we think, is
because the hybrids yield so profusely. The cross is made and the
best plant of the first generation is simply multiplied indefinitely by
division. This method could be applied more generally to bush
fruits, such as gooseberries, raspberries, blackberries, etc., and to the
larger fruits, like apples, pears, and peaches.
FORESTRY.
There is one other class of economic plants where it seems possible
to make a practical use of heterozygosis. We refer to trees used for
lumber. Many plans for breeding forest trees have been suggested,
yet we have never seen the use of first-generation hybrids suggested.
This omission seems strange, for as early as 1855 (Darwin, " Animals
and Plants," vol. 2, p. 107) M. Klotzsch crossed Pinus sylvestris and
nigricans, Quercus robur and pedunculata, Alnus gluiinosa and incana,
Vlmus campestris and effusa and planted the crossed seeds and seeds
of the pure parent species in the same place and at the same time.
The result was that after eight years the hybrids averaged one-third
taller than the parent trees. Further, the quick-growing hybrid
walnuts produced by Luther Burbank undoubtedly owe that valu-
able quality to heterozygosis.
A large amount of experimental work will be necessary before
definite recommendations can be made as to what species can be
crossed, what result may be expected, and what extra cost must be
allowed for the production of hybrid seed. It is perfectly evident
that hybrid seed will be impossible in many cases, and even where
hybrids can be produced comparatively few can be crossed at a small
enough cost to make the scheme a commercial success. On the other
hand, we have no doubt that with many good lumber trees crossing
would be found easy and hybrid seed could be sold with a wide
margin of profit both to producer and to forester.
243
X
BIBLIOGRAPHY.
The following is a complete list of the literature cited in this
bulletin :
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Bulletin 1, vol. 4, 1907, pp. 1-48.
Baur, E. Das Wesen und die Erblichkeitsverhaltnisse der "Varietates albomar-
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und Vererbungslehre, vol. 1, 1909, pp. 330-351.
Berthollet, Sabine. Phenomenes de l'acte mysterieux de la fecondation. Me-
moires de la Societe Linneenne de Paris, vol. 5, 1827, pp. 81-83.
Blyth, E. On the physiological distinctions between man and all other animals.
Magazine of Natural History, n. s., vol. 1, 1837, pp. 1-9, 77-85, 131-141.
Castle, W. E. A Mendelian view of sex heredity. Science, n. s., vol. 29, 1909,
pp. 395-400.
Heredity in relation to evolution and animal breeding, New York, 1911,
184 pp., illustrated.
— — — and Little, C. C. On a modified Mendelian ratio among yellow mice.
Science, n. a., vol. 32, 1910, pp. 868-870.
Carpenter, F. W., et al. The effects of inbreeding, crossbreeding, and selec-
tion upon the fertility and variability of Drosophila. Proceedings of the American
Academy of Arts and Sciences, vol. 41, 1906, pp. 731-786.
Collins, G. N. The value of first-generation hybrids in corn. Bulletin 191, Bureau
of Plant Industry, IT. S. Dept. of Agriculture, 1910, 45 pp.
Crampe, H. Zucht-Versuche mit zahmen Wanderratten. Landwirthschaftliche
Jahrbucher, vol. 12, 1883, pp. 389-458.
Darwin, Charles. The variation of animals and plants under domestication (2d
edition), London, 1875, 2 vols., illustrated.
The effects of cross and self fertilisation in the vegetable kingdom. London,
1876, 482 pp.
De Vries, H. Plant breeding, Chicago, 1907, 360 pp.
East, E. M. The relation of certain biological principles to plant breeding. Bul-
letin 158, Connecticut Agricultural Experiment Station, 1907, 93 pp.
■ Inbreeding in corn. Report, Connecticut Agricultural Experiment Station,
1907, pp. 419-428. (1908.)
The distinction between development and heredity in inbreeding. American
Naturalist, vol. 43, 1909, pp. 173-181.
The role of hybridization in plant breeding. Popular Science Monthly, vol.
77, 1910, pp. 342-355.
The transmission of variations in the potato in asexual reproduction. Report
Connecticut Agricultural Experiment Station, 1909-10, pp. 120-160. (1910a.)
The genotype hypothesis and hybridization. American Naturalist, vol. 45,
1911, pp. 160-174.
- and Hayes, H. K. Inheritance in maize. Bulletin 167, Connecticut Experi-
ment Agricultural Station, 1911, 141 pp., 25 pis. (1911a.)
Emerson, R. A. Inheritance of sizes and shapes in plants. Preliminary note.
American Naturalist, vol. 44, 1910, pp. 739-746.
243 49
50 HETEROZYGOSIS IX EVOLUTION AND PLANT BREEDING.
Fabre-Domengue, P. Unions consanguines chez les Columbins. L'Intermediare
des Biologist es, vol. 1, 1898, pp. 203.
Focke, W. 0. Die Pnanzen-Mischlinge, Berlin, 1881, 569 pp.
Gartner, C. F. Versuche und Beobachtungen iiber die Bastarderzeugung im
Pflanzenreich, Stuttgart, 1849, 790 pp.
Guaita, G. von. Versuche mit Kreuzungen von verschiedenen Rassen der Haus-
niaus. Bericlite der Naturforschenden Gesellschaft zu Freiburg, vol. 10, 1898,
pp. 317-332.
Hanel, E. Vererbung bei ungesehlechtlicher Fortpflanzung von Hydra grisea.
Jenaische Zeitschrift fur Naturwissenschaft, vol. 43, 1907, pp. 321-372.
Harris, J. A. The biometric proof of the pure line theory. American Naturalist,
vol. 45, 1911, pp. 346-363.
Hayes, H. K., and East, E. M. Improvement in corn. Bulletin 168, Connecticut
Agricultural Experiment Station, 1911, 21 pp.
Herbert, W. Amaryllidaceae, London, 1837, 428 pp.
Jennings, H. S. Heredity, variation and evolution in Protozoa. II. Heredity
and variation of size and form in Paramaecium, with studies of growth, environmen-
tal action, and selection. Proceedings of the American Philosophical Society, vol.
47, 1908, pp. 393-546.
— Experimental evidence on the effectiveness of selection. American Natur-
alist, vol. 44, 1910, pp. 136-145.
Jensen, P. Organische Zweckmassigkeit, Entwicklung und Vererbung vom Stand-
punkt der Physiologie, Jena, 1907.
Johannsen, W. Ueber Erblichkeit in Populationen und in reinen Linien, Jena,
1903, 68 pp.
■ Does hybridization increase fluctuating variability? Report of the Third
International Conference on Genetics, London, 1906, pp. 98-113.
Elemente der exakten Erblichkeitslehre, Jena, 1909, 515 pp.
The genotype conception of heredity. American Naturalist, vol. 45, 1911,
pp. 129-159.
Jost, L. Lectures on plant physiology. (English translation by P. J. H. Gibson.)
Oxford, 1907, 564 pp.
Keeble, F., and Pellew, C. The mode of inheritance of stature and of time of flow-
ering in peas (Pisum sativum). Journal of Genetics, vol. 1, 1910, pp. 47-56.
Knight. T. A. Philosophical transactions. See Knight's collected works, London,
(1799) 1841.
Knuth, P. Handbuch der Bliitenbiologie, Leipzig, 1898-1905, 3 vols.
Handbook of flower pollination. (Translated by J. R. Ainsworth Davis.)
Oxford, 1906-1909, 3 vols.
Kolreuter, J. G. Dritte Fortsetzung der vorlaufigen Xachricht von eihigen das
Geschlecht der Pflanzen betreffenden Versuchen und Beobachtungen, Leipzig,
1766, 156 pp. (Reprinted in Ostwald's Klassiker der exakten Wissenschaften, no.
41, Leipzig, 1893).
Lang, A. Die Erblichkeitsverhaltnisse der Ohrenlange der Kaninchen nach Castle
und das Problem der intermediaren Vererbung und Bildung kohstanter Bastar-
drassen. Zeitschrift fur Induktive Abstammungs- und Vererbungslehre, vol. 4,
1911, pp. 1-23.
Lecoq, H. De la fecondation natureile et artificielle des vegetaux et de l'hybridation,
Paris, 1845, 287 pp.
Love, H. H. Are fluctuations inherited? American Naturalist, vol. 44, 1910, pp.
412-423.
Mauz, E. In Correspondenzblatt des Wtirtteniburgischen Landwirthschaftlichen
Vereins, 1825.
243
BIBLIOGRAPHY. 51
Muller, H. Die Befruchtung der Blumen durch Insekten und die gegenseitigen
Anpassungen beider, Leipzig, 1873, 478 pp.
Nillson-Ehle, H. Kreuzungsuntersuchungen an Hafer und Weizen. Lunds Uni-
versitets Arsskrift, n. s., sec. 2, vol. 5, no. 2, 1909, 122 pp.
Pearl, R. Inheritance of fecundity in the domestic fowl. American Naturalist,
vol. 45, 1911, pp. 321-345.
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fur Induktive Abstammungs- und Vererbungslehre, vol. 2, 1909, pp. 257-275.
Data on the inheritance of fecundity obtained from the records of
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Pearson, K. Darwinism, biometry, and some recent biology, I. Biometrika, vol.
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Ritzema Bos, J. Untersuchungen liber die Folgen der Zucht in engster Blutver-
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Sageret, A. Considerations sur la production des hybrides, des variantes et des
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Annales des Sciences Naturelles, vol. 8, 1826, pp. 294-314.
Shull, G. H. The composition of a field of maize. Report, American Breeders Asso-
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Hybridization methods in corn breeding. American Breeders Magazine,
vol. 1, 1910, pp. 98-107.
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Weismann, A. The evolution theory. (Translated by J. A. Thomson and M. R.
Thomson.) London, 1904, 2 vols.
Wiegmann, A. F. Ueber de Bastarderzeugung im Pflanzenreich, Braunschweig,
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vol. 11, 1911, pp. 135-142.
243
INDEX.
Page.
Abnormalities, relation to heterozygosis 20, 22
Allelomorphs in heterozygosis 15-16, 21, 36, 37, 39
Alnus spp., heterozygosis 12, 48
Althaea spp., heterozygosis 9-10
Angiosperms, heterozygosis 8, 11, 14
Animals, heterozygosis 7-8, 13, 17, 33-34, 39-43, 45
See also Birds, Insects, Mammals, Reptiles, etc.
Apple, commercial application of heterozygosis. 48
Bacteria, heterozygosis in relation to heredity 34
Baitsell, G. A., and Woodruff, L. L., on Paramaecium 38-39, 51
Banana, relation to heterozygosis 39
Barber, M. A., on selection in bacteria 34, 49
Barley, heterozygosis in relation to heredity. 34
Baur, E., on relation of heterozygosis to growth 34, 37, 49
Beal, W. J., on commercial utility of heterozygosis 47
Bean, commercial application of heterozygosis 47
heterozygosis in relation to heredity 34
Begonia spp., heterozygosis 12
Berthollet, Sabine, on effects of crossing 9, 49
Bibliography, list of authors cited on heterozygosis 49-51
Birds, heterozygosis 40, 42
See also Capons and Fowls.
Blackberry, commercial application of heterozygosis 48
Blith, Edw. , on effects of crossing 10, 49
Breeding, plant, value of heterozygosis 33, 3 4, 46-48
Burbank, Luther, experiments which utilize heterozygosis 35. 48
Cane, sugar, relation to heterozygosis 39
Capons, development as related to heterozygosis 10
Capsella spp., heterozygosis 12
Carpenter, F. W. , and Castle, W. E. , on inbreeding. 49
Castle, W. E., and Carpenter, F. W., on inbreeding 49
Little, C. C, on yellow mice ....37,49
on inbreeding and crossbreeding 34, 4] , 42, 49
Castration, effects as compared with heterozygosis , 40
Cattle, heterozygosis in its relation to growth 40, 42
Cereus spp. , heterozygosis 12
Collins, G. N., on utility of heterozygosis 46, 47, 49
Color, relation to heterozygosis 17, 20
Conjugation, relation to heterozygosis 38-39, 42
Connecticut, source of plants under heterozygosis test 27
Convolvulus sp. , heterozygosis 12
Corn, Indian, heterozygosis 17-26, 34, 35, 39, 46-47
practical application of heterozygosis 19, 36, 37, 44-47
243 53
54 HETEROZYGOSIS IN EVOLUTION" AND PLANT BREEDING.
Pa&e.
Crampe, H. , on inbreeding and crossbreeding 42, 49
Crinum spp., heterozygosis 12
Crops, truck, practical application of heterozygosis 47-48
Crosses, first-generation. See Hybrids, first-generation.
Cross-fertilization, relation to heterozygosis , 7, 9,
10, 13. 14, 15, 17, 24-26, 32, 35, 36, 38, 40, 42^5
See also Fertilization.
Cypripedium spp. , heterozygosis 12
Daphnia, selection 34
Darwin, Charles, on inbreeding and crossbreeding 8, 11, 48, 49
work relating to heterozygosis 12, 13-17
Datura spp., heterozygosis 9-10, 12, 13
Degeneration, relation to heterozygosis 37-38, 39, 43
See also Deterioration, Dwarfness, etc.
Deterioration, relation to heterozygosis 7, 37
See also Degeneration.
Development, relation to heterozygosis, 8-10,16,21,29,30,36-37,40,42
See also Size, Vigor, etc.
De Yries, H., on species hybrids 35, 49
Dianthus sp., heterozygosis 9-10, 12, 14
Differences, normal, relation to heterozygous behavior of plants 15-16, 19-20, 29
Digitalis spp., heterozygosis 9-10, 12
Dogs, inbreeding 42
Drosophila arnpelophila. effects of inbreeding 42
Dwarfness, relation to heterozygosis -. 12, 20, 39
Earliness, relation to heterozygosis 12, 22, 24
East, E. M., and Hayes, H. K., on heterozygosis in relation to heredity. 17, 35, 47, 49, 50
on heterozygosis in relation to heredity 17, 32, 34, 35, 39, 46, 47, 49
Eggplant, commercial application of heterozygosis 47-48
Ellis, Havelock, on sexual organs in their relation to heterozygosis 40
Emerson, R. A. , on inheritance 35, 49
Eschscholtzia spp., heterozygosis 15
Eurhododendron sp., heterozygosis 12
Evolution, relation to heterozygosis 13, 14, 43-^5
Experiments relating to heterozygosis 7-43, 46-48
Fabre-Dornengue, on inbreeding 42, 50
Fertility, relation to inbreeding 7, 10, 12, 28, 29-30, 35, 36, 40, 42-43
Fertilization, relation to heterozygosis 11', 29-30, 36
See also Cross-fertilization, and Self-fertilization.
First-generation hybrids. See Hybrids, first-generation.
Flies, heterozygosis 42
Flowers, relation of characters to heterozygosis 13, 17, 28, 32, 38, 45
Focke, W. O., on effects of crossing 8, 11-13, 50
Forestry, practical use of heterozygosis 48
Fowls, effects of selection 34
Fruits, relation of characters to heterozygosis 32
Fungi, nuclear fusions . ' i 39
Gardner, F. D., and Morrow, G. E., on commercial utility of heterozygosis 47
Gartner, C. F., on inbreeding and crossbreeding 9-11, 13, 50
Generations, successive, relation to heterozygosis. . . 14, 16-26, 35, 36, 37, 39, 43, 45, 48
243
INDEX. 55
Page.
Genotypes, relation to heterozygosis 18, 20-21, 33, 34, 36, 39, 45
Germination, relation to heterozygosis 28
Gesneraceae, heterozygosis 12
Geum sp., heterozygosis 9
Gladiolus spp., heterozygosis 12
Gooseberry, heterozygosis 48
Grape, heterozygosis 48
Grasses, heterozygosis 39
Guaita, G. von, on inbreeding 42, 50
Gymnosperms, application of heterozygosis 8
Hanel, E. , on selection 34, 50
Hardiness, relation to heterozygosis 12
Harris, J. A . , on the genotype theory 34, 50
Hartley, C. P., on commercial utility of heterozygosis 47
Hayes, H. K., and East, E. M., on heterozygosis in relation to heredity. 17, 35, 47, 49, 50
Helianthemum spp., heterozygosis 12
Herbert, W., on effects of crossing 9, 50
Heredity, relation to heterozygosis 33-34, 36, 43
Hermaphroditism, relation to heterozygosis 41, 42
Heterozygosis, bibliographic list of authors cited 49-51
characters affected 7-8, 31-32
experimental study. See Experiments.
interpretation of results of experiments 7, 32-39
investigations, summary 8-13, 17-19
statement of the problem 8
value in evolution and plant breeding 7, 43-48
work of Darwin 13-17
Hibiscus sp., heterozygosis 12
Hieracium, apogamy 39, 44
Hippeastrum spp., heterozygosis 12
Hogs. See Swine.
Homozygosis, relation to development 8, 16, 17, 20-21, 32, 36, 37, 42-44, 46
Hop, heterozygosis 39
Horses, heterozygosis 42
Human beings. See Mankind.
Hybridization, significance of heterozygosis 7-13, 19, 28, 33, 35, 36, 38, 40, 47, 48
See also Cross-fertilization.
Hybrids, first-generation, utility of heterozygosis 19, 24-25, 31, 39, 40, 46, 48
Hydra, effects of selection 34
Impatiens spp. , heterozygosis 32
Impotence. See Sterility.
Inbreeding, relation to heterozygosis 7, 13, 15, 26, 32-33, 36-38, 40-43, 46
See also Self-fertilization.
Indian corn. See Corn, Indian.
Inheritance, relation to heterozygosis 21
Insects, relation to heterozygosis in plants 14
Introduction to bulletin 7-8
Ipomoea spp., effects of inbreeding 14, 15, 16
Isoloma spp. , heterozygosis 12
Italy, source of plants for study of heterozygosis 27
243
56 HETEROZYGOSIS IN EVOLUTION AND PLANT BREEDING.
Page.
Jennings, H. S., on effects of selection 34, 39, 50
Jensen, P. , on effects of selection 34, 50
Johannsen, W., on the genotype theory 33, 34, 35, 36, 44, 50
Jost, L. , on effects of crossing 19, 50
Keeble, F., and Pellew, C, on crosses 19, 39, 50
Knight, T. A., on crosses. 9, 11, 12, 50
Klotzsch, M., experiments to utilize heterozygosis 48
Knuth, P. , on pollination 11, 50
Kolreuter, J. G. , on crosses 8-10, 50
Lang, A., on heterozygosis in its relation to heredity 35, 50
Lavatera sp. , heterozygosis 9-10
Lecoq, H., on heredity 12, 50
Legumes, heterozygosis 13, 14
Linaria spp. , heterozygosis 12
Little, G.G., and Castle, W. E., on yellow mice 37, 49
Lobelia spp., heterozygosis 9-10
Longevity, relation to heterozygosis 12
Love, H. H., on effects of selection 34, 50
Luxuriance. See Development, Vigor, etc.
Lychnis spp., heterozygosis 9-10
Lycium spp., heterozygosis 9-10, 12
Lycopersicum esculentum. See Tomato.
Maize. See Corn, Indian.
Malva spp., heterozygosis 9-10
Mammals, heterozygosis 42
See also names of different animals; as, Cattle, Swine, etc.
Mankind, heterozygosis 40-41
Matthiola spp. , heterozygosis 10
Mauz, E., on heterozygosis in its relation to heredity 9, 50
Mendelism in its application to heterozygosis 8, 13, 17, 20-21, 31, 33, 34-35, 40, 41
Mice, heterozygosis 37
Miniums spp., heterozygosis 14, 17
Mirabilis spp., heterozygosis 9, 10, 12, 13
Morrow, G. E., and Gardner, F. D., on commercial utility of heterozygosis 47
Miiller, H., on cross-pollination 11, 51
Narcissus spp., heterozygous behavior .' 12
Nathusius, experiments on swine 43
Nicotiana spp., heterozygosis 9-10, 12, 17, 26-32, 38
Nillson-Ehle, H., on inheritance of quantitative characters . 35, 51
Normal differences. See Differences, normal.
Nuphar spp., heterozygosis .. 12
Nymphaea spp., heterozygosis 12
Oats, heterozygosis 35
Orchids, heterozygosis 11
Papaver spp., heterozygosis 12
Paramaecium, selection , 34, 39
Parthenogenesis, relation to heterozygosis 42
243
INDEX. 57
Page.
Passiflora spp., heterozygosis 12
Pea, heterozygosis 15, 34, 39, 47
Peach, heterozygosis 48
Pear, heterozygosis 48
Pearl, R., on effects of selection on fowls 34, 51
Pearson, K., on the genotype theory 34, 51
Pelargoniums, heterozygosis 37
Pellew, C, and Keeble, F., on effects of crossing 19, 39, 50
Penstemon spp. , heterozygosis 9-10
Petunia spp., heterozygosis 9-10, 14
Pinus spp., heterozygosis 12, 48
Pisum spp., heterozygosis 15
Plant breeding. See Breeding, plant.
Plants, asexual reproduction as related to heterozygosis 48
heterozygosis 32-39
See also names of different plants; as, Corn, Tobacco, etc.
utility of heterozygosis 7-8, 46-48
Pollination. See Fertilization.
Potamogeton spp., heterozygosis 12
Potatoes, heterozygosis 34, 48
Productiveness, relation to heterozygosis 17-18, 22, 26
See also Fertility, Vigor, Yield, etc.
Propagation. See Reproduction.
Pteridophytes, application of heterozygosis 8
Pumpkin, heterozygosis „ 47
Quercus spp., heterozygosis , 12, 48
Raspberry, heterozygosis 48
Reproduction, application of heterozygosis 7-8, 9, 12, 36, 39, 43-44, 48
Reptiles, specializations as related to heterozygosis 43
Rhododendron spp., heterozygosis 12
Ritzema Bos, J., on inbreeding 42, 51
Rosa spp., heterozygosis 12
Rubus spp. , heterozygosis _ 12
Sageret, A., on heterozygosis in its relation to heredity 9-10, 51
Salix spp., heterozygosis 12
Sanborn, on commercial utility of heterozygosis 47
Seed, utility of heterozygosis in production 47-48
Segregation, relation to heterozygosis 35, 45
Selection in its relation to heterozygosis 31, 33-35, 38, 42
Self-fertilization in its relation to heterozygosis. 7, 9,
11, 13, 14, 15, 16, 17-32, 38, 41, 42, 45, 46
See also Fertilization.
Sex, differentiation as related to heterozygosis 43^4
Sheep, heterozygosis 40, 42
Shull, G. H., on heterozygosis in its relation to heredity 17-19, 34, 38, 46, 47, 51
Sibs, crossing, relation to heterozygosis 18-19
Size, relation to heterozygosis : 7-10, 12, 16, 17, 18, 19, 28, 32, 39
See also Development, Vigor, etc.
Squash, heterozygosis 47
Sterility, relation to heterozygosis 10, 24, 28-30, 37, 40
243
58 HETEROZYGOSIS IX EVOLUTION AND PLANT BREEDING.
Page.
Strawberry, heterozygosis 39
Structure, floral, relation to heterozygosis 13
See also Flowers.
Sugar cane. See Cane, sugar.
Swine, heterozygosis -... 40, 42, 43
Tallness, relation to heterozygosis 39
Tammes, T., on inheritance of quantitative characters „ _ . . 35, 51
Taraxacum, apogamy 39, 44
Theory, interpretation of heterozygous phenomena 7-8, 32-39
Tobacco, heterozygosis 14, 32
See also Xicotiana spp.
Tomato, heterozygosis 32, 47
Tropaeolum spp., heterozygosis 9-10, 12
Truck crops. See Crops, truck.
Ulmus spp., heterozygosis 12, 48
Vegetative vigor. See Vigor.
Verbascum spp., heterozygosis . 9-10, 12
Vigor, relation to heterozygosis 7-12,
13. 15, 17-19. 21-22, 24, 28, 29, 31-32, 37, 38, 39, 40, 42
See also Development, Size, etc.
Vilmorin , M. L. , on methods of selection 34, 35, 46
Weismann, A., on inbreeding .- 33, 36, 42, 43, 44, 51
Westermarek, on incest 40
Wheat, heterozygosis 14, 35
Wiegniann , A. F. , on crosses 9-10, 51
Wind, relation to heterozygosis in- plants 36
Winkler, on graft hybrids 34
Woltereck, on selection in Daphnia 34
Woodruff, L. L., and Baitsell, G. A., on Paramaecium. 38-39, 51
Yeasts, effects of selection : 34
Yield of corn, relation to heterozvgosis 22-25
'J ov
Zea mays. See Corn, Indian
243
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