THE ESTABLISHMENT OF VARIETIES IN COLEUS BY
THE SELECTION OF SOMATIC VARIATIONS.
BY
A. B. STOUT,
Director of the Laboratories, New York Botanical Garden.
WASHINGTON, D. C.
Published by the Carnegie Institution of Washington
1915
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THE ESTABLISHMENT OF VARIETIES IN COLEUS BY
THE SELECTION OF SOMATIC VARIATIONS.
BY
A. B. STOUT,
Director of the Laboratories, New York Botanical Garden.
WASHINGTON, D.
Published by the Carnegie Instituti
1915
THIS BOOK IS DUE ON THE DATE
INDICATED BELOW AND IS SUB-
JECT TO AN OVERDUE FINE AS
POSTED AT THE CIRCULATION
DESK.
)00M/5-79
Carnegie Institution of Washington
Publication No. 218
Copies of this Book
were first issued
OCT 7 1915
PRESS OF GIBSON BROTHERS, INC.
WASHINGTON, D. C.
TABLE OF CONTENTS.
Page.
Introduction and Historical Review of Literature 3
The Problem in Coleus 12
Method of Recording Results 13
Method of Handling Cultures 16
General Survey of the Variations 16
Constancy of the Various Patterns 20
Plants with yellow-red blotched pattern 21
Plants with green-yellow-red blotched pattern 21
Plants with color pattern green-yellow spotted-red blotched and with uniformly entire
leaves 24
Plants with laciniate leaves 26
Plants with green-red blotched pattern 29
Plants with pattern yellow-green-red blotched 31
Plants of color pattern green-yelloiv-solid red 32
Plants with green-solid red pattern 32
Plants with green-yelloiv-solid red upper center 33
Plants with pattern yellow-green-solid red 34
Plants with pattern green-yelloio 34
Plants with pattern green and pattern green-yellow spotted 34
Plants with pattern yellow-green 35
Range of the Variations 37
Frequency of the Bud Variations 39
A. Changes involving yellow and green 39
I. Increase of yellow and decrease of green.
II. Decrease of yellow and increase of green.
III. Reversal of the relative positions of green and yellow.
B. Changes involving the epidermal red 40
I. Increase of epidermal red.
II. Decrease of epidermal red.
C. Changes involving leaf-shape 41
Summary and comparisons 41
Distribution of the Bud Variations among Different Clones 43
Environmental Influence 47
Seed Progeny 49
History of Coleus 51
Discussion . . . , 58
Summary ' 74
Bibliography 77
Explanation of Figures in Plates 79
1
THE ESTABLISHMENT OF VARIETIES IN COLEUS BY THE
SELECTION OF SOMATIC VARIATIONS.
By a. B. Stout.
INTRODUCTION AND HISTORICAL REVIEW OF LITERATURE.
Judgment as to the genetic constitution of a plant is based on the
expression of characters in a plant itself, in members of a selfed progeny
and in a hybrid progeny. It is largely through the study of the last
named that there has developed the conception that characters are
represented in cells by unit factors. In considering the evidence as to
whether these assumed factors are in any sense units, it is highly
essential that the individual be studied as to the variability of char-
acters and the range of expressions exhibited in homologous parts.
For this purpose so-called bud variations are especially significant,
since they represent, perhaps, the extreme of spontaneous somatic
variability and suggest that quite as marked variations as exist in
hybrid progeny may develop in the parts of a single individual.
In an excellent summary of all cases of bud variation known to him,
Darwin (1868) shows that the phenomenon is widely distributed in
the plant kingdom and that it may affect various parts of a plant. He
drew the highly interesting conclusions that they include: (1) rever-
sions to characters not acquired by crossing, (2) reversions in hybrids
to parental characters, and (3) cases of spontaneous variability.
The latter he considered as of the same sort as appear in seed progeny.
According to Darwin's views, "long-continued and high cultivation"
are conditions that induce bud variations, but he recognized that cer-
tain cases, especially those when only single buds or parts of buds are
changed, do not seem to be due to external conditions. Darwin con-
sidered in general that bud variations are evidences of the almost
unlimited variability that plants manifest, due to the nature of hving
structures and the exciting causes of environment.
Darwin did not believe in fixed hereditary units. He held that
characters may respond directly to the effects of environment, and also
exhibit spontaneous variability both in seed and in vegetative propa-
gation. Furthermore, sexual hybridization was considered to be of
influence in modifying and changing inherited characteristics.
De Vries (1901, vol. i, p. 39) considers that bud variations are spon-
taneous changes most common in varieties with incompletely fixed
characters. He includes (1889, p. 13) these phenomena under the head
of "dichogeny," a general conception proposed for cases in which the
nature of the organ may be determined, as he assumes, by external
3
ESTABLISHMENT OF VARIETIES IN COLEUS
conditions, but he also assumes that the internal constitution may
admit of development in several directions.
De Vries discusses bud variation with special reference to variega-
tion. For the frequent and striking cases of the production of varie-
gated branches on green plants and the development of green branches
on variegated plants, he offers the old explanation of "latent poten-
tiality." They belong in general with mutations in that they appear as
clear-cut discontinuous variations; the former are classed as progressive
mutations and the latter retrogressive (1901, vol. i, p. 606).
De Vries makes a most comprehensive analysis of the nature of
variegation and concludes (1901, vol. i, p. 616) that the capacity for
variegation is more widely distributed in the plant kingdom as a latent
or semilatent character than perhaps any other character. He notes
that true aurea varieties are few and are remarkably constant. Most
variegated races show rather wide fluctuations and constitute what he
calls intermediate races. His scheme (1901, vol i, p. 424) of represent-
ing the relationship is as follows :
Normal
character.
Anomaly.
I. Original species, preen
Active.
Active.
Semilatent.
Latent.
Latent.
Semilatent.
Active.
Active.
II. Half race, rarelv variegated
III. An equilibrium is maintained.
IV. Eversporting variety, variegated ....
V. Constant varietv. aurea
In a later pubhcation (1913) de Vries considers that the pangens,
which he assumes are directly concerned with the transmission and
expression of characters, may be not only active, semilatent, and latent,
but also labile.
The transition from constant green varieties through variegated
varieties to constant aurea varieties is conceived to be dependent on
the degree of activity displayed by antagonistic qualities within the
cells. The essential changes are conceived to be intracellular and not
dependent on qualitative cell-divisions; all the cells are potentially
aUke, but different processes within the cells give differences in expres-
sion. Thus the conception of de Vries does not consider that the hered-
itary physiological units (pangens) are fixed and uniform. They are
subject to effects of environment and thej- exhibit spontaneous change
even to the degree of sudden appearance bj' progressive mutation.
All these phenomena may be exhibited either in vegetative or in seed
progeny. Sexual hybridization may entirely modify or change the
effect and nature of the basic physiological units of heredity. De
Vries's general attitude, however, places the emphasis on discontinuous
or mutational changes as the only really stable variations.
BY THE SELECTION OF SOMATIC VARIATIONS. 5
The exhaustive summary of facts given by Cramer (1907) fully
substantiates the views of Darwin and de Vries as to the main classes
of variegation and the remarkable gradation in degrees of constancy
and inconstancy which are exhibited by the various types and by their
bud variations. Cramer gives the most complete and detailed sum-
mary of the known cases of bud variation that has been published.
He includes (p. 18) under the term " Knospenvariation" all cases where
a character suddenly changes in a plant in a way that can not be attrib-
uted to environmental influence. He considers that there are three
main classes of bud variation: (1) vegetative segregations in hybrids;
(2) intermediate-race bud variations; and (3) vegetative mutations.
Recessive characters in hybrids may, he assumes, separate out by
vegetative cell-divisions, and dominant characters which have been
latent may reappear. His conception of the variability of characters
in an intermediate race is the same as that of de Vries. Characters
concerned in bud variations in intermediate races show, it is considered,
great irregularity in expression. The bud variations which give a seed
progeny quite constant for the character involved are classed as vege-
tative mutations. This classification emphasizes the fact that charac-
ters involved in hybridization may come into expression quite irregu-
larly, and that spontaneous and fluctuating variations are common in
vegetative development.
Cramer devotes a most interesting and instructive chapter to varie-
gated plants, showing, especially, their wide distribution in the plant
kingdom, the wide fluctuation in the degree of constancy of their seed
progeny, the range from vegetatively constant to inconstant types, and
the frequency of bud variations in variegated plants. He considers (pp.
126, 127) that loss of variegation can occur in two ways: (1) by atavistic
bud variations and (2) through influence of external conditions. He
notes that it is very difficult to distinguish fluctuating variability from
mutation.
The general variability of the characters concerned with variegation
is well shown in his discussion of changes that may occur in a single
plant, such as the following: In apparently inherited types of variega-
tion, the seedlings are often green at first. Seedlings may have at
first variegated cotyledons, then a few green leaves, and then variegated
leaves. Some biennial plants are pure green during the first year of
growth, but variegated in the second year. Various plants exhibit
a periodicity in their variegation, being green in spring and variegated
later in the summer, or vice versa. He gives numerous cases of the
appearance of variegation by bud variation on green plants resulting
in new varieties or in the duplication of types already known. These
exhibit various degrees of constancy.
Various adherents of Mendelian doctrines have more recently dis-
cussed the transmission of characters appearing as bud variations,
b ESTABLISHMENT OF VARIETIES IN COLEUS
attempting to explain them, as they do all heritable variation, by the
presence or absence of a unit factor. While this is the general attitude
of those who have investigated the inheritance of variegation and the
nature of bud variations involving variegation, the results of their studies,
as a whole, show great diversity and many necessary modifications of
the general Mendelian doctrine of the integrity of unit characters or
unit factors and the purity of the segregations of such assumed factors.
East (1908, 1910a) considers that a large majority of the known
cases of bud variation are due to the loss of a dominant character and
that 70 per cent of all known cases are color variations. His discussion
and suggestions do not claim to be comprehensive or critical, and he
excludes from his treatment all cases of bud variations in variegated
plants, because some types of these are known to be pathological. It
is interesting to note that he states that no important potato has arisen
as a bud sport. He reports four cases of bud variation in potatoes
giving white from red or pink which appear to be constant; also several
cases giving colored or purple blotched from pure white, concerning
which no data are given except the statement that they are not con-
stant. East's view is that bud variations are due to loss or latency of
hereditary units that stand for characters.
Bateson (1909, p. 273), especially, has advocated, on theoretical
grounds largely, that bud variations are due to qualitative cell-divisions
in somatic tissues, giving somatic segregation of unit factors. The
idea is quite identical in its main features with Weismann's concep-
tion of qualitative divisions, giving tissue differentiation in ontogeny.
In considering the nature of the albomarginate types of variegation,
with special reference to the origin and the development of the green
and white areas, Baur (1909) has made criticial anatomical studies of
a white-margined Pelargonium zonale which indicate very clearly the
relationship of white and green tissues as periclinal chimeras and
explain the appearance on them of branches wholly green or white.
In testing, by crossing experiments, his assumption that the green and
white tissues are each pure, he is forced to the further assumptions that
the loss of green in this case is due solely to the condition of the chro-
matophores and that male sex-cells carry chromatophores. He does
not consider that white cells can arise spontaneously from green, or
vice versa. Yet this probably did occur when the chimera was first
produced. In fact, the numerous varieties that are peripheral chimeras
not only in Pelargonium but also in other genera indicate that such
spontaneous loss of power to produce green is frequent. We may say
that Baur's interesting results, however, do show that if spontaneous
loss occurs in young leaves after they are formed, mottled or striped
variegation is produced, but if the loss occurs in the growing-point itself,
then chimeras will result, their constancy depending largely on the
relative permanency of the change.
BY THE SELECTION OF SOMATIC VARIATIONS. 7
Later, Baur (1910), in studies of seed progeny of variegated types
that appeared spontaneously, assumes that green is the combined
result of three factors. If a certain one (Z) is absent the tissue is color-
less, if Z and Y only are present then a chlorina type is produced, and
if N is present with Z and heterozygous Y, the aurea coloration results.
In Antirrhinum ma jus alhomaculatum, Baur found 8 types of variega-
tion arising spontaneously in cultures of about 30,000 pure-green
plants. His crosses with these show that certain cases appear to be
inherited only from the seed parent. To explain the results, Baur
assumes that hereditary qualities are localized in different parts of the
cell. The nucleus, the cytoplasm, and the chromatophores all possess,
he considers, different but definite factors concerned with variegation.
Such results and conclusions illustrate very well the difficulties and
uncertainties which arise from attempts to analyze variations in terms
of unit factors and suggest most forcibly the need of a more thorough
investigation of such variations in a progeny derived by vegetative
propagation.
The variations among the branches of a single plant which Correns
(1909, a and h) reports in connection with variegated types of Mirabilis
jalapa are apparently quite similar, in degree at least, to those I shall
report for Coleus. It would seem to be highly important that the
inheritance of these variations be studied in vegetative propagation.
Correns, however, made a study of seed progenies only. In the case
of the albomaculata type these were composed of green, white, and
albomaculata plants in quite different ratios for different plants tested.
All these classes appeared when pollen from a pure-green plant was
used, but when pollen from the albomaculata was used on pure green
there was no transmission of the quality of variegation. This case of
matrocliny is due, he assumes, first, to the localization in the cytoplasm
of the factor for variegation, and second, to the condition that male
sex-cells in this case do not carry cytoplasm.
ShuU's (1914) studies with variegated types of Melandrium show
much the same results as those of Baur and Correns. He distinguishes
between chlorina, pallida, and pure-green types of Melandrium on the
basis of presence or absence of three factors which in crosses behaved
quite like units. In types with green-white blotched and with chlori-
nomaculata variegation, however, the variegation seemed to be trans-
mitted only through the seed parent, but not uniformly, for crosses of
variegated branches with pure green gave in the Fi generation plants
ranging from pure green through types of chlorinomaculata to yellowish-
green plants. It is highly interesting that Shull found that the greater
the amount of chlorina coloration in the calyx the greater was the num-
ber of variegated seed progeny. The aurea types, which possessed as
a rule small round flecks, gave such varied results with appearance of
different types in Fi progeny that Shull concludes it must be an infec-
8 ESTABLISHMENT OF VARIETIES IN COLEUS
tious variegation which is transinitted through both germ-cells to a
part of the progeny.
Among the most interesting series of observations especially bearing
on the behavior of red coloration in Coleus and most important in the
consideration of the nature of variegation and the character of somatic
variation and the relations of these to seed progeny, is that of Emerson
(1914). The variegation in question is that of pericarp color in certain
"calico" races of corn. The extent of coloration varies widely, ranging
from solid red through every degree of striping and blotching to entirely
non-red, both for ears as a whole and for single kernels on the same ear.
Emerson made careful studies of the progeny of seeds having different
degrees of coloration. His results show a wide range of variation in
the progeny of kernels that appear to be identical. Selected solid-red
kernels from "freak ears" of unknown parentage gave, in some cases,
progeny with only solid-red or non-red ears, and in other cases the
plants produced solid-red, variegated, or non-red ears. Variegated and
white kernels (data not given separately) gave either variegated and
white, or red, variegated, and white. Again, from two ears, kernels of
white gave progeny pure white, and red kernels gave red and white
only, each of which gave afterward a constant progeny.
In selfed variegated strains, kernels of all classes from solid-red to
non-red gave progeny with ears solid-red or variegated, but none
with no-red ears.
Data are given collectively for progeny of 5 solid-red ears (selfed).
These gave solid-red and variegated (p. 18). Progenies of only two
plants of these solid-reds are reported. One gave again plants with
solid-red or variegated ears, the other gave only solid red. The num-
bers grown in this generation were respectively 9 and 16. Emerson
considers from these data that, in general, red-eared plants behave,
judged by progeny, as if they were hj'brids either between solid-red and
variegated or sohd-red and white races.
The data show quite clearly, as Emerson points out, that the more
red there is in the seed planted the larger the percentage of red ears
in the progeny. The variegated race is therefore far from constant.
Selection from red kernels and from red ears give a strain quite constant
for solid-red, but Emerson's data are far from conclusive that a pure
solid-red strain was obtained in this way.
The results of crossing the variegated race with non-red races are
interesting. WTien the former was the pollen parent, the Fi progeny
gave red, variegated, and non-red ears, but Emerson states that some
of the latter may have been extremely light types of variegation.
From the reciprocal cross selected kernels for the Fi gave red and
variegated or only variegated.
In the selections from hybrid stock, as in that in selfed stock, the
seed that had more red gave greater numbers of pure-red progeny.
BY THE SELECTION OF SOMATIC VARIATIONS. 9
Although there is this irregular expression of solid-red, variegated,
and non-red kernels on the same plant and among different plants of
the same generation, and although selection from solid-red tends to
give greater numbers of solid-red progeny, Emerson concludes that the
factors for solid-red and variegation are "as distinct in inheritance as
any two factors could well be" (p. 33). He points out, however, that
the "factors" concerned are pattern factors, one determining self-
color and the other giving variegation. When a solid-red kernel occurs
in an ear of the variegated race, Emerson assumes that a V factor
changes to S; but when these red kernels do not give solid-red progeny,
he further assumes that in these diploid cells one of the pair of V factors
changes to S and that such somatic changes may affect an area of cells
including macrospores.
Taking this at its full value, we note that as the occurrence of solid-
red kernels is frequent the hereditary factor V is fluctuating and
extremely labile, changing to S readily. For V and S to be distinct in
inheritance under such conditions is hardly conceivable, for they are
not even distinct in cell lineage. Emerson thus reflects the strong ten-
dency of most modern Mendelian investigators to regard their assumed
factors as temporary conditions with quite fluctuating activities.
Such conflicting results as the above, obtained even by careful
pedigree methods of study, may well lead one to question whether our
kaiowledge of the behavior of plant characters in inheritance and
expression has advanced much beyond the views of Darwin.
Such studies may well establish the general breeding values of
certain characters in particular cases which are of practical significance.
The more refined methods of pedigree have shown that plants with
identical appearances may give quite different progeny, and that
selection for relative degrees of constancy should be made in pedigreed
lines rather than in mass selection. Mendelian results also indicate
the possibilities of hybridization followed by selection in pedigreed
lines of hrybrid progeny.
In its theoretical significance, however, Mendel's original work has
two points of special interest. First, it embodied the conception that
all structures of Hke character are due to a single hereditary unit; for
a specific example, all the wrinkles of all the peas on a plant were con-
ceived as represented in the germ-cells by a single unit. This was in
decided contrast to Weismann's general view that each wrinkle is
represented in the germ-cells. The second point of special interest
pertains to the behavior of these assumed units in the formation of
reproductive cells and their behavior in fertilization. Mendel assumed
that they segregated as units independent in behavior and pure in
composition from the unit representing the contrasting character. By
the conception of the purity of the segregations of hereditary units, each
representing only an entire and complete quality or character, Mendel-
10 ESTABLISHMENT OF VARIETIES IN COLEUS
ism must be judged. It is not a question of the appearance of parental
characters in hybrid progeny, but of the purity of those segregations.
Mendel himself greatly modified the conception, gained from his
studies with peas, that an entire quaUty is represented by a single
unit. In Phaseolus he crossed P. nanus having white flowers with
P. multifiorus having purple flowers. The Fo generation of 31 plants
had flowers ranging from white to pale violet and purple red of various
grades. Only one had white flowers like those of P. nanus. He sug-
gests that the color of flowers (and seeds as well) in P. multifiorus
*'is a combination of two or more entirely independent colors" (Mendel,
p. 367). To Mendel, therefore, should be given credit for the con-
ception of multiple factors, later developed especially by Nilsson-Ehle
(1909) and by East (19106).
Very soon after the so-called rediscovery of Mendel's law for the
behavior of certain characters of Pisum in hybridization, it was noted
that new quahties frequently appear in the F2 generation, as Mendel
found was the case in beans. The presence-and-absence theory
developed by Bateson and Punnett (1905) attempted to account for
this increased variabiUty giving the appearance of new types. This
gave chance for dihybrid ratios from what was apparently a simple
pair of contrasting characters, the "absences" segregating out into
expressed characters new at least to the immediate ancestry.
The next important modification of the general view was the apph-
cation of Mendel's idea of what we now call multiple factors. Nilsson-
Ehle (1908 and 1909) found that certain crosses in cereals gave in the
F2 progeny large numbers of one parental type and relatively few of
the other type. For example, the apparently simple cross of white-
kerneled wheat (Predel) with a red-kerneled sort (Swedish Velvet
Chaff) gave an F2 progeny of about 63 red-kerneled plants to 1 white.
Nilsson-Ehle's explanation assumes that the red character in the
Swedish Velvet Chaff is due to three independent factors which are
each of equal value and that any one can produce the same effect as all
three. We note that the variabiUty of the F2 generation was not
increased over that of parents, but that the ratio showed almost com-
plete appearance of one character.
East (19105) applied the term to the same sort of phenomena as
Mendel did, {. e., to increased variabihty. The apparently simple cross
of white with j^ellow corn gave in the F2 a generation with few white but
with a large number of yellow kernels. The latter, which were of all
gradations of intensity, were grouped into two classes by East.
It should be noted that such a group of ''factors" having among
themselves different values but working together to produce a single
character are not necessarily independent in the fertilizations of the
variety concerned. Inside the variety they go together. When certain
crosses are made they separate. This is but another waj' of saying
BY THE SELECTION OF SOMATIC VARIATIONS. 11
that the hereditary bearers of characters appear to be spht up and
modified by crossing, giving in some cases quite new characters.
Thus in the period of less than 20 years the hterature passes from the
confident treatment of characters as units with a simple shorthand
representation to a discussion of '^ factor s^^ that under some conditions
work together as a multiple unit and under other conditions separate,
producing equal or different expressions. The assignment of different
values to the assumed factors as diluters, intensifiers, inhibitors, and
the conception of multiple factors that can separate out, giving aber-
rant ratios or new expressions and even almost endless intermediate
gradations, both of so-called qualitative and quantitative characters all
reduced to descriptive terms, add nothing to the fundamental concep-
tions of Darwin regarding variability in hybridization. The extreme
apphcation of the multiple-factor hypothesis simply means that small
variations are inherited equally as well as are large variations.
The greater number of Mendehans, mutationahsts, and adherents
of the doctrines of pure lines seem to hold that the unit factors are
changeless. Others still accept Darwin's general views of the modi-
fiability of the fundamental units. The latter view is especially well
developed by Castle (1912), who states (p. 356):
"In my experience every unit-character is subject to quantitative variation,
that is, its expression in the body varies, and it is clear that these variations
have a germinal basis because they are inherited."
Morgan (1913) considers that factors are labile aggregates subject
to rearrangement, that processes of mutation and reversion are reversi-
ble, and that in eversporting varieties mutation and reversion are regu-
lar processes.
Bateson (1902, p. 201), in a defense of early Mendelian views, makes
the following admission:
"We have to consider the question whether the purity of the gametes in
respect to one or another antagonistic character is or is likely to be in the case
of any given character a universal truth. The answer is unquestionably No,
but for reasons in which ancestry plays no part."
More recently (1914, p. 322) he has expressed the view that the
conception of multiple factors is in his mind an admission that there
are imperfect segregations. To quote further:
"Segregation is somehow effected by the rhythms of cell-division, if such an
expression be permitted. In some cases the whole factor is so easily separated
that it is swept out at once; in others it is so intermingled that gametes of all
degrees of purity may result."
Thus it seems that the short-hand system of representing hypo-
thetical germ-cell units is often not only cumbersome but inaccurate.
Some phases of it may be useful as descriptive terms, but the method
12 ESTABLISHMENT OF VARIETIES IN COLEUS
has led into purely speculative fields in the attempts to represent and ex-
plain imperfect segregations and variations in the hereditary qualities.
Furthermore, it appears in final analysis that the extended studies
of seed progenies have not contributed anything fundamentally new to
the knowledge of the nature of plant characters. At least, the analysis
of characters in terms of hereditary units has failed.
It would seem that, in considering these problems, the studies of
variation among numbers of a seed progeny is no more important
than studies of variation in progenies derived by vegetative propaga-
tion. The latter should give much more conclusive data regarding
such questions as the constancy of characters (or of assumed factors),
the purity of apparent segregations, and the frequency, constancy, and
nature of spontaneous changes in the expression of characters.
It is evident that the facts regarding bud variation involve the funda-
mental questions of heredity. When such variations occur in a plant
that can be propagated vegetatively, there is opportunity to apply the
pedigree method of experimental study to successive generations pro-
duced by vegetation propagation. The nature, frequency, and perma-
nence of such changes as appear can be studied without the complica-
tions that are associated with alternation of generations and fertilization
as they normally occur even in selfed seed progeny. Special evidence
regarding the "expression" of characters, which also bears on the ques-
tion of their inheritance, may thus be obtained.
THE PROBLEM IN COLEUS.
For the study of variation along the lines indicated above, I have
grown a series of 833 plants, all descended by vegetative propagation
from two plants of a variety of Coleus.
Coleus is particularl)' favorable for such study in that bud variations
are frequent and the plant is readily propagated vegetatively. The
leaves are in pairs which alternate on a square stem, making but four
rows of leaves. Bud variations that appear sectorially can thus be
traced with ease. In a young plant lateral branches usually start to
develop from the axils of all the leaves on the main stem. In a large,
bushy plant many lateral buds remain dormant, but by proper pruning
any bud can be forced to develop or it can be propagated as a cutting
which will give it full chance for development. Large, bushy plants
3 or 4 feet tall grown out-of-doors often have a total of as many as
300 branches. In the greenhouse, also, plants can be grown from cut-
tings, with the production of many leaves and branches. The ease with
which the plant is propagated vegetatively makes it possible to grow a
large series of pedigreed plants from any cutting, and to thus test the
frequency with which variations appear, the constancy of the different
types, and the purity of any vegetative segregations, all bearing on an
analysis of the nature and inheritance of the characters concerned.
BY THE SELECTION OF SOMATIC VARIATIONS. 13
METHOD OF RECORDING RESULTS.
To record fully the series of Coleus plants descended through vege-
tative propagation from any plant or any particular branch used as a
cutting, the pedigree method of culture has been used. It may be men-
tioned that the application of the pedigree-culture method to plants
propagated vegetatively is much simpler than its use in seed progenies,
the difficulties of which have been ably presented by Shull (1908).
Each cutting was given a number which indicated its lineage. The
two plants serving as original parents were numbered 1 and 3, and the
first digit to the left in a number of any plant indicates the original
parent plant. For the progeny of plant No. 1, the second digit indi-
cates the particular main lateral branch from which the first cuttings
were taken, and the third digit is the number of the particular cutting.
Additional digits indicate successive plants grown later in the par-
ticular line of descent. Thus, plant 121 (I read these one-two-one,
etc.), 122, and 123 were grown from three cuttings taken from
branch No. 2 of plant No. 1. Plants 1211, 1212, and 1213 were from
cuttings of plant 121, while the numbers 12111, 12121, 12131, etc.,
were given to plants of the next generation of cuttings. Thus the
number of any plant gives its complete pedigree since the experiments
were begun. Particular data regarding the cuttings and the plants
have been recorded on cards and filed in the manner of a card cata-
logue. This enables one to trace readily the inheritance of any varia-
tion through a series of generations and to compare different lines of
descent from any point in the culture.
Following the suggestions of Webber (1903) and Shull (1912) the term
''clone" will be used in speaking collectively of all plants descended from
any one plant or branch. All the plants derived from plant No. 1
constitute a main clone, itself made up of numerous subclones. The
records of pedigree enable me to designate these as clone 11, clone 13,
clone 117, etc. I shall use the term "fine of descent" to include the
different plants that were the parents of any one plant. The term
"generation" refers to the plants that were grown during the same
period.
The observations here reported were made on successive generations
of pedigreed plants derived by vegetative propagation from two original
plants. These parent plants were alike when young in possessing a
color pattern that can be characterized as a mosaic of green, yellow, and
red, with the colors distributed as shown in figure 2.
In referring to the color patterns, it seems best to the writer to use
terms that are sufficiently descriptive to make the matter concrete and
which at the same time are somewhat compact. The color of the
subepidermal tissues in the center of the leaf will be mentioned first, as
green or yellow; the color of the subepidermal tissues at the margin will
be mentioned next, as green or yellow, and last the character of the
14 ESTABLISHMENT OF VARIETIES IN COLEUS
epidermis as red blotched (both surfaces), solid red (both surfaces), or
solid red upper center. In these compound expressions, hyphens will
be used to separate the terms descriptive of each color element. In
the cases with colorless epidermis no reference is made to this condi-
tion. The colors were determined by Ridgway's Color Standards and
Color Nomenclature. The color pattern of the parent plants, in this
paper referred to as a green-yelloiv-red blotched pattern, is a mosaic
made up of a green center and a yellow border with conspicuous
epidermal blotches of red. The yellow is amber yellow and constitutes
an irregular band about the margin of the leaf. The green is a spinach
green and is chiefly massed in the center of the leaf. Over the green
portions the red appears as violet carmine, but over the underlying yellow
areas it is nopal red. The three color elements are in such sharp con-
trast that any marked variation is readily noted. Increase or loss of
either the yellow, green, or red is conspicuous, as one will appreciate
from a glance at the plates that illustrate this article.
Spontaneous bud variations consisting of marked alterations in color
pattern appear either in single leaves or groups of leaves, or in single
branches or groups of branches, affecting the whole or a part of the
leaves or branches. When appearing in a terminal bud, one or more
leaves have a pattern differing from that of the leaves below. When
appearing in a lateral bud, the first leaves of the branch possess a
pattern different from that of the subtending leaf. Those appearing
in a terminal bud have, in all cases observed by the writer, been sec-
torial in the main branch itself. That is, the change has appeared first
in a part of the branch only. These variations carried on into newly
formed branches give plants bearing two, three, or even four distinct
types of foliage, with differences especially marked in cases of single
branches with sectorial distribution of two patterns. The rather
simple arrangement of the leaves and branches in Coleus enables one to
trace the extent of a variation.
This may be illustrated by a variation that occurred in plant No.
1171. In this variation the relative positions of the green and the
yellow became reversed, as shown in the two leaves reproduced in
figures 2 and 6. When the cutting was made in April 1912, all the
leaves had uniformly green centers. On one of the first pair of branches
to develop, however, all of the leaves had the yellow in the center. As
further branches developed, the new pattern appeared in 5 other
branches. The plant produced 13 pairs of branches on the main branch
before it was necessary to remove the terminal bud to insure proper
development of the lateral branches. All of these branches developed at
least to a size sufficient to show the color pattern of the leaves borne.
The 6 branches with the new pattern were contiguous and were located
as indicated in diagram 1 . The plant was transplanted to a large pot
and kept in a greenhouse over winter and then grown out of doors
BY THE SELECTION OF SOMATIC VARIATIONS.
15
during the summer of 1913. All of the leaves produced by the 6
branches in question in the 17 months of growth were uniform and
constant to the new type and were in marked contrast to the fohage
of the other branches. During the summer of 1913, two bud variations
occurred in secondary branches in the upper part of the plant. One
was a sectorial loss of yellow giving type green-red blotched from green-
yellow-red blotched and the other was a complete loss of green in one
branch giving type yellow-red blotched. In September 1913, the plant
bore four distinct kinds of color pattern, viz,
yellow-red blotched (fig. 1), green-yellow red
blotched (fig. 2), green-red blotched (fig. 5), and
yellow-green-red blotched (fig. 6). The bud
variation to type yellow-green-red blotched was
sectorial in the main stem for a vertical dis-
tance of six nodes, but was not complete for
the entire stem, a condition shown in diagram 1 .
The greater number of bud variations first
appeared in single lateral branches and not in a
series of branches on a main stem, as described
above for 1171. Where such a variation was
sectorial in a branch the continued growth gave
more or less irregular extension of the new type.
The parent plant here designated as No. 1
was one of several Coleus plants which were
grown at the New York Botanical Garden
during the sununer of 1911. This plant pos-
sessed in September 1911, when first observed
by the writer, two branches bearing leaves in
which the yellow was apparently almost
entirely absent. These two branches were in the same rank, one
directly above the other. x\bout one-third of the entire foliage of the
plant was borne by these two branches and the marked green aspect
of this part of the plant was in decided contrast to the conspicuous
yellow in the foliage of the rest of the plant. Upon careful examina-
tion, a few yellow spots could be seen in many of the leaves of one of
the green branches (branch 14) quite like those of the leaf shown in
figure 4. The leaves on the other green branch (branch 13) were
apparently free from all yellow areas (fig. 5).
The decided loss of yellow in these two branches constituted the
only variation in the dozen or more plants in this particular bed of
Coleus. To test the constancy of this variation, as well as the reap-
pearance of it and of other variations, pedigreed cuttings were made
from each of four main lateral branches of the plant.
About the same time random cuttings were made from the bed of
plants for stock for general planting. One of these cuttings produced
2
2
2
2
2
2
2
2
2
2
2
2
2
2
6
2
(j
2
6
2
2
6
6
2
2
6
Diagram 1. — Position of the
six branches on plant
1171 having color-pattern
ycllow-green-red blotched
(6) among those ha\ang
green-yellow-red blotched
patterns (2).
16 ESTABLISHMENT OF VARIETIES IN COLEUS
during the winter of 1911-12 a single branch bearing leaves with the
red completely covering both surfaces. This color pattern designated
as green-yelloiv-solid red is shown in figure 8. All the leaves on the
branch were uniform for this pattern and were in most conspicuous
contrast to the rest of the plant. From this plant a number of pedi-
greed cuttings were also made.
METHOD OF HANDLING CULTURES.
The first generation of plants (series 111, 121, 131, 141) was grown
in a greenhouse during the winter of 1911-12. In April 1912, cuttings
(series 1111, 1211, etc.) were made from these. During the summer all
of the plants, both old and young, were grown out of doors in beds.
In the autumn cuttings were again made. The plants developed
from cuttings taken in the autumn were under observation for a year,
7 months of which they were grown under greenhouse conditions.
Cuttings taken in spring were grown only out of doors. This method
of handling gave opportunity to observe development and behavior
under different conditions and to compare old plants with j^ounger
ones. Except for a few plants that were subjected to special condi-
tions, all the plants of any generation were treated uniformly with
respect to kind of soil, size of pots, and conditions of temperature
and illumination. The plants were cut back somewhat to prevent
early blossoming and to maintain a vigorous vegetative condition.
In the period of three years between September 1911 and September
1914, a total of 833 plants were grown to maturity and discarded. All
of these descended through vegetative propagation from plants Nos. 1
and 3, both of which had originally the green-yellow-red blotched pattern
illustrated by figure 2.
GENERAL SURVEY OF THE VARIATIONS.
Variations in the color patterns of the plants both of the original
and the derived types can be classed as fluctuating variations and as
bud variations. In the former the changes were usually quite gradual
and affected in most cases an entire plant. The changes which are
in this paper included in the term ''bud variations" were those affecting
only a part of a plant and usually appearing as a sudden and conspicu-
ous change. In addition to the variations in color patterns, there
appeared, in several subclones, plants which fluctuated in leaf-shape,
giving in extreme cases leaves deeply cut and laciniate. The variations
that appeared were as follows: (A) changes involving yellow and
green; (B) changes involving the epidermal red, and (C) changes involv-
ing leaf-shape. The bud variations can be grouped as in table 1 with
data regarding the number of plants concerned and the number of
times the different changes appeared as a bud variation.
North Carolina Sti^Library
. i^teigh
<:■
BY THE SELECTION OF SOMATIC VARIATIONS.
17
Table 1. — Types and freqiiency of Ihe bud variations.
Types of bud variation.
a
a .
1^ o
3
c
2.
3.
4.
II.
B.
Changes involving yellow and green:
I. Increase of yellow and decrease of green:
1. Yellow-red blotched (fig. 1) from green-yellow-red blotched (fig. 2)
Yellow-red blotched (fig. 1) from green-yellow spotted-red blotched (fig. 4)..
Yellow-red blotched (fig. 1) from yellow-green-red blotched (fig. 6)
Yellow-solid red from green-yellow-solid red (fig. 8)
Green-yellow-red blotched (fig. 2) horn green-yellow spotted-red blotched (fig. 4)
Green-yellow-red blotched (fig. 2) from laciniate, green-yellow spotted-red
blotched
Green-yellow spotted-red blotched (fig. 4) from green-red blotched (fig. 5) . . . .
Spontaneous yellow from green-red blotched (fig. 5)
Decrease of yellow and increase of green:
1. Green-yellow spotted-red blotched (fig. 4) from green-yellow-red blotched (fig. 2)
Green-yellow spotted-red blotched (fig. 4) from yellow-green-red blotched (fig. 6).
2. Green-red blotched (fig. 5) from green-yellow-red blotched (fig. 2)
Green-red blotched (fig. 5) from green-yellow spotted-red blotched (fig. 4) . . .
Green-red blotched (fig. 5) from laciniate, green-yellow spotted-red blotched . .
Green-red blotched (fig. 5) from yellow-green-red blotched (fig. 6)
Green-solid red (fig. 9) from green-yellow-solid red (fig. 8)
Reversal of position of green and yellow:
1. From yellow border to yellow center
a. Yellow-green-red blotched (fig. 6) from green-yellow-red blotched (fig. 2)
b. Yellow-green-solid red (fig. 11) from green-yellow-solid red (fig. 8) . .
Changes invohnng the epidermal red:
I. Increase of red:
1. Green-yellow-solid red (fig. 8) from green-yellow-red blotched (fig. 2)
2. Green-yellow spotted-solid red from green-yellow spotted-red blotched (fig. 4) .
3. Green-solid red (fig. 9) from green-red blotched (fig. 5)
Decrease of red :
III.
II.
1.
2.
3.
Green-yellow spotted from Uiciniate, green-yellow spotted-red blotched. . . .
Green-yellow spotted from green-yellow spotted-red blotched (fig. 4)
Green-yellow (fig. 12) from green-yellow-red blotched (fig. 2)
Green-yellow (fig. 12) from green-yellow-solid red (fig. 8)
Green (fig. 13) from green-red blotched (fig. 5)
Green (fig. 13) from green-solid red (fig. 9)
Yellow-green (fig. 14) from yellow-green-red blotched (fig. 6)
Decrease of red with concentration in epidermis of upper surface:
1. Green-yellow-solid red upper center (fig. 10) from green-yellow-solid red.
C. Changes involving leaf-shape:
1 . From entire to periodically laciniate
2. From periodically laciniate to constantly entire
5.
III.
337
198
41
54
198
68
90
90
337
41
337
198
68
41
54
337
54
337
198
90
68
198
337
54
90
8
41
54
765
68
7
1
3
1
3
1
2
9
8
1
24
2
1
10
4
6
2
4
2
2
1
2
10
1
3
1
1
13
1
The names given to the different patterns embody the principal
features of coloration on the basis explained above (see pp. 13 and 14).
The patterns selected are with one exception those that appeared as con-
spicuous bud variations and which are sufficiently distinct for ready
identification. Numerous other types that are intermediate between
the types given could also be designated by still more exact classifica-
tion. The following descriptions, together with the colored plates
18 ESTABLISHMENT OF VARIETIES IN COLEUS
illustrating the types as classified, will enable the reader to visualize
the patterns referred to by name.
Color pattern yellow-red blotched (fig. 1): Leaves almost entirely
amber yellow with only very limited and scattered areas of greenish
tissue. The island-like areas of green are surrounded by yellow.
Irregular-shaped blotches of nopal red are scattered over both upper
and lower surfaces. This decidedly yellow pattern was derived from
the several patterns, as shown in table 1, by a sudden and a conspicu-
ous loss of green tissue.
Color pattern green-yelloiv-red blotched (fig. 2) : This is the pattern
possessed originally by the two parents and has already been described.
Color pattern green-yellow spotted-red blotched (fig. 4) : In this pattern
there is no definite border of yellow. The yellow appears in rather
limited and somewhat scattered areas, sometimes nearly limited to the
border zone, but often quite generally distributed throughout the leaf.
The pattern is, therefore, decidedly greener in appearance than that
of the parental type.
Color pattern green-red blotched (fig. 5) : This is a bicolored pattern
of green and red. As there is no underlying yellow the epidermal red
appears uniformly as violet carmine. This t^-^pe arose frequently on
plants with patterns containing yellow by what was apparently a com-
plete loss of yellow.
It may be noted that in the four patterns as arranged above there
is an increase of green and a corresponding decrease of yellow, with
the distribution of the epidermal red quite uniform. The yellow-red
blotched pattern gives the extreme development of j^llow with almost
complete absence of green. The green-red blotched pattern has appar-
ently a complete loss of yellow. The green-yellow-red blotched and the
green-yellow spotted-red blotched patterns are gradations between these
extremes.
Color pattern yellow-green-red blotched (fig. 6) : This is a pattern
of green, yellow, and red as in type green-yellow-red blotched, but the
relative positions of the green and the j^ellow are reversed. The
3^ellow is in the central portion of the leaf.
Color pattern green-yelloir-solid red (fig. 8) : Both surfaces of the
leaf are a solid red. Through the center of the leaf the color is
violet carmine, but the marginal zone underlaid by j^ellow is nopal red.
At the base of the leaves a greenish tint prevails and at the extreme
edge of the margin a fine line of yellow is visible. On the under sur-
face the red seems slightly less intense and does not cover the larger
veins, which stand out prominently^ on this surface. This pattern
differs from green-yellow-red blotched in having the entire epidermis
solid red instead of blotched. Frequently, however, a few isolated
areas are free of epidermal red and the underlying green or yellow
shows clearly.
BY THE SELECTION OF SOMATIC VARIATIONS. 19
Color pattern green-yellow spotted-solid red: This pattern has the
solid red as in the preceding type, but the conditions of yellow and
green are as in type green-yellow spotted-red blotched.
Color pattern green-solid red (fig. 9): This pattern has the entire
leaf above and below of a uniform violet carmine. It differs from type
green-yellow-solid red in the absence of any underlying yellow, and from
type green-red blotched in having the epidermis completely red instead
of red in blotches. The pattern is dull and dark, with a somewhat
metallic luster, in marked contrast to the various patterns with yellow.
Color pattern green-yellow-solid red upper center (figs. 10 and lOo) :
This is a brightly colored and attractive pattern with a rather com-
plicated arrangement of colors. The subepidermal colors of green
center and yellow border are similar to the arrangement in the types
green-yelloiv-red blotched and green-yellow-solid red. The epidermal
red is, however, almost entirely confined to the upper surface; over
the central green it gives a greenish violet carmine cast; over the border-
ing yellow it forms a band of nopal red. At the extreme margin it is
absent, giving a narrow but irregular band of pure yellow. On the
under surface there are only occasional small blotches of red. About
the border the red of the upper epidermis shows through the yellow,
giving pale pinkish tints, as shown in figure 10a. This type was
derived from pattern green-yellow-solid red, with its complete covering
of epidermal red, by the loss of red on the under surface and about
the extreme margin of the upper surface.
Color pattern yelloiv-green-solid red (fig. 11): This pattern w^as
derived from type green-yellow-solid red by a reversal in the relative
position of the underlying green and yellow, the change being the same
that gave type yellow-green-red blotched from pattern green-yellow-red
blotched.
Color pattern green-yellow (fig. 12) : This is a bright pattern of green
center and yellow border with no expression of epidermal red. Some few
internal or vascular strands of red may be seen. The pattern difi'ers
from that of type green-yellow-red blotched in having no epidermal red.
Color pattern green (fig. 13) : A pattern of pure spinach green with
no yellow and no epidermal red, but with a few streaks of red in the
vascular strands or in the mesophyl. This pattern difi'ers from the
parental pattern green-yellow-red blotched in the loss of both yellow^ and
epidermal red. The green-yellow spotted pattern (type 13 a, not illus-
trated) differs only in having yellow spots.
Color pattern yellow-green (fig. 14) : This type has a green border
and a yellow center, with no epidermal red. It differs from the
green-yellow pattern in the reversed position of the two color elements
and from pattern yellow-green-red blotched in having no epidermal red.
As in the case of the green-yellow type, the pattern is bright and
attractive.
20 ESTABLISHMENT OF VARIETIES IN COLEUS
Color pattern green-solid red upper center (fig. 15): A type that
differs from type green-yellow-solid red upper center in having no yellow
and hence is apparently bicolored on the upper surface. The center
of the leaf is violet carmine and the marginal zone is pure green.
Color pattern yellow-solid red: This tj^pe has almost uniform nopal
red color on both surfaces. It differs from type green-yellow-solid red
in not possessing a dark red center and from type yellow-red blotched
in having the epidermis completely red. In both patterns the green
underlying a solid red epidermis is almost entirely absent.
Laciniate leaf shape (fig. 7) : In marked contrast to the type of entire
leaf illustrated in the figures showing the various color patterns is
the deeply and irregularly cut and lobed types of leaf shape, the appear-
ance and behavior of which will be specially discussed later.
All of these color patterns arose as sudden spontaneous bud-varia-
tions, with the single exception of the type green-solid red upper center,
which is a pattern into which plants with the green-yellow-solid red
upper center pattern fluctuated. Throughout this paper, as above
noted, the term "bud variation" is, in all cases not otherwise quahfied,
apphed only to a marked change that appeared suddenly and com-
pletely for a part of a plant, and which was fully in evidence in the
leaves involved when they first unrolled. Gradual fluctuations also
gave in numerous cases types green-yellow spotted-red blotched, green-
yelloiv spotted, and green-red blotched. That is, these types appeared
both by sudden and by gradual variations.
CONSTANCY OF THE VARIOUS PATTERNS.
To test the constancy of the t>T)es, the original as well as those
derived from it by bud variations, successive generations of plants
were grown from pedigreed cuttings. This tested the vegetative con-
stancy of the pattern itself and enables one to make comparisons when
the same pattern was derived from different lines.
The series of plants considered under any type pattern are in large
measure a selected stock. When cuttings were made for the perpetua-
tion of the pattern in a new generation, they w^ere made from the plants
most typical and constant for the pattern concerned. \\Tien a bud
variation appeared, if the conditions were favorable, the parts posses-
sing it were allowed to develop until there were several branches from
which cuttings could be taken simultaneously. In such cases the
selection of branches for the new type was a simple matter, as it
depended on the taking of branches sharply distinct from the main
part of the plant, which in most cases were as different as is shown in
figures 21 and 24. When further cuttings were made for a new genera-
tion to perpetuate the type they were made from plants most uniform
and constant (determined from the records) for the pattern in question.
BY THE SELECTION OF SOMATIC VARIATIONS. 21
Usually but three cuttings were made from a plant, and these were
taken from branches most uniform and clearly conforming to the type.
It has already been noted and it will be very evident in the following
pages that some plants showed fluctuating variations giving irregular
or mixed patterns, or the pattern gradually fluctuated between two
types or changed from one type to another. Except in two cases
no attempt was made to secure new types by such fluctuating varia-
tions. These cases (clone 14 of table 2 and clone 13 of table 3) will
be especially discussed later.
In numerous cases cuttings were made to give two types of patterns
in the same plant. The constancy of the patterns could in these cases
be studied with the two parts growing from the same root system and
submitted to the same environmental factors.
Plants with yellow-red blotched pattern (fig. 1). — Seven cuttings pure
for this pattern were made in the autumn of 1913. Six died soon after
they were placed in the rooting-bench. The other lived and was grown
until the autumn of 1914. This plant was somewhat greener during
the winter, but at all times was decidedly more yellow than any plant
of any other type. It was, also, smaller and less vigorous in its growth.
Eight plants were grown as chimeras with one branch of yellow-red
UotchedpSitteTn and one branch of a pattern with yellow-green-red blotched.
On all these the branches of the part with pattern yellow-red blotched
remained quite constant throughout the year and were at all times in
marked contrast to the pattern of the other part. Two chimeras grown
only during the summer of 1914 were likewise quite constant. While
it is very difficult to obtain plants with this pattern from cuttings, the
type remains quite constant when grown in chimeral association with
branches having green tissue. On account of the difficulties of propa-
gation this type has not been rigorously tested. The few plants grown
gave no marked variations either as bud variations or as fluctuations.
Plants with green-yellow-red blotched pattern (fig. 2). — This is the
pattern originally possessed by the two parent plants, Nos. 1 and 3.
A total of 337 plants were grown from cuttings of this type. The data
given in table 2 are summarized in four main clones. Plants of clone 1 1
all descended from branch 1 on plant 1. Plants of clone 12 were
descended from branch 2 on plant 1. Plants of clone 3 were derived
from the branches of plant 3 that were uniform for this pattern. The
original branches from which the first cuttings were obtained were
uniform for the green-yelloiu-red blotched pattern and all plants used as
parent stock for later generations were selected as typical and most
constant for the pattern. The entire six generations constituted a
series of plants derived by continued selection.
The 45 plants of this pattern in clone 14 are especially interesting,
as they constitute a test for this pattern when derived by a gradual
fluctuation. As already noted, branch 4 of plant 1 possessed a decidedly
22
ESTABLISHMENT OF VARIETIES IN COLEUS
green pattern with yellow blotches designated as green-yellow spotted-red
blotched. In the third generation of plants grown from this branch
8 plants gradually fluctuated during winter until they were uniformly
green-yellow-red blotched. Such an increase of yellow during winter was
unusual and cuttings were made to test the constancy of the type thus
derived. The data for the 45 plants grown in three generations show
that two-thirds of the plants were quite constant for the derived type.
This proportion compares very favorably v/ith that of the clones 11, 12,
and 3, which were from the start selected from plants most typical for
the type.
Table 2. — Summary of plants irith green-yellow-red blotched ■pattern.
Clone
11.
Clone
12.
Clone
14.
Clone
3.
Total.
Number of plants
Number constant
Number fluctuating in green and yellow
Fluctuations to laciniate leaf-shape ....
Plants giving bud variations
Bud variations:
Yellow-red blotched
Green-yellow spotted-red blotched .
Green-red blotched
Yellow-green-red blotched
Green-yellow-solid rod
Green-j'ellow
Green-yellow spotted
151
102
29
22
4
13
3
103
76
22
1
12
1
2
7
1
1
45
21
15
10
3
14
1
2
1
1
2-?
38
19
11
1
9
337
218
77
2
53
7
8
24
6
4
8
o
'All were cases of fluctuation confined to about half of a plant.
^Loss of epidermal red on two plants fluctuating from yellow to yellow spotted.
Of the total number of plants with pattern green-yelloiv-red blotched
there were 218 that were at all times fairh^ constant and true to the
type. They were all somewhat fluctuating in respect to the relative
amounts of green and yellow, but were all constant in possessing at all
times a yellow border.
In figure 2 there is shown a leaf with the average development of
the yellow border, although in this leaf the pattern is somewhat irregular.
Figures 10, 12, 19, 20, and 26 show leaves classed as yellow-bordered;
figure 20, however, shows fluctuation toward tyipe yellow-red blotched, and
26 shows fluctuation toward a green-yellow spotted-red blotched pattern.
Figures 17 and 27 are from leaves classed as having irregular patterns.
The 77 cases classed as fluctuations include : (1) 56 cases of decided
increase of green during winter, followed by increase of yellow in
summer, giving in most cases return to the type of the cutting; two of
these also gave fluctuations to laciniate-leaf shape; (2) 9 cases of
increase of yellow during summer (grown onlj^ during a summer from
cuttings taken from plants that were greener during the preceding
winter) ; (3) 7 cases of fluctuations that were not uniform on a plant,
but gave leaves of the same age with different patterns so mixed that
BY THE SELECTION OF SOMATIC VARIATIONS. 23
no sectorial distribution could be traced; and (4) 5 plants with green
and yellow distributed irregularly.
Aside from the fluctuations in relative amounts of green and yellow,
there was also much fluctuation in number and size of the l)lotches of
epidermal red. On some of the plants there was rather gradual increase
or decrease both in number and size of these blotches, giving such
differences in respect to red as are shown in figures 5, 17, 26, and 28.
Such plants were, however, still considered as blotched in the sum-
maries. Selection of typical red blotched epidermis for various types
has been directed to plants having the epidermal blotching as in figures
2, 5, and 6, rather than as in figures 23 and 28.
53 plants produced bud variations giving loss of green 7 times, loss
of the yellow bordering-band 8 times, complete loss of yellow 24 times,
reversal of the relative positions of the green and yellow 6 times (4 cases
appeared in half of a leaf only, as in fig. 25), increase of red to com-
plete epidermal red 4 times, complete loss of epidermal red 10 times.
In clone 14, 4 plants gradually developed a green-yelloiv spotted-red
blotched pattern in part of the branches. On the basis of my descrip-
tions the change was a fluctuating variation affecting only a few
branches of a plant. These 4 cases are not included in the summaries
of bud variations.
In their extent the bud variations gave extremes in development of
yellow, of green, and of epidermal red. There were cases of nearly
pure yellow and of absolutely pure green ; there were cases of solid red
epidermis and others with no red epidermis.
Furthermore, the changes in green and yellow or in epidermal red
occurred entirely independentl}^ of each other. In general, the different
types of bud variations were quite uniformly distributed in the various
clones.
The type green-yellow spotted was produced on 2 plants by a fluctua-
tional increase of green after the loss of epidermal red had occurred.
In table 2, as in other tables, when the totals given for constant,
fluctuating, and sporting plants exceed the number of plants grown,
it shows that a certain number of the fluctuating plants produced also
sharp, clear-cut bud variations. Also, when the total of cases of bud
variations exceeds the plants giving them, certain plants produced
more than one bud variation.
For the purpose of estabhshing an index of the frequency of bud
variation we may take the ratio of bud variations to the estimated
number of buds developed. Each plant produced an average of at
least 200 branches which made sufficient growth to reveal the pattern
of the leaves. On this basis the index of total bud variation for this
group was about 1 to 1,110. The ratio of constant plants to fluctu-
ating plants was almost exactly 3 to 1, not counting the plants with bud
variations many of which were otherwise constant.
24
ESTABLISHMENT OF VARIETIES IN COLEUS
Plants with color pattern green-yellow spotted-red blotched and with
uniformly entire leaves. — The plants grouped in this class (figs. 4 and
23) present perhaps greater diversity than those of any other type,
embracing (1) plants with considerable yellow in scattered areas in all
leaves, (2) plants with only shght amounts of yellow in scattered areas
in nearly all leaves, and (3) plants with only a few leaves possessing yel-
low spots. Between the extremes there was every degree of variation
and often all degrees would be seen at one time among the leaves of a
single plant.
It is difficult in such plants to determine what constitutes a variation
either as a fluctuating or a bud variation when it involves green and
yellow. The cases given in table 3 are those in which an entire branch
or a sector of a branch showed leaves that were uniform for a new
pattern. Plants having irregular mixtures of leaves of equally different
patterns were common. Such cases are of special interest, as are the
Table 3. — Plants irith entire leanes and pattern green-yellow spotted-red blotched {fig. 4)-
Clone
11.
Total number of plants
Number constant
Fluctuations for green and yellow
To type green-yellow-red blotched . . .
To mixed patterns
To laciniate leaf
Total number plants gi\nng bud variations .
Bud variations:
To yellow-red blotched
To green-yellow-red blotched
To green-red blotched
To green-yellow spotted-solid red . . . .
To green-yellow spotted
Clone Clone
12. 13.1
16
6
2
2
4
2
79
01
16
Clone
14.
89
50
21
8
4
3
3
1
1
2
Clone
3.
22
9
8
4
1
Total.
198
126
47
8
13
7
8
1
3
2
2
2
^ Pattern derived by fluctuating variation.
fluctuating variations in a seed progen5\ They possess many simi-
larities to cases of size inheritance described b}^ Goodspeed (1912) and
raise the question as to whether color heredity is not also quantitatively
rather than qualitatively inherited.
As has already been noted, the parent plant (No. 1) had one branch
(No. 14) with leaves green-yellow spotted-red blotched. All the 89 plants
of clone 14 descended from this branch through 6 generations of selec-
tion. The plants of this pattern here given with clones 11, 12, and 3
were obtained from cases of bud variation from the type green-yellow-
red blotched (see table 2). The 79 plants of clone 13, however, were
derived from 5 plants that gave a fluctuating change from green-red
blotched to green-yellow spotted-red blotched. This was a frequent
fluctuation from the green plants especially of clone 13, as shown in
table 5, and the yellow-spotted condition thus obtained was tested in
four generations, comprising a total of 79 plants.
BY THE SELECTION OF SOMATIC VARIATIONS. 25
As a whole, there was a rather large proportion of the plants that
remained within the type as classified, although there was hardly a
plant grown during the winter that did not become somewhat greener
during that period. 68 plants fluctuated in a marked degree; 47 of
them were almost entirely green during the winter, but were again
quite uniformly yellow spotted in summer, although some of these
remained much greener in summer. None of the latter, however, could
be considered as of the pattern green-red blotched.
13 plants fluctuated irregularly, giving mixed patterns, mostly of
green-yellow-red blotched, green-yellow spotted-red blotched, and green-red
blotched, all more or less intermingled among the various branches and
on the same branch. These were not used as parent plants, but doubt-
less by selection it would be possible to obtain a marked degree of
constancy for the irregular and mixed patterns, with, also, production
of plants that would be uniform for various types.
The most uniform and marked of the fluctuations was the case
of 8 plants of clone 14 which gradually became more yellow during
the winter of 1912-13, until they were quite typical green-yellow-red
blotched; 5 of these were used as parents of the plants of clone 14 already
reported with table 3. The change in pattern arose as a gradual
increase of yellow from various degrees of a yellow spotted condition to
a well-defined yellow bordered pattern that was quite uniform for the
entire plant, and which when tested in progeny was subsequently quite
as constant as cases which arose by sudden variations. An analysis of
the pedigrees of these 8 cases shows that all of these descended from
only 3 of the 7 plants grown from cuttings of the original branch 14.
This phenomenon of the appearance of the same variation in different
plants that were derived from the same more remote ancestor is common
and constitutes what we may call duplicate-reversions or variations.
Besides the fluctuating variations in regard to green and yellow,
there were numerous cases of fluctuation in the red-blotched condition
both of the epidermis and of the subepidermal tissues, giving extremes
of very finely red-blotched or coarsely blotched. No selections were
made to secure types of the red-blotched condition. None of the
plants fluctuated to a no-blotched or to a soUd-red pattern.
The behavior of the 79 plants of this pattern in clone 13 is especially
interesting. They constitute a test for this pattern obtained by the
selection of gradual and accumulated fluctuation. The progeny of
5 plants grown in 3 generations, subjected to the same sort of selection
as the other clones, showed the highest degree of constancy obtained in
the clones of this pattern.
Seven plants gave, during the winter, a marked increase of green, ac-
companied by the production of cut and laciniate leaves (fig. 7), the
appearance and constancy of which are quite fully discussed later.
26 ESTABLISHMENT OF VARIETIES IN COLEUS
There were few cases of sudden, clear-cut bud variation in this
group. One was a very decided and ahnost complete development of
yellow to yellow-red blotched; 3 were sectorial variations to green-yellow-
red blotched; 2 cases gave loss of yellow to green-red blotched; 2 involved
gain of epidermal red to solid red, and in two cases there was loss of
epidermal red. All these cases were sectorial for a plant or in some
cases for a single branch. The ratio giving the frequency of bud varia-
tions involving color in this group is 1 to 3,960.
Plants with laciniate leaves. — Until the winter of 1912-13, all the
plants in the cultures had been constant and very uniform for leaf-
shape, showing no greater variation in this respect that is seen in fig-
ures 2, 6, and 9. During that Mdnter it was noted in 11 cases that as
new leaves developed they were more and more deeply cut and lobed
until in Januar}^ and February the uppermost leaves were in extreme
cases much divided and deeply laciniate, as shown in figure 7. The
plants appeared like the middle plant in plate 4; 3 of these plants
were from cuttings of branches that were pure-green bud variations;
the others were plants which had a somewhat fluctuating green-yellow
spotted pattern. Nine of these plants were grown during the following
summer, when it was noted that without exception the new leaves
gradually became more entire until by late summer all the leaves then
hanging to the plants were entire. At the same time 5 of the plants
became decidedly more yellow, even becoming quite uniform for type
green-yellow-red blotched. Cuttings were made in April from each of
these plants and from these 16 plants were grown during the summer
of 1913. All of these fluctuated to entire leaves, and in regard to color
gave plants some of which were uniform for type green-yellow-red
blotched, others for tj^pe green-yellow spotted-red blotched, while the
foliage of others showed mixtures of these patterns with also pattern
green-red blotched.
The further generations in subclone 12 exhibited, as shown in table
4, the same periodicity in change of leaf-form, except that in late sum-
mer of 1914 a rather large number of the plants showed new leaves
that were laciniate. One plant of this subclone was grown in the winter
of 1913-14 from a pure-green bud-sport that appeared during the
previous summer. This plant remained constant for entire leaves of
pattern green-red blotched during the winter, but as it died early in the
summer no further progeny were grown. Further generations of the
plants with laciniate leaves in subclone 14 were not grown.
The laciniate leaf-form appeared anew in the winter of 1913-14 in
clone 3 in two instances. One was a plant whose line of descent
showed bud variation from type green-yellow-red blotched to type green-
yellow spotted-red blotched; that of the other showed bud variation from
green-yelloio-red blotched to green-yellow solid red, and then from this to
green-yellow-solid red upper center. The latter was the only one of
BY THE SELECTION OF SOMATIC VARIATIONS.
27
8 plants grown from cuttings of a single plant to exhibit the fluctuation
to laciniate leaves.
It is to be noted that this character of laciniate leaf-shape has not
appeared thus far in any of the plants grown in clones 11 and 13. It
has appeared as a fluctuating character that develops most strongly in
winter. With one exception all the plants grown to test the reappear-
ance of the character have exhibited it. This plant was grown from
a bud variation giving a single branch of green-red blotched on a plant
otherwise uniform for pattern green-yellow-red blotched at the time the
cutting was made.
During the time these plants exhibited the laciniate character most
strongly, there were growing among them numerous plants of other
clones of various patterns, especiall}'- of green-yelloir-red blotched and
green-red blotched, all submitted to the same conditions of Ught, tem-
Table 4. — Summary of plants with laciniate leaves.
Total number of plants
Much greener in winter ....
Very laciniate in ^\dnter. . . .
Entire in winter
In summer as tjije 2
as type 4
as type 1
mixed patterns . .
uniformly entire.
slightly laciniate .
Plants giving bud variations
Type green-red blotched and entire .
green-yellow spotted
green-yellow-red blotched
Clone 12.
1912-
1913,
6
f)
G
0
5
0
0
1
6
0
1
1
1913.
10
4
1
0
5
10
0
1913-
1914.
32
32
31
1
4
.5
1
22
5
27
1
1
1914.
1
3
0
3
0
7
Clone 14.
1912-
1913.
1913.
3
3
3
0
0
0
0
3
3
0
0
6
0
0
6
0
Clone 3.
1913-
1914.
1914.
2
2
2
0
0
0
0
2
1
1
0
1
1
0
0
2
perature and soil, yet not in the least degree exhibiting the fluctuation
to laciniate leaves. This is well shown in plate 4, which gives a photo-
graph of 3 plants of the same clone (12), all grown under the same
conditions. The plant to the right (No. 125111) had entire leaves
and a green-yellow-red blotched pattern; the one to the left (No. 1251412)
was of a pure green-red blotched pattern; the one in the middle (No.
123153) shows the transition from entire leaves to deeply laciniate
leaves as it occurs during the winter.
The late summer of 1914 was exceedingly dry. In July there had
been 5.36 inches of rainfall, during which time the plants made an
unusually vigorous growth. From August 12 until October 16 there was
but 1.26 inches of rain. During the dry warm period of September, the
28
ESTABLISHMENT OF VARIETIES IN COLEUS
new leaves on very many of the plants of this series were strongly
laciniate. Cuttings were made from these laciniate-leaved branches.
The new leaves that developed on these young plants during November
were entire; hence it would seem that the laciniate character in these
particular clones of Coleus is in some degree associated with decreased
vigor. When most favorable conditions for growth prevail, or when
rapid growth is brought about in cuttings, the leaves become entire.
It is, however, clear that the first appearance of the laciniate char-
acter was confined to a few plants and that once it originated it
reappeared with marked constancy in the vegetative progeny.
During the winter of 1914-15 the laciniate character appeared in
the manner of a bud variation. A large plant that had been grown out
of doors during the summer was in September severely pruned and
placed in a pot for further development in the greenhouse. It was
intended to use the plant for stock in general border planting and the
plant label was not preserved. From the records of the pattern and
generation it is clear that the plant itself and all the plants in its line
of descent possessed only entire leaves and the plant belonged to the
main clone 1. This plant was given the number 9.
In the course of 3 months numerous new
branches developed on the 10 pairs of main
lateral branches to which the plant had
been pruned. It was noticed that of the
20 main branches, 3 bore branches with
laciniate leaves. The positions of these are
indicated by numbers 1, 2, and 3 of
diagram 2.
All the branches on all other of the main
branches bore only entire leaves. All the
branches arising from 2 bore only laciniate
leaves, but the branches with entire and
with laciniate leaves were sectorially dis-
tributed on the branches 1 and 3. The
contrast was most marked, especially when
opposite branches were different, one hav-
ing laciniate and the other entire leaves. The sectorial differences
appeared in some of the secondary branches and carried the two types
into parts of individual leaves.
The most striking behavior of this series of plants summarized in
table 4 is the wide fluctuations in the leaf-shape and in the amount of
yellow and green, the marked correlation of decrease of yellow with
decrease of leaf area, and the rather pronounced periodicity of these
fluctuations. These fluctuations are so general and rhythmic that
they can almost be considered constant. Of bud variations there were
but 3 cases, giving a ratio of about 1 to 2,530, which, however, shows
10
10'
9
9'
8
8'
7
7'
6
6'
5
5'
4
4'
3
3'
2
2'
1
1'
Diagram 2. — Showing position
of the branches on plant
No. 9.
BY THE SELECTION OF SOMATIC VARIATIONS.
29
greater frequency than that of the pattern green-yellow spotted-red
blotched. Of the bud variations one was a loss of yellow, one gave
increase of yellow to type green-yelloiv-red blotched and one was a loss
of red.
Plants ivith green-red blotched -pattern. — The apparently complete loss
of yellow, giving only green subepidermal tissues, was a frequent bud
variation in plants having green-yellow, yellow-green, or green-yellow
spotted patterns, regardless of the degree of red in the epidermis (fig. 5).
The condition of pure green also developed as a fluctuation on plants
of these same types. In cases the fluctuation was quite general for the
entire plant, while in others it occurred irregularly, giving plants with
mixed patterns.
The 90 plants included in this summary are, however, selected stock,
all descended from cases of bud variation similar to that of figure 21,
in which the part concerned showed no trace of yellow in any leaves.
Table 5. — Summary of plants unth green-red blotched pattern (fig. 5).
Clone j Clone
11. 12.
Clone I Clone
13. 1 14.
Clone
.3.
Total.
Number of plants
Plants constant
Fluctuations:
Green-yellow spotted
Mixed patterns
Laciniate leaf-shape
Plants with bud variations
Bud variations:
Green-yellow spotted-red blotched . .
Spontaneous yellow
Green-solid red
Green
13
5
1
6
18
5
7
1
3
5
1
1
2
1
51
17
31
3
6
3
2
1
90
31
41
4
3
15
2
9
2
3
Selections for further generations were made from plants that had
remained uniformly pure green. The type was maintained bj^ selection
quite as it is practiced in a herd of dairy cattle. In the case of clone 13,
6 generations were grown, all descended from the pure-green branch of
the parent plant, No. 1.
Of the total number of plants in this group, 31 remained pure green,
showing no trace of yellow by either fluctuation or bud variation.
In addition, 14 of the 15 plants with bud variations were otherwise
constant for the pure-green condition. All of these plants were grown
during an entire summer. 41 plants developed varying amounts of
yellow in scattered areas, making a pattern classed as green-yellow
spotted-red blotched. In 11 of these the yelloio spotted condition was
quite uniform and typical and from these were selected parents for
the plants of subclone 13 given in the table 3. Three plants also gave
fluctuations in leaf-shape to the laciniate type and their progeny are
included in the summary of table 4. Four plants gave decidedly
30
ESTABLISHMENT OF VARIETIES IN COLEUS
mixed and irregular patterns like those that appeared from patterns
treated in tables 3 and 4.
There were 16 cases of bud variation; 5 were concerned with the red
epidermis, 2 giving solid red, and 3 giving no red. There were 2 cases
of vspontaneous appearance of yellow, giving branches sectorial for the
green-yellow spotted-red blotched pattern. There were, also, 9 cases of
spontaneous development of j^ellow that were not carried on in suc-
cessive leaves, and with the exception of 1 case were confined to but
one or two leaves. These yellow blotches were large, irregular-shaped
pure-yellow areas covering from one-eighth to one-fourth the entire
area of a leaf. The locations and relative sizes of these yellow blotches
in the leaves of one plant are shown in text-figure 1, the shaded protions
of which indicate yellow areas. All other leaves were pure green and
the branches produced in the axils of the yellow-blotched leaves were
pure green.
Text-figure 1. — Position of the yellow areas that developed spontaneously in these
leaves of plant 11714221.
A study of pedigrees reveals the interesting fact that 6 of the plants
with spontaneous yellow all descended from a branch on plant 11714.
The full record of this clone is given in table 13, but the summary of
the pedigrees of the particular line of descent is here given in table 6.
The pedigree numbers enable one to trace relationship quite readil3^
In this case we note that plant 1171 gave a bud variation with loss of
yellow. Plant 11714 was grown as a chimera with about one-half pure
green. From the green part two cuttings were made for plants 117142
and 117144, both of which remained constant for loss of yellow. In
September 1913, three cuttings were made and from two of the plants
grown three more cuttings were taken in the following spring. From
September 1912 until August 1914 all plants grown in these lines of
descent were constant for loss of yellow, then in one season the spon-
taneous development of yellow occured in 6 closely related plants.
BY THE SELECTION OF SOMATIC VARIATIONS.
31
Table 6. — Pedigree of 6 plants giving spontaneous yellow.
Time.
Plant.
Pattern.
Record.
Apr. 1912 to Sept. 1914
Sept. 1912 to Sept. 1913
Summer 1913
Sept. 1913 to Sept. 1914
Summer 1914 .
117 Green - yellow - red
blotched
117, 1 do
117 14» /••••do
\Green-red blotched .
117, 142 do
117, 144 do
117,142,1.. do
117,142,2 do.
117,144,1 do.
117.142.11. do.
117.142.12. do.
[117,142,21..] do.
Constant.
Bud variation to green-
red blotched.
Constant.
Constant for loss of yellow.
Do.
Do.
Spontaneous yellow in
August.
Do.
Do.
Do.
Do.
Do.
'Chimera.
Including the cases of spontaneous appearance of yellow with sec-
torial bud variations, the ratio of frequency was 1 to 1,120. The ratio
for the sectorial bud variations alone was about 1 to 2,570.
Plants with pattern yellow-green-red blotched. — All of the plants grown
with this pattern (fig. 6) belonged to clone 11, and all, excepting 3, de-
scended from the plant 1171, which produced a series of branches with
this pattern, as already shown in diagram 1. The record of these
plants is given later in table 13, but may be summarized here as given
in table 7.
Table 7. — Summary of plants loith yellow-green-red blotched pattern {fig. 6), all
of clone 11.
Yellow-green-red blotched
p. . fyellow-green-red blotched. .
^yellow-red blotched
/^f Jyellow-green-red blotched. .
Chimera < j ui * u i
1 green-red blotched
CK fyellow-green-red blotched. .
\yellow-green
Total
o
a
o
29
8
a
O
17
5
41 24
a
"■+3
3
o
3
M a
a o
Bud variations.
•n
(D
O -C
-I
9 3
13
1^
cl aj o
g +^ O
Op
a; ^
(U o
10
a
i-i
60
I
"53
32 ESTABLISHMENT OF VARIETIES IN COLEUS
In several instances cuttings were so made that the resulting plants
possessed two patterns. These plants are included here in respect to
the behavior of the part with pattern yellow-green-red blotched.
All of the plants grown during the winter showed more or less
increase of green, but as long as the yellow was present as a definite
central area they were classed as constant. There were, for example,
just such differences in development of yellow as is seen in figures
14 and 14a, the former representing the usual condition during winter
and the latter the development of yellow during the summer. Four
plants fluctuated in marked degree, giving mixed and irregular patterns
with many leaves in which there was much green.
One case of bud variation was concerned with loss of epidermal
red, giving the type yellow-green (fig. 14). The other instances gave
3 cases of extreme development of yellow, 10 cases of entire loss of
yellow, and 1 case of change to the yellow spotted condition. The
latter, however, occurred on a plant with also a bud variation to pure
green-red blotched. The ratio of frequency for bud variation in this
group was about 1 to 540.
Plants with pattern green-yelloiv-solid red. — In respect to the green
and yellow this pattern (fig. 8) is identical with that of green-yellow-
red blotched. It differs in having a solid-red instead of a red-blotched
epidermis. The 54 plants grown with this pattern remained free from
any noticeable variations in respect to the solid red epidermis, except
those that were bud variations. Frequently a leaf appeared with a
few small areas in which the red of the epidermis was absent, but these
were rather isolated. There was some degree of fluctuation in the
relative amounts of green and yellow, with a tendency for plants to be
greener in winter and yellower in summer. On account of the soUd red,
it was more difficult to judge these fluctuations than in plants with
red blotched or with non-red epidermis, hence attention was chiefly
directed to the condition of the epidermis. In making cuttings, plants
most constant and typical for the green-yellow condition were, however,
selected.
Of the 10 cases of bud variation, 4 gave complete loss of yellow,
2 gave a reversal of the relative position of green and yellow, and 1
gave extreme development of yellow. Only 3 cases involved variation
in the amount and distribution of red; 1 was a complete loss and 2 gave
the pattern described above as green-yellow-solid red upper center,
a type which is an interesting intermediate between no red and solid red.
The ratio of frequency for all bud variations for the group is 1 to 1,080.
Plants with green-solid red pattern. — This pattern (fig. 9) first ap-
peared during the summer of 1913 as a bud variation on a plant of
pattern green-red blotched. From this branch cuttings were taken for
6 plants grown during the summer of 1914. All of these remained con-
stant for loss of yellow and for a solid-red epidermis, except 1 plant.
BY THE SELECTION OF SOMATIC VARIATIONS.
33
upon which a sectorial variation in one small branch gave the pattern
green.
With respect to the red epidermis, patterns green-yellow-solid red
(fig. 8) and green-solid red are identical, and it is noteworthy that in the
62 plants of these two patterns which were grown to maturity there
were no noticeable fluctuations and but 4 cases of bud variations involv-
ing the red. For the red epidermis the ratio of frequency of bud varia-
tions was 1 to 3,100.
Table 8. — Summary of plcmls ivilh solid red -patterns, clone S.
Green-yellow-
solid red.
Green-solid
red.
Total number of plants
54
51
9
1
2
4
1
8
7
1
1
Constant for solid red
Number of plants with bud variations. . .
Bud variations:
To vellow-solid red
To vellow-green-solid red
To green-yellow-solid red upper center .
To green-solid red . .
To green-yellow
To green
Plants with pattern green-yellow-solid red upper center. — This bright
and attractive pattern (fig. 10) first developed as a sectorial bud vari-
ation in the winter of 1912 on a plant otherwise uniform and constant
to type green-yellow-solid red. During the summer of 1913, the plant
grew vigorously and numerous branches developed from the part hav-
ing this new pattern, all of which were constant and uniform for the
new type and which were in conspicuous contrast to the rest of the plant.
In the autumn of 1913 cuttings were made from these branches for
5 plants, which were grown until the autumn of 1914. Four of these
remained quite constant, although they were much greener in winter.
One plant became gradually greener during the early part of winter
until it was apparently pure green, giving type green-solid red upper
center. The upper surface of a leaf of this plant painted in January
is shown in figure 15. Toward spring the new leaves produced by this
plant became quite laciniate, but during the following summer the
leaves produced were entire and strongly tinged with yellow. During
the summer of 1914, two plants from cuttings of one of the plants
constant for the type remained true to that pattern.
No bud variations appeared in any of the 7 plants and no noticeable
fluctuations in the amount and distribution of the red ; yet there was
no plant that did not show at some time a few leaves with tiny red spots
scattered on the lower surface, much as is shown in figure 10a.
This pattern also appeared late in the summer of 1914 as a sectorial
variation on a plant which during the summer had been constant and
uniform for type green-yellow-solid red. A cutting was made from this
34 ESTABLISHMENT OF VARIETIES IX COLEUS
branch, and the young plants grown from it are at the present writing
(December 10, 1914) nearly devoid of yellow, but have the change in
pattern for red as a clear-cut sectorial variation. Both surfaces of a
single leaf are shown in figures 24 and 24a. In figure 24 the loss of
epidermal red on the lower surface of half of the leaf illustrates very well
the definiteness with which color variations in Coleus appear. The
upper surface of the corresponding half of this leaf is shown in figure
24a, with the decrease of red about the margin. Such differences are
usually seen in a series of leaves in the same row and in the branches
that develop in the axils of such leaves, gi\dng a marked degree of
sectorial symmetry to the distribution of pigmentation, a condition also
well illustrated with reference to green and yellow in figure 21.
Plants with pattern yellow-green-solid red. — This pattern (fig. 11) first
appeared during the summer of 1913 as a variation complete for a single
lateral branch of a plant with green-yellow-solid red. This branch was
removed for a cutting, but died soon after it was rooted and placed in
a pot. Early in the spring of 1914 this pattern appeared as a sectorial
variation in the main axis of a plant having green-yellow-solid red. This
plant grew vigorously, giving a large, bushj^ plant with the two types
of foliage distinct and constant on the different branches. Numerous
cuttings have been made to test the vegetative constancy of this type.
Plants with -pattern green-yelloiv. — Three plants of this type (fig. 12)
were grown from September 1913 until October 1914 and 4 were grown
during the summer of 1914. All of these remained quite constant for
the loss of epidermal red. They were much less uniform in regard to
the relative amounts of green and yellow, one plant possessing a branch
that was quite green. There was also a strong tendency among the
leaves on one plant to show somewhat irregular distribution by green
and yellow, as is shown in figure 12.
Plants with pattern green and pattern green-yellow spotted. — The loss
of epidermal red occurred as a sectorial bud variation during the late
summer of 1913 on a plant that had fluctuated from type green-yellow-
red blotched to type green-yellow spotted-red blotched. The develop-
ment of yellow was, however, very faint, so that the bud variation
gave a leaf pattern that was almost pure green. This plant was
taken up and grown in a pot during the winter of 1913-14 and during
the following summer again grown out of doors. All branches on the
two parts were quite constant in respect to presence and absence of the
epidermal red, but there were traces of red coloration in the sub-epider-
mal tissues which, as shown in figure 13, were almost entirel}^ confined
to the vascular tissues. There was more or less fluctuation on the entire
plant in the appearance of yellow, but no decided development of it.
In January 1913 a cutting was made from the part of this plant
which showed the bud variation. The plant grew vigorously and was
during the winter mostly free from yellow. In April, 5 cuttings were
BY THE SELECTION OF SOMATIC VARIATIONS.
35
made for plants. During the summer of 1914 the G plants remained
constant for loss of epidermal red, but bore some leaves with yellow
spots. There was, however, a rather weak development of yellow and
from a short distance the plants appeared to be pure green. The
fluctuations in the development of green and yellow are quite like
those in other patterns. The pattern is of special interest in regard
to the development of red pigmentation in the subepidermal tissues,
especially in the vascular elements giving a reticulated effect well
shown in figure 13, and also seen in figure 14a. This condition
also prevails in the red blotched types, but is more or less obscured
by the more conspicuous epidermal coloration.
During the summer of 1914 there was no noticeable variation in
the amount of red in subepidermal tissues. Cuttings made in the
autumn of that year for a new generation exhibited during the winter
marked variations in this respect. The summaries given in this paper
do not include the generation to which these plants belong, but the
behavior of this particular set of plants can be included here. Figure
Table 9. — Summary of plants with non red epidermis.
Pattern.
Green-yellow
Green and green-yellow spotted .
Yellow-green
Total
plants.
7
7
4
Constant
for non red
epidermis.
7
7
4
Plants
giving bud
variations.
0
0
0
13d gives a leaf painted on February 2, 1915, showing the development
of red in the internal tissues. Free-hand sections of such leaves indi-
cated that the epidermal cells are non-red. The coloration appears
dull, as if glazed over rather than velvety as in the epidermal colora-
tion, a contrast due largely to the coloration of the trichomes of
the epidermis and which the reproductions do not adequately show.
Plants with 'pattern yellow-green. — This pattern (fig. 14) identical with
that of type yellow-green-red blotched except for the loss of epidermal
red, appeared as a sectorial bud variation late in the summer of 1913.
The sporting branch was used as a cutting, from which a large plant
grew during the winter of 1913 and the following summer. In the
winter there was an increase of green, but throughout its growth the
part with pattern yellow-green remained constant in respect to the loss
of epidermal red.
Three plants of a new generation were grown during the summer of
1914, and these remained constant and uniform for the loss of epidermal
red. In regard to the relative positions of green and yellow, the plants
were quite constant, but there was a strong tendency for green to
increase in winter and decrease in summer, giving such differences as
are shown in figures 14 and 14a.
36 ESTABLISHMENT OF VARIETIES IN COLEUS
The figures 13, a, b, and c, show the three leaves growing in the
rank above the leaf shown in figure 13d and ilkistrate the increase of
red which very plainly occurs as the leaves mature. In the red-
blotched and solid red patterns an increase in the total amount of red
pigmentation must, it would seem, also occur as the leaves enlarge.
As the last four patterns, green-yellow, green, green-yellow spotted, and
yellow-green are alike in respect to loss of epidermal red, they may be
grouped in this respect. It is noteworthy that there was no case of a
development of epidermal red. No plant was free of some red colora-
tion in stems and in vascular strands of the leaves, as especially well
shown in figures 13 and 14, There were cases where the coloration
seemed to spread out near the ends of the vascular strands, but the
appearance was not the same as that of the blotches. The number of
plants of these patterns grown thus far is small and their behavior is
not taken as fully indicative of the possible variations that may appear
in future cultures.
From the summaries of patterns given above it is quite clear
that the various types noted (with the exception of pattern green-solid
red upper center) have been kept quite constant by a selection of the
parent plants to be used in vegetative propagation, and that every new
type of pattern (excepting the one) that arose either by fluctuating
variation or by bud variation can be propagated as a vegetative type.
It is highly possible that finer distinctions could be made in regard to
pattern types, especially inside of the rather comprehensive groups
classed as red blotched, as yellow bordered, and as yellow spotted, among
which there were many variations that gave all degrees of gradation to
or even into a different pattern. The writer wishes to state that the
keeping of records satisfactory to him was no simple matter, even for
the pattern classes as determined.
BY THE SELECTION OF SOMATIC VARIATIONS.
37
RANGE OF THE VARIATIONS.
At this point the data already presented may be summarized and anal-
yzed in reference to the range or extent of the variations as a whole.
There have already been described 15 different color patterns that
arose by bud variation, 1 very decided color pattern that arose solely
as a fluctuating variation, and the laciniate type of leaf. Several of
the principal types arose also by fluctuating variations. In respect to
the relative amounts of green and yellow there are the two extremes:
(a) almost pure development of yellow (fig. 1), and (b) pure green
(figs. 5 and 9), with almost every possible gradation between. Of the
epidermal red there are the extremes solid red and no red with the
intermediate red-blotched type (including wide variations), and the type
solid red upper center as another intermediate. In respect to the relative
positions of the green and yellow there are the extremes: (a) green center
w^ith yellow border (fig. 2) and (6) yellow center with green border (fig. 6) .
Between these the irregular patterns present numerous intermediates.
,1. Yellow-red
blotched
/A- Green-
/yellowspotled
red blotched
Spontaneous yellow
7 2or4andE-----5and7
8a.Greer>-yellowspcrtted-sclid ned
13a. Greeri -yellow spotted
Z.Grccn-yCllow- .
red blolcheci
. 5. Green-red
blotched
6. Yellow-
green- red
blotched
8. Green-
yellow-
Solid red
Spontaneous yellow
*• ^5 and t
2— -4 and E Send?
9. Green- sol id red '^®
13. Green I3e
I.
4
5.
14. Yellow-green
9. Green -sol id red
10. Green-yellow-solid red upper center'
11. Yellovy-green-solid red
12.
16. Yellow-solid red
.i5eryJ7
12. Green-yellow
Diagram 3. — Derivations of color patterns.
Diagram 3 gives a graphic representation of the extent of variations
in each pattern and shows the derivation of types and the appearance
of the same types of pattern as variations from quite different patterns.
38 ESTABLISHMENT OF VARIETIES IN COLEUS
In diagram 3 the numbers (except 13 a) refer to figures illustrating
the types of color pattern or leaf-shape. The name of the pattern is
written in full only in the line of descent when it first appeared. A
continuous Une indicates origin by bud variation, while a broken line
indicates fluctuation.
From the original pattern of green-yellow-red blotched there arose
directlj' 6 different patterns (see tables 1 and 2 and diagram 3) each
involving a single marked variation. For the development of any
other pattern thus far produced, excepting the green-yellow-solid red
upper center pattern a second change is necessary. On any of these 6
derived patterns a further single change may give a new pattern or
produce a pattern already realized. The bud variations in the derived
pattern green-yellow spotted-red blotched (fig. 4) illustrate this point.
Increase of yellow gave yellow-red blotched (fig. 1) and loss of yellow
gave green-red blotched (fig. 5), both patterns previously derived, and
also a return to the original type of green-yellow-red blotched (fig. 2).
In all these the red-blotched condition of the epidermis is much the
same. Changes in the epidermal coloration, however, give new pat-
terns. The appearance of solid red gives a slightly different pattern
than that of green-yellow-solid red. The pattern green-yellow spotted
appeared as a bud variation by a loss of the epidermal red.
The green-red blotched (fig. 5) pattern gave opportunity for new
patterns by the same changes in epidermal red which have previously
appeared. These are realized in green-solid red (fig. 9) and green-no red
(fig. 13). There is also chance for reappearance of yellow to give return
to old types or possibly to new types. Of these only one appeared, and
this was the pattern green-yellow spotted-red blotched (fig. 4). The cases
of spontaneous appearance of yellow did not result in a definite pattern.
In this line of descent there also developed the character of laciniate
shape of the leaf, with its marked periodicity of expression.
The changes in the pattern yellow-green-red blotched (fig. 6) which
involved amounts of green and yellow gave no new patterns. The
loss of red, however, gave a new pattern yellow-green (fig. 14). At
least 3 possible bud sports did not appear in this pattern: (1) changes
producing a reversal of green and yellow giving return to the parent
type; (2) a solid-red (fig. 11); or (3) solid red upper center (fig. 10).
In the plants with green-yellow-solid red (fig. 8), a loss of yellow gave
the same pattern that was produced by gain of soUd red from pattern
green-red blotched. The two changes involved are identical, but occurred
in reversed order. Reversal of the positions of green and yellow, a
change identical with that giving the yellow-green-red blotched pattern,
gave a different pattern because the tissues were overlaid by solid red.
The same is true of the loss of green. Loss of the epidermal red on the
lower surface and about the margin of the upper surface gave a pattern
green-yellow-solid red upper center (fig. 10) that has not appeared else-
where and is really the only new variation that appeared in this pattern.
BY THE SELECTION OF SOMATIC VARIATIONS. 39
Plants grown from pattern green-yellow-red blotched (fig. 2) derived
from green-yelloiv spotted-red blotched (fig. 4) gave again the same pat-
terns that were directly derived from that pattern.
A general review of the entire series of variations shows that in
respect to the relative positions and total amounts of green and yellow,
and the total amount of red in the epidermis, the extremes of develop-
ment possible are reahzed, with, also, the appearance of a series of inter-
mediate types. Judging the variations in any one pattern by the
range of bud variations that have thus far developed, it appears that
any pattern tested in considerable numbers gives by bud variation the
entire range of changes possible.
FREQUENCY OF THE BUD VARIATIONS.
With the list of the tj^pes of variations given in table 1 there is also
given the data as to the number of times each occurred and the total
number of plants involved (not including plants of patterns which did
not produce the particular variation) . The various tables present the
details of their data, which may be now summarized under the main
types of changes outlined in table 1.
A. CHANGES INVOLVING YELLOW AND GREEN.
I. Increase of Yellow and Decrease of Green.
1. The almost complete loss of green with increase of yellow occurred
as a bud variation 12 times in a total of 630 plants (not including plants
of patterns which did not give this variation). It was derived most
frequently from patterns green-yellow-red blotched (fig. 2), yellow-green-
red blotched (fig. 6), and green-yellow-solid red (fig. 8). It developed once
from pattern green-yellow spotted-red blotched (fig. 4). The pattern
was not realized uniformly on any plant as a fluctuating variation. On
a few plants of the pattern green-yellow-red blotched which developed
mixed patterns with a marked increase of yellow during the summer,
some leaves approached this pattern. One of the most marked of
these is shown in figure 20.
2. It will be remembered that the green-yellow-red blotched pattern
was borne by the parent plants and that all other patterns were derived
directly or indirectly from this. The return to this pattern occurred
as a bud variation from type green-yellow spotted-red blotched in 4
instances on a total of 266 plants and also as fluctuating variations,
especially in clone 14, as shown in table 3.
3. The sudden appearance of scattered areas of yellow in single
branches of plants otherwise having no yellow occurred but twice.
The same type green-yellow spotted-red blotched, however, appeared
quite gradually for entire plants in 38 cases out of the 90 plants which
were grown for the pattern green-red blotched.
40 ESTABLISHMENT OF VARIETIES IN COLEUS
4. The development of rather large conspicuous areas of yellow in
one or more leaves of a plant otherwise pure green occurred 9 times.
In one case three leaves of the same branch showed yellow areas that
arose in this apparently spontaneous manner (see text-fig. 1).
Summary. — There were 27 cases of bud variation giving increase of
yellow and involving directlj^ 788 plants. There was opportunity
for increase of yellow to occur in all plants grown, except those of the
pattern yellow-red blotched, and even in these, 10 of the 11 were grown
as chimeras with at least half of the plant green. In computing a
final ratio for the frequency of bud variations giving increase of yellow,
we may use all but 6 of the plants grown. The ratio of frequency on
this basis is 1 to 6,130.
II. Decrease of Yellow and Increase of Green.
1 . Pattern green-yellow spotted-red blotched was produced from green-
yellow-red blotched and yelloiv-green-red blotched in 9 instances on a total
of 378 plants. In 4 other cases the change to yellow spotted affected
single branches, and although marked for a time after the first appear-
ance, later fluctuated to the parent type green-yellow-red spotted, for
which the plants became quite uniform.
2. Pattern green-red blotched with complete loss of yellow occurred on
plants with green-yellow-red blotched, with green-yellow spotted-red blotched
(entire and laciniate), and with yellow-green-red blotched patterns in
37 instances on a total of 644 plants. The same change gave pattern
green-solid red 4 times on 54 plants of pattern green-yellow-solid red.
Summary. — Bud variations producing increase of green occurred 50
times. The total plants grown with more or less j^ellow were 740.
The ratio of frequency for loss of green by bud variation was 1 to 2,960.
III. Reversal of the Relative Positions of Green and Yellow.
1 . This reversal has only occurred in patterns with the yellow at the
border of the leaf, giving yellow-green-red blotched (fig. 6) from green-
yellow-red blotched (fig. 2) and yellow-green-solid red (fig. 11) from green-
yellow-solid red (fig. 8) in a total of 8 instances on 391 plants. It is also
possible for a reversal to occur in any other patterns having a distinct
border of green or yellow. The total of such plants is 450, which gives
1 to 11,250 as the ratio of frequency for this change.
B. CHANGES INVOLVING THE EPIDERMAL RED.
I. Increase of Epidermal Red.
Eight instances of bud variations giving solid red occurred in red
blotched patterns involving directly a total of 625 plants. None of
the 41 plants of the pattern yellow-green-red blotched gave this variation.
Increase of red was possible in all except the solid-red patterns (62
plants in all). The ratio for this change was 1 to 19,250.
BY THE SELECTION OF SOMATIC VARIATIONS.
41
II. Decrease of Epidekmal Red.
Bud variations of this sort can be graded as follows: an almost com-
plete loss of red on the under surface and about the upper margin,
which occurred 2 times, and an apparently complete loss of epidermal
red in patterns with spotted or with solid red epidermis, which appeared
in 19 instances. A total of 815 plants were grown of patterns having
some degree of red in the epidermis. The ratio for complete loss of red
was 1 to 8,580 and for all cases of decrease of red it was 1 to 7,760.
C. CHANGES INVOLVING LEAF-SHAPE.
The appearance of the laciniate leaf-shape as a fluctuating variation
which marked periodicity of development occurred 13 times. (The
bud variation giving this type late in 1914 is not included.) The
total number of plants grown with entire leaves was 765, hence the ratio
on the basis used hitherto was 1 to 11,770. It seems, however, that
this basis hardly affords the same degree of accuracy for comparison as
it does between the different bud variations in color. Here the change
appeared in an entire plant (except one plant grown during winter
of 1914-15 and not included in these computations), but as several of
these were from a same immediate parent, it may be that the change
really arose as a bud variation, with, however, a delayed effect.
Table 10. — Frequency of changes giving the different types.
Type of change.
Plants.
Frequency.
Ratio.
Increase of yellow and decrease of green ....
Decrease of yellow and increase of green
Reversal of positions of green and yellow ....
Increase of epidermal red to solid red
Decrease of epidermal red, complete loss ....
Decrease of epidermal red, all cases
Appearance of the laciniate character
Entire leaf from laciniate leaf
827
740
450
770
815
815
765
68
27
50
8
8
19
21
13
1
6,1.30
2,960
11,250
19,250
8,580
7,760
11,770
13,600
Of the 68 plants grown with the habit of producing laciniate leaves,
a single case of persistent change to the entire leaf-shape appeared.
There were also 3 cases of clear-cut bud variations involving color
changes in these plants.
SUMMARY AND COMPARISONS.
For the purpose of comparison, the ratios showing the frequency
with which these different types of changes appear are brought together
in table 10. In deriving these ratios the total number of plants in which
there is possibiHty for the change to occur has been considered.
These data indicate the tendencies of the bud variations and give a
clew to the behavior of the characters in question. In the bud vari-
ations, decrease of yellow occurred twice as often as the increase of
yellow. Likewise, the loss of red occurred 2.2 times as often as the
42
ESTABLISHMENT OF VARIETIES IN COLEUS
increase of red. If we consider that the abihty to produce yellow and
red are the more recently acquired characters of the cells, these data
would indicate a tendency toward loss rather than gain of these
characters.
A summary of the data regarding the degree of constancy of the
various patterns and the nature of the variations which they exhibit
is of further interest in a consideration of the tendencies of the vari-
ations. In comparing bud variations which originate in a bud the
comparison on the basis of the total buds produced seems quite ade-
quate. The comparison of fluctuating variations requires a different
treatment. On plants with irregular and mixed patterns it is not
practicable, if possible, to attempt a statistical determination of the
fluctuating branches. Only in few cases when such changes were
Table 1 1 . — Summary of changes occurring in the principal patterns.
Green- Yellow-
yellow- green-
red red
blotched, blotched.
Green-
yellow
spotted-
red
blotched.
Plants
with
laciniate
leaves.
Green-
red
blotched.
Total number plants ' 337
Plants constant for green and yellow. . . 218
Percentage of constant plants 65
Changes in yellow:
Increase:
Frequency 7
Ratio of frequency 9,630
Decrease :
Frequency 32
Ratio of frequency 2,100
Reversal : j
Frequency 6
Ratio of frequency ■ 11,230
Total frequency 45
Ratio of frequency for all bud variations . 1 ,490
41
24
59
3
2,730
11
740
14
590
198
126
63
4
9,900
2
19,800
6
0,600
68
0
0
1
13,600
1
13,600
90
31
34
11
1,6.30
2
6,800
11
1,630
Umited to a branch could there by any degree of accuracy. Further-
more, fluctuations in number and size of the blotches of epidermal red,
although frequent and somewhat persistent, were not recorded. As
long as the pattern was blotched the plants were grouped together and
changes to solid red or to no-red for considerable areas of a leaf were
not considered as a bud variation unless a series of leaves showed that
the change was sectorial for a stem. For this reason the data given in
table 11 are summarized for fluctuations and bud variations involving
yellow and green in patterns with red blotched epidermis.
The percentage of constant plants for yellow and green given in
table 11 is derived by dividing the number of plants which were con-
stant by the total grown of the pattern concerned. This gives an
index of the constancy of a type, although it does not take into account
the varying degrees of the fluctuations which appeared.
BY THE SELECTION OF SOMATIC VARIATIONS.
43
Judging from the data on changes in green and yellow, there appears
to be no general correlation between the number of fluctuating plants
and the number of cases of bud variations. In the green-yellow-red
blotched group there were proportionally more than four times as many
bud variations as in the group green-yellow spotted-red blotched, but the
percentages of constant plants were nearly identical. Not one of the
plants with laciniate leaves was constant for green and yellow, but only
one case of bud variation occurred. In the green-red blotched group
there was chance only for the appearance of yellow, and this change
occurred in a relatively large number of cases, both in fluctuations and
as bud variations.
A very marked contrast appears in a comparison of the two patterns
green-yellow-red blotched and yellow-green-red blotched. Both have about
the same proportions of green and yellow, except that the relative
position is reversed. Both groups agree quite closely in the per-
centage of constant plants. In the latter, however, bud variations
were 2.5 times as frequent. The position of the j^ellow in the center
seemed to increase bud variations involving green and yellow over
that in plants with the yellow at the border.
DISTRIBUTION OF BUD VARIATIONS AMONG DIFFERENT CLONES.
The wide range of variation both of fluctuations and of bud varia-
tions emphasized in the summaries already given was realized in a
series of plants derived by vegetative propagation from two plants
having the same color pattern. The records of pedigrees show that
marked differences appeared among the various clones with respect to
constancy and to the range and the frequency of bud variations.
This is shown quite clearly when the data regarding the main clones
derived from plant 1 are grouped together as arranged in table 12.
Table 12. — General summary of clones.
Clone.
Total
number
plants.
Plants
constant.
P. ct. of
plants
constant.
Number
of bud
variations.
Ratio of
frequency.
11
12
13
14
117
111
211
192
138
155
91
34
132
87
75
80
54
29
62
45
54
51
59
85
49
21
4
18
31
4
860
1,830
6,900
1,720
590
1,700
The main clones 11 and 12 were derived from two branches of plant
1 which had the same color pattern. Although the branches were
identical in appearance, the two progenies were quite different. 62
per cent of clone 11 were constant, while 45 per cent of clone 12 were
constant; but in the more constant clone 11 there were proportionally
44
ESTABLISHMENT OF VARIETIES IN COLEUS
Table 13
—
Record of Clone 117.
Generation
and numbei
of the plant
d
'•1
o
o
a
1
CO
a
o
O
a
03
3
Bud variations.
Remarks.
o
01
tn
=i
O
«
a
03
d
o
ft
T3
-d
o
0)
h
1
".d
d
o
-a
<v
o
_o
"S
d
<u
<v
u
o
01
o
t
Hi
1
"3
-6
"o
a)
1
d
Ol
d
IP
O
-6
O
a
_o
>.
1
d
Ol
o
13
>>
d
01
a
1911-1912
117
1912-1914
117,1
1912-1913
117,11....
117,12....
117,13
117, 14. . . .
117,15....
117, 16
117,17
1913
117,111...
117,112...
117,121...
117,122...
117,132. ..
117, 141. . .
117,142...
117,143...
117,144...
117,151...
117,152...
117,161...
117,162...
1913-1914
117,111,1.
117,111,2.
117, 113.. .
117,122,1.
117, 133.. .
117,134...
117,135...
117,141,1.
117,141,2.
117,142,1.
117,142,2.
117,143,1.
117,143,2.
117,143,3.
117,144,1.
117,151, 1.
117,151,2.
117, 151,3.
. 2
. 2
2
. 2
. 6
■{I
. 6
. 6
6
. 2
2
. 2
. 2
. 6
. 2
. 5
r 2
"I 5
. 5
. 6
. 6
. 6
6
■il
2
2
2
' 6
. 6
. 6
. 2
. 2
. 5
. 5
. 2
. 5
r.
. 5
1
. 6
{I
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
. .
X
X
X
X
X
X
X
X
X
Somewhat greener in winter.
Each part distinct, but somewhat fluctuating.
Somewhat greener in winter.
Do.
Mixed during winter, both greener and yellower.
Constant for pattern, somewhat greener in winter.
Constant for loss of yellow.
Well-defined sectorial bud variations.
Shghtly greener in winter.
Do.
Each part constant.
Slightly more yellow as summer advanced.
Do.
Do.
Very irregular patterns.
Each part constant.
No trace of yellow.
SUght fluctuation in yellow.
No trace of vellow.
Do.
Bud variation verj- marked; parts uniform.
Do.
Do.
Irregular patterns.
Quite uniform and constant.
No trace of yellow.
Constant except for bud variation.
Mixed patterns 2 and 4.
Very uniform and constant.
Slightly greener in winter.
Do.
Constant except for bud variation.
Slightly greener in winter.
Much greener in winter.
No trace of yellow except in two leaves late in summer.
No trace of yellow except in one leaf late in summer.
Shghtly greener in winter.
No trace of yellow.
Very constant.
Constant except for bud variation.
No trace of yellow except in two leaves in late summer.
Died in a few weeks.
Bud variation sharply sectorial.
Constantly very yellow.
Slightly less yellow in winter.
BY THE SELECTION OF SOMATIC VARIATIONS.
Table 13. — Record of clone 117 — Continued.
45
Generation
and number
of the plants
a
si
Bud variations.
m \Xi
TJ
>)
I
a
V
(-1
O
tu
^^
T3
o
I
a>
o o
Remarks.
1913-1914
Cont'd.
117,
152. 1 .
117,
152,2.
117,
152.3.
117,
152,5.
117,
152.7.
117,
153...
117.
154...
117,
155. . .
117.
156. ..
117,
157...
117,
161.1. .
117,
161.2..
117,
161,3..
117,
161,4..
117.
161.5..
117,
162,1..
117,
162,2. .
117,
162,3..
117,
171....
117,
18
117,
19
117,
1X1...
117,
1X3...
117.
1X5...
1914
117,
111,11.
117,
111.12.
117,
123.1. .
117,
133, 1 . .
117,
136, 1. .
117,
136,2..
117,
142,11.
117,
142,12.
117,
142,21.
117,
152,31.
117,
152.32.
117,
152,33.
117,
154,1. .
117,
154,2. .
117,
156, 1 . .
117.
162,11.
/ 1
\ 6
1
6
6
114
6
1
5
6
6
6
6
6
1
6
1
6
1
6
1
6
1
6
6
6
6
6
2
5
5
2
5
2
2
5
5
2
2
6
6
6
5
5
5
14
14
14
5
6
5
6
X
X
X
X
X
X
X
X
X
X
X
Very yellow at all times.
Slightly less yellow in winter.
Very yellow at all times.
Slightly greener in wdnter.
Do.
Do.
Do.
Plant lived, but made poor growth.
Fluctuating to yellow spotted.
Somewhat greener in winter.
Very constant except for bud variations.
Very constant.
Much greener in winter.
Do.
Constantly very yellow, but greener in winter.
Somewhat greener in wanter.
Do.
Very uniform except for bud variation.
Constantly very yellow.
Verj^ uniform except for bud variation.
This part soon died.
Slightly greener in winter.
Do.
Mixed patterns, 4, 5. and 6.
Slightly greener in winter.
Do.
Do.
Do.
Lived only a few weeks.
Soon died.
No trace of yellow.
Very constant.
Verj' constant; no trace of yellow.
Very constant except for bud variation.
Fluctuated to yellow spotted.
No trace of yellow.
Slightly more yellow as summer advanced.
Do.
Slightly more yellow, except for bud variation.
SUght increase of yellow.
Do.
No trace of yellow except in two leaves.
No trace of yellow except in one leaf.
No trace of yellow except in three leaves (seetext-fig.l).
Slight increase of yellow.
Do.
Do.
No trace of yellow except in branch with Inid variation.
Slight increase of yellow.
Do.
Do.
46 ESTABLISHMENT OF VARIETIES IN COLEUS
more than twice as many bud variations. Clone 13, which was derived
from a branch that was green-red blotched, gave a progeny (of several pat-
terns) of which 54 per cent were constant, but bud variations were very
infrequent. Clone 14, with nearly the same percentage of constant
plants, produced four times as many bud variations. This summary
of the data by clones irrespective of patterns shows a general irregular-
ity and lack of correlation between fluctuating variations and bud vari-
ations. The special interest, however, pertains to the clones 11 and 12,
which show that two branches apparently identical may have quite
different potentialities for constancy and for bud variations.
Even more marked differences than these developed among the
various subclones. A study of pedigrees shows that in all patterns
and in all main clones there were certain lines of progeny much more
constant than many others. These could not be detected by any other
than a pedigree method.
Clone 111 can be given as one of the most constant clones. Its
members numbered 34. Four cases of bud variation appeared; 3 were
a loss of yellow and 1 was a reversal of the position of the green and
yeUow occurring in one-half of a leaf only. These 4 plants were other-
wise constant. Only 1 plant showed fluctuating variability, becoming
quite uniform for green-yellow spotted-red blotched. All the bud varia-
tions involved changes in the green and yellow. There were no marked
changes in the amount and distribution of epidermal red. As shown in
table 12, the percentage of constant plants was 85 and the ratio of bud
variations was 1 to 1,700. The clone was highly constant both in regard
to fluctuations and bud variations.
On the other hand, the series of plants derived from plant 117 was
26 per cent lower in number of constant plants and gave nearly three
times as many bud variations, yet plants 111 and 117 were both
uniform and constant for the pattern green-yellow-red blotched and were
apparently identical. Until the autumn of 1912, plant 1171 was the
only one of the 17 plants grown in clone 11 that showed variation. It
gave during the summer, by sectorial variation in the main axis, 6
branches with the position of the green and the yellow reversed. The
plant was grown in a large pot during the winter and then grown out
of doors during the following summer. In the second sunmier two
more bud variations appeared on the part with green-yellow-red blotched
foliage, but on branches quite separated. Both were sectorial; one was a
loss of green, giving the yellow-red blotched pattern, and one was a loss
of yellow, giving the green-red blotched pattern. The plant possessed for
some time four patterns, each uniform for a certain part of the plant.
The record of pedigree for the progeny of the plant can be given as
illustrating a clone in which bud variations occurred with a high ratio
of frequency. In table 13 the plants are arranged in generations
according to number. To trace the progeny or the ancestry of any
BY THE SELECTION OF SOMATIC VARIATIONS. 47
plant, one should look in the generations following or preceding for the
serial numbers. Certain plants were grown with two patterns as a
chimera, and these are indicated by brackets, with a record for each
pattern. In order to make the tabulation more compact, numbers
are used to represent the different patterns, and these correspond with
the numbers of the figures in the plate, as follows:
1 = yellow-red blotched. 5 = green-red blotched.
2 = green-yellow-red blotched. 6 = yellow-green-red blotched.
4 = green-yellow spotted-red blotched. 14 = yellow-green.
A survey of this series of plants shoAvs that on the 91 plants 31 bud
variations appeared, giving a ratio of frequency of 1 to 590 against
1 to 860 for the entire clone 11 and 1 to 1,700 for the sister clone 111.
A further analysis within this progeny shows that in several cases
similar bud variations can be traced to a common ancestry. For
example, 6 of the 9 cases of what has been called spontaneous develop-
ment of 3^ellow occurred during the summer of 1914 in plants descended
from plant 117142 (see table 6), which itself was constant for loss
of yellow during its period of growth. Another case of spontaneous
development of yellow was in plant 1171441. In this clone, therefore,
all cases of spontaneous development of yellow were in plants descended
from plant 11714.
It is quite clear from such pedigrees that distinct differences in
tendencies in regard to the degree of variation may exist among buds
of branches bearing similar foliage.
ENVIRONMENTAL INFLUENCE.
Observations were made and pedigrees of plants examined to deter-
mine whether changes in ordinary environmental conditions influence
fluctuations and bud variations.
To secure accurate data on the relative frequency of bud variations
during summer and winter is hardly feasible. In general only about
half of the summer plants were from cuttings made early in the spring.
The others were grow^n in pots in a greenhouse during the winter and
transplanted to the garden, where the}' grew during the summer,
making larger plants with many more branches than were produced
during the winter. Of the total number of 1 15 bud variations iuA'olving
color, 9 appeared during the winter, which is fairly proportional to the
relative number of branches that developed.
There was a strong tendency' for plants having yellow to become
greener in winter and yellower in summer, and also to become greener
when severely pruned. At any time, however, during the winter,
some plants of each pattern having yellow (for example green-yellow-
red blotched) could be found with the pattern as pronounced as dur-
ing the summer. During the winter of 1913-14, two plants of each of
the following types, green-yellow-red blotched, green-yellow spotted-red
48 ESTABLISHMENT OF VARIETIES IN COLEUS
blotched, and green-yellow-solid red, were grown in a greenhouse with
northern exposure only, which gave scarcely any direct lighting. These
plants were constant for the respective patterns. The green and
yellow, however, were slightly less bright and intense.
During the summer of 1914 several plants of nearly all types of
pattern containing yellow were grown in a greenhouse the glass of
which was whitewashed to decrease the intensity of illumination. No
appreciable differences could be noticed between the patterns of these
and of plants grown out of doors. So far as I have observed, it does
not seem that any of the color variations can be attributed to such fac-
tors as heat or degree of illumination.
Furthermore, the loss of yellow, loss of green, and gain and loss of
red all occurred in single branches and in sections of branches (figs.
21 and 24). Frequently two quite difTerent changes appeared on the
same plant, which was then grown for some time with 2, 3, or even
4 quite distinct types of foliage. Cuttings were made so as to give
plants with two types of foliage, as (a) green and green-red blotched, or
(6) green-yellow-red blotched and green-red blotched, or (c) green-yellow-red
blotched and green-yellow-solid red, etc. In all these cases branches
with two types of foliage were submitted to as uniform conditions
as possible; they grew on the same plant, were subject to the same
degree of heat and illumination, and were supphed by the same root
system. Under this test the different patterns were fully as constant
as if grown on separate plants.
These facts indicate quite clearly that the marked and sudden
variations and differences in expression of color concerned in the
different patterns are not readily attributable to external environ-
mental factors.
Flammarion (1898) used a variety of Coleus with a yellow, green,
and red color pattern in testing the influence of hght on pigmentation
in plants. With red light he secured decrease of red pigmentation
and a broader leaf; under influence of green light the red coloration
mostly disappeared, and under blue light there was somewhat less red.
A series grown out of doors under conditions of diffused hght showed
decrease of red coloring, while those under very dim light gave still
less development of it. With decrease of red the center of the leaves
became quite yellow. Evidently the red pigmentation of his variety
was chiefly located in the epidermis. In marked contrast to these
results it may be noted that the bud variations that I have reported
give more marked changes than those induced by Flammarion and
that these appear suddenly and in a sector of a bud in a manner that
suggests internal readjustments rather than external environmental
influence.
In respect to the laciniate leaf-shape and its periodic development,
however, environmental influences seem to have some effect. As
BY THE SELECTION OF SOMATIC VARIATIONS. 49
already indicated, conditions favoring rapid and vigorous growth lead
to the development of entire leaves. In vegetative propagation,
however, the periodic laciniate condition develops only in certain
subclones. In other subclones it has not appeared. That is, the same
conditions of environment and treatment do not lead to the appear-
ance of the laciniate character in all plants, so it is hardly to be con-
sidered as purely environmental. The laciniate character has also
developed in one plant in the manner of bud variations.
SEED PROGENY.
The data obtained from the seed progeny of my strains of Coleus
have direct bearing on the nature and inheritance of the bud variations
that appear and indicate that bud variations can give rise to as widely
difTerent forms as can be obtained among the various members of a
hj^brid progeny.
Selfed seed was obtained from a plant of the pattern yellow-green-
red blotched and 22 plants were grown during the latter part of the
summer of 1914, In respect to the development of green and yellow,
there was every gradation between green with large yellow blotches
irregularly distributed through the leaf-blade and pure green. In
respect to the development of red in the epidermis, there were grada-
tions from absence of red to a general distribution of large irregular
blotches. As in the case of bud variations, the difTerent types of epi-
dermal red occurred independently of the degree of development of
underlying green and yellow.
In regard to leaf-shape, there was every range of variation. Few
plants could be classed as laciniate, but there was every gradation from
shallow to deep lobing and from coarse to fine lobing. The leaves on
any one plant were quite uniform. It should be noted that the lacini-
ate character had not appeared in the particular subclone from which
these seedlings were derived. It had appeared in sister subclones
as described above. Seven plants possessed leaves quite like those of
the parent type, both in respect to the cuneate base and the crenate
margin. Five plants, however, had large leaves, some measuring 10
inches long, that w^ere broadly obtuse at the base, with the cuneate
character lacking. From plants possessing fiat leaves with a smooth
surface there was gradation to those with leaves much crinkled or
folded.
At the same time 23 plants were grown from selfed seed of a plant
which possessed the laciniate leaf-type as a fluctuating character.
The plant itself also fluctuated in respect to the development of green
and yellow as follows: during the winter of 1913-14, its leaves were
strongly laciniate and devoid of yellow, and during the following sum-
mer the plant was quite yellow, becoming almost like the type green-
yellow-red blotched (fig. 2) and every leaf was entire. In their develop-
50 ESTABLISHMENT OF VARIETIES IN COLEUS
ment of green and yellow the plants ranged from pure yellow plants
that died within a few weeks to those that were pure green. Two
of the plants had much yellow in very irregular and mixed patterns.
Eleven had no trace of yellow for several months, when a few yellow
spots appeared in some of the leaves on several plants. In regard to
epidermal red there were numerous types ranging from solid red to no
red. Of those wdth red-blotched epidermis, some were uniformly
finely blotched, while on others the blotches were large, with a single
blotch sometimes covering one-fourth of a leaf. As to the shape of the
leaves, the series showed the same range of variation exhibited by the
seed progeny described above. There were but 5 that were strongly
laciniate.
Besides the above plants which were grown to maturity, there are
at the present date (February 8, 1915) 20 seedlings of a plant that
had an epidermis of solid red, as shown in figure 8. The plants have
from 2 to 3 pairs of leaves, but it is clear that only one of the seedhngs
has a solid red epidermis. Red-blotched types prevail and few of the
seedlings show any yellow coloration.
Summarizing, it is plain that the plants grown from seed give wide
variations. In respect to color patterns, there were numerous types
which gave very complete gradations between extremes, especially in
regard to epidermal red. Many of the types that had appeared as
bud variations appeared also in the seed progenies, such as yellow-red
blotched, green-yellow spotted-red blotched, green-red blotched (wide range
of variation in respect to size of blotches), green-yellow spotted, and green.
One of the new patterns could be described as yellow-green blotched-
solid red. Another had the red blotches of the epidermis coalescing at
certain points, making the red markings continuous but not solid (see
fig. 29), so that the underlying green showed through in blotches.
In vegetative propagation leaf-shape remained very constant, the
only exception being the clones that developed laciniate leaves. In
the seed progeny of all plants tested, however, new types of entire
leaves have appeared. The laciniate leaf characters and various
intermediates between it and the typical entire leaf appeared in the
first leaves of certain plants and remained as a rather constant character
during the time the plants have been grown.
The wide variations appearing in the seed progeny indicate that this
strain of Coleus is either of mixed parentage or that the processes
concerned with production of color patterns and leaf-shape are them-
selves subject to wide variations. The variations in respect to color
patterns were, however, no greater in range or extent than were those
that appeared in bud variations, and the fluctuations from entire
to laciniate leaf-shape gave extreme types of leaf-shape, with all grades
of intermediates, on a single plant, as is quite well shown in plant 123153
of plate 4.
BY THE SELECTION OF SOMATIC VARIATIONS. 51
HISTORY OF COLEUS.
The early historj^ of the cultivated varieties of Coleus shows that the
original species utilized in cultivation and hj'bridization were few in
number and relatively simple in respect to diversity of color patterns
and leaf-shape. The variations which appeared in vegetative and in
seed propagation within a few years after introduction gave a wide
range of variabiUty, with greater extent than the extremes of the char-
acters in the original species. In general the variations reported in this
paper are quite parallel with those appearing in the development of the
numerous cultivated varieties, both in regard to the apparently spon-
taneous development of new patterns and to the reversion to parent
or ancestral types.
It appears that the first variegated species of Coleus introduced
into European cultivation was Coleus blumei Bentham. The original
description (Blume, 1826) under the name Plectranthus scutellarioides
states that the leaves were spotted above with dark purple (folio
supra maculis atropurpureis picta). This plant was introduced into
Holland in 1851 and the next year Planchon (1852) gave a brief de-
scription of it, accompanied by a colored plate. It was soon intro-
duced into England, and in 1853 a description with colored plate
appeared in an English magazine (Hooker, 1853). These two illustra-
tions agree quite closely, although the latter shows the plant in some-
what a brighter green, with leaves with a somewhat more solid mass of
central red. The central part of the upper surfaces were dark purple
or sanguineous, breaking into spots near the margin. In the descrip-
tion it is stated that the leaves were entire at the base, ''which is gradu-
ally attenuated into a more or less elongated petiole," a character well
shown in an outUne drawing of a leaf.
There are no specimens of this species in the Bentham collections
at the Kew Herbarium. There is, however, a specimen in Herb.
Hookerianum with a label in Sir WiUiam Hooker's handwriting, stating
that the plant w^as grown at Kew Gardens and citing the description
and plate in Botanical Magazine (Hooker, 1853). This was evidently
regarded by Hooker as a typical specimen. The leaf-shape is identical
with that of the illustration referred to and is, with the exception of fig-
ure 7, quite the same as the leaves shown in the plates illustrating this
paper. From the description and illustration the color pattern was
nearly identical with my type green-upper center solid red (fig. 15).
Morren (1856) describes the variety C. blumei pectinatus as somewhat
more richly colored, but differing chiefly in having the leaves deeply
and doubly lobed. The colored plate shows that the base of the
leaf-blade was cuneate, as in the species.
Coleus verschaffeltii was first named by Lemoine (1861), who decided
that it was distinct from C. blumei. It appears (Witte, 1862) that this
plant was introduced into Rotterdam in 1860 from Java. Colored
52 ESTABLISHMENT OF VARIETIES IN COLEUS
plates (111. Hort., 8, pi. 293, and Flor. Mag., 2, pi. 96) show that this
species was richly and deeply colored with crimson on both surfaces
and that the base of the leaf -blades were not cuneate but heart-shaped.
Being richer in color than C. hlumei, this plant attracted considerable
attention as a foliage plant. In 1864 (Gard. Chron., p. 506), a sport
of this species called marmoratus with bright green patches in its leaves,
was described.
Concerning C. verschaffeltii, Herincq (1865) remarks that the expres-
sion of red coloration fluctuates with light conditions and suggests
that one might eliminate the red by keeping the plants in the shade.
Later he (1866) notes fluctuations that are less due to environment,
for he observes variation in the leaves of a single plant and states that
no doubt selection of cuttings would give pure-green plants. He men-
tions that he had seen a young plant having no trace of red coloration.
Two types of Coleus destined to play an important part in the
development of horticultural varieties were introduced into England
from New Caledonia by John G. Veitch. Although briefly mentioned
in 1866, they were first described and illustrated in 1867. C. gibso7iii
(Verlot, 1866 ; Dombrain, 1867a) was of a dwarf bush}^ habit. The leaves
were large and ''of a light-green color, distinctly veined and blotched
with dark crimson-purple." The plate clearly shows that the leaves
were only slightly crenate and that the bases were broadly cordate.
C. veitchii (Dombrain, 18676) possessed leaves quite similar in shape,
but with the entire central portion of the leaf of a deep chocolate color
with the edges green.
These four species, C. hlumei, C. verschaffeltii, C. gibsonii, and C.
veitchii, were used as parents in the production of hybrids by F. Bause,
in the employ of the Royal Horticultural Society of London. 12
hybrids, of which C. verschaffeltii was the seed parent, resulted the first
year. Rather extended descriptions of these are given by Thomas
Moore (1868). The Fi progeny, even from the same parentage, were
widely different, some resembling the seed parent, while others resem-
bled the pollen parent. In regard to leaf-shape, there were two groups,
one with flat crenate leaves, as C. veitchii, and one with frilled-dentate
leaves, as C. verschaffeltii. In colors there were various shades of purple
in solid colors, blotched areas, and in reticulations. At the time this
variation among the Fi progeny aroused considerable interest. One
anonymous writer (Gard. Chron., 33, 407) raises the question how
several kinds of Coleus could originate from the same cross. But
one hybrid with C. hlumei as a parent is reported. This had frilled
leaves and coloration much like that of C. hlumei. It was less deeply
colored than the hybrids resulting for the other pollen parents.
There is no mention of yellow in any of these hybrids. All were
bi-colored, but with striking combinations of the green and various
shades and amounts of purplish or red colorations. They were sold
BY THE SELECTION OF SOMATIC VARIATIONS. 53
at auction (Gard. Chron., 33, 432) for the aggregate sum of £390.
One hybrid brought 59 guineas.
The production of these valuable variegated Coleus varieties stimu-
lated further hybridization work. Wilham Bull produced 18 different
types (reported by Herincq, 1868), and in November of that year he
advertised pedigreed seed from 20 crosses involving 14 varieties grown
that year (Advertisement, Gard. Chron., 33, 1232).
Meanwhile, at the gardens of the Royal Horticultural Society, a
new series of hybrids were produced much finer than those of the
previous year. The parentage of these interesting hybrids is not fully
given, but it is stated by Moore (1869) that certain of the hybrids of
the previous year were crossed with C. hlumei itself. In this second
lot of hybrids, yellow coloration appeared as a new or spontaneous
development. Eight (Gard. Chron., 33, 1210) possessed distinct
yellow, forming in some cases a golden margin. Two (Prince Arthur
and Princess Beatrice) are described as having a yellowish ground-
color or golden green. Most of them had a yellow and green ground-
color overlaid with shades of purple or crimson red. The most brilliant
of the series was named Queen Victoria, a colored plate of which
appeared as a frontispiece in the Florist and Pomologist (volume for
1869). This plate shows that the ground-color was mostly yellow,
overlaid by an epidermal red, appearing crimson over the yellow and
entirely covering the upper surface of the leaves except at the margin.
None of the series possessed frilled leaves. The leaves of all w^ere
flat, with crenate teeth somewhat deeply cut.
Although of much more remarkable variegation than the hybrids of
the previous year, 9 of these new coleuses brought but 65 guineas.
While the yellow element in the variegation appeared strongly in this
second lot of hybrids, it should be noted that in the year 1867 (Gard.
Chron., 33, 460) a golden Coleus arose as a bud-sport from C. hlumei.
It is described as like C. hlumei, but with the green exchanged for a
decided yellow tint. The sport appeared in one-half of a single leaf.
The bud at its base was propagated and gave the new variet3^ It
does not appear that this sport was used in the hybridization work that
produced the golden coleuses.
During 1868 and 1869, the various horticultural publications men-
tion by name no less than 54 new varieties of Coleus. For several
years thereafter few varieties were mentioned, but in 1878 (Garten-
flora, p. 50) 13 forms not previously mentioned are listed. The next
year this journal (pp. 341-346) states that breeding of Coleus had been
carried on in Germany, speaks of new forms that arose, and prints an
uncolored plate illustrating 4 types. New types were also credited to
Bull (Rev. Hort. Belg., 5: 49; Gard. Chron., 45: 748).
At the exposition in Paris in 1879, Morlet exhibited varieties described
by Andre (1879) as far surpassing all previous varieties. These had
54 ESTABLISHMENT OF VARIETIES IN COLEUS
enormous foliage, with remarkable combinations of shades of carmine,
yellow, and green. 20 types are mentioned by name and 12 of these
are described.
Pynaert (1881) states that about 250 varieties of Coleus had been
put upon the market. He notes that it is difficult to establish a nomen-
clature for the various Coleus varieties. He describes and illustrates
in color a variety called Reine des Beiges which arose from seed of the
variety Duchess of Edinburgh, a variety illustrated in the Floral
Magazine in 1874. The leaves of the Duchess of Edinburgh pos-
sessed a yellow border; the Reine des Beiges had the yellow in the
center of the leaves. The relative position of the green and the
yellow was therefore reversed, which is the difference between tj^pes
green-yelloiv-red blotched and yellow-green-red blotched already reported
in this paper.
From the evidence at hand it is clear that a large number of types of
Coleus have been produced. Probably the same type or nearly identical
types have been given different trade names. With the exception of
the first hybrids produced by Bause, there is almost no record of the
parentage of most types. The plants attracted attention solely on
account of their variegated foliage, and for a time were more extensively
used as bedding and foliage plants than they are at the present time.
From the standpoint of genetics, it is suggestive that such wide
variation appeared in the cultivation and hybridization of the 4 species
already discussed, although it should be noted that there is the possibil-
ity that other species were concerned in the parentage of some of the
varieties now in cultivation. The parent species possessed a green
background, or at least were without pronounced yellow. The epider-
mis especially was more or less colored with purple or red in blumei,
verschaffeltii, and veitchii, while in gibsonii the purphsh coloring was
largely confined to the veins.
In the varieties derived from these, yellow appeared as a pronounced
part of the coloration. Some tjqjes were largely yellow, others were
pale yellow, and others were entirely green. In many the yellow was
localized at the border, but in others it was at the center and in others
the yellow blotches were well distributed. These variations in yellow
and green were combined with variations in amount and quality of
epidermal and internal (chiefly in veins) development of purplish and
red tints.
The historical evidence indicates that the form of Coleus used
in the experiments here reported is derived from C. blumei. In
respect to the cuneate base of the leaf-blades and the marginal charac-
ter of the first pair of lateral veins almost making the petiole attenuate,
the leaves of all types but the laciniate agree almost exactly with
that of the figure for C. blumei (Bot. Mag., 1853). This character dif-
ferentiated sharply C. blumei from C. verschaffeltii, C. veitchii, and C.
gibsonii or any other species introduced into cultivation, and seems to
BY THE SELECTION OF SOMATIC VARIATIONS. 55
indicate clearly that the type in question is derived more or less
directly from C. hlumei. The flowers agree with the description of
C. hlumei.
Coleus hlumei, as stated above, was already in cultivation in Java
when described by Blume. Blume (1826) suggests that the plant
named by him Plectranthus laciniatus may have been simply a variety,
as it seems to have differed largely in having laciniate leaves. Blume's
original description speaks of the leaves of C. hlumei as spotted above,
but the colored plates appearing in 1852 and 1853 show the greater
part of the upper surface of a soUd purphsh color. Coleus hlumei was
introduced into the German gardens under the name Plectranthus
concolor var. picta (Gartenflora, 1853, 2: 220). Only in one of Bause's
first lot of hybrids was C. hlumei concerned, but this species was, it is
stated, the seed parent of the second lot of hybrids. If this hybrid was
used in further hybridization work there is no record. It was the least
brilliantl}' colored, possessed no yellow, and sold for the sum of 5 guineas,
which was the lowest sum paid for any one of the 12 hybrids (Gard.
Chron., 3>o, 432). I have been unable to find further mention of this
hybrid, which was named reevesii.
C. hlumei produced in 1868 a bud sport with the green changed to a
decided yellow tint. Through propagation this gave rise to the variety
telfordi aurea (Gard. Chron., 33, 460).
Andre (1880) illustrates and describes 4 new varieties, which he
attributes to C. hlumei. Apparently all have, however, leaves with
cordate bases and not at all cuneate, as is the case in C. hlumei, which
makes his determination of doubtful validity. The same is true of the
type Gloire de Dijon, described and figured by Rodigas (1888). Rodi-
gas later (1892) notes the wide variability obtained from the seed prog-
eny of what was considered as C. hlumei from Chile. The 4 derived
types illustrated possess, however, strongly cordate leaves which make
the identity with C. hlumei doubtful.
At the present time it does not appear that any pure strains of C.
hlumei, C. gihsonii, or C. veitchii are in cultivation. The strain used
in these experiments agrees most closely in regard to leaf-shape with
the original C. hlumei, but the variability of the seed progeny seems
to indicate that it is not a pure strain. The writer inquired about and
observed all types of Coleus available at numerous botanical gardens
and nurseries during a 6 weeks' trip to Germany, Holland, and Eng-
land during the summer of 1914. Only one plant, a plant observed
at the Royal Botanic Gardens at Regent's Park, London, was seen
which had the hlumei type of leaf.
Coleus verschaffeliii is, however, quite generally in cultivation at the
present time and agrees quite closely with the type first described.
I am especially indebted to Mr. F. J. Chittenden, of the Royal
Horticultural Society, for securing a statement (a letter to Mr. Chit-
tenden) from Mr. B. Wynne, in which he states that he spent 3 months
56 ESTABLISHMENT OF VARIETIES IN COLEUS
of his time (as a student) at Chiswick in 1866, under Bause, in the
propagating department. He was quite famihar with the methods used
in the development of the hybrid coleuses and states that "it fell to my
lot to convey the first half dozen coleuses sold to Sleven's rooms."
Mr. Wynne refers to the description (see reference above) of the hybrids
as adequate. In regard to the present existence of these hybrid types,
Mr. Wynne gives the following statement:
"I am unable to say whether there is more than one of the Chiswick-raised
set of Coleus in existence now, but I very much doubt it. It is very interest-
ing, however, that at least one of them has survived and is still grown for
Covent Garden and possibly for other markets. This is the variety originally
named Queen Victoria, which received a first-class certificate (R. H. S.) in the
autumn of 1868, and was bought at the second sale by the now extinct firm of
John and Charles Lee, of Hammersmith. I do not think it is now known in
the market by its original name, but it is the well-known variety with choco-
late leaves and golden segments and I have no doubt about its identity with
our old Chiswick plant."
It should be noted that Wj^nne's description of this form as "choco-
late leaves with golden segments" does not agree with the colored
plate in the Florist and Pomologist for 1869. The latter represents the
pattern as solid crimson in the center, with well-defined yellow border.
While Coleus is now regarded with less favor than it formerly
received, there are many types in cultivation exhibiting great range
of color patterns and leaf character. At Erfurt, Germany, during
the summer of 1914, the writer saw large collections of Coleus grown for
seed for the trade. In the Ernst Benary collections they were chiefly
of large-leaved, small-leaved, and f ringed-leaved tj^pes. In the large-
leaved plants the yellow, if present, was usually in the center, a condi-
tion which was true without exception for the plants with small leaves.
The fringed-leaved types possessed most curious doubly cut prolifera-
tions about the margin. Of the entire collection hardly any two plants
were ahke as to color distribution. There were, however, fewer classes
in regard to leaf-shape. One of the newer tj-pes was a dwarf with
fringed leaves derived by selection.
At the greenhouses of Haage and Schmidt there were 2 rather definite
types with laciniate leaves. One is salicifolia, with narrow, slender,
quite irregularly-lobed leaves. Another is quercifolia, with broad
leaves coarsely cut with round-tipped segments. Their large-leaved
types, entire and fringed, showed great diversity of color patterns.
The plants are grown for seed, and in producing stock for seed parents
selections are made with special reference to leaf-shape and general
habit of growth. In general no selection is made with reference to color
patterns and each type exhibits wide variation in this respect.
From the history of Coleus it seems quite clear that the numerous
and diverse varieties have arisen from few species. These varieties
exhibit many characteristics of coloration and leaf-shape that were
BY THE SELECTION OF SOMATIC VARIATIONS. 57
not possessed by any of the parents; for example, there is no evidence
that a yellow element of variegation was present in any of the parent
species. In leaf-shape also, many new types have arisen. Hybridi-
zation and rather intense and artificial cultivation have been associated
with the development of such diversity from plants comparatively
simple.
Considering the various types of Coleus as a whole, we may note
that the wide range of variability which they exhibit is in large measure
realized in the bud variations that have appeared from the single
types here reported. The green, green-yellow, yellow-red blotched,
and green-solid red are extreme forms that are in degree and quality
counterparts of the extremes seen in the different varieties. In the
development of the laciniate character of the leaf-shape in which the
leaves on a single plant fluctuate from entire to extremely laciniate,
the counterparts of cut-leaved quercifolia and salicifolia types are
in large degree realized.
In seed progeny and in bud variations the single strain of Coleus
investigated has in the 3 years of observation shown the same types of
variation that have developed in the entire series of cultivated varie-
ties derived by both seminal and vegetative variations.
58 ESTABLISHMENT OF VARIETIES IN COLEUS
DISCUSSION.
The appearance and subsequent behavior of bud variations in Coleus
present numerous analogies to various phenomena of variation exhib-
ited by members of a seed progeny of hybrid origin.
In respect to the definiteness of the characters contrasted, green and
yellow, red and non-red, the color patterns arising by bud variations
are as different, at least in their extremes, as one could expect in the
members of a seed progeny even of hybrid origin. This is especially
noticeable in comparison with the seed progenies of Coleus itself. The
solid-red epidermis and the no-red epidermis represent two extremes
fully commensurate with the so-called presence and absence of a
character and the bud variations giving these were fully as different as
the types arising in seed progeny. The intermediates red blotched and
solid red upper center are pattern characters equally distinct both in
manner of appearance and in vegetative constancy. The same is true
of the extremes of development of both green and yellow.
Plants of the same pattern in a single line of descent, both in the
same generation and in successive generations, frequently produced
the same type of variation independently, a behavior quite analogous
to the segregations that reappear in each generation of a hybrid line
or in the successive generations of the progeny of a mutating plant.
Many of these variations show a return to a parental pattern, just as
a recessive parental quality reappears as a result of segregation in
hybrid progeny. There is much in such reappearance of patterns that
is quite identical with the phenomena of alternative inheritance.
Two plants identical in appearance and derived from adjacent
branches on the same plant may give quite different progenies in suc-
sessive generations. One line may be very constant and uniform, the
other may give numerous bud variations of wide range. This is a
familiar phenomenon in hybrid seed progenies where certain of the
plants of any generation, although apparently identical, give quite
different progenies. In Mendel's experiments with Pisumjor example,
although the F2 yellow peas of the cross green X yellow were similar,
they gave different progenies. Some produced only yellow peas, while
the progeny of others gave both yellow and green.
In the strains of Coleus studied b}^ the writer, certain types of
bud variation occur more frequently than others. Loss of yellow was
more frequent than loss of green, and loss or decrease of red was
more frequent than increase of red. In the entire series of plants
derived by vegetative propagation there was decided predominance of
green over yellow, of red blotched and no red over solid red, of the blumei
character of leaf-base, and of the shallow or crenate lobing of the
leaves. These same characters show marked tendencies for domi-
nace among the members of the seed progeny.
BY THE SELECTION OF SOMATIC VARIATIONS. 59
Such phenomena of variation appearing in hybridization experiments
are usually considered as due to segregation and recombination of
hereditary units during the processes of self- or cross-fertilization.
Bud variations in vegetatively propagated plants are, of course, inde-
pendent of such recombinations.
That bud variations are generally due to a complete loss during cell
division is not substantiated by the results here reported or by the
bulk of other experimental work. In the majority of cases the
character concerned does not breed true. Mendelian students have
interpreted this to mean that such bud variations are produced by a
loss of only one factor of a diploid pair, giving heterozygocity. This
illustrates the tendency of the Mendelian interpretation to assign the
numerous cases of fluctuation in characters to heterozygocity rather
than to fluctuations in a factor or to irregular mutational changes
spontaneous in the organism.
In regard to the range of expression in a single plant the laciniate
leaf-shape is a more striking character even than the color patterns.
It arose, as already noted, in 13 individuals obtained by vegetative
propagation, but these were all derived from a few plants of the next
preceding generation. This character was inherited through vegeta-
tive propagation by all plants grown but one, but the leaves on each
individual plant varied from deeply laciniate to fully entire. Plants
raised from seed gave all types from extremely laciniate to fully entire,
the particular type appearing in the first leaves that developed and
remaining quite constant for all leaves developed in the 6 months that
the plants have been grown. The special point of interest is that a
single individual of the laciniate group passed through a series of
fluctuations, giving all grades of leaf-shape from entire to fully laciniate.
The range in a single plant is greatly more marked than the dilTerences
between the Urtica hybrids (Correns, 1905, 1912), in which the serrated
type of leaf was dominant. In the hybrids of the normal and the
laciniate types of Chelidonium majus (de Vries, 1900) there seems to be
no published data regarding the range of variation in the F2 generation.
Of the hybrids between palm-leaf and fern-leaf types of Primula sinen-
sis, Bateson (1909) states that ''dominance is usually complete,"
but that he has seen two strains with intermediate leaf-shape. Greg-
ory (1911) states that 'Hhe palmate character is dominant, though a
slight difference can sometimes be recognized between pure and hetero-
zygous palmate types." Crosses between an ivy-leaf (a palmate
shape with margins crenate) with the fern-leaf gave the normal palmate
leaf as an Fi hybrid. The F2 generation exhibited a wide variation,
which Gregory groups into 4 classes and assumes that shape of the
entire leaf and crenation of the margin are two independent characters.
On the basis of character of lobing of climax leaves, Shull (1911)
distinguishes four biotypes in Bursa hursa-pastoris. The view that
60 ESTABLISHMENT OF VARIETIES IN COLEUS
hybrid forms always segregate out into only these types is somewhat
in doubt, for Shull (p. 9) finds that a plant classed as simplex gave
unexpectedly a mixed progeny with defective ratio, so that it is clear
that the assumed hereditary "gene" became less potent. Hus (1914)
distinguished in a culture of Capsella hursa-pastoris 4 forms different
from those of Shull and added another factor which he considers
determines the narrow character of early leaves in certain forms.
In none of these studies have the individuals of the F2 exhibited
greater variation among themselves than have single plants of Coleus
with the laciniate leaf, nor have the individual parent plants been
more distinct and uniform as a plant than the plants derived from
seed progeny of Coleus, and grown for a period of several months.
Furthermore, there has been evidently no attempt to select persistently
intermediate types for modified potency of characters. Furthermore,
emphasis has not been laid to selection of variations in a seed progeny
of a single individual or in a line propagated vegetatively. The
more intensive Mendelian studies, such as those by Shull, Hus, and
Gregory, indicate that the character of leaf-shape is complex and that
selection studies along the lines indicated ma\' reveal further data
on variations in the potency of characters.
The historical evidence and the studies of seed progeny reported
above indicate that the strain of Coleus studied is most certainly of
mixed parentage. If this strain had been studied solely in its seed
progeny the variations obtained would be attributed by many modern
geneticists, I venture to imagine, to chance combinations of hereditary
units. Yet, as has been fully reported above, these variations are
analogous to and even identical in nearly all cases with those arising
by bud variation.
At this point we may note that modern genetics have furnished no
evidence as to the real nature of the characters considered. What
we may call the genetical or breeding value of characters has alone
been emphasized. Characters have been considered solely in regard
to their unity of expression in successive generations of plants of
selfed or hybrid origin. Special emphasis has been placed on the
reappearance of characters and upon their phylogenetic significance.
From this standpoint we may further consider specifically the char-
acters concerned in these studies of Coleus.
Considering first the color characters, we note that the pattern of
the parent plants was a mosaic of green, yellow, red (or blue), and non-
red cells. In the mature leaf the cells are apparently qualitatively
different, and furthermore, the color differences between the various
cells are identical with the color differences between entire leaves
and branches derived by bud variation. The contrast between pure
j^ellow and pure green leaves in the bud sports and on plants of seed
origin is the same that exists between green and yellow cells that may
BY THE SELECTION OF SOMATIC VARIATIONS. 61
be adjacent in the same leaf. Likewise, the differences between
solid-red epidermis and the non-red epidermis is the same that exists
between red and non-red cells in the epidermis of a single leaf having a
red-blotched pattern. In the blotched patterns the cells of the epi-
dermis are either red or non-red. The number of similar cells that
are adjacent to each other determine the size of the blotches.
The facts regarding development and distribution of the colors raise
sharply the question whether the possibility for development of red,
yellow, and green is possessed by every cell that is formed or whether
these possibilities have been sorted out by qualitative divisions in a
Weismannian sense. A sharp distinction should be made between a
character that is metidentical and one that can only belong to a group
of cells. In Detto's (1907) sense, power to produce green, yellow, and
red seems to be metidentical; that is, this abiUty may be strictly a
property of all cells, while the different patterns appearing so strikingly
as characters of entire leaves can not in any sense be metidentical.
Most important evidence regarding the development of leaves with
red-blotched patterns was obtained by the cytological studies conducted
by Mr. E. G. Arzberger. In the early stages of leaves that later will
become red-blotched, all the epidermal cells possess red pigmentation,
forming a complete periclinal layer of red cells, and in this respect the
early stages of leaves with solid-red and red-blotched patterns are alike.
In solid-red patterns all cells continue to possess red sap color, while
in blotched types the red disappears in certain cells. The evidence is
clear in these cases that all the epidermal cells arise from cells having
red cell sap and in this respect are potentially alike.
The differences in patterns in respect to red are largely those of
quantity; the total of red cell sap in a leaf is comparatively large
in solid-red patterns and much less in blotched patterns, of which there
is every grade to the no-red epidermal patterns. Differences in dis-
tribution are also involved. There is the tendency for red to be in the
epidermis rather than in the subepidermal tissues. Usually it is in
both upper and lower surface, but in one pattern (fig. 10) the red is
almost entirely massed in the upper epidermis and in the center of the
leaf. In the blotched types the number, size, and degree of coalescing
of the blotches present every gradation from a finely blotched condition
to a sohd red.
While the total amoimt of red and its distribution in the epidermis
determines quite definite and constant patterns, it should be said that no
plant has been obtained, either in seed or vegetative propagation, that
was entirely devoid of red. Subepidermal cells may also possess red
cell sap; red sap is quite pronounced in veins of leaves (see figs. 13
and 14), and varying amounts can be seen in stems. In some plants,
especially those of seed origin, no trace of red can be seen except at
the nodes, and here the amount for different plants varies from faint
62 ESTABLISHMENT OF VARIETIES IN COLEUS
rings to quite a definite band or zone of red cells extending through the
stem. Also the stems of plants, especiallj^ those with blotched patterns,
may have large irregular streaks of red-colored tissue and we may say
are internally blotched. In general, plants with a decided nodal
zone of red were those with epidermis free of coloration, and on the
other hand, plants with solid epidermal red or heavily blotched epider-
mal red had as a rule slight localization of red in zones. There was also
the marked localization of red in upper center of leaves (see fig. 10), with
almost complete absence of red below, although in many leaves of this
type small blotches of red were evident on the under side (see fig. 10a).
All these conditions indicate that the possibility of producing and
possessing red cell sap is a specific property of all cells and that the
distribution giving localization at nodes, in streaked areas in stems, and
in subepidermal tissues, and in the epidermis either as a uniform red or
a blotched red are dependent on two fundamental conditions : (a) total
amount of pigmentation, and (6) the appearance of it in certain centers
of concentration. The facts as described for Coleus seem to indicate
the such conditions are determined largely by intercellular relations.
Chemical studies, in general, show that differences in quality,
quantity, and distribution of pigments in flowers and leaves are cor-
related with changes in quality, amount, or distribution of any one
of such substances as chromogens (glucosides, phenols, tannic acid, etc.),
oxydases, enzymes, oxygen, etc.
That marked changes in color quite comparable to those I have de-
scribed for Coleus may result from slight chemical changes is well shown
by the chemical studies of various members of the genus Monarda.
The results of these studies are fully summarized by Wakeman (1911).
The isolation and determination of yellow and red pigments and a study
of their chemical relatives have given rise to the quinhydrone hypoth-
esis of plant pigmentation. It is considered that the plant oxidizes
thymol or carvacrol to a series of oxidation products of yellow, orange,
and red colors, but all closely related to each other. Of these, hydro-
thymoquinone, thymoquinone, and dihydroxythymoquinone have been
definitely isolated and determined. These in turn have the capacity of
adding phenols yielding highly colored phenoquinones and quinhy drones.
Wakeman (1911, p. Ill) states:
"Taking into consideration only those compounds that have been isolated
or whose presence has been indicated in the monardas thus far, the number of
possible pigments becomes truly bewildering."
Furthermore, some of the pigments are phenolic in character and
can combine with metallic constituents of the plants, giving rise to
different shades of the original pigment.
The highly colored red and purple pigments of the stems and leaves,
and the yellow and purphsh pigments of the flowers in Monarda fistulosa
are thus quite definitely identified as mixtures of quinhydrones which
BY THE SELECTION OF SOMATIC VARIATIONS. 63
are shown to be direct products of the plant. The important point is
that while the pigments are themselves of highly complex chemical
substances, the changes which produce marked differences in color are
very slight.
It is also significant that marked differences may exist between the
parts of a single flower. The simplest of color patterns in Antirrhinum
treated by Wheldale (1914, p. 110) illustrates this condition. Flowers
of the yellow-flowered variety possess a pale yellow pigment in the tube
of the corolla, a deep yellow pigment in the lips, and a still deeper
patch on the palate. In attempting to harmonize these varied con-
ditions with a Mendelian factorial analysis, the power to produce these
pigments in spatial relation is assigned to a factor Y. In the "ivory"
flower a pale pigment is found in the tube and Ups and a yellow pigment
only in the epidermis of the palate. Power to produce pale pigment
quite generally in the flower, and to inhibit the formation of deep yellow
every^^here but in the epidermis over the palate is assigned to a single
factor, /. The conditions of color distribution are in themselves color
indicators that different processes occur in different parts of the same
flower, giving different kinds of substances in different amounts and
with different distribution.
Studies of oxydase reactions in different tissues indicate that inter-
cellular relations are of much importance in determining the distri-
bution of pigments. By means of micro-chemical tests Keeble and
Armstrong (1912) obtained evidence that the distribution of oxydase in
various types of Primula is closel}^ correlated with the development of
anthocyanin. In general they find the oxydase most abundant in
epidermal and in vascular tissue. The extent of ox3'dase distribution
differs much in different varieties. They make the highly interesting
observation that action of chloroplasts seems to act as an inhibitor
of oxidase formation or of the production of chromogen. Varied types
of color pattern in flowers, and even anthoc^-anin development in
stems and leaves, is, according to these investigations, closely related
to distribution of oxydases and chromogens.
It is of further interest to note that patterns resulting from such
qualitative and quantitative reactions which depend in considerable
degree on flow of substances in a plant may be quite uniform not only
in the flowers of a plant, but among the various members of its seed
progeny. Such phenomena have led to the assumption that patterns
are represented as such in germ-cells by hereditar}- units. A^Tien, how-
ever, hybridization occurs between varieties having even the simplest
of patterns, the F2 generation more often than otherwise presents a most
remarkable range of types. Usually, this sort of diversity results in
crosses betw^een varieties, especially when the color patterns of flowers
and leaves are concerned and gives results that could not be predicted
with any degree of accuracy.
64 ESTABLISHMENT OF VARIETIES IN COLEUS
Riddle (1909), in a comprehensive survey of the chemical and phys-
iological facts regarding the origin and nature of melanin pigments,
points out (p. 323) that "a single chromogen acted upon by a single
enzyme usually produces several colors depending upon the degree of
oxidation involved." He gives conclusive evidence that "the power to
oxidize tyrosin compounds is not dependent primarily upon germinal
segregation, but rather upon active tissues, relations, and conditions,"
and that local conditions, especially in pathological cases, determine the
production of melanin. In general Riddle shows the inadequacy of a
strict interpretation of color inheritance in animals on the basis of unit
factors and gametic purity.
In the light of all the chemical studies on pigmentation, it seems
clear that qualitative reactions are concerned which involve the pro-
duction, flow, and assembling of substances through the relations and
interactions between cells. That these interactions should be so widely
readjusted in a hybrid progenj^ resulting in such varied expression of
color in quantity, quality, and distribution is suggestive that funda-
mental readjustments may occur more readily with characters that are
dependent on cellular interactions than with those that are strictly meti-
dentical. As already pointed out, the variations in Coleus propagated
vegetatively give numerous patterns differing widely in regard to quan-
tity and distribution of the pigments concerned and present the same
sort of phenomena of readjustment seen in seed progeny of hybrids.
In questions relating to the development of color patterns, the
Liesegang precipitation phenomena, especially as developed by
Gebhardt (1912) and Kiister (1913), seem to me most illuminating. By
the various phenomena associated with rhj^thinic precipitation and
crystallization of mineral solutions in gelatin plates, Gebhardt was able
to produce simple and multiple eye, line, and flaked patterns strikingly
similar to various markings in butterfl}- wings. By varjdng the sub-
stances used, modifjdng the amounts, the distribution, the degree of
concentration, and providing for interaction between areas of different
concentration, a wide range of markings were produced. Gebhardt
points out that such physical and chemical phenomena indicate that
the distribution of pigment even in the intricate markings of butterfly
wings may be due to an epigenetic regulation of the quality and quan-
tity of such substances as chromogen, oxydase, and oxygen and the
reciprocal influences of different centers of action. It is pointed out that
the cell boundaries and especially the position of veins may determine
the distribution of the substances involved and determine the relative
locations of centers of action as well as areas and centers of no action.
Kiister (1913) extended the study of Liesegang precipitation pheno-
mena to the effects produced in capillary tubes, obtaining various types
of banded precipitation patterns which, as he emphasizes, suggest that
similar chemical and physical processes may determine many tj-pes of
variegation in both monocots and dicots.
BY THE SELECTION OF SOMATIC VARIATIONS. 65
In Coleus the development of patterns is considered by the writer
to be due largely to cellular and tissue interactions influencing general
and metidentical qualities with results quite analogous to the Liesogang
phenomena. Changes involving red are on this basis rather simple
cases of readjustment influencing the total amount of pigment produced
and the distribution in centers or areas. The ability to produce the
different chemical substances concerned with the final development of
the red pigmentation is assumed to be a general property or potentiality
of all Coleus cells. The assembling of all the products necessary for
the final stages in its development, however, are determined by the
amounts produced and their flow to centers of activity and interaction.
The development of red, especially in the subepidermal tissues, indi-
cates that this is the case and suggests strongly that, as shown by
Overton (1899), changes in the amounts of red pigmentation may be
closely related to changes in the sugar-content of the sap. In Coleus,
however, it is clear that such changes arise quite spontaneously in the
cells and tissues.
In respect to the development of green and yellow in particular cells,
the processes seem to be antagonistic. Plastids are present in both
green and yellow^ cells, but in yellow cells they are fewer in number,
smaller in size, and somewhat distorted in shape. The green and
yellow cells are subepidermal, extending from upper to lower epidermis.
In a pure-green leaf all these cells remain green. In the most extreme
cases of yellow development nearly all the cells fail in the production of
chlorophyl. The different patterns result from variations in the
relative number of green and yellow cells and in the grouping of the
cells of like color. In some a green field is blotched with island-hke
areas of yellow cells, in others the central area of green is bordered
by an irregular band of yellow, and again the yellow may be situated in
the center with a green band at the border.
In a leaf with blotched or banded green-yellow patterns the inter-
mingled areas of green and yellow cells indicate quite clearly that both
types of cells are derived from the same cells in the growing-points.
In the development of leaves it can be observed that while the yellow
areas are in evidence when the young leaves unfold, the yellow seems
tinged with green, and that as the leaf grows the yellow becomes
more intense. Furthermore, the yellow bleaches until in old leaves
the yellow areas change to a pale yellow or white, while the green areas
are still bright. As a leaf dies the green areas become pale greenish
yellow. These observations indicate that many of the cells which
later become yellow are actually green at first.
The fluctuations that appear substantiate this view. A plant with
yellow-bordered leaves ma}^ produce, especially in winter, new leaves
entirely green, and thus possess for some time old leaves of green-yellow-
red blotched pattern and younger leaves of green-red blotched pattern.
66 ESTABLISHMENT OF VARIETIES IN COLEUS
Later the leaves formed may show increased amounts of yellow, until
by midsummer the plant is uniform for the green-yellow-red blotched
pattern. Figures 14 and 14a show winter and summer conditions quite
general for the leaves of the yellow-green pattern, which indicates that
cells may be either green or yellow. Furthermore, all degrees of varia-
tion can appear as bud variations affecting segments of a bud.
Such conditions indicate that all the cells are potentially green.
If this be accepted, a further point is raised regarding the source of
the influences leading to loss of green and to development of yellow in
its place. In respect to the final action in the cells, change from green
to yellow is itself apparently a local phenomenon in that local action of
plastids is affected.
The fluctuations and variations in the extent and position of the
green and yellow tissues, however, indicate that here, as in the develop-
ment of red, certain intercellular influences are operating. The con-
figurations of the yellow and green areas, as well as that of the red, are
in marked degree bounded by veins. For the green and yellow this
is most strikingly shown in figures 12 and 14. For the red similar
conditions are seen in any of the blotched patterns, as, for example,
figures 2 and 5. This also indicates that the flow of substances giving
different centers of distribution and concentration is the important
factor in the production of patterns.
In respect to the extent and degree of the variations, it has already
been pointed out that no plant in my cultures of Coleus has been
obtained without some red coloration in some part of the plant. In
regard to yellow, however, there were frequently fluctuations and bud
variations giving branches with no yellow. The loss of yellow appears
to be complete in a manner that suggests loss of hereditary qualities
through segregations, but even in constant selection of pure-green stock
for vegetative propagation about half of the offspring show return to
patterns containing yellow both by fluctuations and by bud variations,
with, also, cases of marked spontaneous appearance of yellow in a few
or in single leaves. To say that the power to produce yellow has been
latent is to say that the conditions causing its development can arise
in an apprently spontaneous manner.
AVhile the evidence indicates that the ability to produce green,
yellow, and red is a metidentical property of the cells, it is equally
clear that these metidentical properties do not exist as units. They are
subject to interaction between cells. They are more or less perma-
nently modified either by the intercellular relations or by spontaneous
intracellular changes. The records of pedigree show very clearly that
tendencies to give vegetative progeny of different degrees of constancy
and variation arise or exist in sister branches that are apparently
quite identical. Such tendencies detected by the pedigreed cultures
have already been mentioned. The production of branches which give
BY THE SELECTION OF SOMATIC VARIATIONS. 07
clones of plants with decreased amounts of red indicates a speoific
decrease in power to develop red and the results of pedigree culture
indicate that continued selection in this direction would give lines
with only slight amounts of red or perhaps that are entirely free of
all red.
The metidentical characters of green, red, or yellow are themselves
fluctuating not only in expression but in inheritance through cell
Hneage. They do not appear to be independent. I have never yet
obtained a plant by seed or by vegetative propagation that did not
possess some degree of red coloration, and as yet no strain has been
isolated that was pure for loss of yellow.
On the whole, vegetative propagation of any new type that arises
gives a progeny that exhibits a rather marked degree of constancy with
fluctuations and variations about a new mode. Selections for pure
green do not give a progeny of pure-green plants, but do give a greater
number of green plants than does any other pattern. Selections for
decreased red or for increased red likewise give clones with this
tendency prevailing. In other words, readjustments of the processes
concerned with total production of pigments and their distribution
tend strongly to occur in growing-points or to so affect them as to
secure a certain degree of permanency.
The assumptions of de Vries already noted in the introduction seem
to apply quite adequately to the behavior of the metidentical char-
acters as far as expression in individual cells is concerned, but does
not fully explain the phenomena of pattern changes as well as the con-
ception of a further influence of intercellular relations, modifications
of which may in time affect more or less permanently the expression of
metidentical qualities. x\ny Mendelian conception of pattern factors
that are units in heredity is quite inadequate, as is also such a con-
ception for even the metidentical characters.
This analysis of the nature of variegation and the significance of
bud variation in Coleus has a direct bearing on the nature of certain
other types of variegation.
One of the most clearly marked types of variegation is that of the
infectious chlorosis. The best known cases are those of tobacco
(Beijerinck, 1899; Woods, 1899) and Abutilon (Baur, 1904 and 1906;
Lindemuth 1897, 1899, 1901, 1905, and 1907). All investigations
agree that in these types the variegation is not transmitted to seed
progeny. The searching investigations of Beijerinck and Baur lead
to the conclusion that a living fluid or virus carries the contagion. In
the case of tobacco the infection is readily accomplished by various
agencies. In Abutiloti grafting is necessary, and by this means the
variegation has been transmitted to numerous species of Abutilon and
related genera. Similar types of infectious variegation exist in other
groups of plants, as Fraxinus, Jasminum, Liburnum, and Ligustrum.
68 ESTABLISHMENT OF VARIETIES IN COLEUS
In the infectious variegation of the type seen in Abutilon striatum
thompsonii the pattern is a mottled one, with irregular yellowish areas
mingled with the green. The amount of yellow varies considerably,
especially according to intensity of illumination. In Abutilon mega-
potamicum variegatum especially the distribution on a single plant is
very irregular (see Reid, 1914). Only a few blotches may appear on
a leaf and often entire leaves or all leaves on an entire branch may
be pure green. Baur (1906a) found green branches on Abutilon striatum
thompsonii and was able fully to establish that they were immune.
Immune branches arise on a plant as bud variations, but the leaves
differ from those with variegation only in having all cells immune, for
in the latter a part of the cells perhaps remain immune. It may be
noted that blotched variegation in this case either results from irregular
immunity or to irregular distribution of virus, and hence emphasizes
the intercellular relations concerned with distribution.
V^Tiatever the nature of the ''virus" may be, it is fully demonstrated
by Baur that it is a product of the diseased cells of the old leaves and
is transported to young leaves in which certain areas of cells succumb
to the influence while others do not. This immunity exhibited by
some cells, however, may suddenly extend to entire leaves or to all leaves
on a branch. As to the flow of the "virus," Baur (1906a) found that
it could pass through tissues of immune strains of Abutilon arboreum,
causing infection to non-immune parts beyond, but that such infection
was not produced if immune tissue of Lavatera arborea intervened.
It should be noted that this type of variegation can not, as far as we
know now, be distinguished by appearance from other types of mottled
variegation. Its infectious nature and the failure to transmit to seed
progeny are the characteristics of these cases. As noted above,
ShuU (1914) suggests that certain yellow-flecked types of variegation
giving very irregular transmission to seed progenj^ may be of such
infectious nature that it can be carried in some of the germ-cells.
It may well be that in many cases of variegation (especially of the
blotched t>"pes), the disturbing cause producing loss of chlorophyl
may be quite similar in nature to that of the \agorously infectious types.
In the latter it is quite clear that the production of a "virus" in varie-
gated leaves and its flow to 3'oung leaves does occur.
Differences in the extent of influence of such a "virus" may give in
some types an apparent inheritance through seed progeny. In the
infectious types we know nothing definite concerning the appearance
of variegation in the plant first showing it. It appears (Reid, 1914)
that many, if not all, of the abutilons with infectious variegation arose
through grafting with one original strain, Abutilon striatum thompsonii.
The presence of infectious variegation in such widely differing genera
as Nicotiana, Fraxinus, Abutilon, and Ligustrum indicates that the
condition should arise spontaneously. Frequent and almost continued
BY THE SELECTION OF SOMATIC VARIATIONS. 69
spontaneous development of a less vigorous virus may be very common,
and many cases of variegation, even of those that are apparently seed
constant, may be due to such a condition.
Lindemuth (1905) determined that the variegation in Coleus was
not infectious like that of Ahutilon. Just what types he used is not
clear from the data given. Conclusive evidence regarding this point
has not been obtained by the writer. Thus far grafting experiments
between green-yellow and pure green types have shown no cases of
development of yellow, and the writer has assumed that the variegation
is not at least vigorously infectious.
We may also note that spontaneous loss of ability to produce green
in a part of the cells of the growing-point may result in a chimeral
variegation such as Baur (1909) reports in white-margined types of
Pelargonium zonale. His anatomical studies showed that in growing-
points the white-colored tissues may lie over the green, forming histo-
genic layers and giving remarkable permanency of the pericUnal
chimera in vegetative propagation.
In maintaining this relation white cells give rise to white cells and
green to green, but mechanical readjustments in the growing-points
may give branches with quite different distribution of the two kinds
of cells. Branches may thus arise with sectorical distribution of green
and white, with only white or only green cells, or even with reversed
positions (Baur, 1909; Stout, 1913). These readjustments give no
new qualities to cells, nor do they appear to involve changes of either
kind to the character of the other. Yet the occurrence of numerous
types of variegation with this chimeral relationship indicates that such
spontaneous loss is not infrequent.
Certain types of bud variation in Coleus present features quite
similar to the readjustments that appear in Pelargonium, and raise the
question whether there is possibility of spatial readjustments of dis-
tinct tissue elements. The sudden and apparently complete loss of
epidermal red suggests that this layer may exist as peripheral in a more
or less chimeral relationship, but the development of red and non-red in
the adjacent cells of the epidermis of a single leaf indicates clearly that
these differences can arise within cells of the same immediate progeny.
If for any reason a part of the epidermal cells fail to develop red, the
red might be absent in the entire epidermis for the same reason.
The loss of yellow giving pure green might seem to be due to the
exclusion of yellow cells. Also, the sharp contrast between patterns
of green-yellow (figs. 2, 8, and 12) and pattern of yellow-green (figs. 6,
11, and 14) suggest the possibility of a spatial readjustment in the
growing-point of two distinct cell elements. But certain plants in
all these patterns have fluctuated, getting greener to a green-yellow
blotched pattern during winter and returning to the type pattern in the
next summer, and often exhibiting at one time in a single row of
70 ESTABLISHMENT OF VARIETIES IN COLEUS
leaves various gradations between the extremes. The extreme yellow
types show some green areas of tissue. The extreme green type, which
appears to have no j^ellow, gives numerous fluctuations, and cases of
well-marked spontaneous development of yellow. The distribution
of green and yellow does not in any pattern show anatomically a
chimeral distribution, as both colors are much intermingled in the
subepidermal tissue.
That a certain degree of chimeral relationship exists in certain
patterns of Coleus is evident. In the patterns with solid-red epidermis
the epidermal layer is rather specialized in respect to concentration
of red cell sap. These apparent chimeral relationships in Coleus are
due to intercellular development of patterns rather than to specific
and qualitative differences in cells as such.
Numerous cases of variegation are induced by environmental con-
ditions. Cramer (1907, chap, xi) summarizes cases of the influence
of parasites, of soil conditions, light, and temperature in producing
certain types of chlorosis and variegation. These are, we may say,
direct reactions to external conditions, which in most cases are quite
apparent. At first thought this class may seem quite distinct from
what are considered as true hereditary types, but the difference is
chiefly one of degree, for there are few types of cases of variegation
that do not fluctuate in response to certain environmental conditions.
The infectious types of variegation fluctuate verj^ much according to
degree of illumination and may entirely disappear from a plant if it
is kept in darkness for sufficient length of time. The types with green
and white as periclinal chimeras show, perhaps, least fluctuation in
regard to environmental influence.
In this respect what I have called fluctuations in Coleus are of inter-
est. Fluctuations in amount of red present in blotched types is con-
stantly occurring. One plant, at the present writing, has some branches
with leaves sparsely blotched, as in figure 5, others with the red blotches
strongly coalescing, as in figure 29, and still others with nearly a sohd
epidermal red. These fluctuations do not seem related to external
environmental conditions. In the amounts of green and j-ellow, the
various patterns possessing these two elements showed a strong ten-
dency to be more green in winter and more yellow in summer. The
degree of fluctuation was, however, not uniform for the different sub-
clones, for the several plants of a single generation, or even for all the
branches of a single plant. A few plants maintained a maximum of
yellow in their leaves throughout the winter. Still it is very clear that
many cells in the leaves are green in winter which would have been
yellow had the particular leaves developed during summer.
In certain plants variegation appears periodically. These types
are, perhaps, in a strict sense, to be classed with the preceding, but
differ at least in showing marked periodicity which may well be due
BY THE SELECTION OF SOMATKJ VARIATIONS. 71
in large measure to inner conditions. Cramer (1907, p. 128) sum-
marizes numerous cases of those showing marked differences in beha-
vior. Certain varieties of Quercus have pure-green foliage each spring,
but later produce variegated leaves. Uhnus scabra var. viminalis is
yellow during midsummer, but pure green at close of the summer.
Linaria biennis is pure green during the first year of growth, but varie-
gated in the second.
These phenomena illustrate again that cells of the same lineage may
fluctuate in development from a maximum of green to various degrees
of loss of chlorophyl, with often development of yellow coloration, and
that such phenomena may reappear with marked constancy in progeny.
Aside from these classes there is a wide range of types, including
many cases in which the variegation reappears more or less generally
and constantly in the individual plants and in the variety. The
variegation appears to be inherited, at least in a certain degree.
In the propagation of variegated plants, much general data has been
obtained regarding the degree of constancy both in vegetation and seed
propagation. Cramer (1907, p. 129) gives a summary of numerous
cases where the variegation disappears in certain methods of vegetative
propagation. In Cornus mascula variegata (T. M., Gard. Chron.,
32: 952), root-cuttings give pure-green individuals, but plants from
layers retain the variegation. Such cases, however, should be investi-
gated wdth special regard to the nature of the variegation. In many of
the cases noted, especially those in Pelargonium, the variegation may be
chimeral, and when root-cuttings are made the green cells have greater
power of regeneration. The inconstancy exhibited in vegetative propa-
gation, however, is no greater or more marked than that which develops
on a single individual. Exact evidence of the inheritance and con-
stancy of variegation through pedigreed vegetative progenies seems to
be lacking. \ATiile in some varieties the variegation appears to be
quite constant, in others it is widel}^ variable.
The Mendelian studies that have been made of the seed progeny of
variegated plants and of the bud variations which involve changes of
pattern show likewise a wide range of behavior. Hybridization studies
involving variegation, as has been pointed out in the introduction,
indicate clearly the wide range of variability, and w^hat, from the
Mendelian viewpoint of unit characters or unit factors, is most erratic
inheritance. These studies have contributed interesting and valuable
data on the sort of variations one may espect in hybridization studies
of this kind, but they indicate verj^ clearly that the assumed factors are
themselves fluctuating. In these studies the variabilit}- of plants used
as seed parents has not been determined by vegetative propagation.
This is not possible in all cases, but whenever it is possible the emphasis
should be placed on this line of investigation if one is to speak with
certainty regarding the nature of the inheritance.
72 ESTABLISHMENT OF VARIETIES IN COLEUS
If we turn our attention to the various phenomena associated with
the laciniate character of leaves and petals, we find the same degree of
irregularity and diversity of origin, expression, and inheritance that is
exhibited by variegations. Cramer (1907, chap. 21) devotes a chapter
to an excellent survey of the facts regarding the behavior of the char-
acter. It is interesting to note that Cramer observes that varieties
quite constant in seed progeny are likewise very constant in vegetative
propagation. There are numerous cases known of the spontaneous
development of a laciniate type from one with entire leaves, both as
seed mutations and as vegetative mutations; likewise of return to the
types with entire leaves. While some cases are quite constant, others
are widely fluctuating, even exhibiting a marked periodicity.
The behavior of this character in Coleus is most striking in its varia-
bility of expression and of its inheritance as a periodic variation through
vegetative propagation.
Fundamentally, the processes involved in the development of leaf-
shape are quite different from those involved in the production of
pigments such as green, yellow, and red. The shape of the leaf in
general depends on the rate, number, and regularity of cell divisions
in the different planes of growth. It would seem that a general and
quite uniform series of cell-divisions would give a leaf of more regular
outline, and that if the cell-divisions in the growing leaf occur irregu-
larly, giving, so to speak, points or lines of more rapid growth somewhat
analogous to apical growth, with a more or less multiple dichotomy,
then cut, lobed, or laciniate leaves would result. The shape of the leaf,
it would seem, is determined by intercellular relations concerned with
the manner of cell-divisions.
A comparison of the variability that develops in vegetative propa-
gation with that occurring in the seed progeny reveals some essential
differences between the inheritance of the characters involved. While
the range of variation is quite the same, there is a marked difference
in what we may call the intensity of variation. In vegetative propaga-
tion the degree of the intensity was low, with reference to the frequency
of the appearance of new color patterns or to the development of the
laciniate character. A large number of plants were grown by vege-
tative propagation. The bud variations were comparatively infrequent,
occurring something like once for every 10 plants grown, giving in the
course of 6 generations the different types of pattern described. In a
seed progeny, however, practically the entire range of variation which
appeared in vegetative propagation was seen in a single seed genera-
tion comprising 50 plants. The processes concerned with reduction and
fertilization increased the intensity of variation and brought out in a
single progeny of no great number the full extent of variations.
Color patterns, which are intercellular characters and in a sense
vegetative types, are inherited through vegetative propagation in
marked degree, while in seed propagation there is no evidence that they
BY THE SELECTION OF SOMATIC VARIATIONS. 73
are inherited as such. The intercellular relations involving amounts
and distribution of pigments are widely and suddenly disorganized
and readjusted during the processes concerned with seed formation.
On the other hand, the metidentical characters green, yellow, and red
are quite uniformly transmitted in both sorts of cell-divisions.'
The behavior of the laciniate character is significant in this con-
nection. After this character made its appearance in a plant it was a
constant feature in the development of all plants (but one) derived
by vegetative propagation. The seed progeny, however, exhibited
wide differences, ranging from deeply laciniate through all degrees or
grades to the entire type of leaf. Furthermore, the laciniate leaf
appeared in all seed progenies thus far grown, even when derived from
plants of a Hne in which the character had never appeared. The proc-
esses of reduction and self-fertihzation in a Coleus plant seem to bring
out latent possibilities for various developments of leaf-shapes.
It remains to be seen if any of the types appearing in seed progeny
are more constant in vegetative or in seed propagation than are the
similar types that develop by bud variation. Already several bud
variations have appeared in plants of seed progeny, indicating vegeta-
tive changes in the processes concerned with pigment and pattern
formation. The colors involved in the variegation of Coleus represent
every type of coloration (green, yellow, white, and red or blue) con-
cerned with the variegation and coloration of plants. There is no
evidence that the essential nature of these characters in Coleus differs
from that of the characters concerned with variegation and pigmenta-
tion in corn (Emerson, 1914) , in Mirabilis (Correns, 1909) , in Melandrium
(Shull, 1914), in Pelargonium (Baur, 1909), in Antirrhinum (Baur, 1910).
The explanations here given regarding the spontaneous variability
of the characters concerned in the development of pigments and of the
changes in intercellular and intertissue relations influencing develop-
ment of color patterns in Coleus apply equally well to such cases as
those just noted. The evidence indicates that the characters in ques-
tion, and most especially the pattern characters, are not represented by
units or factors, unless these assumed factors are to be considered in a
general sense as temporary conditions descriptive of tj'pes of develop-
ment and not as particular localized units of germ-plasm, which is the
conception that gave the Mendelian interpretation its definiteness and
simplicity.
The knowledge of the nature and the heredity of color character-
istics will be advanced more by studies of the natural variability of
the characters involved and by chemical and phj'sical investigations of
the processes concerned in the formation and distribution of such sub-
stances as melanin, flavone, and phenol compounds rather than by
further elaboration of complicated formulae involving multiple factors
that attempt to explain fluctuations, inherited variations, and cases
of increased variability that appear in hybrid seed progenies.
74 ESTABLISHMENT OF VARIETIES IN COLEUS
SUMMARY.
(1) A single variety of Coleus propagated vegetatively by cuttings
in two main clones has shown (a) gradual fluctuations and (6) sudden
mutations, giving a total of 16 distinct and characteristically different
color patterns.
(2) (a) A total of 15 patterns (see diagram 2) arose by sudden muta-
tion affecting a part of a leaf or a branch, or a series of associated leaves
or branches; (b) 6 of these 15 patterns (figs. 2, 4, 5, 10, 13, and 13a)
also appeared among the fluctuating variations; (c) one type of color
pattern (fig. 15) has thus far appeared only as a fluctuating variation.
(3) (a) Six (diagram 2, and figs. 1, 4, 5, 6, 8, and 12) of the 15 color
patterns arose directly from the parent type bj^ sudden bud variation.
One of these 6 (fig. 4) also appeared as a fluctuating variation. (6) Five
of these 6 tj^pes (figs. 4, 5, 6, 8, and 12) propagated by cuttings showed
further fluctuations and bud variations, giving (a) the parent type
(fig. 2), (6) 4 of the 6 types already directly derived (figs. 1, 2, 5, and 12),
and (c) 8 new types (figs. 9, 10, 11, 13, 14, and 16, also 8a, 13a, not
illustrated).
(4) The variations in the development of color patterns mentioned
above involve (a) increase and decrease of green and yellow, (b) increase
and decrease of red pigmentation, (c) reversals of the relative positions
of the green and yellow by which a type with green center and yellow
border (fig. 2) gave one with yellow center and green border (fig, 6),
and (d) changes in the distribution of the red pigmentation especially
giving concentration in the epidermis of the upper surface of the leaves.
(5) Progeny of 11 types of color pattern have been grown through
from 2 to 6 generations, as follows:
Type of figure 1 2 4 5 6 8 9 10 12 13 14
Number of generations 2 6 6 6 6 5 2 2 2 2 2
Total number of plants 11 337 198 90 41 54 8 7 7 7 4
Some of these types have shown themselves more constant than the
parent type (fig. 2), others were less constant. All varied about a new
mode and all would be considered good horticultural races.
(6) The relative constancy of color-pattern types derived by the
accumulation of fluctuating variations was tested in two cases: Type
of figure 2 (see clone 14 of table 2), 3 generations, total, 45 plants;
type of figure 4 (see clone 13 of table 3) 3 generations, total, 79 plants.
In both cases the constancy of the progeny showed no essential differ-
ence from that of the same types obtained by sudden bud variations.
(7) Variations in leaf form, even more striking than changes in color
patterns, from entire to deeply laciniate-leaved forms, appeared in 13
instances as fluctuations affecting an entire plant and in one case
(during the winter of 1914-14) as a bud variation. The striking
BY THE SELECTION OF SOMATIC VARIATIONS. 75
feature of this variation is the marked tendency to the production of
laciniate leaves during winter or after a period of particular drought
during summer.
(8) The behavior of the laciniate character was observed in three
subclones (see table 4) of 4, 2, and 2 generations, totaling 68 plants.
In 67 of these plants the laciniate type of leaf appeared as a definite
character, with a marked tendency to a periodic summer and winter
fluctuation,
(9) Plants with the laciniate leaf character also showed wide fluctu-
ations in regard to the degree of green or yellow coloration. When
grown for a period of a year from cuttings made in autumn, the leaves
were as a rule entire and slightly yellov/ in autumn, deeply laciniate
and pure green in winter, and entire and very yellow during the
following summer.
(10) In sexual reproduction all the principal types of variegation and
leaf-shape appear at once in an Fi generation, with also numerous
types that were intermediate and fluctuating. The extremes of vari-
ation are no greater than those obtained in vegetative propagation,
although some new types of entire leaves were thus obtained.
(11) Between any two types numerous intermediates arose, showing
that we have here no evidence of the somatic segregation of invariable
pattern factors.
(12) In the bud variations, decrease of red occurred with about twice
the frequency as did increase of red; likewise decrease of yellow
occurred about twice as often as the increase of yellow, indicating a
definite tendency for variations in the direction of the increase of green
and the decrease of red. These facts are doubtless due to fundamental
relations between the chemical compounds involved.
(13) The types of color changes involving (a) green and yellow and
(6) red and non-red occurred entirely independently of each other.
(14) The types produced by bud variations are the equivalents of
the "Kleinarten" or the ''biotypes" commonly occurring in cultivated
species propagated by seed.
(15) Selection within clones is effective in securing progenies of new
types with as high degrees of constancy as is possessed by ordinary
cultivated races.
(16) The results indicate that slight variations arising either as
sudden mutations or as gradual fluctuations can perpetuate themselves.
(17) The green, yellow, red, and non-red colorations in Coleus can
best be characterized as metidentical characters; that is, they are the
same in the cells as in the tissue and their appearance is possible in
the development of any cell.
(18) The distribution of the colors giving pattern characters are
properties of groups of cells and tissues as such. Pattern characters
are probably due entirely to tissue and cellular interactions.
76 ESTABLISHMENT OF VARIETIES IN COLEUS
(19) The explanation suggested by the production of patterns in
colloids by the Liesegang precipitation phenomena, especially as applied
by Gebhardt to the markings of butterfly wings and by Kiister to the
development of many types of variegation in plants, seems to apply
to the production of color patterns in Coleus. On this view color changes
may be considered as due to the formation of different diffusion centers
for the development and concentration of pigments.
(20) Bud variations in Coleus are (a) common; (6) give numerous
different types which may be vegetatively quite constant from the first
or can be made so by selection; (c) show development of certain
types more commonly than others; (d) produce reversions to parental
t3qDes; (e) give development of different degrees of variability among
sister clones; (/) exhibit spontaneous changes in the fundamental color
characters (metidentical) and in the cellular and tissue processes result-
ing in color patterns.
(21) The results show that in Coleus asexual and sexual reproduction
are not fundamentally different in respect to the extent and range of
variation.
A. B. Stout,
Director of the Laboratories.
New York Botanical Garden,
New York City, February 10, 1915.
BY THE SELECTION OF SOMATIC VARIATIONS. 77
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BY THE SELECTION OF SOMATIC VARIATIONS. 79
EXPLANATION OF FIGURES IN PLATES 1. 2. 3. AND 4.
The paintings here reproduced were made by Misa Mary Eaton, artist of the New York
Botanical Garden. All figures are reduced to about three-fourths natural size.
Plate 1.
Fig. 1. A typical leaf of the pattern yellow-red blotched taken from plant 1171013 on
February 21, 1914.
2. The pattern green-yellow-red blotched. Taken from plant 121 on April 1.1, 1912.
The yellow border is somewhat irregular and is not so fully developed as in
summer.
4. Leaf classed as green-yellow spotted-red blotched. Painted on January .30, 191.3.
Shows rather few scattered yellow spots and a few rather large epidermal
blotches.
5. Pattern green-red blotched from plant 131, taken April 1.5, 1912. Shows the com-
plete loss of yellow as it occurred in the first bud variation observed and as it
frequently appears in plants having yellow.
5a. Young leaf of the pattern green-red blotched.
6. The pattern yellow-green-red blotched. Taken from plant 1171 on November 2,
1912. Shows the type which appeared on a plant with the pattern of figure 2
by a reversal of the relative positions of the green and yellow.
7. A good example of the laciniate type of leaf. Taken on January 31, 1912, from
a plant quite identical to plant 123153 shown in plate 4.
8. The green-yellow-solid red type. Differs from figure 2 in having a solid red instead
of a red blotched epidermis. Taken on January 4, 1913, from plant 32 and
shows the typical development of the yellow, which was very constant and
uniform on the plant diu-ing the winter.
9. A typical leaf of the green-solid red type. This differs from figure 8 only in having
no yellow in the subepidermal tissues and from figure .5 in possessing a solid
red epidermis.
9a. A young leaf of the pattern green-solid red. The red completely covers the leaf
and is of the same intensity as in a mature leaf.
10. This figure shows the upper surface of a leaf of the i^attern green-yellow-solid red
upper center. Figure lOa is of the under surface of the same leaf. The dis-
tribution of the green and yellow is as in figure 2. The red pigmentation is
almost entirely massed in the epidermis of the upper surface. There are a
few small areas of red in the lower epidermis. Painted on December 9, 1913.
1 1 . The type yelloiv-green-solid red painted on October 26, 1914. This tj^pe developed
from type of figure 8 by a reversal of the relati\'o positions of the green and
yellow, a change which also gave figure 6 from figure 2.
Plate 2.
Fig. 12. Type green-yellow, painted May 17, 1914. The i)attein is slightly irregular in
this leaf, with a yellow segment extending to the midrib. This pattern was
derived from type of figure 2 by a loss of epidermal red. There are a few
streaks of red in the vascular tissues.
13. The green pattern showing some epidermal red in the vascular tissues and a pale
and diffuse shade of yellow. Painted on June 5, 191 4.
13, a, b, c, d. Four successive leaves of a plant of the same line of descent as the plant
of figme 13. Painted on February 2, 191.5. The series shows an increase of
red in the subepidermal tissues as the leaf matures, with tlie maxinunn develop-
ment in figure d. Comparison with figure 13 shows the increase of red in a
hne of descent by gradual fluctuation. The distribution of red indicates a
relation involving vascular tissues.
14. Pattern yellow-green. This was derived from type G by a loss of epidermal red.
The streaks of red are subepidermal. Painted on February 11, 1914, showing
the decreased development of yellow frequent in this type during winter.
14a. The typical summer condition of the type yellow-green. Painted on May 29,
1914. The yellow areas are bounded in marked degree by veins and there is a
much less development of green cells in the central area.
80 ESTABLISHMENT OF VARIETIES IN COLEUS.
Plate 2 — Continued.
Fig. 15. A leaf with no yellow and with almost no red on the under surface. A type quite
like that described for the original Coleus bluniei. This type appeared thus
far only as a fluctuating variation as a winter condition of a few plants of the
type of figure 10. At the time this was painted, January 24, 1914, the oldest
leaves of the plant were typical for type 10 and the younger leaves were uni-
form for the coloration here shown. The under surface of this leaf had only
a few red blotches similar to those of figure lOo.
17. A leaf with irregular pattern, developed from type of figure 2.
18. One of the leaves of the plant shown in figure 21, showing the absence of yellow in
half of a leaf.
19. Leaf painted on January 18, 1914, showing rather marked increase of yeUow in the
type of figure 2 during winter. Painting was made after the yellow areas had
begun to turn to white.
20. Painted on July 5, 1913. Typical of the most extreme fluctuation of type figure 2
in regard to increase of yellow.
Plate 3.
Fig. 21. Young plant 12514, grown from a bud showing sectorial loss of yellow, by bud
variation in half of the bud. Photographed and painted on November 9, 1912.
22. One of the leaves in a branch showing sectorial loss of green in the pattern of
figure 6. In this leaf the loss appeared in one side of the leaf. Painted on
December 4, 1913.
23. Leaf classed as green-yellow spotted-red blotched. The red epidermal blotches are
large and much coalesced, which is a frequent variation from the condition of
figure 5.
24 and 24c. Lower and upper surfaces of the same leaf from a young plant grown
from a bud variation which involved a sectorial loss of red, giving solid red
upper center from solid red on both surfaces. In this leaf these two types are
sharply limited to one half of the leaf. The young plant exhibited loss of
yellow by fluctuating variation. The leaf painted was an upper leaf showing
no yellow, but at the same time the older basal leaves possessed much yellow,
as in type of figure 10.
25. Leaf painted on March 25, 1913, showing sudden bud variation affecting only the
half of a leaf. The plant possessed leaves of the type figure 2 shown in the
left side. A reversal of the relative position of the green and the yellow gave
the pattern of figure 6 in the right side of the leaf, as shown.
26. Taken on March 28, 1913, showing the fluctuational decrease of yellow, giving a
poorly defined yellow border. During the f ollowmg summer the plant returned
to the typical form of type 2. This leaf is quite typical of the increase of green
during winter seen in numerous plants of type 2, as mentioned on page 22 of
the text.
27. Painted on July 27, 1913, showing an in-egular pattern with yellow at the border
about the apex of the leaf and illustrating a fluctuation produced from both
types 2 and 4. Leaf also shows irregular distribution of epidermal red.
28. Leaf classed as green-red blotched, but with few large blotches somewhat run
together.
29. Leaf also classed as green-red blotched, but with fine blotches much coalesced.
Plate 4.
Three plants photographed on January 16, 1914, all descended from branch 2 of plant 1.
Plant 125111 has leaves uniformly entire and of the pattern green-yellow-red blotched (fig. 2).
Tips pinched off to prevent early flowering in greenhouse.
Plant 1251412 has leaves uniformly entire and of the pattern green-red blotched.
Plant 123153. Old leaves entire, youngest leaves deeply laciniate. Typical condition
during early winter for plants showing the fluctuation in leaf shape.
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