GENETICS OF SESAME

D. G. LANGHAM

V. Some Morphological Differences of the Sesame Flower (S. mdicum L.)

N a previous paper8 in which color
I differences in sesame flowers were
discussed it was pointed out that
2. Internal pubescence vs. its ab-
sence.—Near the foveola of some varie-
ties is found a small brush of plant hairs;

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many structural differences also existed. these are absent in other varieties. The
These are too numerous to describe in number of hairs varies from one to ap-
detail, but their principal characteristics proximately 50 per flower, but the num-
will be the subject of the present paper. ber for any given variety is relatively
Material for this study consists of 35 constant. The ease of classification of
varieties of sesame with numerous hy- this character in segregating populations
brids between many of them. The off- made possible the determination of seven
spring of crosses of varieties from wide- different qualitative factors for the pres-
ly separated geographical regions have ence or absence of hair. Second genera-
expressed many characters not.found in tion ratios of 3:1, 9:7, 13:3, 15:1, 1:3,
either parental phenotype. The expres- 7:9 and 1:15 have been confirmed by
sion of some other characters was F 3 and F 4 populations.
changed by transferring the genes re- 3. Tubular flower vs. split flower.—
sponsible for them to other genotypes. In contrast to the usual tubular flower,
The study of this wealth of material has a type with five petals completely sepa-
been extremely useful in preparing a rated has been isolated. It is structural-
linkage map for sesame and also in ex- ly sterile because the absence of the tube
plaining certain cases of sterility and permits the style to curve upwards there-
semi-sterility among varieties. by separating the stigma from the an-
Description of Characters thers. Application of pollen, however,
results in seed set. In the F2 progenies
The extreme variation in the sesame studied, recessive duplicate genes have
flower revealed by an intensive «tudy of been found responsible for "star" flower.
different varieties makes the selection of Other types of split flower occur in
a typical flower difficult. The most fre-
which the tube may be opened by divi-
quently appearing type is shown in Fig-'
ure \AA and is described in Bailey's sion of the two dorsal petals, or along
Cyclopedia1. one side, or near the base. All of these
types have been isolated in pure lines.
Some of the variations of this typical Segregations of normal to split flower
flower as seen in our breeding plots are recorded in different F 2 generations are
described below: 3:1, 9:7, 1:3.
1. Foveola vs. absence of foveola.— 4. Open flower vs. closed flower.—
The typical sesarne flower has a small Ordinarily about seven o'clock in the
V-shaped pit in the petal, which forms morning, the lip of the lower petal of
the base of the floral tube. Neither the the flower bends downwards, leaving an
origin nor the purpose of this small in- opening in the floral tube. Some lines
dentation is known at this time, but it have been isolated in which the tube re-
is useful in a genetical study of the flow- mains closed. There are three different
er because of its relation to color distri- types of closed flowers:
bution and to certain morphological
A. Lip normal, but remains closed.
characters. A few plants without this
B. One half of flower tube, including lip,
foveola occurred and have been selec- remains green instead of becoming violet
tively bred to give pure lines. in color.
347
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FLOWER MORPHOLOGY
Figure 14
A—"Normal" sesame flowers on day of opening. The curvature in the center of the vertical
portion of the flower tube is the U-shaped foveolar indentation. B—Portion of flower with the
dorsal part of corolla removed, showing tuft of flat hair near foveola as compared with ab-
sence of this hair. C—"Star" flower, with petals separated. D—Extra enations on exterior of
the corolla, a characteristic associated with non-shattering seed capsules. E—Flower with fused
stamens, as compared with normal flower (Z7), and flower with extra tissue on inside of corolla
CO.
 Langham: Genetics of Sesame
C. Tips of petals become necrotic the day
the flower should normally open.
A and C are recessive and B are domi-
nant in Fi hybrids with normal flowered
types. The F 2 generations have not yet
been studied.
5. Extra citations on the exterior of
the corolla tube vs. wild type.—Flowers
with these extra growths have been iso-
lated three times and in each case the
plant differed from the others in that the
seed capsules were indehiscent in con-
trast to ordinary sesame. This valuable
mutant form behaves as a simple reces-
sive.6
6. Extra tissues on the inside of the
sesame flower vs. wild type.—Ordinarily
the five petals are smoothly joined to
form the corolla tube; the borders of
adjacent petals unite edge to edge. A
new type of flower in which adjacent
petals, particularly the lower ones, dou-
ble inward and unite with extra tissues
on the inside of the tube, has been iso-
lated in a pure line.
7. Fused filaments vs. separate fila-
ments.—Usually the filaments are com-
pletely separated one from another, but
new lines have been obtained by selec-
tion among hybrids in which several or
all of the filaments are fused. This type
is recessive to the normal.
8. Flattened vs. curled lip.—A type
of flower in which the lip curls down-
wards instead of remaining straight, oc-
curred in a variety from Brazil. It is
recessive in Fi generations and difficult
to classify in F2.
9. "Rough" foveola vs "smooth".—
By looking in profile at the foveola of
some varieties, the elongated cells in the
pit of the foveola have a brush-like ap-
pearance, while in others the surface is
smooth. In crosses between the two
types smooth is dominant and segre-
gates as a simple Mendelian factor.
10. Defective foveola vs. normal
foveola.—In some segregating popula-
tions individual flowers occurred in
which the foveola does not form a com-
plete U, but rather a portion of it, or a
split U. Seeds from these plants give
349
progeny with similar type flowers. In
crosses between this and ordinary flow-
ers the complete U dominates in Fi gen-
eration and classification is difficult in
the second generation, although the de-
fective types occur in approximately 25
per cent of the progeny.
11. Single vs. double lip.—Selections
in which the majority of the flowers
have an extra petal in the base, giving
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two lips and usually two foveoli, have
been made in segregating populations of
certain hybrids.
12. Single vs. multiple flowers.—Se-
lections have been made in which dou-
ble flowers occur frequently. The flow-
ers may be nearly separated, or fused
at different angles. Triple and other
multiple forms occur in these same
lines.
13. Large, normal, and small flow-
ers. — The average flower is approxi-
mately two and one-half centimeters
long and one centimeter in diameter.
But flowers three and one-half centi-
meters and others one centimeter long
with a corresponding variation in diam-
eter have been isolated. The size of the.
flower segregates as a multiple factor
character in hybrids.
14. Thick, normal, and thin petals.—
The flowers of some varieties are so thin-
walled that they become soaked by a
heavy dew, while others are so thick
they are quite firm to the touch. The
most common flower type is between
these two extremes.
15. Wide, normal, and narrow fove-
ola.—In the study of color distribution
in the sesame flower, a wide foveola is
desired because the limits of the differ-
ent pattern factors are more easily deter-
mined than in narrow types. Individual
plants in which flowers with a broad
foveola were obtained in a variety from
China. The narrow type occurs in a va-
riety from Nicaragua.
16. Reduced lip of the petal vs. nor-
mal lips.—In a variety from Brazil cer-
tain types were isolated in which the tips
of petals were shortened and the lip
hardly existed. In this type natural
350 The Journal of Heredity
crosses are reduced because the bees can-
not find a resting place to enter the flow-
er. Reduced petal segregates as a simple
recessive in F2 generation.
Discussion
The morphological differences in the
sesame flower described in this article
occurred in breeding plots in which
many varieties, hybrids and selections
were under study for commercial use.
The importance of flower type in the
percentage of natural cross-pollination,
dehiscence of seed pod, relative fertility
and other factors demonstrate the need
VI. Some Genetic Variations in Plant Color in Sesame
T
H I S is a preliminary report of a
study on inheritance of color dif-
ferences in the sesame plant. Some
of these plant characters are related to
flower color and to seed color. The com-
plete relationship between foliage, flow-
er, and seed will be discussed in a fu-
ture article.
Seedling Colors
1. Albino seedlings. — For the past
six years a few albino seedlings have
been observed in commercial plantings,
but it was impossible to isolate this char-
acter without testing thousands of plants
in an effort to find the heterozygous
form. This year, however, one line from
an individual plant selection segregated
green and white seedlings in the ratio of
3:1, and a heterozygous strain is now
available.
2. Virescent seedlings.—In spite of
the fact that sesame is approximately
96 per cent self pollinated2'8, virescent
seedlings are found frequently. Those
observed in our breeding plots are sim-
ple recessives and three have been dem-
onstrated to be genetically distinct.
3. Pale yellow. — There is extreme
variation in the degree of color in sesame
plants, ranging from pale yellow to dark
green. One of the yellow types segre-
gates as a simple recessive but the oth-
ers are difficult to classify in segregating
populations due to intermediate grades.
of a detailed study of genetics of flower
type in order to facilitate plant breeding
with this crop. The results presented
were obtained from a study of approxi-
mately fifteen thousand segregating pop-
ulations.
Summary
1. Brief description of some morph-
ological differences occurring in the
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sesame flower are given.
2. The relation of the differences to
the sesame breeding program are dis-
cussed.
4. Purple, green, and yellow leaf tips.
—When the tips of the first pair of true
leaves are barely visible between the
cotyledons, they may be purple, green,
or yellow. Both the purple and the yel-
low change within a day or so to green
and the classification must be made at
the proper time. In all cases studied to
date, purple dominates green or yellow.
Two types of purple have been ob-
served : one which is independent of
whether the plant germinates in the sun
or in the shade and another which germi-
nates green in the shade and purple in
the sunlight. These are complementary
dominant genes and one is linked with a
virescent. Classification is simple.
Among many F2 progenies from nu-
merous crosses, green leaf tip dominates
yellow in some cases, and yellow is domi-
nant in others. Two complementary
dominants have been isolated for green
leaf tip (yellow leaf tip being recessive
in these cases). Yellow leaf tip segre-
gates as a simple dominant in certain
other crosses.
5. Purple stem.—The color of the
seedling stalk immediately below the
cotyledons may be purple or green. Pur-
ple stem is dominant to green and segre-
gates sharply in the F2 generation. Seed-
lings with the combination of purple leaf
tip and purple stems, green leaf tips and
green stems, purple tip and green stems,
have been found, but the combination of
 Langham: Genetics of Sesame 351

Cp X Gp

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op

Op Gp Op
COLORED LEAF TIPS
Figur. 13

The tips of the first pair of true leaves may be purple, green, or yellow as they emerge
between the cotyledons. Later the purple and yellow varieties turn green, so that the character
can only be recognized during a brief period.

green tips and purple stems has not yet This type in which the color is localized
been recorded. in dots, occurred in a variety from Bra-
zil and in our Selection No. 5. It is re-
Plant Colors cessive to green stem and also to purple
6. Purple stems and leaf petioles.— and bronze.
Two types of purple have been encoun- 9. Green stem and leaf petiole.—As
tered : can be seen from the above discussion
a. Develops in the shade or in the sunlight. green acts as a recessive to purple and
Determined by a simple dominant gene. bronze, and as a dominant to dotted
b. Develops only in the sunlight Dominant
complementary genes differentiate this stem. The combination of the flower
purple from green. color known as flake and green stem has
All of these genes for purple are es- not yet been found; but the flower color
sential for the expression of the flower smear occurs in combination with green
color known as flake.8 stem and bronze stem, but not in purple
7. Bronze stem and leaf petioles.— or dotted.
This color is usually faint and is visible 10. Mottled leaf. — This character
only in the young leaves and stems of was described in a previous paper4 and
the nearly mature plants. It is deter- consists of yellow blotches in the green
mined by a simple dominant. leaves of nearly mature plants. It is a
8. Dotted stem and leaf petioles.— simple recessive.
352 The Journal of Heredity

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GREEN AND YELLOW COTYLEDONS
Figure 16

Green and virescent cotyledons represent an allelic pair, with green dominant.' In F , a
3:1 segregation of green and yellow is observed.

Discussion care is exercised in classification, all

Most of the seedling and plant colors
described in this paper can be easily
classified in segregating populations, if
care is taken to make the observation at
the proper stage of growth and under
favorable conditions of light. Purple
leaf tips, for example, are visible just as
they emerge between the two cotyledons,
but a day or so later they cannot be dis-
tinguished from green. Bronze stems
may be confused with green stems un-
less classification is made in the later
development of the plant. If proper
these characters are useful in a genetical
study of sesame. They are particularly
valuable in reference to flower and seed
color. A complete discussion of the re-
lations of the three will be made at a
future date.
Summary
1. Brief descriptions of seven seed-
ling colors and five plant colors are given.
2. The mode of inheritance of most
of these is discussed.
3. The importance of the time factor
in making classification is emphasized.

Literature Cited
1. BAILEY. L. H. The Standard Cyclopedia -Science 103:2670. p. 280.
of Horticulture. Vol. I l l , pp. 3157. 1946.
2. LANCHAM, D. G. Circular Xo. 4, Insti- 6. (abstract). Genetics.
tuto Experimental de Agricultura y Zoolecnia 1946.
(Depto. de Genetica), Venezuela. 1943. 7. -Circular No. 18, Minis-
3. Jour, of Hcrei. 35:254- terio de Agricultura y Cria (Depto. de Ge-
256. 1944. netica), Venezuela. 1947.
4. Jour. of Hcrcd. 37:149- 8. Jour, of Hered. 38:221-
152. 1946. 224. 1947.
378 The Journal of Heredity
None of these investigators recorded
nausea caused by P.T.C. in the amounts
used for testing, although some of their
subjects must have had far greater doses
than the two concerned in this note.
(From the unused portion returned, it
was calculated that those tested in this
family each had used about one-third of
a square inch of the test paper. This
contained whatever amount of P.T.C.
It is worth noting that neither of these
persons became sick at the time of chew-
ing the paper, but that both did so while
asleep two or three hours later. It is
doubtful if many adult persons tested
with P.T.C. have been asleep within a
short time thereafter, as most such tests
are made during working hours. Could
sensitivity be heightened merely by in-
activity, by sleeping, or by recumbent
posture? Although this last suggestion

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was absorbed during immersion in a 0.5
per cent solution in acetone). Two per-
sons tested by Blakeslee and Fox did
become nauseated, but both had taken
by mistake many times the proper dos-
age of the actual crystals. Dr. L. H .
Snyder (private communication) writes
that he has known people to complain
that P.T.C. almost nauseated them, but
never to the point of actually vomiting.
It is easy to say that nausea is not
unusual and that the occurrence of these
two cases in one family is merely a coin-
cidence, but it would be difficult to con-
vince the family concerned with that
argument. Obviously, the experiment
should be repeated, but, while it is all
very well for Haldane to drink dilute
hydrochloric acid, or for the writer to
gorge himself on chicken said to be load-
ed with estrogenic hormones, there are
limits to the experimentation that one
may ask of a six-year-old child and her
grandfather.
seems far-fetched, it may be recalled that
one genetic character in man is mani-
fested more readily in patients flat on
their backs than otherwise (asthma)
and that another is suppressed simply
by lying down (urobilinuria).
As far as they go, these two cases in
one family only suggest that there may
be an extreme sensitivity to phenylthio-
carbamide and that it may be genetic.
They are noted here merely so that other
investigators will be encouraged to re-
port similar instances if they occur.
Literature Gted
1. BLAKESLEE, A. F. and A. L. Fox. /.
Hered. 23:96-107. 1932.
2. Editor. / . Hered. 23:107-110. 1932.
3. FALCONEB, D. S. Ann. Eugen. 13:211-
222. 1947.
4. HARTMAN, G. Ann. Eugen. 9:123-135.
1939.
5. SALMON, T. N. and A. F. BLAKESLEE.
Proc. Nat. Acad. U.S.A. 21:78-83. 1935.
6. SNYDER, L. H. Ohio J. Sci. 32:436-
440. 1932.

Postscript on the Diabetes-Non-Tasting Correlation

O E V E R A L interesting questions were raised by the correlation between diabetes
O melitus and inability to taste PTC, as reported by Terry and Segall in the
May 1947 issue of this JOURNAL. The authors suggested the possibility that non-
tasting diabetics may be genotypic tasters. Can it be that non-tasting in diabetes
is non-specific for PTC ? Have any tests been made to determine whether a lessen-
ing of taste acuity in general accompanies diabetes ? Has the possibility been ex-
plored that there may be a correlation between non-tasting and insulin ? The
following reply was received from the senior author.
TO THE EDITOR: saw, viz., 46 per cent of non-tasters
The data from which our paper was among the diabetics. All I have got, for
written contain no answers, I am sure, sure, from the present much larger num-
to these questions. I am no wiser as to ber of observations is assurance that the
those particulars than I was several undertaking was worth while.
years ago when, after testing 36 persons, I have not inquired how far out of
diabetic and non-diabetic, I saw what I bounds the following ideas may be with
 Diabetes—PTC Postscript
respect to genotype and phenotype; but
I have thought that an experiment with
alloxan and rabbits, white rats, or dogs,
might turn up something new. White
rats would be best because of rapid re-
production (and because they were
available to me in the Department of
Anatomy), dogs best for a better reason,
viz., their known susceptibility to dia-
betes.
I tested a few dogs (and even a few
birds!) and found that, with one excep-
tion, the dogs showed a strong dislike
for PTC. The one exception was a very
fat, very old, Cocker Spaniel bitch who
is much of the time under a veterin-
arian's care. She does not object to the
taste of any medicine he gives her. She
may be diabetic for all I know.
Blakeslee et al (at Smith) made a
study on threshold recognition of the
taste of P T C ; but, of course, without
any thought of diabetes.
Alloxan diabetes, they say, may be
controlled indefinitely by insulin. If so,
experiments could be devised.
There are exogenous factors in the
incidence of human diabetes. The Met-
ropolitan Life Insurance Company men-
tions some of these in their pamphlet on
diabetes and undertakes to give advice.
In a 1945 number of the Nezv England
Medical Journal, (233[13] :376-78),
Dr. H. C. Miller presents some evi-
dence for a "prediabetic state" operative,
in his cases, well before the appearance
of the disease but not discovered un-
til long thereafter. Perhaps PTC might
help to identify such "prediabetics." Dr.
Miller's data consist in the following
mortality rates for "prediabetic" and
non-diabetic mothers at childbirth.
Number Number Number
Status of of of of infant Mortality
mother mothers births deaths rate—%
Xon-diabetic 59 253 5 2.0
"Prediabetic" 57 264 22 8.3
•See the control chart for poliomyelitis in California (Rich and Terry: Public Health
Reports Vol. 61, No. 42. 1946). Also the report on diabetes in a New England town, J.A.M.A.
Sept 27, 1947. There are two other recent articles of some relevancy: 1) C. W. Jungablut:
Annals of Internal Medicine 26:1, 1947, and 2) M. B. Frank: Proc. Soc. Exp. Bio/, and Med.
62:1, p. 17, 1947. The first correlates blood group A3 with paralytic poliomyelitis, and the
second correlates it with filariasis.
379
The difference is highly significant
statistically. All of the 57 "prediabetic"
mothers were non-diabetic at the time of
delivery. All 57, it was discovered, de-
veloped diabetes later, but none before
40 years old.
Dr. Best, of Toronto, has a chapter
in a new book entitled, Currents in Bio-
chemical Research, in which he says
(page 431) in speaking of the possible
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prophylactic use of insulin: "applica-
tion will not be easy until the potential
human diabetic can be recognized much
earlier than is possible at present." It
is barely possible that PTC or the like
might be of some service here.
There is a diabetic woman here who
says she can tell by a bitter taste in her
mouth when she must take her shot of
insulin. Neither just before taking the
insulin nor at any time before or after
does she detect the taste of PTC.
It looks as if the Negroes may con-
stitute a special case.
M. C. TERKY, M.D.
P.P.S. As to the possibility that insulin
therapy (or diabetes itself) may change a tast-
er to a non-taster, we may have to wait until
the youngest diabetic known to be a taster is
old enough to have the trophic ulcers of the
disease. I have but one case to the point—a
patient who was a taster—and so remains.
The following was contributed by a physi-
ologist who knows some of them: "There are
diabetics who do not take insulin; if they were
tested they would show a low I.Q." Projec-
tion: Insulin is taken now by all diabetics
smart enough to get it Wider use would tend
to preserve an unwanted mutant gene for sus-
ceptibility to diabetes, to lower I.Q. and to
more diabetes 1
It isn't insulin that makes smart diabetics
smart, it keeps them smart longer. If any con-
trollable exogenous factor in the causation of
diabetes is found,* the use of insulin may be
expected to raise the I.Q. and decrease dia-
betes ! Therefore: On with surveys, experi-
ment, and theory.
M.GT.