NATIONAL INSTITUTE OF GENETICS(JAPAN),ANNUAL REPORT
No.1.

(1949"'" 1950),--_·~oo----

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PUBLISHED BY

THE NATIONAL INSTITUTE OF GENETICS(MISIMA, SIZUOKA.KEN, JAPAN)

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Published November 5, 1951

NATIONAL INSTITUTE OF GENETICS (JAPAN)
ANNUAL REPORT
No. 1 (1949-1950)

Contents
Page

History.
.
.
Laws and regulations.
Location, buildings and campus
Experimental fields. . . . . . .
Abstract of diary from June 1949 to December 1950
Staff.
Council.
Researches carried out in 1950 .
A. Human genetics.
1. Microcephaly. . . . . .
2. An apparently new disease with splenomegaly and anemia
as main symptoms . . . . . . . . . . . .
3. Some hereditary characters of the tongue.
B. Cat genetics. . . . . . . . . . . . . . . .
C. Silkworm genetics
I. Studies On unstable genes in the silkworm.
1. Inheritance of multistar marking .
2. Inheritance of multilunar marking . . .
3. Inheritance of knobs. . . . . . . . . .
II. Embryological and genetical studies of a kind of malf()rmation in the silkworm.
1. Role of nucleus and cytoplasm in the appearance of themalformation. . . . . . . . . .
2. Environmental effects Upon expression of the malformation. . . . . . . . . . . .
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III. Studies on new genes belonging to the E-series on sixth
chromosome.
IV. Effect of B.H.C. on meiosis.
1. Abnormality in spermatogenesis.
2. Non-disjunction of chromosomes
D. Population genetics of some invertebrates.
I. The lady-beetle, Harmonia axyridis.
1. Geographic and temporal variation in the relative frequency of pattern types and in the frequency of elytral
ridge. . . .
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2. Microgeographic variation . . . . . . . . . •II. The land-snail, Bradybaena similaris. . . . . . .
III. The butterflies, Zephyrus taxila and Colias hyale .
E. Physiological genetics of Drosophila
F. Endocrinology of some insects.
G. Genetics of some cereals.
1. Radiation genetics conducted on cereals.
2. Nullisomics deficient in a chromosome pair of the D-genome
found in pentaploid wheat hybrids.
3. Genome·analysis in Argopyrum, a genus related to Triticum.
4. Other studies on wheats. . . .
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5. Cytogenetic studies on wheat-rye hybrids.
6. Determination of right- and left-handedness in the spikeletof Triticum monococcum vulgare . . . . . . . . . . . . .
H. Genetics of fertility and sexuality in plants.
1. Sterility of triploid plants. . . . . . . . . . . . .
2. Sterility and autotoxicosis of Colchicum autumnale
3. Karyotypes and sterility in Hemerocallis . . .
4. Studies on sex-differentiation in higher plants.
5. Sterility of Phaseolus multiflorus under high temperature . . • . . . . . . . . . . . . . . . . . . . . . . .
J. Genetics and cytology of some flowering plants.
1. Genetics of flowers in Zinnia elegans. .
2. Genetic studies on Solanum melongena and Capsicum

annuum

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3. Karyological studies of Lycon's radiata .
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4. Cytogenetic studies on the genus Nicotz"ana. . . . . . . 35
5. Some effects of sodium ribose nucleate on somatic mitosis
of some plants. . . . . . . . . . . . . . . . . . . . . 35
K. Studies on lower organisms.
1. A method of measuring the rate of growth of Neurospora

sitojJhila.
2. Studies on the virus infecting silkworm.
L. Improvement of some useful plants.
1. Improvement of sugar beet by means of induced triploidy.
2. Genetics and breeding of Citrus plants .
3. Induced polyploidy in useful plants.
M. Theoretical genetics.
1. Semiallelic genes. . . . . . . . . .
2. On variance due to competition between plants of different
types in plant populations . . . . . . . . . . . . . . .
3. Theoretical studies on individual selection in plant-breeding with special reference to gene recombination
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4. Effect of the random fluctuation of selective value on the
distribution of gene frequencies in natural populations. .
5. Physiological theory of polygenic actions. . . . . . . .
6. Recombination of chromosome segments through continued
self-fertilization. . . . . . . . . . . . . . . . . . . . .
7. Studies on the process of chromosome substitution . . .
8. Theoretical relations among map-distance, recombination
value and coincidence ratio. . . . . .
Papers and books written by the members of the staff. . . . . . .

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HISTORYtf•

The National Institute of Genetics was officially organized onJune 1st, 1949. Previous. to this, the demand for a national institute devoted to the study of genetics in Japan had been expressedby Japanese geneticists on various occasions. It had been taken
for granted from the beginning that such an institute should be
an independent one, instead of being a part of a university orother institution. It had also been conceded that, in view of theextensive field now covered by this science, and the intricate
relation between this science and many other sciences, the institute should be of a fairly large scale. The first formal announce~
ment of such a demand was made by the resolution passed
unanimously at the Thirteenth General Meeting of the Genetic.Society of Japan held in August 1940 in Seoul. A little later, a.special committee for the study of genetics was organized in the'Foundation for the Promotion of Scientific Researches in Japan.'
Most of the senior geneticists became members of this committee,
and shared its activities. The formation.of this committee paved
the way for the establishment of a national center for the studyof genetics in Japan. Besides, a small foundation for the promotion of theoretical and applied genetics, called the Genetics
Research Institution, was organized in May 1947 by the donationof Mr. S. MATSUMURA, Member of the House of Councillors.
Practically the same group of geneticists joined this foundation
also and cooperated in their research activities. This foundation
became the forerunner of the National Institute of Genetics.
In July 1948, the bill for the establishment of a national instituteof genetics was presented by the Ministry of Education to theDiet, and passed. The formal start of the institute was made on
July 1st, 1949, by the Government Law No. 146. On the sameday, the office of the Institute was openned in the Ministry of
Education with Mr. K. KENNOKI, Director of the Higher Education and Science Bureau as the Acting Director. Moreover, acouncil responsible for setting up the basic principles of theorganization and function of the Institute was formed on thesame day. At its first meeting held on July 30th, the council
nominated Dr. KAN OGUMA, Emeritus Professor of Hokkaido
1University, as the Director of the Institute, who was officiallyappointed on August lOth, 1949. Within a few months the restof the staff was chosen, and by the middle of 1950, nearly all of
the present members were appointed.
Thus the new Institute was started. For the establishmentof the Institute under the current difficult circumstances, we areindebted to a number of interested scientists and officials. Especially, the valuable help accorded by pro H. C. KELLY of C.I.E.,
G.H.Q. and Dr. S. KAYA, formerly Director of the Higher Education and Science Bureau in the Ministry of Education, should
not be passed without mentioning. Among the geneticists, Dr.
H. KIHARA and Dr. K. MASUI were most active in realizing the
project laid out by geneticists' group.
The whole Institute moved into the present building in Misima
on October 29,th, 1949. Shortly after this, the Institute received
a request from the President of the Japan Monopoly Corporation
to help their breeding experiments aimed at the improvement of
the varieties of the tobacco plant cultivated in Japan. The proposal was accepted, and a branch of Tobacco Experiment Stationof the Corporation was started in the Institute on May 25th, 1950.
The botanical members of the Institute headed by KIHARA havevolunteered their assistance in these' experiments.
In November 1950, the Institute received another request froma group of poultry breeders to improve the white Leghorn strain
in Japan for higher egg production. A new association called'the Whole-Japan Association of Poultry Genetics' was organized forthis purpose, and contributed the funds for this untertaking.
The experiments are to be carried out in aviaries to be constructedon the campus of the Institute, under the supervision of Y. TANAKA.
In July 1950, Prof. R. GOLDSCHMIDT presented an offer to theInstitute to submit his own library for the use of the Institute.
On February 2nd, 1951, more than 50,000 reprints and about 600
books arrived in good condition. These reprints and books will
become the nucleus of a library for the Institute, the need of which
had previously been seriously felt.

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LAWS AND REGULATIONS (Abstracts)

The Japanese Government Law No. 146
The National Institute of Genetics shall be an organ of studies
on the principles of heredity, and for providing a theoreticalbasis to the solution of practical problems in heredity; also it
shall guide and promote genetical researches in Japan.

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The Ministry of Education Ordinance No. 30
The National Institute of Genetics shall have the following
Departments :-the Department of Administration, the First Department of Research, the Second Department of Research, theThird Depm-tment of Research.
The Department of Administration shall consist of the Sectionof General Business and the Section of Finance.
The First Department of Research shall persue research works• on: a. phenogenetics in man, b. phenogenetics in animals andc. phenogenetics in plants.
The Second Department of Research shall pursue research works
on: a. problems concerning cytoplasmic inheritance and b. problems concerning chromosomal inheritance.
The Third Department of Research shall pursue research works
on: a. physiological genetics and b. biochemical genetics.

Cabinet Order No. 247•

The Board of Councillors shall advise the Director concerning
the matters stated below:-a. planning of researches and otheractivities in the National Institute of Genetics for the coming
year, b. budget for the coming year, c. appointment and dismissalof staff member and d. other important administrative business.
The Board of Councillors shall consist of members not morethan sixteen in number.
The members shall be appointed by the Minister of Educationfrom among competent persons in educational, academic andeconomic circles.

3Regulations concerning Research Associates
Research Associates shall be appointed by the. Director fromamong persons who have full training in genetics and intend to
do research works in the Institute.
A Research Associate may use the equipment of the Institutefor his research work; he may also use the organ of the Institutefor the publication of the results obtained thereby.

Regulations concerning Research Students•

Research Students shall be appointed by the Director fromamong applicants who have completed their course in genetics
in some university or college, or who have qualifications of acorresponding grade, and who intend to pursue genetic researchin the Institute under the direction of the staff.

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Regulations concerning Practice Students
Practice Students shall be appointed by the Director from among
applicants who have graduated from a Middle School of the Old
School System, or from a High ~chool of the New School System,
and who intend to study genetics in the Institute under the
direction of the staff.
LOCATION, BUILDINGS AND CAMPUS
It had been an important problem from the beginning of thecampaign for the establishment of a national institute of genetics,
to secure a suitable site and building for the institute. It had
been agreed upon that, the site should be located within a reasonable distance from an academic center, and in a district of mild
climate. There should be standing a building of proper size andconstruction, in view of the great difficulty in erecting a new
one under the financially most difficult circumstances. Also an
adequate amount of fertile lap.d should be available in the neighborhood. After a great deal of searching within and around Tokyo,
the present site was finally chosen. This site seemed to meetfairly satisfactorily all the requirements enumerated above. It
is located in a suburb of the city of Misima, on a small foot hill
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of Mt. Hakone. Buildings formerly used as a factory for manufacturing aeroplane accessories were standing virtmi.lly intact. A
levelled area of nearly 19 acres was attached to these buildings.
The city of Misima is located 118 kilometers west of Tokyo, andcan be reached in two hours by express trains and in two andhalf hours by local trains. The climate is mild, and the neighboring district is noted for various kinds of vegetables of goodquality, as well as for richness of dairy products.
One of the buildings, a large two-storey wooden building, 1,165.66
tubo (1 tubo=6 feet square) was taken for the use of the Institute.
It was repaired, partitions were set up inside, and it was turned
into a building containing 20 laboratories, 3 offices, a lecture room,a library, a dark room, store-rooms, etc. Two smaller buildings
attached to the main building were remade into living apartments
for the staff as well as a garage.
EXPERIMENTAL FIELDS•Of the whole land 19.23 acres in area, 11.46 acres were turned
into a field suitable for growing plants for experimental purposes.
At the start it was necessary to make the land arable. In January
1950 about the half the land was made suitable for cultivation,by breaking the ground with a tractor and by adding :;l. large
quantity of organic matters and neutralizing chemicals to the soil.
The other half will be treated in the same way during 1951.
The principal plants grown for experimental purpose in 1950
were as follows:Several varieties of Triticum, Aegilops, Solanum melongena
var. esculentum, Capsicum annuum, Nicotiana tabacum, Zinnia
elegans, Cosmos bipinnatus, Aster spp., Saccharum o.fficinarum andthe tetraploid and triploid water-melon Citrullus vulgaris, tetraploid radishes Raphanus sativus, Ricinus communis and Morus·bombycis.

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ABSTRACT OF DIARY FROM JUNE ]949
TO DECEMBER 1950
1949
June 1. The National Institute of Genetics was officially organized;
Mr. TOSHIHIRO KENNOKI was appointed its Acting Director;
and the Board of Councillors was formed.
July 30. First meeting of the Board of Councillors.
August 10. Dr. KAN OGUMA was appointed the Director of theInstitute.
September 14. The land 23,526 tubo (19.23 acres) in area locatedat Yata in Misima was bought from the Fuji Industrial
Company for the campus of the Institute.
October 1. Buildings 1,347 tubo in floor dimension were rented
for the laboratories, offices, etc. of the Institute.
.October 29. The Institute moved to the buildings in Misima.

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1950
January 23. Second meeting of the Board of Councillors.
February 11. Public tecture in the Public Hall at Shizuoka.
12. Public lecture in Laborer's Hall at Misima.
March 29. Staff meeting.
April 29. Board meeting of the Genetics Research Institution.
Joint meeting of the National Committee of Genetic
Researches of the Japan Science Council and the National
Committee of Researches in Plant and Animal Breeding of
the Japan Science Council.
May 25. Inauguration of the Misima Branch of the Hatano
Tobacco Experiment Station of the Japan Monopoly Corporation.
May 26. Third meeting of the Board of Councillors.
.September 3. Committee meeting of the Genetic Society of Japan.
September 4. Joint meeting of the National Committee of Genetic
Researches of the Japan Science Council and the National
Committee of Researches in Plant and Animal Breeding of
the Japan Science Council.
September 5. General meeting of the Society of Chromosome
Research.
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September 24. Inauguration meeting of the Whole-Japan Association of Poultry Genetics.
November 18. Inauguration meeting of Misima Geneticists' Club.
December 15. Board meeting of the Genetics Research Institution.
December 16. Meeting of the Misima Geneticists' Club.
STAFF

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Department and Laboratory Heads
Kan OauMA, D. Agr., Director, and Head of the Second Department
Yoshimaro TANAKA, D. Agr., D. Sc., Head of the First Department
Taku KOMAl, D. Se., Head of the Third Department
Part-time Staff
Hitoshi KIHARA, D. Sc.
Yosito SINOTO, D. Sc.
-Laboratory Heads
YO TAKENAKA, D. Sc.
Kan-Iehi SAKAI, D. Agr.
Seiji MATSUMURA, D. Agr.
Mitsuo TSUJITA, D. Agr.
Kazuo FURUSATO
Research Assistants
Masahiro OGAKIMoto KIMURA
Kanji GOTOH
Taro ITO
Bungo SAKAGUCHI
Toru ENDO
Sanae KAJI
Akira MIYAZAWA
Temporary Assistants-9
Research Associate
Yoshinari KuwADA, D. Sc.
Department of Administration
Morihei TSUKAMOTO, Head of Department
Sumiyoshi Sumo, Head of General Business Section
Masao MIYAZAWA, Head of Finance Section
N~omi MATSUBARA
Hiroko NAKANO
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Junzo KADOWAKI
Typist, Telephone operator, Chauffeur, Field-laborers, Janitors,
etc.-9.
Misima Branch of Rutano Tobacco Experiment Station
Masao TANAKA, Head
Flora LILIENFELD, Ph. D.
Seiji IMAI
Assistants - 3
COUNCIL

YO K. OKADA, Professor of Tokyo University, Chairman
Seishi KAYA, Professor of Tokyo University, Vice-chairman
Fujio EGAMI, Professor of Nagoya University
Tanemoto FURUHATA, Professor of Tokyo University
Seizo KATSUNUMA, President of Nagoya University
Takeji KOBAYASHI, Governer of Sizuoka Prefecture
Makita KOGURE, Professor of the Tokyo College of Agriculture
and Technology
Yoshio KOYA, Director of the Institute of Public Health
Kiyoshi MASUI, Emeritus Professor of Tokyo University
Bungo MIYAZAWA, President of Woman's College of Anjo University
Toshitaro MORINAGA, Head of Department of Physiology andGenetics of Agricultural Technique Research Institute
Daigoro MORIWAKI, Professor of Tokyo Metropolitan University
Masanori NAKAIZUMI, Professor of Tokyo University
Dichi NODA, Member of House of Councillors
Bungo WADA, Professor of Tokyo University
Yasuke YAMAGUTI, Professor of Ibaraki University

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RESEARCHES CARRIED OUT IN 1950

A.

HUMAN GENETICS(Report by Taku KOMAI)

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1. Microcephaly

Microcephaly is one of the most severe congenital defects in man. It
causes high-grade idiocy. Various views have been propounded concerning its origin and heredity. This difference in opinion is at least partly
due to the rarity of this abnormality. KOMAI, in cooperation with thepsychiatrist Dr. K. KISHIMOTO of Nagoya University, is pursuing a genetic
study of this abnormality. So far, they have collected nearly 50 caseswith family histories. Psychiatric and anthropometric examinations
have been made for most of them. Most of the cases have a genetic
back-ground, and are apparently due to a completely recessive gene.

2. An Apparently New Disease with SPlenomegaly andAnemia as Main Symptoms•,

A family of farmers living in a suburb of Kyoto have a lesion withsplenomegaly and anemia as the main symptoms. KOMAI, with thecollaboration of Dr. M. MASUDA and some students in the Prefectural
Medical College of Kyoto, is studying the genetics of this disease. The
disease resembles Banti's syndrome in many respects. It has the great
peculiarity, however, that only women of the family are affected, andall the women connected to the pedigree on the mother's side have either
splenomegaly or anemia or both.

3. Some Heredz"tary Characters of the Tongue
The presence or absence of the ability to roll up the lateral edges of
the tongue in man is controlled by a pair of Mendelian genes. The
manifestation of the character is completed at about the age 12. Amongthe 6-year old children, the incidence of the absence of the ability is
about 50 per cent in both sexes. The incidence diminishes with age,
until it becomes 29.5 per cent in males and 25.5 per cent in females atabout 12, from which time onward the values remain practically constant. These values obtained from Japanese are somewhat different fromthe values so far recorded for Americans and Chinese, although thefigures for these peoples are based on relatively meager material andneed confirmation. Family studies on this ability have shown conclu9

sively its genetic back-ground.
Three families with a slight abnormality in the tongue, having thesublingual frenula abnormally extending to the front and hindering thefree movement of the tongue, have been discovered. This peculiarity
seemes to be due to a dominant gene.

B.

CAT GENETICS(Report by Taku KOMAI)

Opinions are varied concerning the inheritance of common coat colors
in domestic cats, black, tabby and yellow. A census of cats carried out
in some districts of Japan has revealed that black and tabby are due toautosomal genes, while yellow is produced by a sex-linked gene. As forthe rare tortoise-shell male, KOMAI has expressed the opinion that thiskind of male results from unequal crossing over between the X-choromosomes of a tortoiseshell mother cat. The son that receives the X-choromosome containing both the yellow gene and its allele will become atortoiseshell. He will also become sterile owing to the presence of anabnormal X-chromosome containing too much of the femaleness factor fora normal male, but too little of the femaleness factor to become a normal
female, and he becomes a kind of male intersex and sterile. According
to this hypothesis, the mother of a tortoiseshell male should be tortoiseshell herself. All the four indisputable and one probable cases collectedby KOMAI and the seven cases observed by Dr. C. KEELER conform tothis postulate. The decisive evidence for the hypothesis will be obtainedby cytological observa.tions. Prof. S. MA.K:INO'S examinations of the sections of testis of one of these cats did not show any mitotic figures, but
his study on this line is still under way.

C.
1.

SILKWORM GENETICS

Studies on Unstable Genes in the Silkworm.
(Report by Yoshimaro TANAKA)

1.

Inheritance of Multistar Marking

In the larva with normal marking, there are regt.darly found threepairs of spots; i. e. the "eye" spots on the second segment, the crescentspots on the fifth segment and the star spots on the eighth segment,
the segment numbers being counted all through the thoracic and abdominal segments. When a larva is provided with paired or unpaired spots
on one or more segments other than the above-mentioned, it is called
multistar. The multistar marking is recessive to the normal and represented by the symbol ms.

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The material of the present .study consists of descendants of a single
egg-batch of a Chinese bivoltine race Shao-shing, which the author
obtained forty years ago from a breeder. The culture has been continued
ever since, and several types have been isolated chiefly by selection
through more than fifty generations. Of these types, the more importantones are as follows:
1) 0 type, which has no star spots, except the normal "eye" spots
and the crescents.
2) 8-10 type, which has star spots on the 8th, 9th, and 10th segments.
3) 6-10 type, which has stars on the 6th, 7th, 8th, 9th, and 10th
segments.
4) 4-10 type, which has stars on the 4th, 6th, 7th, 8th, 9th, and10th segments.
5) 6-9 type, which has stars on 6th, 7th, 8th and 9th segments.
To explain the process of breeding of various star types, the 0 typemay be taken as an example. At the start 33 individuals of this typeappeared in a lot containing 160 larvae. These individuals were selected
and inbred. The percentage of the invididuals showing the 0 type went
up generation after generation, though not steadily, and 4 lots consistingexclusively of this type were obtained among 7 lots in the sixth generation. Since then, the author has been breeding only the 0 type in thisstrain. However, this type has never been made pure, throwing in every
generation a few exceptional types.
Other star types behave similarly in many respects, although thevariability is much different from one another, and also from the 0 typedescribed above. The stability of the star spots is also different according to segment; e. g., the spots on the 8th segment are most stable,
succeeded in stability by those on the 9th and 10th segments, while thoseon the 6th and 7th are most variable.
The author assumes the presence of several modifiers which reduce or
increase the number of the spots. These modifiers do not act independently, but are 'correlated with one another to some extent. For
instance, the stars on the 4th segment are comparatively stable when
they are combined with those on 6-10th segments, but they cannot exist
independently.
Crossing different star types conspicuously increases the range of variation in F 1 and F 2 , but distinct segregation has never been observed.
From these facts, the author is inclined to conclude that the majorgene, ms, does not change, but the modifiers are unstable, and theireffect can be sifted by continued selection in both plus and minU3
directions. The genotype of the normal marking is +p+q+m s (p andq designate plain and quail marking respectively), plain p+q+m s, striped
ps+q+".., quail +P q +"", pale quail p q +"", multistars +p+q ms.
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When ms co-exists with P and q, the pale quail multistar (P q ms) isformed.

2. Inheritance of the Multilunar Marking
The muItilunar marking consists of a series of roundish brownish-yelIowor dark-brown spots on the subdorsal lines of the larva. The gene formultiIunar, L, is dominant to non-multilunar, and is equistatic to othermarkings, normal, striped, moricaud, quail and zebra. Only when it is.
combined with plain marking, P, does the pigment not develop on theskin, so that the presence of the L gene is hardly perceptible except to,
skilled eyes. The gene for multilunar is linked with the gene for "stick,"and is located at the 0 point of chromosome IV.
In the L marking, the characteristic spots are observed on at least four
segments, "i. e. 2nd, 3rd, 5th and 8th, the segment numbers being counted
from the thorax to the abdomen throughout. For the sake of description,
the spots on the 2nd and 3rd segments are disregarded, and only those.
on other segments are mentioned.
The selection has been practised for more than thirty years covering
about forty generations, and the following L types have been isolated:1) 5.8. type, in which L spots aTe limited to 5th and 8th segments.
2) 5.6.8. type, which has L spots on 5th, 6th, and 8th segments.
3) 4-8 type, which has L spots on 4th, 5th 6th, 7th and 8th
segments.
4) 4-9 type, in which six pairs of L spots are developed.
5) 4-10 type, which is provided with seven pairs of L spots on all
segments from 4th to 10th.
The author assumes several modifying genes which control their respective distributions of spots, in the presence of the major gene L. The.
5.8 and 5.6.8. types, are always heterozygous for L-gene.
In spite of long continued selection, none of these spot-types has bred
true; they proouce some (10-30%) exceptional forms in each generation.
Even in the most stable type 4-10, a few exception~l individuals appear.·almost invariably. When the 4-10 type is crossed with some other race,as many as 10-20% exceptional types appear even in the F 7 generation
after rigorous selection fer a pure 4-10 type.
According to the author's opinion, these breeding results can be explained by the unstability of modifying genes which can be changed inplus or minus direction by selection. There must be a limit, of course,
beyond which the selection is ineffective. The major gene apparently
remains unchanged from the beginning.
It seems to the author that the present case as well as the case of
the multistars can hardly be explained by the assumption either ofmutable genes or polymerism, or of the polygene hypothesis of MATHER.
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3. Inheritance of Knobs,••"Knobbed" is a remarkable characteristic of the Chinese race, "Dragon
horn." These larvae are provided with four pairs of large skin protuberances on the second, third, fifth and eighth larval segments (thoracic andabdominal segments being included in the count). Usually the knobs of
the fifth segment are the largest, succeeded in size by those of the eighth
segment, those of the second segment being the smallest. Knobbed isdominant to normal, and is represented by the symbol K.
The facts that all the four segments on which knobs appear are eachprovided With a pair of spots (the eye-spots, crescent spots, or star spots),and that the ·positions of the spots and knobs coincide exactly, speak fora close relation between the swelling of the skin and the developmentof dermal pigment. This has been proved by crossing K-strain with L(multilunar) or ms (muItistar) strain; knobs appear on all segments where
L or ms spots are present. On the contrary, the striped marking factor
inhibits almost entirely, when homozygous, the development of knobs,
while L-spots and striped marking are both fully developed when theyco-exist.
After a long-continued selection, the author obtained the following
strains with regard to the distribution of knobs;
1) 5-8 type, which has two extra pairs of knobs on the sixth andseventh segments;
2) 5--9 type, which is provided with three extra pairs of knobs on
the sixth, seventh, and ninth segments;
3) 5-10 type, which has knobs on all segments from the second to
the tenth, except the fourth;4) 5.0 type, in which knobs are absent from the eighth segment,
and are present on the second, and fifth segments only;5) 4-10 type, which has knobs on all segments from the second to
the tenth;6) 4.5.0 type, in which knobs are present on four segments fromthe second to the fifth, but absent from the eighth.
Extra knobs are as a rule, smaller than those regularly present on thefifth and eighth segments.
Selections have been practised with respect to the number of knobs,
and they have proved to be highly effective in both directions, plus andminus. For example, those larvae provided with extra knobs on the
fourth segment increased in number after selection extending over nine
generations; then the selection was turned to the minus direction, andthe individuals destitute of knobs on the fourth segment only were allowed
to reproduce. The knobs on the fourth segment decreased rapidly, andafter two or three generations, no larvae had knobs on that segment.

13

In another strain, only a few larvae had knobs on the tenth segment.
Before the selection was practised, the percentage of such individuals
had remained nearly the same, When once the selection was started
for the extra knobs on the tenth segment, such larvae increased rapidly,
and after several generations, they became'more numerous than the larvae
having no knob on the tenth segment.
ALhough the selection has been effective in both directions, no pureextra-knob type has been obtained, some aberrant forms appearing fromtime to time. These aberrant forms may be increased or decreased by
selection.
I conclude, therefore, that the modifying genes which control thenumber of extra knobs are unstable, and vary in both the "plus" and"minus" directions while the major gene K remains unaffected.

II.

•

Embryological and Genetical Studies of a Kindof Malformation in the Silkworm(Report by Mitsuo TSUJITA)

1.

Role of Nucleus and Cytoplasm in the Appearanceof the Malforma#on

OMURA (1948) has reported on a strain of abdominal segment malformation in the silkworm larva. The present report deals with embryologicaland genetical studies of this strain. The malformation appears on thedorsal and ventral sides' of several segments ranging both anteriorly andposteriorly from the 5th segment. The shape and degree of the malformation vary markedly from individual to individual. In extreme cases
one or two segments are missing; often one or two segments on
one side of the larval body are gone, and the body is curved like ahook. In the cases where the deformity is slight, the larva looks almost
normal. Between these two extremities there are various degrees ofabnormality. The number of caudal horns ranges from one to several,and these are irregularly located on the dorsal side. Besides, deformity
may be found in various organs; for example, trachea and spiracles,
dorsal vessel, muscles, ganglia, Malpighian tubules, silk glands, gonads
and midgut. Among these gonad malformation is very conspicuous, thenumber of gonads on one or both sides may increase, to three or four; in.some individuals, the gonad is missing on one side. Furthermore, the ovarian
tubules and the testicular follicles may vary in number, from one to six.
.Sometimes the segmental structures appear normal, while the internal·organs are abnormal. The most distinctive feature of this strain is that many
·deformed individuals emerge without getting any treatment with dilutehydrochloric acid. The penetrance of the gene for this abnormality is

14

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about 50-70%. The cross between this strain and the normal strain
yields no abnormal individuals in Flo no matter which strain is used as
mother, and a few mutants segregate in F 2 and F 3 , with a much lower
frequencey than expected on the monohybrid basis. Some apparently
normal females were mated with abnormal males, and the eggs were
exposed to high temperature (of 40°-41°C) within one hour after their
being laid, i.e., about the time of the maturation division of the egg.
By this treatment numerous exceptional individuals were obtained throughmerogony, the egg nucleus degenerating and two sperm nuclei conjugating together. In this case almost all individuals became abnormal,
showing the characteristic features mentioned above. It has been concluded, from the results obtained by experiments ensuring various combinations of nucleus and ooplasm of the normal and abnormal strains,that the appearance of the malformations is largely governed by genes
in the nucleus, irrespective of whether the cytoplasm comes from the,abnormal strain or from the normal strain.

2. Environmental Effects upon the Expression of the Malformat£on••

As stated by OMURA (1949), when the early-stage eggs of the strain,ghowing the abdominal segment malformation are immersed in a dilutehydrochloric acid sCllution, abnomal individuals appear more frequently
than otherwise, amounting to 80-l00%. Thus, almost all individuals,can be made abnormal by this treatment. It has been ascertained that noor few abnormal individuals are produced by change in the environment
in early stages. Eggs of this strain were kept for lO~15 hours, directly
after being laid, at the high temperature of 25°C, and then brought intoa low temperature room (l5°C) where they were kept for several days.
This period corresponds to the stage extending from the migration of
the egg nuclei toward the periphery to the diapause of the embryo.
Also, eggs which had been treated with the low temperature mentioned-above were kept during 40-60 days at the refrigerating temperature of
5°C and then immersed in dilute hydrochloric acid. Strikingly, almost all-of the larvae hatched from these eggs were normal. On the contrary,
the eggs of the same strain produced 80-85% crippled larvae, when they
were kept at temperatures of 25°-30°C for 36 to 48 hours after being
laid, refrigerated for 40-60 days at 5°C, and then subjected to the usualhydrochloric-acid treatment.
Judging from these results, it may be said that, at 25°C, the stage •from 15 hours to 36 hours after oviposition is the period during which
the body segments are differentiated. Thus, the frequency of crippledlarvae can be -controlled to a fairly large extent by the adjustment of-certain environmental conditions, notably temperature.
As stated above, the segment formation in the silkworm is under the15

control of genic activity. The present experiments, however, have madeit clear that the expression of this activity is strongly affected by theenvironmental conditions. However, when eggs of this strain, laid bymoths which had emerged from apparently normal silkworms, had been
subjected to low temperature treatment at an early embryonal stage
through two or three generations, and placed in an ordinary temperatureroom, abnormal individuals emerged at a ratio of about 50-7096. Thisshows that the experimental results described above cannot be attributed
to mutation. The differences observed are rather due to a varying pe·
netrance of the same genes.

III.

Studies on New Genes Belonging to the E-Series on.
Sixth Chromosome•t(Report by l\IitslJO TSUJITA)The ancestor of this strain was discovered as a mutant among the
hybrids between a Japanese race and a Chinese race. The mutant has
duplicated star-shaped patterns. This character is dominant, and when
the F 1 of this mutant and the normal is backcrossed to the normal,
individuals with supernumerary stars and normal individuals segregate
according to the 1 : 1 ratio. Among the laTvae of the former type SClme
have star-patterns on the 7th and 8th segments, or on the 8th and 9th
segments; also there are individuals with star-patterns on the 7th, 8th
and 9th segments. The embryos homozygous for this gene also J;1ave
supernumerary abdominal legs on the 10th and 11th segments, and theydie in the later embryonic stage or immediately after hatching. It is a
striking fact that in this strain, larvae with duplicated crescent patternson the dorsal side of the 5th and 6th segments occasionally appear,
apparently as mutants. As this character is also dominant, normals andmutants segregate in the back-cross of F 1 according to the 1 : 1 ratio.
Sometimes both larvae with crescent patterns on two segments andlarvae with crescent-patterns on three segments emerged from the samebatch of eggs. This strain is characterized by having no star-pattern.
Embryos homozygous for this gene also die in a later stage; they areprovided with thoracic legs and thoracic setae on all segments, and areincapable of blastokinesis.
From the strain of the multi-star pattern, some larvae entirely devoidof the star-pattern were obtained. These larvae were viable even though
they were homozygous for the gene 'in question. Genetic analysis of thischaracter is now under way.
The number of the spots, i. e. the stars and crescents, can be increased
by selection through sib-mating.

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Judging from these breeding results, the gene for the supernumerarystars and the gene for the supernumerary crescents are in multipleallelic relation, both of them being located at the 0.0 point of the 6thchromosome, to which locus ten genes have already been assigned.
The gene for the multi-star pattern is undoubtedly different from Ee7
reported by TAKASAKI ('47), while the gene of non-star-spots apparenly
resembles Ec r , but the latter can be easily distinguished from the former
by its lethal action. The multi-crescent character is similar in appearance
to EN discovered by IcHIKAWA ('43), although different in some respects.

IV.

Effect of B. H. C. on Meiosis(Report by Mitsuo TSUJITA)

•

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•

1.

Abnormality in Spermatogenesis

NYIOM and K~UTSSON ('47) have reported that mitosis in the cells of
the onion root is interrupted by the action of the r-isomer of B.H.C.
TSUJITA and SAKAGUCHI have observed that the mitosis of the spermatocytes in silkworm poisoned by this chemical is also markedly disturbed.
Larvae 3 days after the 4th moulting were dusted with B.H.C. powder,which contains 0.5% of the r-isomer. The larvae soon afterwards showedsigns of poisoning. Ten, twenty, thirty and fourty hours after the treatment, cytological observations of the poisoned larvae were carried out. The
spermatocytes in the testes of healthy larvae were examined as controls.
In the spermatocytes of the poisoned larvae ten hours after the treatment, 26, 27, 29 and 30 chromosomes were counted on the equatorialplate of the 1st spermatocyte, although in some of them the normal
28 chromosomes were observed. Often aggregation of several choromosomes occurred. Moreover, lagging of one or several choromosomes wasobserved at the anaphase. These abnormalities increase with the advanceof time and stage.
Among the materials from poisoned larvae thirty or fourty hours after
the treatment with B.H.C., some showed about half of the chromosomesalready at the poles, while the rest remained on the equatorial plate.
Thus, the spermatocytic division may be thrown into confusion by thistreatment: also fusion of chromosomes occurs. This change seems to bedue, partly, to the effect of B.H.C. on the protoplasm followed by someobstruction in the spindle-formation. Also, it is possible that the chromosomes themselves are affected.

2. Non-disjunction of Chromosomes
NYLOM and KNUTSSON ('47) suggest that the r-isomer of B.H.C., likecolchicine can induce polyploidy. The present experiment was carried17

out to test this possibility. Although no polyploid has been obtained.
trisomies due to non-disjunction of chromosomes were observed withfairly high frequency. The outline of the results is presented below.
Male and female larvae having the genic composition pSpM were treatedwith 0.596 B. H.C. powder 3 days after the 4th moulting. Among the
200 treated larvae, 11 females and 5 males survived to emergence. These
were mated with moths of a strain with the normal pattern (+1' +1').
The number and hatching rate of the F eggs were almost the same as.
those of the control lots.
In the lots in which the male larvae had been treated, six larvae ofan exceptional type appeared. These had both striped and moricaud
patterns. Four of these (three females and one male) survived to emergence. These were mated with a normal strain. The eggs were notlarger than those of the controls. The segregation of the larvae withrespect to colour pattern is shown in the following table:j•

Table 1.
Mating type

Stripedpattern

Moricaud_patternSMpattern

Normal

Total

44
44
31

4G
50
5u

46
51
47

51
59
70

187
204
203

46

71

83

(jO

260

pSpMx +p
No. 50m
5032
5033+ p x P'" p.~f
No. 5034

From the table shown above, we learn that the female was trisomic
Consequently, among the larvae
with the striped pattern, there are two types, i.e. ps+1' andps+p+p, andsimilarly, the larvae with moricaud pattern are either pJf+1' or pJf+1'+1'.
'This was substantiated by cytological observations. Thus, it seems
certain that B.H.C. has induced irregularity in the chromosome distribution, and a supernumerary chromosome has been added to the ordinary
2N chromosome set at the maturation division of the spermatocyte.
(PSpM+l') as to the second chromosome.

D.

POPULATION GENETICS OF SOME INVERTEBRATES(Report by Taku KOMAI)I.

The Lady-beetle, Harmonia axyridis

1. Geographic and temporal variation in the relative frequencyof pattern types and in the frequency of elytral ridge
The various types of elytral pattern are due to a set of multipleallelic genes. The relative incidence of these genes considerably varies
18

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in different localities. Samples were obtained from 26 localities in Japan
as well as in Korea and Manchuria. The geographic variation shows agradual change so as to present a distinct cline in respect to thischaracter. An abrupt gap, however, may be found at the Strait ofof Tsugaru (the so-called Blakiston Line) and also at the Kore'l Strait.
A similar geographic variation is found in the frequency of the individuals provided with a ridge near the distal end of the elytra. The
presence or absence of this ridge is governed by a set of Mendeliangenes, the gene for its presence being dominant. Furthermore, temporalvariation (variation according to the difference in time) has been observedin materials collected in some localities. Materials collected on different
occasions with 20 to 40 years' interval have been compared. The
materials from Sapporo and Suwa at least, show a distinct change which
took place during 20 years and 40 years respectively. This change may
be seen in the incidence of both elytral' pattern and the elytral ridge.
2. Microgeographic variation••

There have been found, in the range of· distribution of this beetle inJapan, at least two 'islands' which comprise characteristic populations
very different from the neighboring populations. One is· at Sanage in
Aiti Prefecture, visited last year by KOMAI and HOSINo. The beetle
populations inhabiting pine-tress in this district are characterized byhaving a surprisingly high percentage of the otherwise extremely rare
type, axyridis, while the populations inhabiting other plants, wheat,fruit-trees, etc., are practically identical in composition to the populations in the neighboring localities It has been known that the aphids
infecting the pine are different from those infecting other plants. We
are, however, ignorant of how this difference in food aphids is connectedwith the difference in the beetle population. Another example of thisbeetle showing very different composition from those in the neighboring
localities is that of Kamikawa in Hokkaido. Here, the composition as awhole is very different from the samples coming from other localities in
Hokkaido. However, I have no knowledge of the foad aphids of thisexample. Further inquiry is under way.
(T. KOMAI, and Y. ROSINO: Contributions to the evolutionary genetics ofthe lady-beetle Harmonia axyridis. II. Microgeographic variation.
in the July 1951 Number of 'Genetics')

II.

To appear

The Land-snail, Bradybaena similaris

This common land-snail has four distict color and banding types,
namely, yellow plain, yellow banded, purple plain and purple banded.
These are known to be due to a set of triple-allelic genes. 83 popula-19

tion samples of this snail from 62 different localities in Japan have been
examined for the relative incidence of these genes. Generally speaking,:samples from localities of the same district in Japan show similarity,
There are occasional cases, however, in which samples from adjacentlocalities have very different composition. Also, distant colonies may
have very similar composition. When compared with the study on thelady-beetle stated above, or the study on Lymantria by GOLDSCHMIDT,
the absence of any cline in the land-snail in contrast with its presencein those insects appears striking. It is obvious that this difference iscorrelated with the difference in the capacity of migration between thetwo kind of animals.

III.

The Butterflies, Zephyrus taxila and Colias hyale

The female of the Lycaenid butterfly Zephyrus iaxila common in themountainous districts in Japan has four distinct color types, namely,
plain dark-brown, dark-brown with a light-brown marking on the forewing, dark-brown with a bluish marking on the fore-wi1)g, and darkbrown with both the light-brown and bluish markings on the fore-wing.
It has been postulated that these color types are produced by a set of
triple-allelic genes. Four population samples or this butterfly fromdifferent localities of Nagano Prefecture comprise these four types, andthe relative incidence comforms well with the postulate given above.
No simihlr case has been recorded for butterflies.
Genetic studies on the Pierid butterfly Calias hyale are being pursued
by S. AE, a student in the Kyoto University under KaMAl'S direction. SO'
far it has been confirmed that the white wing color character is due tcf
a sex-controlled dominant gene W, and the homozygote WW seems tobe viable at least in some cases. Population samples of this butterfly
from different localities of Japan are being gathered. It is expected thatthe relative incidence of the genes for yellow and white color will show
geographic variation.

E.

•

PHYSIOLOGICAL GENETICS OF DROSOPHILA(Report by Taku KaMAl)

OGAKI, with J. KITADA of Kyoto University, is working on the powerof resistance to cold of various species and strains of Drosophila.
. Various strains of D. melanogaster, D. virilis and D. americana which had
been kept at 25°C for a number of generations, were used. Practicallyall individuals of D. melanagasfer, when kept at O°C died within 30 hours.
However, some individuals of D. virilis could withstand OC ,for 200
hours. Difference in power to withstand cold may be found among

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various strains in both species. The strains showing mutant characters
are not necessarily inferior in this respect to wild strains. F 1 usually
has a higher power of resistance than either parent strain.
OGAKI is also working on the power of resistance to the effect of
CUS04 in various strains of D. melanogaster. Different ~trains showed
different powers of resistance to this chemical. The ranking of different
strains as to this power does not coincide with the ranking as to theresistance to coid, indicating that the powers of resistance to cold andchemical are not merely manifestations of general vitality.
OGAKI, with T. YAGI of Kyoto University, is working on the inductionof mutation by means of the photo-sensitive substance 'Illuminol'.
Larvae of D. melanogaster were dipped in 0.01 aqueous solution of illuminol R II for one hour, and then illuminated with 100 w electric lamp
through a Mazda VR 3 infra-red filter for 5 hours. A number of abnormal
flies-mouth less, abnormal abdomen, Notch, half-thorax-appeared.
Crucial genetic examination of these abnormal individuals is under way_F.

ENDOCRINOLOGY OF SOME INSECTS(R~port•by Taku KO_V1AI)

KAJI has worked OD the metamorphosis hormone of the moth Phyrosamia cynthia ricini. When a thoracic leg of the larva before the criticalstage is transplanted to an older larva, the leg undergoes metamorphosis
in consonance with the host. However, when a leg of the larva whichhas passed the critical stage is used, it undergoes metamorphosis byitself. For instance, a thoracic leg of a third-instar larva, transplanted
to a fifth-instar larva, became a pupal leg. skipping the fourth- andfifth-instar stages. Also, a thoracic leg of a fifth-instar larva transplanted
to a third-instar larva, underwent metamorphosis. after making two
excess molts, in unison with the metamorphosis of the host, Theseexperiments seem to indicate that, in the former case the larval leg ofthe transplant degenerated under the influence of the pupation hormone,
whereas the development of the adult leg rudiment contained in thatleg was accelerated; in the latter case the development of the adult leg
rudiment was interrupted by the molting hormane.
In another experiment, in which a thoracic leg of a fourth-instar larva
which had passed the critical stage was transplanted to a fifth-instar
larva. the transplant molted when the host pupated, and the former
became a pupal leg when the latter became an adult. This experimentapparently shows the identity of the pupating hormone and the imaginating hormone.
KAJI, with K. HAYASHIYA of the College of Textiles Fibers, Kyoto
Technical University is working on the effect of metamorphoasis hormone on21

different kinds of insect. These workers injected an extract from silkworm pupae or prepupae into larvae of the fly Lucilia. They -also
injected an extract of Lucilia prepupae or pupae into silkworm larvae.
In both cases, the extract seemed to induce pupation in the operated_larvae. The extract of Lucilia was apparently more effective on thesilkworm, than the extract of the silkworm on the Lucilia larvae. The
effective substance is solubie in water, acetone or alcohol, but insolublein chloroform or petroleum ether, and is relativelv resistant to hightemperature or acid. Further work is bein~ pursued.
KAJr'. with Prof. M. ICHIKAWA of Kyoto University, has examined thefunction of the corpus allatum, by transplantation and ligature methods,
using various stages of silkworms as material. The corpus allatum,when transpl''TIted singly, did not induce molting; but when it wastransplanted with the prothoracic gland, it induced molting. This result
indicates that the prothoracic hormone can probably induce both pupation and imagination, and this activity is converted by the corpus-allatum
hormone into molt-inducing.
KAJI, with Prof. M. ICHIKAWA and two students in Kyoto University,
h'ls examined the endocrine activity of the brain in Lepidoptera. Brainsof fifth-instar silkworms were transplanted into fifth-instar larvae or
prepupae of the butterfly Luehdorfia japonica, which has a very long(eleven months) pupal period. This operation evidently accelerated themetamorphosis of the pupal host, which developed very nearly to abutterfly in 3 to 4 months. The hormone from the transplanted silkworm
brain apparently activated the prothoracic gland, and induced meta..
morphosis.

G.

•

GENETICS OF SOME CEREALS(Report by Seiji MATSUMURA)

1. Radiatz'on Genetics Conducted on Cereals

Chromosome aberrations in einkorn wheat. Dormant seeds of Triticum
monococcum L. var. vulgare KORN. were exposed to unfiltered X-ray
irradiation, which was applied at 90KVP, 3 mA tube current and 15 em
distance for 12.5 to 62.5 minutes. The doses ranged from 2,700 to 13,500 r
units. The frequency of chromosome aberrations (mostly reciprocal translocations) in the PMC's increased in a parabolic rehtion to the dose.
The relatively small increase of the frequency at high doses has been
assumed to be partly due to limitations by the "saturation effect". At
the same dose (8,100r) and target distance (15 cm) the aberration frequency
decreased with an increase of wave lengths of X-rays obtained by varying
the kilovoltage and the time of exposure (comp. MATSUMURA 1946: PreL
22

I

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•

Rep. in lap. lour. Gen. 21).
If. the two-hit aberrations are due to two separate breaks, their new
recombination should be limited in time and space. The effect of different
wave length was studied by keeping both the dosage (8,100 r) and thetime of exposure constant (296 min.), and changing the target distance.
There has been found similar relation between frequency and wave
length. These facts can be explained on the basis of the difference inionization distribution within nuclei and chromosomes. In the case of
hard X-rays, the majority of the total ionization produced by the electrons
is more scattered than in the case of soft X-rays. The chromosome
breaks induced by hard-ray radiation are generally scattered on different
parts of chromosomes or on different chromosomes, so that the interchange between different parts occurs more readily than in the case ofsoft radiation, where the breaks are closely adjacent and the restitutionof broken ends more commonly results (comp. MATSUMURA 1949: Pre!.
Rep. in lap. lour. Gen. 24).
In most of the cases of induced chromosome aberrations found in
T. monococcum ®+5n, often 6][+2r and seldom ®+ lrv+3 n or ®+4n
have been observed, while in tetraploid 'plants, cases of loss of a wholechromosome or the deficiencies or duplications of chromosome pieces
besides translocations have been confirmed. These findings may be understood by assuming the necessity of at least two complete genomes inthe diplophase.
Chromosome aberrations of the same kind were also induced by fastneutrons and r-rays. The preliminary accounts on dosage and aberration
frequency relation found in these experiments have been published previously (comp. "Cytologia Vol. 16, 1951," MATSUMURA: Radiation genetics
in wheat. I. Chromosome aberrations in einkorn wheat induced byirradiation).
Mutations in barley. Dormant seeds of Hordeum distichum were subjected to X-ray treatment (30-100 KVP, 8,100-16,200 r). Various kinds ofmutants were found among the F 2 progeny of the irradiated seeds. Thesemutants showed chiefly chlorophyll abnormalities: albinistic, yellow,
virescent, striped, shrivelled, fused,. etc. There also appeared mutations
in morphological and physiolosical characters of mature plants, such as:
sterile ear, compact ear, dwarf, narrow leaf, early and late maturity.
Most of these mutants behaved like simple Mendelian recessives. Of
these the ear'ly maturity and compact ear (short culm) are apparently
useful for breeding purposes (MATSUMURA, MOCHIZUKI and SUZUKA 1950:
Rep. KIHARA Inst. BioI. Res. 4).

23

2. Nullisomics Deficient in a Chromosome Pair 0/ theD-genome Found in Pentaploid Wheat Hybrids
Dwarf plants possessing 40 chromosomes occasionally appear among
the offspring of the pentaploid wheat hybrid, Triticum polonicum (AABB)
x T. Spelta (AABBDD). These are nullisomics with the sterile combination 20 rr, deficient in a chromosome pair among the D-genome. If the
7 chromosomes of the D-genome are designated as a, b, c, d, e, f and g,
7 different nullisomics which might be called a-~g-dwarfs, a~cording to
the missing D-chromosome pair, are theoretically possible. MATSUMURA(1947) has actually been able to find all these 7 different dwarfs. Amongthe total of 25 different dwarf lines so far secured, 6 have been recognized
as go, 5 as f-, 6 as eo, 3 as do, 2 as CO, 1 as b- and 2 as a-dwarfs. Theselines differ in their fertility and morphological characters, according towhich chromosome pair of the 7 is missing.
Among the offspring obtained by selfing these dwarfs, some unexpected
gigas-plants with high fertility and more than normal height have beenfound. Such giants appear most frequently in the b-dwarf line: as amatter of fact, one-third of the 'offspring of this line was gigas. These
gzgas-plants are respectively called a-~g-gzgas, according to the original
7 dwarfs (comp. MATSUMURA 1950: Cytogenetics and Breeding of Wheat,
Tokyo).
The hybrids of these 7 different gzgas strains with T. polonicurn mostly
show the chromosome conjugation 1m + 13 n+6!, unlike the combination
14n +7r found in F 1 T. Spelta x T. polonicum. In the hybrids with T.
Spelta, the configurations I m + 19)[+ lr and 20n+2 r were most commonly
met with, and in the case of b- and f-gzgas also 21 n and lrv+ 19n were
frequently observed. In the different gzgas-plants, therefore, the excess
chromosome pair must have been derived from the A- or B-genome.
It has further been assumed that b- and f-chromosomes have many
segmental similarities with the excess pair derived from the A- or
B-genome.
The hybrids between the different gzgas-plants originating in the samedwarf line s30wed mostly the configuration 21rr, and sometimes 11\'+ 19!1
(Im+ 19u + h, 20n +2r). Thus, the excess chromosome pair of the different gigas-plants from the same dwarf line is always identical. On
the other hand, the hybrids between different gigas-plants derived fromdifferent dwarf lines (e.g. a-gzgas x b-gzgas), had the conjugation 20 n +2 1
and I m + 19n + h (seldom ltv). Therefore, there is no doubt that thesupernumerary chromosome in different gzgas-lines is different. Suchchromosomes could be called a, {j, r, 0, e, t and 1 according to whether
they have originated in a-, b-, CO, do, eo, f- or g-gzgas plants The
chromosome a is semi-homologous to a, b to {j, c to r, d to 0, e to e, f
to t and g to 1, a-1 belonging to the A- or B-genome.
24

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•

Dr. SEARS recently obtained all the 21 possible nullisomics in Triticum
vulgare var. Chinese Spring. The writer is now studying the hybrids ofthese nullisomics with his a-~g-dwarfs. The results will be reported
later.

3. Genome-analysis in Agropyrum, a Genus Related to Triticum

•
a

I

,

The genus Agropy1'um comprises 3/ sections; namely, Euagropyrum,
Intrasectional hybrids are easily obtained, but
it is difficult to get intersectional hybrids (comp. MATSUMURA 1948: DGUMA
Comm. Vol. Cyt. Gen. 1).
Among the hybridization experiments between Triticum and Agropyrum,
that between Triticum and the E~ytrigia-section is feasible, while thecrossing of Triticum with Roegneria, an allied plant commonly distributedin Japan, has been unsuccessful. In the hybrid, T. turgidum (AABB) x
A. glaucul1i (2n =42), there appear 5-10 gemini with the mode at 7. Also
the hybrid T. Spelta (AABBDD) x A. glaucum has 2-11 gemirti with themode at 6. A. glaucum, therefore, apparently has a genome homologous
with one of the Triticum-genomes, either A or B. It is difficult to decide
whether this genome is A or B, because the hybridization of A. glaucumwith einkorn wheat (AA) has been unsuccessful.
In the hybrid, T. Timopheevi (AAGG) x A. glaucum, 2-10 gemini (modeat 5) appear. As T. Timopheevi has the A-genome of Triticum, the modeof the gemini number is expected to be at 7. It is inferred from thesefacts that one of the genomes in A. glaucum, which is homologous with
one of the Triticum genomes, is B (comp. MATSUMURA 1949: Jap. Jour.
Gen. SuppI. 2).
According to these results the following genorre formulae are assumed:
E~ytrigia and Roegneria.

Euagropyrum

A. cristatumr A. glaucum

Elytrigia

Roegneria

1 A. obtusiusculum, A. elongatum
A. ciliare
A. semicostatum{
A. Mayebaranum

HH
BBEEFFBBEEFFBBEEEEFFFFIIKKIlKKLLII ? ? ? ?

4. Other Studies on Wheats
a. Triploid hybrids. Hybridizations have been carried out in all possible
triploid combinations between emmer wheat (AABB) or T. Timopheevi(AAGG) and einkorn wheat (AA). Crossing of einkorn wheat with T.
Timopheevi has yielded distinctly lower setting and germination thanthe cross with emmers. The Ft-plants, however, grew more vigorously

25than either parent, with the exception of the case of the crosses with
T. dicoccoides or Khapli (T. dicoccum). The F 1 from T. dicoccoz'des orKhapli x T. monococcum died in the seedling stage, while the F I from T.
dicoccoides or Khapli x T. aegilopoides were dwarfs, only 30 cm in height.
Chromosome conjugation in the hybrid T. Timopheevi x einkorn isrelatively more frequent, and apparently more complicated than that in
the hybrids between emmers and einkorn. This is rather surprising inview of the presence of a tetravalent occasioned by a translocation in
the former. In T. Timopheevix einkorn, the chromosome conjugation
hv+5 n +7r was the commonest; even a penta- or hexavalent chromosome
was sometimes observed, and the number of conjugated pairs varied from
5 to 7. In emmer x einkorn, on the other hand, the number of bivalents
varied from 3 to 7, the mode being at 6. There have been found as
many as three trivalents (once 4 trivalents), occasionally a tetravalent.
Presumably 6 chromosomes from the A-genome of the emmer conjugate
with those of A-genome in the einkorn, and at least one chromosome inboth the A-genomes is semi-homologous. Furthermore, the A-genome ofT. Timopheevi is apparently closer to the A-genome of the einkorn, than
to the genome A of the emmer. The G-genome of T. Timopheevi also
seems to be semi-homologous with the B-genome of emmer, and somewhat akin to the A-genome of the einkorn.
All the F,-hybrids of these crosses were entirely or nearly sterile. All
the 8 F 2 -plants, obtained from open pollination, had nearly 28 or morechromosomes. Some of these had undoubtedly been derived from pollination by dinkel. But no mature offspring was produced by artificial
back-pollination or pollination of dinkel. In Fa and later generations theoffspring with 28 chromosomes had gradually recovered their fertility,
and they s:10wed the appearance of the emmer type (MATSUMURA 1950:
Rep. KIHARA Inst. BioI. Res. 4).
b. The manifold effects of the lone-glume gene P. An awnless tetraploid with long glumes (PP) was crossed to T. dicoccoides (2n=28), andanother tetraploid with short glumes (PP) to T. persicum (2n =28). In
the F 2 of these crosses long glume (P), awnless (A), hairy glume (Hg)and red coleoptile (Rc), in contrast respectively to short glume (P),awned (a), glabrous glume (hg) and green coleoptile (rc), were recongnized;
the difference in each case is obviously due to a single gene. The difference between waxy and waxless is produced by a pair of the basic
alleles Wow and their inhibitors !w-iw. The genotype of the waxless
T. dicoccoides is ww!w!w, and in the F 2 the ratio 13 waxless : 3 waxy
is ordinarily observed. Wand!w show segregation independent of thegenes P, A and Hg. p-p is linked to Rc-rc with a cross-over value of
20.3%. P gene for the long glume of T. polonicum with manifold effects
inhibits in its homozygous state awn length, glume hair, coleoptile color

26

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I

•

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•

and awn color.
Two abnormalities have been observed in crosses between tetraploid
wheats. The first, apparent failure to segregate, was reported by BIF~'EN(1916) in a cross between a white-chaffed T. polonz"cum and a grey-chaffed
T. turgidum. A similar phenomenon involving chaff color and hair was
found by BACKHOUSE (1918) and ENGLEDOW (1920) in the cross T. polonicum
x T. durum. The writer, on the basis of his own finding outlined above,is inclined to believe that these abnormalities arc due to the inhibition of
the P-gene. The second case of abnormality was found in glume lengrh
by ENGLEDOW (1920, 1923) in a cross T. polonicum x T. dururn, who termed
the phenomenon "shift." This abnormality could be explained also by
postulating a major gene P and multiple modifiers. DARLING1'ON (1928)
and WATKINS (1940) consider that these two abnormalities reported ior
tetraploid wheats are examples of the outcome of autosyndesis in polyploid plants. My own experiments, however, have revealed that this is
not the case (MATSUMURA 1950: Jap. Jour. Gen. 25).

5. Cytogenetic Studies on Wheat-Rye Hybrids(Report by Toru ENDO)I

It has often been suggested that the wheat- and rye-genomes havebeen differentiated from a common ancestral genome. This hypothesis
is based on the so-called semi-homology between the genomes of thetwo genera.
It was in 1889 that RIMPAU established the first Triticale strain inGermany. This finding was followed by a number of similar hybrid
strains raised by many investigators who also carried out cyotologicalinvestigations on these strains.
ENDO pursued some cytological studies on RIMPAU'S Triticale, and reported his findings in 1949. These findings are essentially the same as
those by the previous authors. The same kind of abnormalities in meiosis
such as non-pairing and some other irregularities of chromosome behavior
have been observed.
ENDO has suggested that a considerable number of segmental exchanges
have taken place between genomes, as well as within the same genome
during some sixty years since RIMPAU'S work and also that the ryegenome within the wheat plasm may have received some influence fromthe latter.
To examine whether this kind of alteration of genomes might befound, crosses were made in the spring of 1950 between rye and RIMPAU'S
or other Triticale .strains which were originally established in Russia aslong ago as RIMPAU'S time. While many seeds were obtained from suchcrosses, most of them failed to germinate, and only a few seedling27

developed. ENDO is now intending to make cytological examinations ofthese hybrid plants, and also to raise back-cross progeny.

6. Determination of Right- and Left-handedness in the Spikeletof Triticum monococcum vulgare*(Report by Moto KIMURA)

In Einkorn wheat, every ear has two sides. namely, one in which lefthanded spikelets are predominant and the other in which right-handeds;>ikelets are predominant. We can define the "handedness" of the
spikelet by the situation of the first flower viewed from the dorsal eide,
designating it as left-handed if the first flower is on the left side and asright-handed if it is on the right-side. Each spikelet may conform orcontradict, as regards its handedness, the prevailing tendency in the r:ow
to which the spikelet belongs. and the term "concord'ance" or "discordance" is used to designate the propensity of the spikelet.
Assuming that the determination of the handedness is controlled by
the threshold reaction and that external influences as well as the specificpotency are responsible for its determination, the relative values of thepotency 'denoted by Y will be calculated by finding the "probit" of concordance proportion from the FISHER and YATES' statistical tables andsubtracting 5 from it. 1) From the examination of 821 ears, we haveconstructed by this method curves showing the relative values of thepotency in various parts of the ear. It has been found that the determination in any spikelet is not indifferent to that of the state of theneighboring spikelets. If Y.,., Yo
Y oo .x, etc. respectively be thevalues of Y in the xth spikelet (numbered from the lowest one), when
the spikelet immediately below it ((x-2)th), the second one below ((x-4)th),
the third one below ((x-6)th) and so on in the same row, become discordant, it has been found that:
OX>

(1)

Y.x<::Yo.x<:YO(~.x<~.~ . . <Y,,: )where Y x is the value of Y caiculated independently for the xth spikelet.
This shows that the more distant the discordant spikelet becomes, thesmaller is its effect on the decrease of potency. On the other hand, if• Y" is the value of Y in the xth spikelet when the spikelet immediately
below but belonging to a different row (Cr-1)th spikelet) becomes discordant, then>( 2) • Y x< Y x (:1's.14), • Y x Y" (x> 14) .
This fact might be explained by assuming that the tendency of bipolar
polarization as regards the left- and right-handedness gradually decreases.
upwards on the ear until replaced by that of the unipolar one.

* Work by H. KIHARA, M. KIMURA and H. ONO.
28

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•
1

Also it has been found that the influence produced by the simultaneous
discordance in (x-1)th and (x-2)th spikelets is equal to the sum of theeffects produced by the discordance of each spikelet independently:(3)..

.Y."=.Y,,+Y..,-Y,,,2) The relative values of the potency for the determination of leftand right-handedness have decreased as a result of X-ray irradiation inan early stage of the ear development. 3) In the cross between Einkorn
wheat having a high potency and. Emmer (T. turgidum nigrobarbatum)
wheat having a low potency, it has been found that the relative valueof the potency in F 1 is approximately equal to the mean value of thepotencies in the two parents.

H.

GENETICS OF FERTILITY AND SEXUALITYIN PLANTS(Report by YO TAKENAKA)

1.

•

Sterility of Triploid Plants

Among the wild and cultivated triploid plants which have been collected
and examined cytologically by the writer up to this time, there havebeen found four types to be distinguished on the basis of the degree andform of sterility. Of these types three give a few seeds either 'by naturalpollination, by water culture of the cut flower-stem, or by crossing with
its ancestral diploid, but the last type never gives seed by any treatment,
as far as I have been able to determine.
a) Hemerocallis typeThis type has been found in the triploid plants of the genus Hemerocallis in which the cross inter se and also the cross 'with diploid plants
including their reciprocals are perfectly sterile.
,
The sterility may be due to some of the following causes: (1) irregularities in meiosis, (2) exceedingly short life of each flower, which is
insufficient for allowing the style to wait for the delayed maturation of
the embryosac as is usual in triploidy, (3) a part of the embryosac, mainly
the egg nucleus, often becoming necrotic during its development, as is
usually the case in polyploid plants. (4) It is also possible that theirancestral diploids have already perished, or even if they are present, thephylogenetic relations between the assumed diploid ancestor and thepresent triploids have become too remote in the course of evolutionary
process.
Among the plants cytologically investigated by the writer, Daphne
odora THUNB., Crocus sativus L. and Zingiber Mioga Rose. (pentaploid)
probably belong to this type.

29b) Lycorz"s typeThis type is found in triploids of the genus Lycorz"s, such as L. squamz"gera MAXIM. and L. radz"ata HERB. Although these plants apparently
are completely sterile in the.field, a quantity of seed can be gathered by
the water culture of the cut flower-stem.
The sterility of these triploid plants under natural conditions is probably
due to one or both of the following causes: (1) irregula\I" meiosis, (2) asituation such that ferltilization may occur during the long life of eachflower, but the proportion of fertilized embryosacs is too small to produce
a ripe capsule, because of a struggle for nourishment between the fertilized embryosacs and the subterranean bulb, in which the few fertilized
embryosacs fail to compete successfully with the bulb.
c) Lz"Hum type
In L. tz'grznum KE:R-GAWL, hardly any ripe seeds can be secured either
by self-pollination or by intraspecific hybridization; only the hybridization with pollen of L. Maxz"mowz"czz"z" REGEL. (diploid species) gives somegerminating seeds, while the reciprocal cross produces none.
As the cause of this phenomenon may be assumed: (1) irr~gular meiotic
division; (2) the life of each flower beip.g so long as to permit embryosacs
or ovules in the flower to be capable of maturing though in delayed
condition; or (3) the presence of the ancestral diploid species. Accordingly, only a small proportion of such a cross gives some seeds.
d) Crocus vernus typeAs is seen in the triploid garden varieties of Crocus vernus, same seeds
may be obtained by natural pollination.
This rather high fertility for a triploid plant seems to be due to either,(1) the meiosis being less irregular than that usually found in othertriploid plants, or (2) the genome of these plants consisting of several
subgenomes, so that the plants are actually polyploids of a higher orderthan ordinary triploids.
.
The comparatively long life of each flower may also be a contributing
factor.
2.

Sterz"lz"ty and Autotoxz'cosz"s of Colchz'cum autumnale

Colchz"cum autumnale is generally found to be a sterile plant in the
middle and southern parts of Japan, Its somatic cells contain 38 mitotic
chromosomes, as shown by LEVAN. These chromosomes are not only ofvarious sizes, but also show some features resembling fragmented or
attached chromosomes.
At ordinary temperatures, the maturation division of PMC's shows
not only swelling, stickiness, abnormal conjugation and abnormal distribution of chromosomes, but also delayed nuclear division, and in somecases, even the dissolution of the nucleus. Especially, the formation of

30

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•

•

•
•

•

the cell plate is very incomplete or does not occur at all. Accordingly,
monad and dyad microspores in3tead of tetrads, are found in a large
majority of cases. In extreme cases, very small microspores without any
chromosomes are occasionally found. These abnormalities are strongly
suggestive of the change found in cells after the colchicine treatment
for a rather long time or at a high concentration.
, In material cooled at SoC for 24 hours, however, the abnormal chromosome features and behaviors mentioned above become less frequent,
the most remarkable sign of recovery from the abnormality being theformation of a cell plate in the phragmoplast of PMC, which is probably
already in the tetrad stage. Thus, low temperature apparently weakens
the effect of colchicine on the plant.
In material which had been kept for the first 2 days at about lOoC,
and for the following 2 days at about 20°C, the maturation division
was apparently nearly normal. However, many C-pairs appeared on the
nuclear plate probably owing to the effect of colchicine toxication received
before being subject to low temperature.
This phenomenon probably shows the threshold of the colchicine effect.
Almost all the pollen grains formed at either ordinary temperatures,or the low temperature mentioned above belonged to the monad or dyad
type, the fragmentary spores being probably devoid of any chromosome3,
chromosome fragments or micronuclei.
Thus, it has been demonstrated that the meiosis becomes very irregular
jn Colchicum even under the climatic conditions at Misima, and that theirregularities differ significantly from those observed in hybrids or heteroploid plants and those produced by high- or low-temperature treatment.
Rather, they seem to be analogous with the meiotic irregularities whichappear under the influence of colchicine-treatment.
Under the influence of high temperature the swelling of cells occurs,
which induces some change in the nature of the nuclear membrane, so
as to allow colchicine molecules to penetrate through the cytoplasm into
the nucleus and to combine with the polypeptid chains of the attractoplasm. Accordingly, the deleterious effect characteristic of colchicine
toxication results, as well as abnormal features and chromosome behavior
jn nuclear and cell division.
Thus, Colchicum autumnale apparently becomes a sterile plant as aresult of the relatively high atmospheric temperature which is prevalent
jn the middle and southern parts of Japan.

3. Karyotypes and Sterility in Hemerocallis
Under the title, "Karyologicai studies in Hemerocallis (1929, Cytologia)"
1 reported many years ago my findings of the cytology of many species()f day lily. Further karyological and genetic studies on some other31•

materials are under way.
So far, karyotypic analyses have been carried out on 28 species, varieties, forms and garden varieties of this genus. The fundamental chromosome number in this genus is 11. These chromosomes can be classified
into seven distinct types according to their shapes. All the karyotypes
investigated contain either some or all of these types in various combinations.
The taxonomy of this genus is considered very difficult on account of
the variability in their morphological characters, which obscures thespecific distinction. The karyotype characters show the same kind ofvariability.
All triploid plants in this genus show complete sterility not only in
self- and sister-pollination, but also in crosses between different species.
or varieties including many diploids.
This sterility of the triploid plants may be largely attributed to irregularities in meiosis which result in the formation of incomplete pollen
grains and embryosacs. The embryosac, especially, is considerably
delayed in maturation, and frequently contains necrotic tissue.
~- As for chromosome numbers, on ly diploid (2n =22) and triploid (2n = 33)
numbers have been ascertained. SrouT (1932) has assumed 6 to be thebasic chromosome number. As stated above. I hold a different view,
based on my own observations on karyotypes and chromosome behaviorin PMC's of a large number of diploid and triploid plants.

••

4. Studies on Sex-differentiation in Higher Plants
From 1938 to 1943, experimental studies were conducted on sexual'
characters in higher plants, in the cases of colchicine-induced polyploidy,
aneuploidy and other chromosome aberrations with various dioecious
plants including Rumex acetosa 1. as material. The preparations andrecords used in this study were left in the Biological Institute of Keijo(Seoul) University in Korea, where I had a position. As there is no hopeof restoring them at least for the time being, I wish to give here abrief note of my observations based on some fragmental records at hand.
Among the crosses between tetraploid females induced by colchicine
treatment and normal diploid males of Cannabis sativa, one strain
contained among the offspring 7 females, 5 intersexes and 2 males. Thechromosome constitutions were 3A+3X in the female, and 3A+2X+Y
or some of its derivativ~s in the intersex, as well as in the male. BothYAMADA'S finding (1943) that the Y-chromosome was larger in size thanthe X-chromosome, and W ARMKE and DAVIDSON'S assumption that theY-chromosome influences the appearance of male characters, are considered to be largely correct.
32

4

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,

In the crosses between tetraploid females induced by the same treat·'ment and original diploid males of Humulus japonz"cus, one strain gave
'Offspring consisting of 18 females and 7 intersexes. These females andintersexes showed respectively 3A+3X and 3A+2X+ Y 1 + Y~ in their'chromosome constitutions. This observation agrees well with ONO'S:assumption (1941) that the sexuality of this plant is determined mainlyby the ratio of the X-chromosome to the autosome. Similar studies
were conducted on Humulus Lupulus, H. Lupulus var. cord~roUus, Asparagus offidnalis var. altz"lis and Spinacia oleracea.
The same kind of experiments were resumed in 1950, with the hopeof discovering the relationship between chromosome- or choromosome-set
aberrations and sexuality in various dioecious plants, by using the colchidne and radiation methods, and to reexamine my previous findings andthe results obtained by other workers.
5. Sterility of Phaseolus multijlorus under High Temperature(Report by Akira MIYAZAWA),

Phaseolus multz"florus commonly bears fruits in cool regions in Japan,
but in milder regions the yield is much lower. This fact suggests in'Complete development of reproductive organs in this plant as a result ofhigh temperature. It has been found that the flower buds which pass
the metaphase stage of the first maturation division from the end of
August to the middle of September, bloom in about 14 days. Measurement was taken of the size of pollen grains during the period from
September 2 to October 11. It has been found that the pollen grains
range between 35 and 70 fl. in diameter. The percentage of empty pollen
grains amounted to about 20 percent of the whole number examined
under the micr03cope on September 2. The size of such pollen grainsvaried from 40 to 70 fl.. The percentage of empty grains gradually fell
when the season advanced, to 3.9 percent on September 18, and 2.3 percent
on October 1. The percentage fluctuated to some extent, but never
became as high as in the first part of September. The range of fluctuation in size also declined significantly. These facts become understandable,
if correlated with the change in the maximum temperature during theperiod. It may be surmised that those poHen -gftlins which included
the highest percentage of empty and variable (in size) ones, had beenin the maturation stage when the temperature was as high as 31°e.

33

J.

GENETICS AND CYTOLOGY OF SOMEFLOWERING PLANTS
1.

Genetics of Flowers in Zinnia elegans(Report by Kanji GaToR)

The capitate flowers of Zinnia elegans are highly variable in respect
to the number of petalous florets on the head. Between the two extremes,
i. e. the completely single flower and the completely double flower, there
are many transitional types which are difficult to assign to clear-cut
classes.
This experiment has been undertaken to make clear the genetic behavior of petalody in Zinnia elegans. In 1950, several crosses between
various flower types as well as selfing experiments were made.
2. Genetic Studies on Solanum melongena and Capsicum annuum(Report by Kan-Ichi SAKAI)

The principal object of the present investigation is twofold :-first,
the study of the quantitative-character genes and second, the study ofheterosis. A number of selfings and inter- and intravarietal crossings
within each of the two species were made during the past year, andthey are now in the process of being grown preparatory to further investiga tion .
3. Karyological Studies of Lycoris radiata*(Report by Hitoshi KIHARA)

Many individuals of the wild plant, Lycoris radiata, have been collectedfrom various localities in Japan for morphological and ecological investigations. The chief purpose of this study is to find out the karyotypicaldifferences among those individuals which, if present, would be originatedthrough somatic mutations, since this species constitutes a single clone.
The results so far obtained show that the chromosome complement of
this species consists of thirty-three i-shaped chromosomes.
From the pollination by its own pollen as well as by the pollen of L.
sanguinea (2x), many ripe seeds were obtained, which were then sownfor the purpose of obtaining diploid individuals.

* Work by H. KIHARA and F. LILIENFELD.
34

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,

•
4. Cytogenetic Studies on the Genus Nicotiana(Report by YO TAKENAKA)••

Although tobacco plants have long been investigated by plant breeders
in Japan, cytogenetic studies are very few. The writer has been carrying out cytogenetic studies on the genus Nicotiana since the summer of
1950.
The number and behavior of chromosomes in the meiosis of PMC's
in 5 wild species have been studied. Of these N. plumbaginzfolz"a, N.
repanda, N. glauca and N. sylvestris, have respectively 10, 24, 12 and 12
gemini, and all of them show completely regular meiotic cycles. One exceptional species among the five, N. glutinosa, shows some irregularity,
i. e. a sign cf secondary association among its 12 paired chromosomes.
The chromosome number and behavior of 3 artificial tetraploids, inducedby the colchicine method at the Hatano Tabacco Exp. Station in 1944 andtransplanted into our garden last spring, were also studied. The tetraploid plant of the 'Odaruma' variety of N. tabacum was found to havecompletely reverted to the original diploid state at some time during the
7 years of cultivation. The tetraploid plant of the 'Nambu' variety of
N. tabacum has returned to forms with various chromosome numbers
ranging from approximately tetraploid to diploid. The amphidiploid plantfrom N. plumbaginzfolia x N. alata showed 19 bivalents in the 1st meiotic
division, just as expected, and some irregular distributions in the 2nd
anaphases.
'

5. Some Effects of Sodium Ribose Nucleate on Somatic Mitosis
of some Plants(Report by Kanji GOTOH)

Detailed interpretations of the so-called reductional-type divisions induced
by sodium ribose nucleate in tissues of plants have already been given
by HUSKINS (1948) and WILSON and CHENG (1949) on Allium cepa and Tril_ _ _ _---::lium species.
According to these authors, in the reductional-type divisions eachmember of the homologous chromosome pairs goes towards opposite poles
in most of the cases. The present experiment was undertaken to reexamine the findings by these workers, and to clarify the mechanisms ofsynapsis and reduction in mitosis.
Root tips of Tradescantia paludosa (2n =12) were treated with 1 % sodium
ribose nucleate solution for 12-24 hours, and w~re fixed in NAVASRIN'S
fluid. Sections were cut 12-18.u in thickness and stained with ·HEIDENRAIN'S iron-alum hematoxylin.
The results obtained are as follows:• .

35

1.

In a few cells two separate chromosome groups, each with six
shortened chromosomes, the number corresponding to those of the
haploid set in the plants, were found lying not far from each other,or a group of twelve chromosomes about to divide into two groups.
In a few other cells two clusters of chromosomes were found.
These seem to have resulted from the separation of these chromosome
groups.
2. In a very few cases abnormalities in the arrangement of chrosomes were observed at the metaphase. Further experiments are in
progress.

K.
1.

•

STUDIES ON LOWER ORGANISMS

A Method of Meseasuring the Rate of Growthof Nuerospora sitophila(Report by Taro ITO)

For the investigation of growth of this mold in response certain
amino-acids or vitamins, it is important to find an exact method of
measuring the amount and rate of growth.
At present the following methods are commonly used. First, the testtube method (BONNER'43); this consists in observing the growth responseof the mold to substances added to the medium. Second, the dry weight
method (HOROWI'i'Z '46), of weighing mycelia cultured for 55 to 72 hours
and dried at lOOoe. The mycelia are killed at this stage, and there isno subsequent growth. Third, the horizontal-tube method, which consists
in measuring the rate of elongation of mycelia along a glass tube duringa certain time. By this method, only the mycelia which grow in contact
with the inner wall of the tube can be measured, so that precision can
not be expected. All these methods have some defects, and I have devised
the following technique, in order to meet the two conditions; first, tobe able to weigh at some regular intervals. and second, to secure thesubsequent development of the organism.
A glass tube 8 em long of known weight, is stuffed with cotton at
both the upper and lower ends and sterilized at 120 o e. The mycelia areinoculated into the lower part of the tube, which is inserted in a testtube containing liquid media. For measurement, the glass-tube containing the mycelia is drawn out of the test-tube, and is weighed after being
dried in a desicator for 10 minutes.
By this method, I have investigated the growth of N. sitophila inresponse to the basic medium to which various quantities of two ammonium salt, ammonium tartaric acid and a. nitrate, have been added.
Thus, it has been estimated that the maximum growth occurs at CjN=
36"

12.2, and that the lower the concentration of these substances, the greater
is the rate of growth in the early stage of culture. It has also beenshown that, when a medium inoculated with the mycelia, left from 32
to 64 hours, and filtered to remove all traces of mycelia, is used for the
new culture of mycelia, the growth is accelerated as compared withcases in which fresh medium is used.
2.

•

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Studies of the Virus Infecting Silkworm(Report by Mitsuo TSUJITA)

a. The Virus Particles and their Growth
The virus particles used in the present study were obtained by theultra-centrifugation for from 1.5 to 2 hours at 35,000 or 40,000 R. P. M.
of the polyhedra-free biood of jaundice-diseased si kworms. Electron
micrographs show that the diameter of these particles ranges from 10
to 30 m!~. To clarify the formation process of the so-called polyhedral
body, living material and fixed preparations of the tissue of the diseased
insect were observed. In the first stage a number of minute particles
appear. These grow into definite-shaped particles, which aggregate
into poiyhedral bodies. In the couJ:se of this process, diplococcus-like,
ellipsoidal or bead-shaped bodies are formed-these seem to be the forerUnners of the polyhedral bodies.
The dipiococcus-shaped or ellipsoidal bodies emerge from th..:. polyhedralbodies, when these bodies are dissolved in a dilute (0.3-0.5%) solutionof alkali (sodium carbonate).
A great number of particles emerge from the diplococcus-shaped granules
which are enclosed in a proteid membrane when the granules are dissJlved in a diiute aikali solution. It has been found by electron-microscopic examination that some of these particles are square-shaped rather
than spheroidal, with diameters varying from 50 to 100 mt!; there are also
particles of linear shape, with length varying from 300 to 350 mt! andwith width from 50 to 100 mt!. In some cases these particles emerge
directly from the polyhedral bodies when the latter are dissolved in adilute alkali solution.
Inoculation experiments have shown that the particles have a strong
virulent capacity, suggesting that they represent a developmental stageof the virus in the cell.
The facts mentioned above show that the so-called polyhedral bodies
are apparently made up of 3 components: i) an outer envelope, ii) noninfectious protein and iii) intermediate bodies which contain a numberof virus particles.
It seems that the non-infectious protein which constitutes the greaterpart of the polyhedral body is a product of the intermediate bodies.

37

b. Resemblance between the Function of Genes and the Activity ofInsect Virus
We have been able to observe under the microscope polyhedral bodies
of various shape in the cells of certain lepidopterous larvae suffering fromjaundice disease. These polyhedral bodies are known to be formed byvirus. In the silkworm hexagonal inclusions are formed. In the larvaof the tussur, Antheraea prenyi, infected by virus, pentagonal, squareshaped or triangular bodies are formed in the tissue. In the Eri-silkworm,
Attacus· ricini, similarly infected, granular bodies are formed instead,while in the larva of the moth Ivela auripes, comparatively large triangular bodies appear. Inoculation experiments with these four kinds ofviruses have shown that the shape' of the polyhedral bodies is largely
determined by the kind of virus, regardless of the host. It is accordingly clear that the different shape of the polyhedral body is due to thevariety of the virus itself.
It seems to me that this fact shows the similarity of the activity ofvirus to the activity of the gene. For example, the shape and natureof the pigment granules in the cells of the Malpighian tubules are controlled by the nuclear gene. After examining various races of thesilkworm, Dr. KIKKA w A and I have classified the four types of pigment
granules in the Malpighian tubules as shown in the table.

••

Table II.
Four types of pigment granules in the Malpighian tubule cells
in the silkworm
Nature of pigment granule
Type

Race
ColourShape

1st typedense yellow

granular,diameter 1--:5 p.

KH 19, KH 21

2nd type

yellowneedle- or
rod-shaped; Normal strain, W2' bW2
and some eye- or eggcolour mutants

3rd type

yellow

010,
011,
No. 106

4th typ~almost colourless

granular,diameter 1-3 It
minute granularwithin 1/L~-

Japanesed-oily, a-oily

My cytological observations have revealed that these pigment granules
are the derivatives of mitochondria. Moreover, these mitochondria seemto arise from minute granules in the cytoplasm. Probably, under thecontrol of a nuclear gene, these minute granules are formed by the selfduplicating particles in the cytoplasm, and develop into coarser granules
or filamentous substances to be called mitochondria. In this manner
the formation of the polyhedral body by virus simulates the mode offormation of pigment granules in normal cells.

38

•

•

L.
1.

IMPROVEMENT OF SOME USEFUL PLANTS

Improvement of Sugar Beet by Means of Induced Triploidy(Report by Seiji MATSUMURA)••

Improvement of sugar beet by means of induced p0lyploidy, has been
continued since 1941 in the KIHARA Institute for Biological Researchunder the guidance of Prof. H. KIHARA. Recently, this work has gained
the cooperation of the National Institute of Genetics, the Laboratory of
Plant Breeding of the Hokkaido University, the Hokkaido Agricultural
Experiment Station and the Japan Beet-Sugar Manufacturing Company.
The maximum yielding capacity of autotetraploid sugar beets is hardto estimate accurately, because of the late maturity of 4x plant espec.ially since unfavorable weather conditions during the late period of growthor an inadequate term of cultivation greatly influence their yielding
capacity. It seems thus reasonable to estimate the yield of 4x plant
sometimes lower than that of 2x (MATSUMURA, MOCHIZuKI and SuzuKA
1950: Rep. KIHARA Inst. BioI. Res. 4). On the other hand, 3x beets are
more vigorous, grow better, and always show a higher yield than theother beets, according to the experiments conducted in the Hokkaido
Agric. Exp. Station during the years 1945-47. In 1949 and 1950, comparative studies on 2x, 3x and 4x sugar beets were carried out on a large
scale in various districts in Hokkaido. The results show that, as expected,
the 3x beets are superior to either 2x or 4x. It is further expected thatthe intervarietal 3x hybrids, for instance, between the high yielding 4x
variety and the disease resistant 2x, will bring still better yields. However, the inferior germination rate of 3x seeds comes into qu~stion; thisshould be solved as soon as possible.
For a large scale production of 3x seeds, the self-incompatibility ofthis plant can be utilized. If 2x and 4x beets are planted in alternaterows, 3x seeds can be obtained through natural pollination either fromthe 2x or 4x plants. Microscopic examinations of root tips have revealed
that about 30% of the progeny raised from seeds of the 2x motherplants are triploid. Also, the seeds from the 4x mother plants produce
more than 70% 3x plants. The frequency of the 3x plants, however, isvaried to some extent by some factors, such as the ratio between the
2x and 4x plants, the difference in blooming season, the difference inpollen-tube growt h rate of the parents (MOCHIZUKI and MATSUMURA 1950 :Rep. KIHARA Inst. BioI. Res. 4).

39

2. Genetics and Breeding of Citrus Plants(Re{:ort by Kazuo FURUSATO)

a. Cytological observa~ions
In order to elucidate the relationships between various Citrus species,
chromosome numbers, chromosome behavior at meiosis, and karyotypes
are being investigated. Most of the Citrus species are diploid with 9
gametic chromosomes. Some are tetraploid with 18 gametic chromosomes,
or triploid with 27 somatic chromosomes.
b. Polyembryony and comparative studies of individual nucellar
embryo plants
The phenomenon of polyembryony is very common in Citrus. My task
has been to elucidate this phenmoenon in various hYbrids. Methods havebeen devised to make possible the growth of all embryos. The work is
still under way.
c. Albino seedlings
In Citrus a certain percentage of albino seedlings usually appears,
sometimes amounting to 10 percent or more. Almost all of them are ofnucellar origin. Cytological observations and experiments are under way
in order to explain the development of these albino seedlings.

•

3. Induced PolYPloidy in Useful Plants
By colchicine treatment tetraploids of some useful plants such as watermelon, radish, sesame and others were obtained. Morphological andphysiological studies have been carried out to examine the possible utilityof these tetraploids.
a. Tetraploid water-melon
A tetraploid water-melon plant produces triploid and tetraploid progeny
when it is pollinated by the mixture of pollen-grains of tetraploid anddiploid plants, in the ratio d about 90 percent triploids and 10 percent
tetraploids. Probably this ratio results from certation between haploidand dipoid pollen-grains.
b. Grafting of polyploid water-melons
In preliminary experiments young buds of a diploid water-melon plant
were grafted on diploid, triploid or tetraploid stocks. The grafts on
triploid stocks grew more vigorously and gave higher yields than thegrafts on tetraploid stocks. Grafting of buds of diploid, triploid or tetraploid water-melon on calavash stocks gave very high yields. These
results show that the grafting of tetraploid buds on calavash may beutilized for producing triploid seeds for commercial purpose. Such grafts
give much higher yields than ordinary tetraploids.

40

•

•

c. Triploid water-melons of. different combinations of varieties
Attempts.are being made to get triploids of various combinations ofvarieties of water-melon from tetraploids. In 1950 tetraploids of thevariety Kaho were obtained, and from them triploid plants were bred.
d. Triploid fruit trees
Attempts are also being made to obtain triploids of various kinds offruit-trees, in the hope of getting seedless fruits.
e. Polyploid forage, green-manure crops and vegetables
Tetraploids of some leguminous plants such as lupin and alfalfa, as
well as of fodder radish, have been obtained. Also tetraploids have beenraised for some vegetables like carrot, edible burdock, Luffa, calabash, etc.

M.

THEORETICAL GENETICS
1.

Semiallelic Genes(Report by Taku KOMAI)•The relation between two chromosomal genes falls into one of the following categories :-They may be located either, a. on different chromosome sets, or b. at different loci on the same or homologous chromosomes,
or c. at the corresponding loci of the homologous chromosomes. In thefirst case one finds in the subsequent generation independent assortmentof the genes; in the second case, linkage between the genes; and in thethird case, segregation of the genes. In some cases a state between theabove band c is seen. These are semiallelic genes. The characteristicsof this kind of genes are: 1. They have similar but distinct phenotypes;
2. they are arranged in direct sequence, with no other intervening gene;
3. when they are located on homologous chromosomes, their manifestation
is partially covered: 4. they never or very rarely recombine. Examples ofsemiallelic gene sets can be given from various animals. Especially, thegenes for the human blood-types, A-B-O, M-N-S, Rho-Rh'-Rh", probably
beJong to this category. If so, the rare and much disputed recombination
cases should be admitted to be possible.
(cf.

KaMAl T. American Naturalist 84: 381-392, 1950)

2. On Variance due to Competition between Plants of Different
Types in Plant Populations(Report by Kan-Ichi SAKAI)

Plants of two varieties of rice, apparently different in plant color,
tillering and grain yield were mix-planted in a difinite manner to examine
the effect of neighbors of different types on the weight and number ofpanicles per plant and the mean weight of a panicle, in each of the two41

varieties. The experiment was conducted by the randomized block desIgn
with six replications. Results of the analysis of variance showed thedifference between varieties to be statistically significant so far as theweight and the number of panicles per plant were concerned. So wasalso the difference due to the interaction between variety and treatment
regarding the total weight of panicles per plant and the mean weightof a panicle. That is, the weight of panicles per plant, and the mean
weight of a panicle are significantly affected by the neighbors. resulting
in phenotypes respectively exaggerated or arrested as compared with the
expectati0n from the genotype.
The variance in hybrid populations of plants may be due to either
genetic or environmental causes. It has generally been accepted that thelatter in segregating populations is identical to that in the parental or F 1
population. But the experimental results demonstrated above suggest thatthe variance in segregating populations comprises not only the environmental element in the usual sense, but also the element due to thecompetition between plants of different types. The theoretical estimationsof the latter element are to follow.

•( 1) Variance due to competition within population after
backcrossing F 1 to parents
In case of monohybird segregation, the sum of all increments in both
backcross populations, which one plant will receive thrcugh competition
with n plants surrounding it, will be,'))'It -' t

2.: C,,2 + ~

h~~l

'1'~

~

n

'U -1

CiCj + 2.: a,,2 + 2.:-l=l,j=t+i

h=l

?1

2.: a:iaj(1)'i=l j=l+'iwhere C or a stands for the increment given to the plant through competition with the homozygous (AA or aa) or the heterozygous (Aa) hth
individual.
If the plants are singly-spaced in rows, the variance exhibited by eachplant due to competition with the surrounding eight individuals will be,t:T,,2(BP1 +BP2 )=(c2+a2)K ..........•...... (2)where

K=3k I 2 +3k2 2 + 10k.?+4k 1k"+8k 1 k 3 +8k"k:;,k 1 and k 2 are the constants given by the distance between hills and rows
respectively, and k" those given by the distance between two neighboring
plants at diagonal positions of the rectangle formed by four plants on
two adjacent rows.
If r pairs of alleles (Aa, Bb,, Rr) are concerned,
22
t:T c (BP,+BH) = (ca +c,P++Cr 2+ a,,2+ a,? + ." .+ar 2)K ... (3)LetC,,2+C. 2+ .... +cr 2 be C, and a,,2+ ab2+ .... +ar 2 be A,
t:Tc 2(B)=(C+A)K ...•...••...••••.... (4)42•

(2) Variance due to competition within the population of
successive hybrid generations of self-fertilization
In F2 , F 3 , and F.. + 1 hybrid populations, it may be computed as,u c 2 (F2 ) =( c+ ~ A)Ku c 2(FJ )=(5),~ ( C+~A)K(6),

u c 2 (F.. + 1 )= ~:..-_ ~ (2 n C+ A)K

and

•

(7).

(3) The mean of the variances due to competitionin different F J families
These are computed asuc~(F3 families) = ~ ( c+ ~ A)K(8).

Variances caused by competition between plants of different genntypes
in segregating populations as computed above shOUld be taken into accountin the analysis of variance of hybrid populations in biometrical genetics
as well as in the study of population genetics of autogamous plants.

3.

Theoretical Studies on Individual Selection in Plant-breedingwith Special Reference to Gene Recombination(Report by Kan-Ichi SAKAI)•

It is now well accepted that the quantitative characters of plants andanimals are often controlled by a large number of genes distributed ona definite number of chromosomes. Since a majority of economic characters of plants, such as, grain yield or time of heading in cereal plants
are considered most likely to be due to genes of this kind, it. will be ofinterest to plant breeders to give some theoretical considerations on theeffect of recombination on the efficiency of individual selection in autogamous plants.

( 1) Proportion of homozygotes in segregating hybrid populations
Assuming that two genes are located on every chromosome, and theaverage recombination value between the two is a, the proportion of all
homozygotes in any segregating. generations may be represented as2"-1+ 1 [a 2+(l-a)2{-2n~1 + _. 2---

]n}R

(1),

where R is the number of pairs of chromosomes and n the number of thesegregating generations.
43

(2) Selective efficiency of dominant alleles in segregating

hybrid populations
If we call the proportionate number of dominant homozygotes al10ng
selected phenotypic dominants the 'selective efficiency', then, i) the selec~
tive efficiency (Er ) of dominant homozygotes in the case of independent
alleles may be expressed by the equation,2"-1

E r == (,2"+ 1

)r

(2),

where r is the number of pairs of independent alleles. ii) The selective
efficiency (E R ) of dominant homozygotes in the case where these two
respective genes are linked' on every chromosome in the repulsion phase
may be computed by the equation,
2)R
E R =={1-1+zn- 1[l+AI"--1.2(1-A;")/(1-).2)] J•••••• (3),where
,1.1= ~ [(l-a)2+ a 2] ,A2=~ [(l-a)2- a 2] ,and R stands for the number of pairs of chromosomes.
( 3) Interrelationship between the number of phenotypic
dominants and the selective efficiency
In breeding work, it is possible for the breeder to know the frequencyof occurrence of desired dominants in the hybrid populations. Hence itwill be of much interest to him if he can estimate the selective efficiency(E) from the number of dominants (D) which he desires to select andbreed. This estimation can be given from the solution of the following
equations :-i) In the case of independent alleles, it may be expressed as.
,

E==DK,

where

(4),

,2" 1)

log,o( 2"+i
K,==

(1

l'

log,o ,2 + 2"+1 )ii) If each two dominant alleles are linked on every chromosome in the,repulsion phase, it may be expressed as
E=D K 2where

44•••• , • • • • • • • • • • • • • • • • • • •

(5),

•

•

and

,

(4) The size of hybrid populations for the individual selection andthe effective number of individuals to be selected
The number of dominant individuals to be selected in order to include
at least one homozygous dominant, the maximum frequency of failure to
be 1/100, will be computed by,n> lo~:'(IO'01)
or

."

(6)

log!o 0 . 0 1 ·(6')
L.log!o (l-DK) . . .. . . . . . .. . . ... ...
•n-'"and the size (N) of the population should be-.., IQg'loO.Ol_
N L.log1o (I-DE)

or

N-..,~g!O O.()I
L.log (I-DK+1)
1o.
..................

(7)

(7')

•

All the formulae given above will help us, in the practice of plant
breeding, in deciding the best time for exerting the individual selection
out of the successive hybrid generations, and the number of individuals
to be selected, as well as the size of the hybrid population on which
the individual selection will be made.

4. Effect of Random Fluctuation of Selective Value on the Distributionof Gene Frequencies in Natural Populations•(Report by Moto KIMURA)

Recently, in the field of population genetics, there is a sharp antagonism
between WRIGHT on one side and FISHER and FORD on the other. WRIGHT
maintains the evolutionary significance of the "WRIGHT effect", while
FISHER and FORD reject it, and emphasize the extreme importance of therole played by natural selection. The present study is to examine thisantagonism by means of precise mathematical analyses.
The long term distribution of gene frequencies in natural population
under influence of the fluctuation in the force of natural selection (Proposed by FISHER and FORD) can be calculated by
cp(x)=(C/V6x)exp {2~(M6X/V6X)dx},

which is a modified expression of WRIGHT'S formula (1938, '39, '42). HereM 6x and V 6x respectively denote the mean and the variance of(rate ofchange in gene frequency per generation) for a given gene frequencyx.

ox45

If there is no dominance, the formula for the long term distribution
becomeswhere
A =-{s/(2NV.)+A1U-A2V}/(Al-).2) ,B= {S/(2NV.)-AIV+).2U}/(AI-).2),and ).1 and ).2 are respectively the positive and the negative root of thequadratic equation:x2-x-I/(2NV.) =0 .
In this formula, N is the effective size, U and v are respectively themutation rate of the gene A to and from its allele A', ~ and V. arerespectively the mean and the variance of the selection coefficient (s) ofA, and C is a constant such that
\ Itp(x)dx =1 .

•
•

"0

On the other hand, for recessive genes, if the selection coefficient of
the recessive individuals is t, the distribution can be given by
tp(x) = Cx~Nv-l(1_x)4NU--l(Al_X)4N.A -1(X-A2)4NB-lx [(x_a)2+pr ND - l eXP J -8NEtan-- l x-a}
[3

l

where
A = {t/(2NVt Al)+(U+V)Al-V}/(3-4A. 1) ,B={l/(2NVt A2)+(U+v)A. 2-v}/(3-4).2) ,D=(ab+a lb/)/(!J2+b'2) ,E=(alb-ab/)/(b2+b'2) ,and
a=-A1A2at+(u+v)a-v,
a ' = A1A2i~t+(u+v)[3,b=3-4a,b'=-4l~ .
In this formula tan-- l X is defined by·x

tan-1X=\ (l+X 2 )-ldx,-"0

and Ah A2 and a±[3i are the four roots of the quartic equation:x4-x3 -1/(2NVt ) =0,-of which Al is real and larger than 1, ).2 is negative and a+ [3i and a-I~iare both imaginary. From the study of the distribution curves it has
become clear that although the effect produced by the randvm fluctuationin natural selection is relatively unimportant for small populations, it has.a remarkable effect for large ones for the case in which the frequency-of either allele is low (where dominance is lacking) or the frequency of
the recessive gene lies inside a certain range of high frequencies (in the46•

case of complete dominance). Therefore the biological importance of the"WRIGHT effect" should be judged by the frequency distribution of theeffective size (N) of populations in nature, as well as of the quantity
NV.. which is the product between the effective size and the variance of
selection coefficient, for alleies contained in them.
ef. KIMURA, M. 1951. Effective Size of Populations and Random Fluctuationsof Selective Values (in preparation).

5.

Physiological Theory of Polygenic Actions•,•(Report by Mota KIMURA)

A polygenic system may be constructed by a group of genes, if eachof them controls the different parts of the physiological reaction chain
leading to the expression of a given character, and the effect produced bysubstituting one of them by its allele is so small that it can be easily
counterbalanced by the environmental variation. To explain the mode ofaction of these genes, the writer has attempted a mathematical analysisof the chain reactions, using the model adopted by WRIGHT (1934), in
which the catalytic transformation of the substance takes place by steps
to attain the final product, the quantity of which corresponds with thecharacter in question.
Let the chain be constructed by the linear linkage of n-l reactions
each of which is assumed to be monomolecular, and let Ph P 2 , •••• , p ..- 1
be respectively the amount of intermediary substances transformed in serial
order, and Po" be that of the final product. If the concentration of the
enzyme contributing to the change: P,-+Pi + 1 is e; and that of Pi-+Di 1(other destructive change) is ft, the relation:
,",-1

P"oc

n {k;ed(a+k;ei+lift)} (ki , Ii : constants) .(1)i=l,

is obtained under the assumption "that the amount of intermediary substances increases at a given rate a( ==(dPddi)/Pi ), keeping a constantratio among one another. If it is assumed that the concentration of eachenzyme is proportional to the dose of genes (AA, Aa, aa) controlling it,
many important results obtained by experiments of previous authors (e. g.
arithmetic effect, geometric effect, dominance relation etc.) become understandable:
For example, if all plus genes accumulate in the larger parent and alI
minus genes in the smaller parent, the concentration of the enzyme in agiven link of the chain may be represented as e+2.de, e and e+.de forthe larger parent, the smaller parent and the F 1 individual respectively,
so that the measure (Q) of an F 1 individual approaches the geometricmean of those of the two parents.
Qt =,/(QQ')+€,

47

(2)

where e is a small and positive quantity.
The dominance relatbn between .alleles may also be explained by using
the relation (1).
In the studies of biometrical genetics, "transformation of scale" is often. indispensable, and it is adopted through empirical tests. The study presented -above gives the theoretical basis of such transformation.

6. RecombinaNon of Chromosome Segments through. Continued Self-ferNlization(Raport by Moto KIMURA),

In this study the writer has mathematically investigated the processof the recombination of chromosome segments which takes place generation by generation through continued self-fertilization, and he has derived
several formulae regarding the frequency distribution of the length of theheterozygous segment, the rate of increasing homozygosis, the numberof effective factors and the distribution of segments finally attained in
the population.
Let AAI be an initial pair of homologous chromosomes differing inmany alleles as in the case of the species cross, and designate the lengthof homozygous segments derived from A and AI as IOOL A and IDOLA' units
respectively, and that of heterozygous segments as IDOL" units, so thatin any generation
LA + L A,+ L" ""Xo ,where X o is the whole genetical length of the chromosome pair
In a population undergoing self-fertilization n times, the frequency of
the AA' pair is. (l-xo)2"/2'" •The frequency (f,,(x)dx) with which L" lies between x and x+dx (O<x<xo) can be given by solving the following integral equation, the initialcondition of which is fl(x) =Z-xo :f,.(x) = O-;X)2f.._ 1(X) + C(2-~)f",-M)d~+(Z-Xo)(l-xo)2("'- 0/2"'-1 .

The frequency of heterozygous pairs at the nth generation:

This is approximately equal to (1 +Znxo)/Z"', where X o is small; it willapproximate nxo/Z"'-t, when n is large. In the following table the valuesof H •• are given for the initialZO generations, assuming that the chromosome length is 100 units.
48

•

,

)(. This is the corollary of the assumption that the initial length of theheterozygous segment is 100 units and only a single crossing-over is allowed
to occur in that region.

,..

,

From these values, the curves showing the decrease of heterozygosis
for plant hybrids with m pairs of such chromosomes can be easily constructed. If m =7, the frequency of heterozygous plants is lower than
3/10,000. After sufficiently many generations. when all the chromosome
segments have reached the state of fixation, the population contains
three kinds of pairs, namely AA, A'A' and the recombined homozygote.
The frequencies of AA and A' A' are both equal to e- 2%0/2. and in therecombined homozygotes the frequency with which LA/xo lies between t:and t+dt is

qJ(t)dt= 2Xoe-2xo{I0(4Xo Vt(l-t) ) + M4Xoj t(lDJ_} dt ,2\ t(I-t)

where 10 and I. are BESSEL functions.
cf. KIMURA, M. 1951. The Recombination of Chromosome Segments throughContinued Self-fertilization (in preparation).

7. Studies on the Process of Chromosome Subst£tut£on(Report by Mot6 KIMURA)

In order to find out the influence to be exerted by a foreign cytoplasm
on the character expression of the gene, a chromosome substitution experiment is often practised between two different species or strains.
The methods involved in such an experiment are to repeat the backcrossing of F 1 and the descendant individuals to the pollen parent, with(the first method) or without (the second method) the duplication of F 1
chromosome. (d. KIHARA 1948)49

For such an experiment it is important to estimate how many generation are needed to attain a given degree of substitution under various
conditions.
In t he case of the second met hod, if there are m pairs of chromosomes,of which the ith pair have genetical length of lOOxo(i) units, the proportions of individuals for which the substitution is completed will be.jj(l_l±n~X-'i)_)

'-I

2

,

in a n times backcroc;sed population.
For example, the proportion of such individuals will be some 95% afternine times of backcrossing, if the hybrid has seven pairs of chromosomes
each having a cross-over value of thirty per cent.
Similar formulae can de derived in the case of the first method.
If in the course of the experiment the fertility decreases bacause of
the presence of plasmon sensitive genes, the formulae become much
more complicated. The writer's analysis of Prof. KIHARA'S data of a
substitution experiment with Aegilops longissima and A. Aucheri hasrevealed that the effect produced by such plasmon sensitive genes on
fertility might be geometrically cumulative.

1,.~

cf. KIMURA, M. 1950. The Theory of the Chromosome Sub3titution between
two Different Species. Cytologia 15: 281-294.

8.

Theoretical Relations among Map-distance, Recombination
Value and Coincidence Ratio(Report by Moto KiMURA)

The map-distance, the recombination value and the coincidence ratio
are the three important conceptions in the studies of crossing-over, andmany formulae have been proposed concerning the relations among them.
The present writer has attempted a general consideration concerningthese relations and obtained some results which are reported here.
Let ¢(x, y) be the recombination fraction between two points in one
chromosome map, and let CN(X, y) be the coincidence ratio between two
infinitesimal regions (x, x+dx) and (y, y+dy), then it will be proved that2¢(x,,,- y)-] x-a,CJo.{a, b) --.:. [ -c 2"
.

UXUY

.v-b

•••••••••••••••• (I)assuming that the frequencies of triple and multiple crossovers are·negligibly small between two points a and b located on a chromosome.
Therefore, if the recombination fraction and the coincidence ratio aremere functions of the map-distance, we obtain the relation:d 2¢(x)CJo:(x)=-- 2dx2
50

'

(2)

•
j

,

•
where ¢(x) is the recombination fraction between two p:>ints which are
x distance apart, and C/!'{x) is the coincidence ratio for the two points.
Applying this to mapping functions which show the relation between
the map-distance and the recombination fraction, we can obtain a function
showing the relation between the recombination value and the coincidence
ratio.
For example, from the mapping functions:¢(x) = ~ sin (2x) ,.•........ ,(3)(LUDWIG 1935)

1
1
x = n-loge {I +4¢(x)}- 3 log, {1-2¢(x)} ,(DE WINTON and HALDANE.•.... (4)

1935)

1

if' (x) = 2 tanh (2x) , .................. (5)(KOSAMBI 1944)

we can derive the following functions :CR(x)=sin(2x),(3)'

CR(X) =8¢(x){1 +¢(x)}{1-2¢(x)}{1 +4¢(x)}/{1 +2¢(X)}3, .•. (4)'
CR(X) = 2 tanh (2x) sech 2 (2x) ,•............. (5)''Or when x is very small

cf. KIMURA, M. 1951.
paration).

p,,

C g (x)=..2x,...•..••.••....•.••.•. (3)"C g (x)"78x,"CR(x)"74x.

.

(5)"

Coincidence Ratio and Mapping Functions.

(in pre-,(4)"

PAPERS AND BOOKS WRITTENBY THE MEMBFRS OF THE STAFF
KAJI, Sanae and M. ICHIKAWA. 1950. Function of the corpus allatum in the silkworm Bombyx mor;. Annot. Zool. Jap. 24: 1-12.
KAJI, Sanae and M. OGAKI. 1951. In:iuction of phenocopies by me~ns of nitrogenmustard. (Jap.). Zool. Mag. 60: 7.
KIMURA, Moti>. 1950. Mathematical genetics. (Jap.). in "Recent Advances in
Biology." Vol. 1, Tokyo: 178-186.
- 1950. Mathematical population genetics. (Jap.). Ibid. Vol. 2, Tokyo:289-341.
- 1950. The theory of the chromosome substitution between two different
species. Cytologia 15: .281-294.
- - 1950. A note on the prediction of the frequency of genotype expectedin Mendelian hybrids. (Jap.). Jap. Jour. Genet. 25: 118-120.
1951. Recombination of chromosome segments through continued self-

51

fertilization and the degree of homozygosis to be expected. (Jap.). Jap. Jour.
Genet. 26: 53~2.
KIMURA, Mota. 1951 Recombination of chromosome segments in populations.
(Jap.). Ibid. (in press).
KOMAI, Taku. 1950. Linkage of genes and chromosome maps in man. (Jap.).
in "Recent Advances in Biology." Vol. 1, Tokyo: 1-13.
1950. Semi-allelic genes. Amer. Nat. 84: 381-392.
1950. Photograph of a man with a tail. Jour. Hered. 41: 247-248.
1951. Geographic variation in the land-snail Bradybaena simi/aris. (Jap.).
Jap. Jour. Malac. 16: 87-103.
1951. Genetics of butterflies. (Jap.). Shin Konchu 4: No.1; 3~32,
No.2; 8-10.
KOMAI, Taku, M. CHINO and Y. HOSINO. 1950. Contributions to the evolutionary
genetics of the lady-beetle, Harmonia. I. Geographic and temporal variationsin the relative frequency of elytral pattern types and in the frequency of
elytral ridge. Genetics 35: 589~01.
MA'l'SUMURA, Seiji. 1950. "Cytogenet.ics and Breeding of Wheat." (J ap.). Tokyo.
1950. Mutations induced by physical means. (Jap.). in "Recent Advances in Biology." Vol. 1, Tokyo: 73-133.
1950. Chromosome conjugation and fertility of triploid wheat hybrids
and their offspring. (J ap.). Rep. Kihara Inst. BioI. Res. (Seiken Ziha) No.
4: 31-43.
1950. Manifold effects of a long glume gene in Polish wheat. (Jap.).
Jap. Jour. Genet. 25: 36.
1950. Linkage studies in wheat. II. P-linkage group and the manifold
effects of P-gene. (Jap.). Ibid. 25: 111-118.
MATSUMURA, S~iji, A. MOCHIZUKI und O. SUZUKA. 1950. Genetische und zytologische Untersuchungen bei Beta-Arten. III. tiber den Zuckergehalt bei den
Varieti1tbastarden und polyploiden Pflanzen. (Jap.). Rep. Kihara Inst. BioI.
Res. (Seiken Ziho) No.4: I-II.
MATSUMURA, Seiji, A. MOCHIZUKI and O. SUZUKA. 1950. Mutations in barleyinduced by X-rays. I. (Jap.). Ibid. No.4: 22-30.
MOCHIZUKI, Akira and S. MATSUMURA. 1950. Genetic and cytological studies on thegenus Beta. IV. Triploid sugar beet and its yields. (Jap.). Ibid. No.4: 12-16.
OGAKI, Masahiro. 1950. Mutations induced by treatment with chemical substances. (Jap.). in "Recent Advances in Biology." Vol. 1, Tokyo: 56-72.
OGAKI, Masahiro and Y. OJIMA. 1950. The influence of sodium ribose nucleate
on meiotic divisions of the grasshopper. Nature 165.
OGUMA, Kan. 1951. The chromosomes of the dragon-fly Epiophlebia superstes
SELYS. (Jap.). Papers fro Coordin. Comm. Res. Genet. No.2. (in press).
SAKAI, Kan-Ichi. 1950. Heterosis and heterosis-breeding in plants. (Jap.). in"Recent Advances in Biology." Vol. 3, Tokyo: 267-300.
1951. Breeding rice varieties resistant to unseasonable low temperatures.
in summer. (J ap.). in "Recent Advances in Rice Culture." Tokyo: 45--£4.
1951. Experimental studies on significance C?f selection in plant-breeding.
1. Can the performances of normally transplanted and directly seeded riceplants be predicted from those singly space-planted? (Jap.). Proc. Crop Sci.
Soc. Jap. (in press).

52

J

SAKAI, Kan-Ichi. 1951. Ibid. II. Changes in performance of rice varieties due
to neighboring plants of different varieties. (Jap.). Ibid. (in press).
TAKF..NAKA. YO. 1950. On the sterility of triploids.(Jap.). Papers fro Coordin.
Comm. Res. Genet. No.1: 139-143.
- 1950. Note on cytological "observations in Colchicum, with reference toautotoxicosis and sterility. Cytologia 16: 95-99.
- 1951. Inheritance and determination of sex. (Jap.). in "Modern Biology." Vol. 3, Tokyo: 33-86.
1951. Karyotypes and sterility in Hemerocallis. Papers fro Coordin.
Comma Res. Genet. No.2. (in press).
1951. Cytogenetic studies on the genus Nicotiana. I. La Kromosomo(in press).
TANAKA, Yoshimaro. 1951. Genetical studies of the star markings. Papers
fro Coordin. Comm. Res. Genet. No.2. (in press).
1950......1951. Studies on hibernation with special reference to photoperiodicity and breeding of Chinese tussar-silkworm. I-V (Jap.) Jour. Bull.
Jap. Seric. Sci. 19: 358-371, 429--446, 58~590, 20: 1-9, 132-138.
1951. Breeding of tussor (Autheraea pernyi GUER.). (Jap.). (in press).
. 1951. Genetical studies of the multi-lunar marking in the silkworm.
(Jap.). (in press).
TSUJITA, Mitsuo. 1950. Development and heredity in the silkworm. (Jap). in"The Recent Advances in Gene-Analysis of the Silkworm." Tokyo: 35-65.
~~--- 1951. 1. Morphology and embryology of the silkworm.
II. Cytophysiological genetics. III. Development and heredity. (Jap.). in "Geneticsof the Silkworm" (in press).
--- 1951. Studies on the factors connectmg genes with characters in thesilkworm. (Jap.). Papers fro Coordin. Comm. Res. Genet. No.2. (in press).
UMEY A, Yoshichiro and M. TSUJITA 1951. Genetical and histo-cytological studies
on the lethal yellow larva in the silkworm. Bull. Seric. Exp. Stat. (Jap.).
13 (7): 329-345.

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