Bicho Seda Takayana

IHDI
.I. Ui:.3:WJ lClI

.G l:[OLJ IlI'U .ILl,
l' 111 U 1'1, J'JEN DElHI
in Aid [or Scientifjc Research.
Ministry of Education
SILKWORM GENETICS

SC4::!9
r 111111 m 11111111
fARI
SILKWORM GENETICS
ILLUSTRATED
BY
Tadao YOKOYAMA, D. Agr.
Director llie S'rricu/lltml E\j)erimelll S'lation.
Millislry of i\gricil/lurc lind Foresl;y,
81t.'!:illmni-lm, 'l'61<),(), ]aj)({/1
Japan Socidy for the Promotion of Science

Published by Japan Society for Promotion
of Science, Ueno Park, TiJkyo, Japan
erin/cll in l1cratd ill):!;, Co" Ltd" ,'-,'/li/l(/Ilra,
lllj)(lil
Jj
2
7 9
13 14 15
10 11
16 17
I
--
..
6
12
18
Silkworm PattcrilS
1. Bombyx i/I{{/u!ariIlCl
2. normal patterll (-I-I! II-O.O)
3. hlack (I,ll II-(J.O)
1). moricalld (/,11 II-D.O)
r,. sl rilled (j,s 11-0.0)
G. ami dominanl c1\m:ubk.
(1)8 II-lUI, I-a IX-O.O)
7. ll1ul1i1unar (L IV-O.O)
8. lllllltilllJ1ar and speckled
(L IV-1I.0, SjJ,; IV-33.l)
tluail (q VII-O,O)
10. e1il \1l" blaL:k (bd IX-I).7)
11. knl1hbed and ursa
(T\ XI-O.O, [J Xl V-S.O)
(liny en; XIV-lO.?)
1:1. stollY (st VIll-5.7)
H Kins!lirYli translucenl V-'l.7)
Iii. icmon (fant H1-22.:-1)
Iii. red blood (rb)
17. l.cbra fueled (2"f JlI-D.O)
18. hereditary mosaic
(normal amI
Preface
The International Genetics Symposia were held in Tokyo and Kyoto ro111
the 6th to the 12th of September, 1956. The organizing committee of the
Symposia, Dr. H. KIHARA presiding, planned to exhibit genetical objects char-
acteristic of Japan. The Exhibits Subcommittee, Dr. Y. NOGUCHI presiding,
was organized to bring together 8 working groups, each group under the
leadership of the person designated in the parentheses below:
1. Japanese Morning Glory (Dr. Yo TAKEN AKA)
2. The Biggest and the Longest Radishes in the World (Dr. Ichizo
NISHIYAMA)
3. Wheat and its Relatives (Dr. Hitoshi KIHARA)
'1. Rice (Dr. Toshitaro MORINAGA)
5. The Silkworm (Dr. Taelao YOKOYAMA)
G. The Long-tailed Fowl (Dr. Kiyoshi MAScTJ)
7. The Gold-fish (Dr. Yoshiichi MATSUI)
8. Olel Documents (Dr. Yosito SINOTO)
The purpose of the silkworm exhibition was to enable the visitors to
understand as well and as widely as possible the silkworm genetics and the
raw silk industry in Japan. All silkworm geneticists were asked to partici-
pate in it. The exhibition was extended so as to include mulberry breeding,
pathogenic micro-organisms attacking silkworms and mulberry trees, and wild
silkworms, because it was thought that these would bring a better understand-
ing of the whole industry.
Special efforts were made by the research group under Dr. Y. TANAKA 'to
complete all chromosome maps on the silkworm in time for the exhibition with
the result th;;1.t theYJ found twenty-eight presumptive linkage groups of genes
corresponding to the haploid number of the silkworm chromosomes. These
efforts were aided financially by the Ministry of Education.
No special emphasis was put on any particular field of silkworm genetics.
However, many visitors inspected with great interest such exhibitions as 80
different kinds of Jiving larvae; silkworms of artificially induced mutation,
which ieed on various plants other than mulberry leaf; theory and practical
application of sex-determination; developmental genetics advanced by utilizing
various mutants; various hormonal studies in relation to the development; and
studies on the coloring substances of silkworms and cocoons.
The Silkworm Exhibition was opened on the 8th of September, 1956, and
there were 200 visitors, a few of whom remained there for several hours
eagerly looking at and discllssing the exhibitions. Not only the"e specialists
but also many geneticists of recent acquaintance ad vised us to publish the
pictnres and tables exhibited. Their advice is now to be realized by the co-
ii Preface
operation of the members of the working group aided by the grant of the
Ministry of Education.
All members of the working group were co-orc1inated in every aspect of
the task. Especially Dr. S. SIIIMlZU, Dr. M. HARIZl.mA, Mr. T. nmOIl!( and Mr.
T. TAKAMI supported the group. Painstaking efforts by all of them were
highly appreciated. Dr. H. KrnARA, Chairman of the Organil,ing Committee,
Dr. Y. TANAKA, Chairman of the Lecture Subcommittee, Dr. Y. NOGllCH[,
Chairman of the Exhibition Subcommittee, gave us valuable advice, encourage
ment and help whenever needed. We were also aided by thc Science Council
of Japan, the Ministry of Education and the Ministry of Agriculturc and
Forestry. Mr. H. MIYAYAMA of the Ministry of Eclucalioll gave m; very woe-
ful advice and aided us in many ways in the pUblication of this book. On
behalf of the working .lSroup of the Silkworm Exhibition, I express here my
hearty thanko; to all pe1'sons and organizations mentioned ahove.
My sincere gratitude also goes to Dr. Ycniito SINOTO or the International
Christian University f()r his kincl help as well as to Mes:H.s. Edward J\
and Yasuyuki OWADA for correcting the English manuscripts.
April 29, 1%8
Tac!ao Y OKOY.A.MA
Leader, The Working Group of the Silkworm Exhibition
Members of the Worldng Group of the Silkworm Exhihition
Mr. AIDA, Fumio
Dr. AOI<I, Kiyoshi
Dr. Aln.JGA, Hisao
Mr. ClIIKusm, Baruo
Mr. HA.MAllA, Shigeyoshi
Mr. HARADA, Chuji
Dr. HAlHZUI(A, Masaki
Dr. HASIMOTO, HarLlo
Mr. EmOllE, Tatsumichi
Dr. hIKAWA, Nobuka7.ll
Dr. KOGURE, Makita
Dr. Kuw ANA, Zyuiti
Dr. SASAI{f, Shizuka
Dr. SIlIMIZU, Shigeru
Dr. TAZIMA, Yataro
Mr. TAI{AMI, Takeo
Mr. TAKASAKI, Tsuneo
Dr. YOKOYAMA, Tac1ao
St:ricultural Experiment Stati()ll, M. A. F.
Tokyo Univen:ity
Kyushu University
Do.
Sericulturhl Experiment Stat ion, M. A. F.
Do.
Do.
Du.
Do.
Do.
Tokyo University of Agriculture amI Textile
Industry
Sericllitural Experiment Station, M. A. F.
Do.
Do.
Silk Science Research I nstiittle
(Now, National Institute of Gl'neliu;)
Sericu [tural Experiment Stalion; M. 1\. F.
Do.
Do, (Leader of the Gruup)
Silkworm Genetics Illustrated
By
Tadao YOKOYAMA
Contents
Pages
Introduction ..................................................................... '" .... ..... 1
I. Life History (including the Silk Secretion) '" ............ ... ... .......... ..... 4
II. Hereditary Traits and Chromosome Maps .................................... 12
III. Cytology and Reproduction...... ... ......... ..... .... ... ...... ......... ............... 40
IV. Sex Detennillation ........................................................................ 44
V. Induced Mutations and the Practical Application.... .......... ............. 52
VI. Mosaics .............................. '" ...... ...... ....................... ............ ..... .. 66
VII. Developmental Genetics ............................................................... 73
VIII. Physiological Genetics .................................................................. 91
IX. Biochenlical Genetics ..................................................................... 108
X. Breeding (including Heterosis) ............................... : ...................... 138
XI. Relation between the Silkworm Races and the Pathogenicity of
Muscardines ................................................................................. 157
XII. Wild Silkworms ........................................................................... 161
XIII. Genetics of Mulberry (including Food Plants of the Silkworm Other
than Mulberry) .............................................................................. 165
Literature .................................................................................... 183
Silkworm Geneticfl IIlustrated 1
Introduction
The silkworm has been reared in Japan for mote than 1000 years. Since
the latter half of the 19th century raw silk has become one of the most im-
portant export goods of Japan. has developed into an important,
representative industry of this country.
Owing to the governmental encouragement for the advance of scientific
research and extension of techniques, remarkable and research results
have been obtained, contributing much to higher development of the industry.
The most important contribution among l11any is the adoption of hybrid silk-
worm.
The Sericultural Experiment Station of the Ministry of Agriculture and
Forestry was established in 1911. Toyama of the station had already been
engaged in genetical study of the silkworm since about 1900, and published
the results of his research in 1900. In thc course of his research hc recog-
nized the heterosis. Following his advice, the government had decided to
encourage hybrid silkworm instead of aboriginal pure races and started the
distribution of hybrid eggs in 1'914. The rearing of a vigorous and produc-
tive hybrid silkworm stabilized the crop of the summer and autumn silkworm
culture. The production of silk from a unit area of mulberry field remarka-
bly increased by raising two or three crops instead of usual one spring crop
from the same field.
Silkworm genetics in Japan started and has developed in connection with
silkworm breeding, gradually enlarging the domain of its research sphere.
Keeping pace with the advancement of genetics, the related sciences in seri-
culture have also made progress thus contributing to the development of the
industry. The industry on the other hand presented many scientific problems
to be solved, which gave the opportunity for scientific discoveries. In such
a way the industry and the science aided mutually in the course of their own
advances, thus the largest country of the silk industry made itself also the
largest country in the study of silkworm genetics.
There are about GO recognized scientists in Japan in the field of genetics
and breeding of silkworm, the papers published from 1945 to 1955 amounting
to 590. Silkworm genetics in Japan owes much for its progress to the seri-
cultural industry, but the silkworm itself is considered to be a very appro-
priate animal for the study of genetics, making possible the advance in
research in various aspects of the field. In the connection the following
points may merit special attention:
1) There are numerous heritable traits in the silkworm and the number
of such traits can be increased comparatively easily by the induction of arti .
ticial mutations.
2 T. Yokoyama
2) Fi ve to six gcneratiOllE> may he repeated in a year by artiftcial hatch-
ing of eggs, and the larvae thereof are easy to rear.
The number of eggs laid by one moth is about 500 and they are
numerous enough to justify statistical treatment of the results in genetical
stuely.
4) The related sciences to genetics such as morphology, physiology or
oecology have attained a high level of achievement, and the adjustment of
developlllent or rearing is easy. Therefore silkworm is a very convenient
animal to follow the course of development of traits contl'olkd by genes.
5) Also it is easy to obtain large number of animals of uniform charac-
teristics.
For this exhibition an attempt is made to reveal as much knowledge of
silkworm genetics as possible, which has been accumulated in [l\lout: [iO years
by TOYAMA, TANAKA and their followers.
It will be worthwhile to give lwre an outline of the raw silk incluHtqt of
Japan.
The amount of raw silk produced in HlG(i hy 21 countries in the world is
28,808.4 metric i. e. 0.2% of the Lotal textile libel'S. Japan product'S about
63% of the total output of raw silk, China 15;)6, Russia H;),;, rUdy 4.2%, Hud
India 4%. Silkworms arc reared in every pre.i'ecttlrc in Japan, especially thri
ving in the middle region. There me about 7(j.1,OO() scricultural farmers who
grow about 193,000 hectars of mulberry, the ligures corre:->])oncling Lo 12.596
of the total farmers and of the total lanel respectively.
The income from sericulture is ahout Hl% of the total household inl'(nne
in the serlcultural. farmers anel ahout 4;)6 of the toial agricultural income of
Japan.
During World War 1[, tIll' sl'riculture dccrl'<I:->ecl to the levels, hut
since the end of the war it has gradually heen In it increased
to 2.:1 times of the war-time lowest level.
The raw silk produced in Japan amounts to metric t()ns, about
two thirds of which is consumed in the country and the n'st is l'xported as raw
silk or as silk fabrics. The situation of silk among export goods was so im-
portant in the past as amollnting to 1]8.l);i;; of the total export in 1022, but it:
is only :1,4% in 1957, sUll it remaining one of the lllost important agTicl.llturai
export goods in Japan.
Exported silk was used lTIoslly for hosiery befoi"e, but now replaced by
nylon. Silk is now more used for textile fabrics. Silk has many
nnparallelly good for high quality textiles such as smoothnesH,
lightness, dyeing ability, luster, draping-quality and hyg'ienic properties. On
the other hand there are some undesirable aspects in silk such as browl1ing
and fragility by weathering and lousiness. To remove the defect of browning
Silkworm Genetics Illustwtcd 3
and fragility chemical studies are going to be carried (Jut, while lousiness
problem has. been partly solved by breeding good varit;;ties of silkworm and
by a new processing method.
T. Yokoyama
I. Life History (including the Sill, Secretion)
in Japan the silkworm eggs hatch from the end of April to the
beginning; or May under natural conditions. If the eggs which passed the
. diapawic are kept at the temperature of they hatch in lO-14 days,
and the incubation period of the nOll-hibernating eggs is of about the same
duration.
The nl'wly hatched larvae are black of ahout :l111m. in length, about OAS
mg. in weight. These worms spin e(J('nons in 2fi clays when reared al 2fiOC. alter
molting ,1 times.
The larvae grow to thp maximum weight in one or one anel a halt clay
he fore spinning cocoons, the body weight heing about 12,000 times of the
newly hatched worm.
When the larvae have grown tn the extent lhat they are reacly to spin
cocoons, they are called ripe or "mat.ure ". In the mature larva, silkg-Ianc1s
are so large that they Ol'CU[JY ahou! ,10% of the hody weight. ,Farmers llsual-
Iy put the lllature worms in a Rpinning I1CSt, sllch procedure being. called
"mounting ". It takes about a week after mounting [or tlw larvae to pupate,
and 8 to lS days more for emergence of which come out of the cocoons
softening the cocoon 1:;he11 by an alkaline fluid secreted in the crop. The moths
come out mmally in the morning, and soon after that male '.llld female moths
mate. About !lOO eggs arc laid by u female moth hy the IWXt morning.
If the artificial hatching treatment is applied to tlw hiilc'rnating eggs in
about 20 hours after lnying, the eggs hatch in alJlllll III rlays afte], the trc'al-
me nt, thus enabling llw repetition of generations. One gC'l1eration, I'roTll cg.l'; to
egg, is completed in ahout GO clays including the incuhation period. For t'coJlomi-
cal reason, the silkworms are. reared usually times a ye'ar in .lapan, hut
fr0111 gl'llctical point of view, it is possible to repeat six gl'IWratiolls a year
utilizing; a warm place like Okinawa where they get llIull1l'rry leave'S in
winter .
.. The silk substance is secreted hy a pair of silkgJands each originating
from the invagination of epithelial cells in the embryonic labial segment. The
silkgland is divided into three different parts, anterior, middle anc! posterior.
The silk substance is a protein called fibroin of linearly arrang'ed amino aciel
molecules consisting of glycine, alanine, tyrosine, serine and B others, which
is secreted in the posterior part of the gland and accumulated in the middle
part and spun out from spinneret in the labial segment. The fihroin which is
a viscous gelatinous substance in the gland becomes fibro1l3 when spun out.
The fibroin fiber is coated with a "gum" called sericin which is secreted in
the middle part of the silkgland. Recently it was found by M. (19GS)
that cellulose substance is usually secreted in the middle part of the silkglancl.
Silkworm Genetics III 5
I-I. Life history of the silkworm
The exhibition displays living silkworms of all instars together with
pupae, moths and hibernating and nonhibernating eggs as well as a schcmaUc
expression of the life history.
I-I Life history uf the silkworm ,.
& Winterir,g :i
i
4-9 mOI1Ii1s

/ Terminati011 of
I diapausc can I", '\
tlY cold .tomlte i.

A ./' Artificial .. Il'ltchinl1
'"
diupausing egg Eggs can be stored /'
for about 90 days before. 1st instar
. artiliclal treatment at / \ 3"'""4 da}'$
Egg La;nng ,
" low temperature 9-i2 <laysilncubation) 1&./ I-Mellt
0/
'- nan.d,apause eGg ? 0 I '''OJ'
",I" II-Molt
moth Temp''''aturrs
... < \ lbtchinl' El,wLuying & '<JJ:r.!:r!,
Emergence at 3rd
,(ays .,. ''',. )
Dia.
P
31JSe anll \Vmtenng 3-...:J d(f\'s
II
under Natural Condition -
in Japan III-Molt
.... pupa in cocoon
/ 4th illstar
P 1..
1
pat 1011 days)
;:/j days /
}QQ J4---- IV-Melt
'060 dil)'" /
,. f'----------
! 5th instur
Ripening
Newly hatched larvae
1st j l1stars
G
T. Yokoyama
fit Il
pupae
Duration of one generation varies according to genclkal nature and to
treatment. The season of hatching can he adjusted ::>0 that man can rCHl'
silkworms in any seaSOn provided there are available mulberry leaves.
7
1-2. Anatomy of the silkworm
Intestine occupie::; almost all the body cavity of the larva, and is divided
into three parts, viz., fore-, mid- and hind-gut. The mid-gut is the largest
extending over eight of the whole thirteen body segments.
Three pairs of the Malpighian vessels sLart from the rectum, run forwards
along the outer surface of intestine turning back at around the third or t]1e
fourth abdominal segmcnto; and J!nally each three unites into one vessel, opening
inside the intestine at the hind part of the small intestine.
Larva respires through their spiracles opening at both sides of each body
segment except the secone! and the third thoracic segments, on the posterior
1-3a Dissection view along dorsal
side uf the silkworm
1, intestine, 2. trachea, ::I. Malpighian.
vesoel, 4. teslb, 5. [Jui:itcdur i:iilkglalld
1-2b Showing afler removing intestine
opened along ventral side
1. l11u:;c]e, 2. dorsal ves:o]cl, 3. testis, '1. tnt-
chea, 5. middle :;ilkgland 5th
inslar), G, pOl:ltel'ior sill(glund, 7. foregut,
8. midgut, \:). hindgut
T, Ylltwyalll<l
horder of which a [lair til' fUlll'liullll'ss spiraclL'\';, so-calll'd I Ill' rudillll'lll spirack,
exists, It is aSflLllllL'd that t Iw rudiment }ipiral'll' i:; a disc of an adull
Olle, 'flw Illaill trullk; Ill' trachea rUIl tlmlll,l\lt Illl' lHldy Illllgitudinally at holll
siC\cs, Each vair (If slliradl' ill a sugllH'lIl is ('OlllICl'il'd wilh it vl'lltral trans-
verse lral'lJeil,
Dorsal Vl'ssd, Iwart, ie; tlmlligh Ihl! (]lIn.;al p;lrl IIf till' hody frol1l
head t() lail. cllrd nlll,; through tltl' vl'lllral :-;illl' 1'1'11111 hl'ad to I'auclal
l'nd, (,Olllll'clill,l!; till' Ilraill and g',lll,l!;lilHlllt' l'<lell Fal blldies lit- ullcl(,I:
til,) integ-lIml'llt and around tlIt' ilItl'slilll' playing an ililpllrtalll rtll(' ill Ilw meta-
bolism slIch a:i st(lrn,l;t' (If mtllsUUll'(':" for ('lll'i'P;y SlIur!'(', ('x\'I'('1 illn Ill' uric acid
lll' SYlltlll'sh1 or pnll.dn, TIlt' p,'llllacl lie,; Illll1w dm:;nl :;idv Ill' llH' t'i).;hth hudy-
SC!-(llll'llt, IIIW (Ill ('adl si(k Ill' till' dor:;al Vl':;';('1.
FurL[ll'l' des('l'iptiOll or I hv illl;tllllllY Ill"('(:;;:i<lI'Y foj' gl'IIl'1 i('HI l'xplitnali()Jl oI
the silkw()rm will 1)(' )1;ivl'll ill Illl' fllll()willg' l'()ITI':qJllll<lillg ;';l'di()tls.
l-':!l' l'l'!I!;:'; :A'dillil at :ti)i["Illt'11
(If th(:
1.. midgut, 4, Illiddll' divi!., of llliddk !:ilk.
gland, :.l. llu:;(.c'riul' "r middlu >:ill,/;lalld,
4, jJo:;Il:rior 5. Iradl(::a, Ii. Mal-
pi,lillian 7, H. fat hody
Schematic ligures of silkgland
I-:.'d ill lit" full
.. larva
Silkgiaud of lIlal.lll'('([ larva
7 day old, 11lh im;Lar
Silkg-lancl is the second largest organ in lite silkworm body. It originatcH
with the paired invagination in the labial segment. In the full-grown larva it.
Silkworm Genetics lllustratcli 9
occupies most part of the ventra-lateral side of the body from fourth to the
eighlh segment. It is divided into three distinct parts, the anterior, the
middle and the posterior part, while the middle part is again divided into
three functionally different divisions. The fibroin, silk protein, is secreted in
the posterior part; sericin I, the innermost sericin, in the posterior division
of the middle part; sericin II, middle-layered sericin, in the middle division;
sericin Ill, outermost sericin, in the anterior division of the middle part.
There are several kinds of coloring substances of silk, and the secretion
of each is correlated with particular division of the silkgland.
Paired silkglands lead into aile spiner'ette in labium, and about the point
of union of both glands two Filippi's glands, one to each silkgland, are attached.
Several functions of Filippi's gland are assumed but not proved.
A
1-3 Schenmtk IigUl'c of sill,gland
Sm

. F
cross section
Side view of
silkgland in a body
right silkgland
A, anleriur silkgland; 13, anterior division of middle silkglancl;
C, middle division of the same; D, posterior division of the same;
E, posterior silkgland; Sr , sericin I; Su, sericin II;
Sm, seticin III; F, fibroin; SGC, silkgland cell.
Sericitl III is secreted from I'l, ::;eridll II from C,
sericin I from D and jibroin from E.
10
T. Yokuyalll<l
1-.1. Amino add compositioll of fiiJroin, sericin and COCOOIl
A tab Ie is exhihiled in which rceellt r('sLlI t s uf I he s( udy hy T. FUKUDA
Hnel J. KmIMtlHA hy means of bioassay arl' sh()wn.
I-I i\lllinu Acid COllljloCiilioll ,,{ FiiJruin 11 alld CII"lIUIi Fii>l.'1'
Amillu add Fillrllill Sl'ril'ill
l'''l'OOil (l(WI'
(Silk prpll'ill)
,II ,\11 i ae
.1. 110"',,
(I()n,;
argiuine
II. H:l ,[. liD l.ljG
Hl>parl k add
1. (iO 1 Ii. Mil ri. fiO
gllil lImk aei,\ 1.:1', III. Ii;; 1 II
glycilll'
HO H. (ill :::1. Ii II
itlint' H. 1. ::\1 11.'i'::
i:>1.1 Ie uL"i Ill' 1.::0 0.1)0 1. Ifi

[. 1111 II. HH 1I.!)(i
I Ill) II.I:.! ri. H,I I.:.!:.!
llwtlti'Hlilll' 11.11 O. I ri
phL'llylalallilW I. 117 1),li:i 1.11
vrul i Ill'
II, ,IH 1I.It>
ri Ill' 11.711 ::1). III IlL 1111
thrcUlli Ill) I. I:!
,.
d. I"
,I :1,1111
tryptophan O.:.!!i II,I!I
tYl'ut;i 1:1. 7!J :1. 1)0 I :
valinI.' HX L. Hti :t 1 r;
1 !i. Cdlulw;c substallce oC cm:()UIl JilJl!1'
While slurlyinp; llature or 1)1' silk, [\,1.
found L1mt a cLlllskkrall[l> part oj" l\)usy Ilhri !lac clIII\-\h\\s oj" l'l'ilulo:,;\'
(l!lfi!il. It is in Ow posterior divisiull or tilL' l1liddll' part or till!
gl"lncl. The urder of the polYlllQriza\ ion or litis sLllmlalll'(! b ('olll[Jaralllc to /1-
eellulllsl! uf plant i. c, l'1I111jJOSl!cI of ahout :.:()() lllllll'nilcs or
I-G. MeLahulic paLhway (If alllinu acids in silkworlll larva in J'l'lal i011 III silk
production.
The illustratiulls of tIll' urig'in of rnain compOllent l1lillO ad(h; of silk-
protein arc shown based U,l the H!Ccn! sludy o[ T. FUKliDA using radiuactive
0
1

It was proved Lhat phenylalanine of leaf ch,tIll,(cS inlu tYl'O::-;iIW
of silk and the inierrdations between glucose, glyoxylic acid alld
serine arc ::-;1 III wn. Alanine is Pposl!cl to .be c()llnl'ctcd wi l il variouH pal hwaYH
with pyruvic acid, <lHpal'tic add and oxulal:etic acid,
1-5 Cellulosic substance of cocoon fiber
M,
[-() Metabolic pathway of amino acids in silkv,'(Tr:l
larva in relation to silk production

!-IOCl!
IICOII
II(;OH
CH,OIl
gJucose
COOH I
(IINIl, dccorho)lyiuse ClI,NlI,
COOII coon
i1I1l\!10malonir. add glycine
\H,OII

COOH
serine
t

COOH
glyoxylic acid
COOH
COOI-! ell, eH,
bl /J.clcc"rhoxyiase CHNH, ell;
CIINH, "---- COOl! yIlNII,
tOOl( , C9
0H
,
il:.paI'Uc acid alanille aCid
i
trnllsaminasc III . Itr<lIlSaminasc
llmillation transaminase COOll
COOll Gr.l ell,
hI, co tn,
60' coo II
COOI1 COOI-!
ic add (pyruvic acid) acid
()
Cll'9"COOll CII'9IlCOOJI
Nil. --> I NIl
, IIO / '
phcnrlalaninc tyrosine
c: radio;Jcli I'C <':"
T. FUKUDA
11
12 T. Yokoyama
II. Hereditary Traits and Chromosome Maps
Genetical of the silkworm have been studied by many investi-
gators for a long time. The criteria of stuclies consist of various newly
discovered mutants and varioLls noticeable characterislics inherited fr0111
olel times.
[Jombyx IJlrtlularill(( (lIl(.>OjJIli/1( mallc/arillal which is regarded as tIle anceslral
form of Bow")'x mort resembles the laiter in morphological characteristics anc!
can freely be mated with R. 1}lOri. The number in n. mandarinll
is 11=27, while in B. }}lori 11=28. In 11. )}lori there is a skin marking called
ll1oric(luc!, which is genetically proved to be identical with that in L!. lJlmzdal'intl.
The hereclitary traits in /lumbyx mori amount to 211, the sec()ncl largest
number of such characteristics among insects next to that of Drosop/tila. The
difference bet ween Hrlllzhyx anrl Drosop/ti/a is that in DOIll/JYx the heriLable
characterislics are observed in various stages or espe.clally in
egg and larval stages, while ill lJrosoj)/iila mostly in imaginal slage.
T,lbk 1. Hl,reclilary I rail:; ill BO//I{Jyx lIlori
Chara<:ieri:;lics in
Larva
Pupa
COCOOll
Moth
Total
Nu. uf

Ii
1[1
14
:m
J\ certain spontaneoLis mutant makes a special characterbt ic uf a raCl', but
in another case .il is common in various races. For example, LlIU fulluwing
characterist ics are sped lie to different races:
striped marking (j)S) to Chinese race
quail marking Iql to Japanese race
dominant white cocoon (I) to European race
In the following a general explanation of the hereditary traits of Bombyx
mori from egg to moth wi I [ be given:
1. Egg.
1) Egg color. The color of egg is determined by three components, i. e.
the colors of the yol k, the serosa and the chorion.
The yolk color is usually light yellow, but in strains is deep and in
others light in color. The moth which belongs to the yellow blooded-white
cocoon strain lays eggs with very eleep yellow yolk. The color of serosa de-
Silkworm Gcnctks 13
velops after fertilization, In the nUll-hibernating egg the col-
but in the hibernating egg the pi,l!;mcnts in serusa begin to appear on
the secone! clay 01' laying and clevc1uLJ into tlle fixecl color in 3 10 "1 clays, There
are several kinds in serosa color, white, red, bru\\"l1, and dark brownish purple,
II-I Egg color
oorllIol egg _, I
dark colour
IlOfJIl(JI
light (olour
,.-
greeJl llire)
grey t'1}J - hoaa
(/Jr )
grey egg-beIero" greyegq E-16 -I1cIJlO grey egg E -16-bderc
COr) I Gr-16) (6r-1d)
light grey egg
(L'J )
unslable. grey egg' bird .. eyed egg III ur Iig-hl !
({jfll ) {(itt!) grey egg (/ -(,!l )
II --1 normal egg-clark colc.Jl'
..
I
..
II .. --3 normal egg-light color
green egg (Gre 1-4GA)i'
II---4 grey egg--homo \Gr 11-6,\))
II--S grey egg-hetcro IGr 1l-(l,9i
II-(j grey egg E-Hi-holllo (Gr-In II- 6.91
II--7 grey egg E-IG-heteru IGr-lG IICj,D)
II-- 0 light grey egg (Lg 1I-6,l-G,9)
II--9 unstable grey egg (Gm JI-6.1--G.DJ
II--IO bird-eyed egg (Grb II--6,9)
II -11 inhibitor of light grey egg ([-L,g II-G,l-G,\JJ
" Italics, Roman and Arabic numerals ill (he parentheses show gene symbols,
chromosomes and gene loci, respectively,
14 T. Yokoyama
the last being ordinary color. The color of serosa usually coincides with eye-
color. The color of chorion is white or light yellow and in a few cases it is
pinkish.
II-12--27 Egg color and the shape
red el.19 ire)' cocoa Icuel brown egg-Z (1J..)
piled f99 ":, white ew2 (11,,);0 white egg-J (lVJ)' I Aojuku whilcc!l!J' whilr'l"I}j Irons
tmnslucenl (W"') lucent (Qtlll)
- unferlilileli eiJ9
(linn! ((it" 1, dottillllPI SIKlIl t9\l
(S.),
11-12 pink-eyed white egg (pc V-O.O) .,
11-13 brown egg-2 and red egg VI-S.U; re V-31.7)
Il-I4 red egg (re V-31.7)
II-IS cocoa egg (we)
II-I6 brown egg-2 VI-S.O)
II-I7 brown egg IVII
II-IS white egg-I (Wj X-O.O)
II-E) piled egg
II-2U white egg-2 (W2 X-3A)
II-2I white egg-3 (W3 X-G.g)
II-22 Aojuku white-egg translucent-larval skin is translucent (zool X-G.g)
II--23 white-egg translucent-larval skin is translucent ({Jew X)
II-24 white-side egg (se XV-D.O)
1I-25 giant egg (Ge 1-14.0)
II-2G dominant small egg-an X-ray mutant (Sm III-21.U)
II-27 recessive trimolting (rl VlI-9.0)
15
21 Egg shape. The normal egg is short ellipsoid, but there are mutants
in which the egg is spindle-shaped or long ellipsoid. The kidney-shaped egg,
the lethal spindle egg and some other eggs are accompanied by lethal genes_
Egg shape and egg.shell color
C!IlJ . ellipSGid egg (rIp) kidneyshaped egq
fJPJ (/({)
II - 28 spindle-shaped egg (sp)
II -29 ellipsoid egg (elp)
II-3D kidney-shaped egg Iki)
II-31 lethal spindle egg (hPj
II-32 noglue (Ng XII-D.O)
II-33 brown egg lethal ({-be)
II-34 lethal nonhibernating egg (/'n XII-21.0)
II -35 white egg lethal (l-we)
II-36 white egg-shell
II -37 reddish eggshell
II -38 pale green egg-shell
II-39 green egg-shell
16
T. Yokoyama
2. Ant or newly hatched larva.
In ordinary varieties it is black showing a certain degree of fluctuation
in the shade of color. There are two mutants hoth choc(llate in color, but one
dominant and the other recessive.
II -'lG Shori bristle (sb)
Newly hatched larva
1I -110 normal color
II -41 chocolate I.ef/ X III-OJ))
II--12 chocolate and white egg-2 (c11 XIII-O.n; Ui, X-3.4)
11--43 dominant chocolate (I-a IX--5.9)
II --44 lethal non-molting-l (Ilnll)
II-45 lethal non-molting-2 (mIl
2
)
II-46 :short bristle (so)
3. Larva.
-i- sb
1) Marking::>. The larval markings are not clear in younger worms till
the third instal', but clearly distinguished after the fourth. There are large
17
number of different larval markings which are comparable to the imaginal
characteristics in Drosophila.
:.!.) Color of the skin. It is white with greenish, yellowish, or reddish
tint, but there are mutants in which the skin color is greenish yellow or the
skin is translucent owing to the small amount or nothing of uric acid in the
epithelial cells, called" oily". Several genes locating in different chromosomes
are known which express almost the same characteristics, "oily!t.
mood color. It is colorless or yellow, and the color of blood corres-
ponds to tIl(' color of COCODn, but thC're is a mutant which has yellow blood
and spins white cocoon.
4.1 Body shape. There arc wide differences in size, and shape. There
are mutants in which there arc sllpernumerolls prolcgs, knobs on the dorsal
surface of the body, or with the skin highly tensed, hard, ancI "stony",
In the silkworm there are 3--, .4-,5- and 0-- molting strains and 1-, 2-, and
I11ultivoltine races. European race is always univoltine ancl tropical race is
J11 uIt i voltine.
II -47 sex -linked translucent
(os 1-0.01
JI-49 Chinese translucent
(oc V-40.0)
II-48 distinct translucent
(od 1-49.6)
II-50 Kinshiryu translucent
V-4.7)
18
T. Yokoyama
II--51 a translucent and black pupa
(bP XI-I7.I)
II--53 oal-mottlcc1 translucent
(oat II-26,7)
II -55 plain eye-spotted
(p' II-O.O)
II -52 od-mottlec1 translucent
(od"' 1-49,6)
II --[j4 plain (j) Il--O.O)
II -56 normal pattern
(+P II-O.O)
II-57 moricaud and lemon-the
pattern resembles that of B.
malldal'ina (PM II-O.O; lem III-22.3.:
II--59 striped I P' II-D.O.!
II-61 S-mottled-a chromosomal
aberrant
II -58 moricatld and zebra
(PM II-O'!); 2(1 1lI-1.5!
II-GO striped and dominant
chocolate (p' II-O.O; I-a IX-5.D)
II-52 striped 2 and moricaud
(S2 II-6.l; pM II-D.D)
19
20
T. Yokoyama
II-G3 dilute striped (Sa
II-65 black (PB II-O.DJ
. II-67 apodal and zebra
(ap III-O.O; Ze III-1.5)
IT (i4 ventralstriped (PO no,O)
black and dominant chocolate
Ii)}) II n.n; l-rr 1\ [i.HI
II -G8 zebra faded (Z(/ 1.5)
II -69 lemon (lem III-22.3)
II -II multilunar 1:5. Ii, 8 segments)
IL lV-O.(l1
Il1ultilunar 1'1---S
(L lV-O.O)
II-IO multilunar (5, 8 segments)
(L IV-O.O)
II-72 multilunar 15-8 segments)
(L IV-D.G)
21
22 T. Yokoyama
II-75 multi1unar (4-9 segments)
(L IV-O.D)
multilunar (4-10 segments'!
(L IV-O.O)
II-79 stick (sk IV-25,S)
II--76 multilunar (5-10 segments)
(L IV-fLO)
II-78 multilunar and speckled-an
X-ray mutant (L IV-O.O; ,Spt' IV--33.1)
U---SO Kp supernumerary legs
(EKPVI-O.O)
extra-crescents and legs
(EE:' VI-D.O)
JI--83 double crescents
VI-O.O)
II -85 new additional crescents
(EN VI-O.O)
II-84 additional crescents
(ECf! Vl-O.O)
II -86 no crescent marking
(Nc Vr--l.4)
23
24
1'. Yol):oyalll<l

_. .....
1
II --87 quail and plain, 1
(q VII-O.(]; j) II-O.())
U-SB quail and cheek and tail
spots (q VII-OJ); cis)
II-91 stony (sf VIII)
Il-S8 quail (;l'lc1 :2
II} VII-O,(); j) lI-O.())
Il--90 quail (II VII-O,O: V'II-O.l))
II-93 dilute black (bd lX-G,7)
Silkworm Illustrated
II -93 knobbed I K XI-O.Q)
II-95 dirly Wi XIV-O.O)
II-97 sooty (so)
JI-94 burnt-an X-ray mutant
Wu XI-5.5)
II -9G red skin
II-98 extra spiracle (es XII)
25
26
ursa (U XIV-2.7)
T. Yokoyama
r '.
II-IOl solid black-eye spot (blind)
II-I 00 ursa and knobbed
(U XIV-2.7; K XI-O.O)
II-I02 red blood (rb)
II-I03 brown head and
tall SlJots (bts)
II-IOl! narrow breast (nb)
II-I05 dominant retarded
II-I07 second-segment monster
(mse II)
II-lOg hereditary mosaic, cit P'
and + P' (mo)
II-lOG recessi ve retarded
II-lOS hereditary mosaic,
Ze and -I- (rno)
II-110 mosaic (l-Sp)
27
28 T. Y okoyaI1la
r 1- ! I.:: (SI))
II-1l5 apodal I (fP lIl-1I.0)
II-117 burnt--an X-ray mutant
(lilt XI-G.G)
l[ black pupa (bjJ XI-I7.1)
11- I-I! !EkruptLrllLls 1111/) XI--2'-1.0)
(i knobbed Ii( XI-O.D)
lI--1l8 crayfish (c/ XIlI--l1.8)
Silkworm G":I1t:!tics Illustrated
II-1El .eurltc1 Will:,; ,ele, 1I-1Z0 wingless (fl ;(-0.03;
4. Pupa.
1) Shape. In g'eneral the form of silkworm pupa is the same as uSLlal
LepidopleroLls one, but there are a few mutants which show abnormality in
wings or in other Imrts of the body.
2) Colur. Usually the colur of pupa is brown, but: there are mutants in
which the color is black, e. g. black pupa, bj).
5. COCOOI1.
1 i Shape. Ellipsoid is very common, but in Chinese race there arc many
varieties which spin uK'OtmS, and in Japanese race it is peanut-shaped.
21 Size. There is a wick range of difference in size from 20 mm to 60
111111 in length and ,'rom (J.t g to ::LO g in wdght.
3.1 Color. There are two main colors, yellow and white, and besides these
there are green, or reddish colors.
II--121 normal white (-i
r

II-122 yellow inhibitor II IX, 0.0)
Il-12:l sooty plain white '.(( IX-O.O)
Jl-124 green a ami green [J (Ga; GfJ VrI--7.0)
II-125 green (' IGI: XV--7.8)
II-12G flesh-culored and inner-white IF V1-13.G; +c XII-14.0)
1I-127 cinnamon-buff and inner-white ICiI; F VI-'l::l.G; 1-" XII-H.O)
II-128 pin];: anc! inner-white ! Fh; F VI-l:Ui; +c XJI-14.m
pink and inner-white (Pli; F -I ,. XlI-14JJ)
Il-130 golden yellow (C XII-I-1.UI
II-131 dilute yellow leI XI! 14.11)
II-132 Ie" XII--H.O.I
II-133 inr,Cl'-y(.llow ,e Y.I 1- U.O 1
II-13-! yellow blood and wllite cocoun I 1-'" VI-13.li; -f.' XU--U.O!
II-l35 flesh -colored and inner-yellll\v (F V 1-13,(j; C XII-14.0)
1I-13G cinnamoll-buff and inner-yellow \Cb; F VI-l:1.G; e
i
XII-l4.0)
30 T. Yokoyama
Cocoons
+ y 11-122 I a CaCh
1I-12S Gt; 1I-12G PI G 1I-127 CbF+G II-128 PkCbP+ C
II-129 PkF+C II-UO C II-131 Cd 11-132 C"
11-133 Ct
1I-1:J5 FCi II-136 CblCt
][-137 PkFC' Il-US ]i1pilne,;e type Il-139 Eurupean type II-HU type
11-141 tropical type mandarin<l lype Il-143 smoolh surface II-IH rough surface
1I-145 fluff y cocoon
1I-150 polypupal
Silkworm Genetics Illu,tratcd
II-137 pink and inner-yellow (Pk; F VI-13.G; C
i
XIH4.0)
II-138 Japanese type-cocoon shape
II-I 39 European type
II-140 Chinese type
II-141 tropical type
II-142 mandarina type
II-143 smooth surface
II-I44 rough surface
II-145 fluffy cocoon
II-146 sericin cocoon-naked pupa (Nel)
II-I/17 peaked cocoon
II-148 flattened cocoon
II-149 double cocoon
II-I 50 polypupaI cocoon
G. Moth.
31
1) Shape. There are mutants in wing such as vestigial or short wing
like Drosophila, but the number of varieties is known far smaller in Bombyx
than in Dmsoplzila.
2) Color. In B. mandarina the color of wing is dark brown with black patch
on the tip of wing, but in B. mori it is white except a variety" Papillon nair":
II-15I Bombyx mai,tdarina
II-152 normal
S2
1[-153 bimolting (rt XlI-9.0; M:J VI-3.0)
1I--15
c
l recessive trilllolting itt
II-Hi5 1l1icropterous Imp
II-1S6 black moth (Em)
Silbvorm Genetics Illustrated 33
II-I 57 wild wing-spot (Ws)
II-15S white banded black wing (wn V)
1I-1!19 lmobbccl (!( XI-O.())
II-160 burnt-an X-ray mutant (Eu XI-5.5)
34
T. Yokoyama
vestigial-an (Vg 1-38.7)
II-162 wingless (fl X-O.08)
II-163 mosaic (l-sP)
3) Color of eyes. It is black, red or white corresponding to the color of
eggs as mentioned before.
The chromo30me maps which have been made clear till now are shown
in II-164.
Silkworm Genetics Illustrated 35

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'determinor
36 T. Yokoyama
Chromosome Maps of Bombyx mori L.
Explanation of the symbols
I-chromosome: os, sex-linked translucent (skin character)
I-a, lethal a
II -chromosome
palleles
Lm, late maturing (voltinism)
Ge, giant egg
e, elongate (body shape)
I-e, lethal e
Vg, vestigial (wing)
I-b, lethal b
Gre, green egg (egg shell color)
od, distinct translucent (skin character)
odm, od mottled translucent (skin character)
p, plain (skin marking)
pI, semi plain (skin marking)
+P, normal marking (skin marking)
+P', lightest normal marking (skin marking)
+P', light normal marking (skin marking)
+P', standard normal marking (skin marking)
+P', dark normal marking (skin marking)
pB, black (skin marking)
pD, donml spot (skin marking)
pll, ventralstriped (skin marldng)
pL, light crescent (skin marking)
pM, moricaud (skin marking)
pMa, mandal"lna moricaud (skin marking)
pS, striped (skin marking)
pSt, pale striped (skin marking)
psa, sable (skin marldng)
pSa-2, sable-2 (skin marking)
8-alleles:
normal marking (skin marking)
8, new striped (skin marking)
8,t, dilute striped (skin marking)
S1-8, inhibitor of new striped (skin marking)
8"', white-thorax striped (skin marking)
52, striped 2 (skin marking)
Gr-alleles
III -chromosome
IV --chromosome
V -chromosome
+(1r, normal color (egg-shell colOr)
Gr, grey egg (egg-shell color)
Gr
E
, grey egg E-IG (egg-shell color)
Grl, light grey (egg-shell color)
Gr
B
, bird-eyed egg (egg-shell color and shape)
Y, yellow blood (blood color)
oal, oal mottled translucent (skin character)
Re, rusty (cocoon color)
ap, apodal (degenerated thoracic legs)
Ze, zebra (skin marking)
ze
r
, zebra faded (skin marking)
L-III, lethal III (embryonic lethal)
lem, lemon (body color)
Sm, small egg (egg size)
ts, tail spot (pigment)
L, multilunar (skin marking)
l-w, lethal white-rot egg
mal, larval malformation
sk, stick (body shape)
S'pc, speckled (skin marking)
37
pe, pink eyed white egg (serosa color associated with pink
eye)
ok, Kinshiryu translucent (skin character)
re, red (serosa color)
oc, Chinese translucent (skin character)
bw, black-banded wing (wing color)
VI -chromosome
E-allcies
+B, normal (body shape and skin marking)
E, plain extra legs iE-allelic group)
F
'
'', additional crescents (E-allelic group)
ED, double crescents (E-allelic group)
ED", double siars (E-allelic group)
Ell, II extra crescents (E-allelic group)
EKP, Kp supernumerary leg.s (E-allelic group)
EEL, extra crescents and legs (E-allelic group)
38
VII -chromosome
T. Yokoyama
EN, new additional crescents (E-allelic group)
ENe, no-crescent supernumerary legs (It'-allelic group)
ENP, Np, supernumerary legs (E-allelic group)
Nt, No crescent marking (skin marking)
Ma, trimolting (moltinism)
I_AI, tetramoliing (moltinism)
liP, pental110lting (moltinism)
brown egg-2 (serosa color)
kidney (egg shape)
F, f1esh (cocoon {'olor)
l-Il, lethal-Il (embryonic lethal)
Ys, straw (cocoon color)
bo, ordinary browll (serosa color, ordinarily inherited)
q, quail (skin marking)
Gb, green b (cocoon color)
1t, recessive trimolting (molt.inisll1)
oM, Bs mottled translucent (skin character)
VIII -chromosome
IX -chromosome
X -chromosome
XI -chromosome
ae, digestive juice amylase negaLive (digestive fluid)
be, body fluid amylase negative (blood)
st, stony (body shape)
I, yellow inhibitor (blooc] color)
If, sooty plain white (lacking fundamental gene for yellow
cocoon)
I-a, dominant chocolate (body color)
1Id, dilute black (body color)
og, Giallo As('oli translucent (skin character)
WI, white egg-l (serosa color)
II, wingless (wings degenerated)
white egg-2 (serosa color)
111
3
white egg-::l (serosa color)
Will, AOjllku white egg and translucent. (serosa color and
skin character)
oew, white egg translucent (skin character)
K, knobbed (skin character)
BlI, burnt (skin character)
XII -chromosome
I-10, Iethal-IO (embryonic lethal)
bp, black pupa (skin color of pupa)
mp, micropterous (small wing)
Ng, no glue (non-adhesive of egg)
C, golden yellow (cocoon colOr)
C', yellow inner layer (cocoon color)
ton, lethal non-hibernating egg
I'd, round (egg shape)
es, extra spiracle
ms, multistars (skin marking)
XIII -chromosome
XIV ---chromosome
XV -chromosome
cit, chocolate (body color)
1/, crayfish (body shape of pupa)
odk, E15 translucent (skin character)
Nt, no-IUller (skin marking)
U, ursa (bear) (skin marking)
UlJ" brown ursa (skin marking)
oa, Aojuku translucent (skin character)
Di, dirty (skin marking)
S'e, white side egg (serosa color)
Gc, green-c (cocoon color)
Other chromosomes represented tentatively by each of the following genes.
Bnz, black moth (body color of moth)
39
Ym, yellow molting (color of excretes of Malpighian vessels
while ecdisis)
bt, blind (skin marking)
Ill, lustrous (eye color)
Nil, naked pupa (sericin cocoon)
::ill, spindle-shaped (egg shape)
ge, ge01l1ctrid (body shape)
cts, cheek and tail spot (skin marking)
rb, red blood (blood color)
so, sooty, (body color)
otm, t-motted translucent (skin character)
N-Ns, new no-stars (skin marking)
nb, narrow breast (body shape)
4.0
T. Yokoyama
III. Cytology and Ueproduction
The number of chromosomes of Bombyx mori is 28 in haploid. The form
of chromosome resembles each other and each chromosome cannot be disc
criminated. In the case of silkworm, the genetical conclusion is only partly
supported by cytological observations.
The following items are worth exhibiting in the cytology of the silkworm:
heteropycnosis of sex-chromosome,
dimorphism in spermatozoa: eupyrene and apyrene,
difference in chromosome between Bombyx mori and H lnanda1"ina.
Besides the above, the exhibition of sLlch interesting phenomena as
mosaics, sex-determination or polyploid will be made under different. headings.
(cll r01110S0111es )
III-1 Diploid, triploid, tetraploid in spermatocyte and Oocyte.
The silkworm, Bombyx mori, has 28 pairs of chromosomes in diploid. It
is easy to count them at the first metaphase of the maturation division in
both oOcyte and spermatocyte. The eli vision takes place in an egg immecli-
ately after egg-laying and in testis during the fifth larval instal'.
Although polyploiclies have rarely been observed in animals, polyploiclies
in the silkworm have been very frequently reported. Most of them are
triploids and tetraploids. Only one case was reported of hexaploid by KAWA-
GUCHI (1938). Polyploids are easily obtained by means of centrifugation
(KAWAGUCHI, temperature shock (a high temperature, HASIMOTO 1933;
a low temperature, MUROGA 1948) and colchicine coating (I-IIROBE, 193n) applied
to yound eggs at the maturation division which occurs in one hour after
oviposition.
Polyploid first reported in the silkworm was the one which occurred
spontaneously in the heritable mosaic strain. TANAKA (Ul281 found a few
exceptional females of sex-linked inheritance in the mosaic strain and attri-
buted them to polyploidy. This was suggested also in cytological study of
the mosaic strain by GOLDSCHMIDT and KATSUKI (1928) through their obser-
vation that one of a triploid egg nuclei was fertilized by a normal sperm
nucleus. Later I-IASIMOTO (ln34) confirmed genetically and cytologically the
polyploid in the mosaic strain.
SATO (1929) observed polyploids of parthenogenetic origin. Polyploids
which occurred in the strains with chromosomal aberration were reported by
ARUGA (1939) and TAZIMA (1943). Polyploids sometimes occur also in progeny
of interspecific hybrid between B. mori and B. 1nanciarina.
KAWAGUCHI (1934, 1936, 1938) obtained polyploicls by centrifuging eggs
and studied the cause of their sterility which seemed to be a parallel pheno-
menon with the pc::culiar behavior of sex-chromosomes in the course of
gametogenesis of both sexes especially in triploid. His studies, together with
I-lAsIMoTo's, have largely contributed to the elucidation of sex determination
mechanism of this insect.
III-2 Chromosome in spermatocyte in the Fl hybrid of B. 11Wri and B.
11landarina.
KAWAGUCHI (1928) is of opinion that the domestic silkworm, B. mod,
might be derived from the wild one, B. mandarina. This is because it is
easy to cross each other and at the first metaphase of spermatocyte division
of the 1"1 there appear 26 bivalents and 1 trivalent, the latter of which is
reasonably considered to consist of three chromosomes, two from mon and
one from mandarina.
III-3 Heteropycnosis of presumed sexchromosome.
Sex-chromosome can not be discriminated morphologically in the course
ot gametogenesis of the silkworm. KAWAGUCHI (1(:)28) reported that one or
two chromosomes which showed heterQpycnosis were connected with nucleolus
of the nurse cell and that of the ovar:ian cell bel are differentiation into egg

and nurse cells. He suggested thaf'the chromosome might be sex-chromosome.
III-I
III-2
IlI-3
Chromosome and nutleus in
Bomby:x; mod and others
,I
1_<1. Diploid (n = 2R) in
slJermatoq'lc
Triploid (n:12)
in
I-b. Diploid in
{JDCytc
I-d. Tetraploid (n-56)
in spermatocyte
lri\'alent:
one from ""maarina and
two from mad
.
'Ii'
2. Chromosome in sperma[otVle
in thl!' F I o( B. m01'l ..: B. mando,-ina.
Heteropycnosis oC presumed sex.chromosome
C:J GJ
Nucleus in oocyte before Nucleus In The same The same in
differentiation Into egg nutrJii\;e cell in triplOid Lymantr;a
4nd nutritive cells in in. 8. mor; dis/Klr'
B. morl
E. KAWAGUCHI
T. Yokoyama
(Reproduction)
III-4 Location of gonad in larva.
Position of gonads is shown by a photograph. They are located uncler
the dOl'sal integument of the [)th abdominal segment.
III--5 Imaginal eliscs of reproductive organs: Ishi wata;s spots in female
and Herold's spot in male.
During the larval stage, male and female are distinguished with some
spots located at the ventral side of the 8th and the Dth abdominal segments.
Female has two pairs uf spots called Ishiwata's spots; one pair in the 8th
s2gment and the other in the !JLh segment. On the other hanel, male has one
Herold's spot in tlw median point on the border of hoth segments. These
are clearly in early sLage of the last larval instar. This was
llsed as criteria for distinguishing male ane! female, contributing very much
to the production of hybrid ep;gs.
nI--6 Pupae and moths of both sexes.
Though the both sexes are distinguishable during the larval stage, the
difference between the both sexes becomes clearer in adult stages. They can
be discriminated clearly not only by the external genitals but also by the
shape of antennae and .wings and size of the body.
III-g
III-7
I 1I-5
L GCloacb of die lOll va ;' u;t(ing
under the inll'gUntL!l1t Ilf fullowing
purti!lll. ,fr', /

3. Pupae, 4. Mut Ii:,_
(Ie/I) and mating
(rig"t)
uf the
pu::.tcriur .. d Ihe last
larv;l SUlI1I afler Illulling,
":l'A'fi in
'l' (It'/t) and gland in
:t; (right). Even in larval
if pnd 8 ilfl' easily dhilinglli::;hablc
widl imaginal dh;CR of
gon;hl. The d ... t:1 y has
lilrgc!y conlriiluL{!u lhe egg
pruducing indu::;ilry.
.,,01
5. lig. uf
Just before ovillositiuII.
1II-7 (!?)
be bursa copulat.rix,
lIb ductus bursae,
cls ducL. seminal is,
cit duct. tur(:uusLls,
gr glandula
gIll gla. I11UC[)!;;[,
ip intesLinus postcrius,
uri, ovidtlcttls p'cminl1s
C'onll11unls,
(IV iubus uvaril1s,
rs reccptcaculum seminis,
v vagina wilhout egg,
vt vestibulm with an egg.
1II-7 (c',)
aeld ampulla ductus
dcferentis,
dd ductus deferens,
ga gl. alba,
gl gl. lacteola,
gp :v;l.
gpl g I. pell ueida,
gs gl. spermatophorae,
]l penis,
t testis,
VR vesicula scminalis.
5 & G after S. OMURA
Silkwurm Genetics Illustrated 43
1II-7 Anatomy of adult reproductive system in both female and male.
Adult reproductive system in both sexes were carefully studied by ()MURA
(1938). He elucidated not only the anatomy of these organs but also the
physiological function of every part of reproductive systems of both sexes
for fertilization employing a technique of artificial insemination devised by
him.
lII-8 Dimorphism in spermatozoa; eupyrene and apyrene.
MACHIDA (H)29) reported of apyrene spermatozoa yielded in follicle of
normal testis mixed with eupyrene spermatozoa. The completed eupyrene
spermatozoa appear at the end of larval feeding stage and gradually increase
in number with age, while the completed apyrene hegins to appear from the
day of pupation. On the last day of pupal stage, the apyrene exceeds the
eupyrene in number. No apyrene spermatozoon is found in vas deferens and
any other outer part of testis. Therefore, it seems that they may be dissolved
at the basal part of the follicle at which the cyst membranes of eupyrene
spermatozoa are broken. In fact, the apyrene spermatozoa degenerate in
aqueous solution of acid, base and salt in a certain concentration, while the
eupyrene do not with the exception of t.heir cyst membranes which are more
or less dissol ved.
III-8. Dimorphism in spermatozoa
Bundlt' of in mille SpL'TIIl:thlWit l;umlle
reproductive organ in rf'pruducti\'(: I,Jrgan
A few of spermatozoa entering
an egg through the micftJpylc
MJ\ctuOA
44 T. Yokoyama
IV. Sex Determination
A sex linked inheritance was first 10und by TANAKA (191G). The type of
sexchromosome in the silkworm is ZZ in male and ZW in female. I-IASIMOTO
(1933) has ascertained that the W-chromosome nearly monopolizes the power
of determination of femaleness through the findings that even ZZW and ZZZW
individuals in the triploid and tetraploid statcs respectively become normal
females. TAZIMA (19M) has given further proof to thi:s conclusion by his dis-
covery of a strain which has a piece of an auLo:some bearing a marker gene
translocated on W. He has succeeded in finding the approximate locus of the
sex-determiner on W by using strains wilh deflciendes of various lengths and
involving different regions of the chromosome. Furlhermore, he examined
the part played by the Z-chr01110S0tl1C in the determination of sex by using
strains having various deletions each involving different regions of the chromo-
some. Thus, he has definitely made out that in the silkworm the determina-
tion of female sex depends solely upon the presence of the W-chrol110some.
whereas the power of the Z-chroI1losol11C in c1elel'mining maleness is apparently
much weaker. He has also demonstrated that l.he determiner of femaleness
is localized in a particular region of W-chromos(nue without being diffusely
distribuled 011 it.
The exhibition is arranged as follows:
1. The chromosome maps of Z and W chromosomes.
2. Diagram of the mechanism of sex-determination in the silkworm.
The chromosomal constitulioll of both sexes had first been genetically
presumed by TANAKA in 1916, namely the female is ZW and the male ZZ. It
was, howcver, very c1ifiiclllt to identify cytologically the sex chromosome.
Several morphological gene loci (seven loci) have been found on Z chrolllo-
some, but on W chrolllosome not a single morphological gene locus has been
known yet. By the irradiation of X-rays several kinds of deficiencies have
been induced on Z chromosome. Males heterozygolls for any of these defi-
ciencies can sllrvi ve, while hemizygolls females for the same deficiencies are
VI-I. The maps uf Z and W chrumusome
z
II I I I I Lm
r., It Vg (Jrc n( I-a Ge
0.01. 6 H.O
:-lG. 4:38, 7 46.4 49.6 . (Ior:us unlmown)

female determinor
'------_..,.-_____
U W Z
45
not viable. Therefore, even a small portion of Z chromosome is considered to
be necessary to maintain the normal physiological function in the female,
while W chromosome seems to lack of all these functional genes. Thus W is
thought to be physiologically useless.
In 1933 HASIMOTO obtained a very interesting result on the role of W
chromosome. He exposed the F I eggs from the crossing of Z+ IW l' X ZOd IZ
M
0
to a high temperature of 40C. for one hour immediately after laying. Among
F I individuals he obtained several females exceptional to the rule of the criss-
cross inheritance of sex-linked ad gene, of which the most remarkable were
triploids and tetraploids.
As he could not obtain tetraploid male, all of the tetraploid females were
left to mate to diploid male (od/od) and they gave only triploid individuals
in the next generation. Though segregation of + and od was observed with
the back-cross ratio of 1: 1, the sex ratio was altogether different from 1: 1
yielding more than 80.% of female. The result was explained by HASIMOTO
by postulating a preferential homotypic conjugation among two kinds of sex
chromosome and existence of a strong positive female determining gene on
W chromosome (lV-2). Thus he first emphasized the role of W chromosome
in determination of the sex of the silkworm. There were, however, no methods
to prove this, because all of the offspring were triploids and abortive.
IV-2. Abnormal sexratio in triploids and presumption
of female determining gene
.
AAZZ AAZW
----.: __ !f
0t AAA;WW high treatment
gametes
4n female
meiosis in 4n female
Cd
AAZW
+
AAZW
"
sperm 1
"".
fused nuclei of egg
and po lar bodys
AAAAZZWW
1
+ +
AAZWW AAZ
---od--
'4
AAZ AAZWW
1 1
In offspring from 4n !f x2n(od) S 1
rood od+ od+
3AZZW 3AZZW
!f I'
4
3AZZWW
'i'
1
3AZZ

3AZZWW
!jl
""
3AZZ

1
46
T. Yokoyama
In 19,11, TAZIMA discovered a translocation between W ancl II chromosome
in which W chromosome was marked with two noticeable genes tor larv,ll
charader locating on the I [ chromosome attached at one end of W. By using
this translocated W he verified clearly lL"SIMOTO'S view and concluded that
the role of W in sex determination is cledsi ve () V--3).
IV-3. Verilication of femalegene presumptiull on
the W chromosome by using lranslocalion
os
[l_-==:I
r:= 1.1
ad
+
u ____ .::::J l::::::--=
os od os ad
+ II fr':itl, os!fl, + 11 P,'ia I os, + 6 +. oJ Q

fi( __ J
C=:::-=:-..::::J
+p

i _______ =
. .
........ .......... ...
_,---------.,
nChcmi.l tlf lllUlui.d
lranslucatiun
The relat.ion iJet ween the ratio of s('x chrom()s()mes to tlte llumher of set
of aut.osomes and sex expression was studied by HASIl\(OTO, KAWAGUC[[I and
'L'l.ZII>.L'I.. The results are sLlmmerized ill the Tablc (IV-,ll. Whenever W
chromosome is present the individuals becomes invariably female, while its
absence always prom iscs to male. It is, therefore, clearly known that
female is det.ermined by the prescnce of only one 'vV chromosome
of the number of autosomal sets or of Z chromosomes.
Apart from the role of W, the effect of Z on sex expression should be
tested. Partial duplication and deficiency of Z chromosome were examined of
their effectiveness 011 sex determination by using several strains, but no sexual
alteration were observed under any supernumcral or deficient condition of Z
chromosome CIV-5). Thus it was concluded that sex is decisively determined
by the presence or absence of VV chromosome in the silkworm.
Silkworm Genetks
47
IV-4. Sex determination in with special reference
to the ratio of sex chromosomes to autosomes
ploidy
2n
3n
sex
chromo.,
SDme
_fragemcIlt
left
deficiency
-m-iddle-- ---
(leletioll
right
deficiency
number of
autosomal sets
All..
AAA
AAAA
number of
Zch.
zz
Z
Z
ZZ
ZZZ
ZZ
zz
ZZZ
zz
number of
W-eh.
W
W
WW
W
WW
sex
expression
IV-5. Effect of partial duplication or deficiency of the
Z chromosome upon the sex expression
bypoploidy hyperploidy
n-ull1ber
of i
numher oT
strains famale male strains famale male
tested
tested
no change JlO ch,inge no change
9
dies in sex 3 in sex in sex
expression expression expressi(ln
5
dies
no l:hange nll change
,1 dies
2 in sex in sex
expression,
expression
(Vg)
dies 01.\"
Suggested by 'l'_"'ZIM.\'S case of translocation IL%IMD'l'o attempted to induce
translocation between Wand one of the bearing noticeable gene
Zebra. The procedure of his experiment is illustrated in IV"-6.
In Dreier to find the location of female determining genes on W chromo-
_,.--.".
some, Tazima obtained complex translocation Z-''''{ 'vV +1'.pS", which is probably
,..--.... ,,--..,.
a product of crossing over between Z and W in a W.+
1
,.psu/Z,od female (IV-
I) IV-7). If this assumption is correct, W chromosome which has lost a part
48 T. Yokoyama
IV -6. Experimental induction of translocation
involving Wand an autosome
X-ray
irradiation
in pupal x
, stage
'?
zebra
o
zebra
normal 1311)
o
untreated

Q a
zebra od-transll1ccnt
aberrant (I)
oil iebra normal
8
x
O
o zebru-od normal
I X
o
normal

zehra
if
zebra
(,
normal
(j
normal
by crossing over still retains its female determining power (IV-7--11.
He also obtained several mosaics and exceptional types [r0111 the
,.---....,.---.... ,.---....
Z+W+I'ps!tfemaleas dissociation product by X-raying. The mosaic has
,.---....
a _1_
11
.pSa constitution and simultaneously was a gynandromorph, which was
,.---....,.---.... ,.---....
considered to be Z'I. w -J-!l.psa to start with, but the mosaicism was brought
,.---....,.---.... ,.--.....
about by the elimination of w. +p .pStL or Z'I-. W parts in somatic cells
(IV-7-2).
IV-7. Experiments to determine the location of female
determining genes DO W chromosome
(11 rnduction of complex translocation

!- n __
p - :;-...__
"'-=
translocation 01 1I11 to W ---.

==0

+0$ .,.fI od
.P .
rmalI!mlI.
;,01 TO 'for}
high temperature Ireatme"t
(2) Dissociation of complex chromosome by X-ray irradiation
//X.ral's

./J ",ad
=

left side
dissociation
observed
number
23917
=
p,Dd
-------..
9 .3
7
mid right side
deletion dissociation
.t..a
other
types
L1JIIJJCl mosaics for
.P fr".ad I .p "".od .P ",. 'and ,00
T'?
10 iJ
Wl1ll
IV-9_ Functional differentiation among regions of the W chromosome

- III -v Z Z 1lI III
Z III \ oJ ttan$lucentl
.IIT
Will 7. III lIi Will (. rrrwm z Z IllWIll 7. Z III III
zebra od od !f. I
(non. lethal strain) (lethal strain) .;lIe normal q
49
50
T. Yokoyama
Fig. IV-S shows gynandromorphism in a gonad, obtained in a complicated
mosaic female, which probably originat.ed from partial elimination of complex
translocated chromosome involving W. In one of the follicles of a testis of
this female a mass of oogonial cells is seen which is developing independently
from the surrounding spermatocytes, bu!: transitory forms towards the opposite
sex are hardly visible.
As for the functional clifferentiation among the regions of W chromosome,
the following exhibition 'vvere made. When a certain part oJ W chromosome
exists in ZW or ZZ, constitution lethal effect is seen either in t.he female or
in the male.
a) (W) ZZ is lethal, while (W)WZ is viable (IL,.,.sIMoTn) (IV-D).
,.-..._ ".--.... ".--....
b) ZI-.( ).+l'.pS", ZW is lethal, while Z-I-.( ).+)'./;8" is viable (TAZIMA).
Here (W) or ( ) means a part of chromosome originated from W. The former
still contains female genes while the latter does not retain female determin-
ing genes. Hence, the female determining genes do 110t seem to he distributed
evenly over the whole W chromosome but considered to be localized in a cer-
tain section.
IV-S. Gynan(lromorpilism in u gonml of a
type-mosaic induced by Xray irr,ldialioll
.'
._

I
".,
t;,,,, ; ..
.';:1;

'.
"
:/
Explanation of photomicrograph
,
l

' ,
,,,.,,,1'

, /
I
.'
J
':'1
'1

.,-:

This picture shows gynandromorphism of a gonad of a complicated mosaic female.
probably originated from an egg in which elimination of complex translocated chromo-
some, involving 'vV. occurred partially.
In one of follicles of a testis of tflis female a mass of o(jgollial cells which are de-
veloping inclepenclently fr0111 the surrounding spermatocytes il> seen, but tn\nsitory
forms towards the opposite sex are hardly visible. Y. TAZIMA
The mechanism of sex determination in the silkworm IS different from
that in Lymanlria dispar, which was thoroughly studied by GOLDSCHMIDT.
Contrary to Lymantria with its abundance of inter!lexes, no real intersexcs
have been discovered in Bombyx, but all abnormal ones in relation to sex are
found to be gynandromorph.
52
V. Induced Mutations lLlld th.e Practical Application
A mutation was induced in the silkworm for the first time by HJ\SIMOTO
11929), treating eggs and pupae with high tempei-ature, X-rays, centrifugal
force, etc. Afterwards numerous investigators obtained various mutants by
using various methods. The silkworm has been proved to be a material in
which mutations are able to be easily induced and 32 induced IIlutants have
alteady been reported.
1. Mutability in relattQtl to vario\.\s agents.
I) X-rays
The. sensitivity to and mutability by X-ray irradiation of the silkworm
are different according to the developmental stage and the sex. The change
of sensitivity to X-rays of female and male germ-cells is shown ill LD50 curve
in V-l, which shows that female is more sensitive than male in pupal stage
(TA,ZIMA, 1951). Sensitivity of varies also remarkably according to the
developmental stage of germ-cells. Figure V-2 shows lhat male germ,cells
are quite sensitive to X-rays before meiosis, but they become fairly resistant
after meiosis. The difference of mutability in various stages of gametogenesis
was studied by TAZIMA in specific lod and was made clear Lhat mutability
is highest just before the first metaphase of maturation divLlion both in
spermatocyte ancl oocyte, while in gonia mutability remains at rather low
level of frE!quency (V -3).
V-1. Sensith'ity to X-ray of male and female germ-cells
ill developmental stage
2'
15
-

to kill 50% oi embryO)

....
'.
5 6 7
ngc o( pJ.fcnt pupae
10 dOl}' old
emergence
..:\!{i'=> \<;>. $tlg't!
BOth
It

D
>
0
.:

:
t1

_,
o.

, 6
age of parent pupae
Silkworm Genetics Illustratecl
V -2, Xray; Change uf of male germcells
before and after meiosis
%
10
tn:utr:tl <tge and
1st fl'cding
/
,/
,
'
,:
,
.
,
.
.
.
.
agc of the 5th larva
DI.H'clopmcllt of germ-cdt" in !:ith stadium
'>Pl'rm3l11gonia Sllcrm.atocyte spermatid
50
100
m:nuring
100
53
V-8. Xray; Mutability of germcells in
various developmcntal stages
clocs rate: 8t4 1 per minute
V-4. X-ray; Difference in sensitivity and
mutability among different strains
luti: and ch
femal"
GO','
I
.
.
I
.
"
I
I
(lId
fi
100
t' GO

40
"
'0
20
2.U
1.0
Mortality
rb ,/ 9 1.5 days afler pupation
3 dar old, 5th inst?

, ............. .
1(}()(J 30(1()
TOW r
;;,,{i_ (II immediately before emergence
-t-/
1000 30UO
dosis
54 T. Yokoyama
A remarkable difference of sensitivity to X-rays was also found among
the various mutarit strains of silkworm (T AZIMA, 1955) (V-4).
2) High teniperature
When newly laid eggs are treated with high temperature (40'C.) for one
hour, various abnormal worms are to be obtained at a remarkably high fre-
quency. Among the abnormal worms there are mosaics, polyploids and those
expressing reressive characteristics w11ich are in non-treatc(l casc covcred by
the allelic dominant. TAZIMA compared the frequency of those abnormals
according to the age of eggs treated, and obtained the results as sh6wn in
V-5.
The appearance of recessives in this case are presumably clue to the ab-
normal fertilization of sperm nuclei, i. e, fertilization betwc'en two sperm
nuclei without necessitating egg nucleus in c()njugation. This was explained
genetically by HASIMO'fO (1934) in a very skilful way as shown in V-5,
V -5. High temperature
It's effuctivent::ss and specificity! l3y keeping newly
!ai(l eggs at I[O'C., 1 hour, 80 nlHny mosaicH.
recessives and polyploic1s are yielded!
Frequency of appearance (Tuzima '47)
Elucidatiun of dispermic merugony (Hasirnoto ':H)
e

I "'" s
40"C ] hr ;;;osaics

+/od

Exreptjoocils
od/oil

EKP/EKP E"'/t- +/+
o all 0 1 : 2 : I
3) Centrifugal force
By centrifuging 0-3 hour old eggs, polyploids are easily obtained, as a
result of the disturbance of the behavior of polar nuclei. This was first
studies by TANAKA and KAWAGUCHI (1932) and later by KAWAGUCHI (1934, '35,
'36) in detail (V-6) ,
V-G. Centrifugal Force
By centrifuzing 0-3 hour old eggs lots of polyploids are obtainable
PM+lpl'J? 1; (centrifuged)
expected
4n (21. 6.%')
3n (67.7.%')
2n (10.7%)
Mechanism of 3n formation
1 Egg+l Polar body+1 Sperm
(1) PM+/V/PS+
(2)
(3) pY/pr/ps+
4) Colchicine
1 Egg+25perms
(4) pl"/p' +
(5) pJl/P'+/frS+
55
HIROBE (1939) succeeded in obtainin!t polyploid silkworms by painting
dilute (0.05-0.496) colchicine solution on tWe surface of the eggs within 25
hours after being laid. Polyploid individuals were obtained as mixoploids
with diploid and polyploid parts (V-7).
V -7. Polyploid silkworms induced by colchicine treatment of eggs
colchicine hatchability number of moths which
(%) C.%')
laid polyploid eggs
U.4 \l 19
.. ______ , __ ..
U 'J 46 6
0.1
C'J
3
U.05 72 4
control
go
0
-._-.-._- -_-..__ .. _
Polyploid individuals can easily be obtained by colchicine (0.U5-0.4.%')
treatment. of eggs within 25 hours after laid. But, progeny test shows
that these: polyploids are mixoploid, containing diploid and polyploid eggs
in an ovary, T. HIROBE
5) Nitrogen mustard and other chemicals
Nitrogen mustard is also known to be effective inducing mosaics and de-
ficiencies in the silkworm. Nitromine and N-bis alanine are also effective for
this purpose (NAKAO et aI, 1952). NAKAO (1954) also tried to examine the pro-
tective action of cysteine and mercaptoethyl amine, which had been ,known to
56 T. Yokoyama
V-B. of incluCl)c\ lllutanls
New gene mutations
Symbol Mlltation
Ac Abnormal l:lJ]'sekt
A/) Abnormal appdih'
As Abnurmal
Dr Brow11 (lIlurking)
Bu Burnt
Df
Dominant llwarf
Eel/,
Additiollal
E-Lg Epislal.ic g(Clle [u 1[,g
Icr
ILg
nUl
MI)
pll
j) I.
PS(l:.!
S'/IlU
,<:)lP
SI
Spc
Crcsn)nl suppressur
Inhibilul' of Lg
Light gray
mLlstache a
Mustadw b
Venlral striped
Light
Sable (marking)
Sable (Illarl:ing)
Pale
Dominant small C.'m
Wh i lct s t r i )led
SeCOll!\ stripc.c\
Supernumerary legs

PIWl1ulYPl)
Ll'thal
Ldhal
Lethal
Lethal
Lethal
Ldhal
Ldhal
by j'S"
ddiciellt
lIypm;l.at k to ",/,
Lethal
Ll'lhal
Ldhal, ([e.ficiency,
semi sterile, esp. l;
Leihal, ;;l'crill'.
Author
Aruga '3D
'Tazi lna "!i2
i\ruga '.1:]
Axuga
i\ruga ',1:1
C(/ of SU7.uki '8H
lIasiilwll) '41
"'iiI' uf '47
TatilllOl '1:\
Sak:ll.a '1:1
'17
Pf (II' Chikushi
Tazi llla ,,).,
Takasaki '17
S-fI ut Takasaki ',17
J\.ruga '4:-3
Tazim<l '!i2
Swi Short \ving
Chrul1lllsumal alJl'.rralilill Takasaki '117
Swl Swollen (!Judy shape)
Lethal jJsw oj' Takasaki '47
Tb Transparent back
Lethal
i\rllga '111
Vestigial
Sex-linked lcihal
ya yellow antenna
Tanaka
1. Fragmentation
Rusty
od-mottling
os-mottling
S-mottJing
M-mottling
Ze-mottling
2. Deficiency
loch.
II-ch.
III-ch.
etc.
3. Inversion
pv
Sa
4_ Duplication
V-IO. Lists of induced mutants
Chromosomal aberrations
Kawaguchi
Hasimoto, Aruga, Tazima
Morohoshi, Tazima
Tanaka, Kogure, Aruga, Takasaki
_-,----_.- .. "'"----._ ... --.
Aruga
Aruga,
Tazima
Kogure, Takasaki, Tazima
Aruga
Tanaka
Tazima
DS-2, DS-3, paM-I, paS-I
Aruga
TI02 Tazima
5. Attachment
rfh
G. Translocation
pi-mottled
T CPS)
T (PJ!)
T (Ze)
T (II-W)
Takasaki
Tazima
._---
T (III-W)
T (X-W)
T (VI-XIV)
Hasimoto
Kawaguchi
Aruga
Aruga
- .. _ _-_-._0'- .. _. ____________ .. ____._,,,_. _____ . __ _
Tazima
Hasimoto
Tazima
Itikawa
57
58
T. Yokoyama
be effective in other organisms against radiation effect. His res\.!lts, however,
were negative. A fairly positive result was obtained by him only with pyna-
cryptal yellow (V-8).
v-s. Nitrogen mustard [lnd other chemicals
Effective for inducing mutations
Ni trogen mustard (Nakao '!i[))
Nitromi ne (mcthy!-bisi'lchlurllcLhyl'>1ml ne N'llxitle) (Nakao et al. '52)

CII'J-N"
' I "CII"-CI-I
2
-Cl
o
Nitrogen mustard alanine (Nbis-,l dtlorocthylalaninc) (Nakao ct al. '52)

HOOC--CH-W
I "'-CH
2
--CIl"--Cl
CHi!
Protective against the mlltagenic action oj' radiatiun
pynacryptol yellow YICS (Nakao '5,1)
lllcrcaptocthyl amine No (Nakao et al. '54)
No ( )
Gene mutations and chromosomal aberrations induced artiftcial1y in the
silkworm are listed in V-9 and V-1U.
The summarized 1 ist of spontaneous ane! induced mutatiuns is shown in
V-12.
V-12. Frequcncy of spontaneous und illduced mutations
charac:lers
kinds
I
, I pupa &'1
;cmbryu, larva ! C.UCUUll I lIlul It '
tutals
spuntan(!otls duminants
f)
1-7*
I
() <g)
Hi Il
,1 92
!
recessi ves H1+S* 1 :22 GO 2 11 1 lUi
induced dominants
,1
1
U 21 0 1 1 27
I
recessives ()
5
toLals
6 240
.. .. --
*
genes concerning to hibernation
An interesting mutant is especially worth mentioning. TAZIMA (UJ53) ob-
tained 32 mutant silkworms among 1,126,414 individuals, the parent, either
female or male, of which were treated with X-rays. The mutants took the
leaf of beet and other plants as food which normal worms never did (V-lla).
They show abnormality in the structure of maxilla, especially in the sensory
hairs on the lobe, which is supposed to be the food selecting organ. It is,
however, difficult to rear the mutants with leaves other than mulberry
throughout whole larval life probably due to the inadequate constitution of
leaf. V-Ub shows cocoons spun by the mutant individuals fed on leaves of
RyuzetslIsai, Soy bean and red been etc. during the 5th stage. Selection on
the frequency of appearance of mutant individuals gave remarkably positive
result as is sbown in V-ll.
V-l1 (a) Larvae, the mutants, are eating the leaves of
Alianthus g{andulosa :and beet
V-ll (b) Cocoons spun by the mut,tnt "abnormal appetite" individuals
60 T. Yokoyama
year
1952
1953
V-ll. X-ray; Induced mutation in food preference
number of larvae fecl on beet leaves
number
of
;lctuul
FI
2nd
i
3rcl '1th rill!
examined
instal'
I instal' instal' instal'
no. of
I
I11U tants
937, 916
f) II 5 ri 11
198,498
3S;1 :cllJ (ill :-H.l:1 :H
mutant
HC:"lUII nf 1nr 'I alJnol'lllal apPl'litc"
Y. TAZIMA
V-18. Folypluidy
a) Effective agents fur imlLle'ing pulypluidy
1. Centrifugal force Tanaka ct Kawaguchi
2, Cilichicine lIirolJe ';J9
High temperature
,1. gene
lJ) From ecunomic sland [JUill\:
Uasimuto ':'],1
llm;imulu ':H
II enlarges cldl size hul' cietTI!a:;e:; cell lIumbl:r
It's 1101 pl'llIuising frOIll t't'l)lll)lI1i<: stand puiut
freq. of
appearance
O. 001
0.183
------_._--_
cell size
ploidy
weight of
]Jupa
I 11 lImher
1,t sllcrlll<lto- i
L',y te C\OO I
211 1. (i,j em"
311
4n
2. Polyploidy
:-ii I k-gl<llld
cell
0.77 mm
1. 03
1. 4!J g
1. riO
.
' . silkgland
l,'ell
(iHU. ::1
I
370.0
E. KAWAGUCHI '36
Though colchicine is also effective in inducing polyploids in the silkworm
as in pther animals and plants, far more effectual method in the silkworm is
centrifugal force (TANAKA and KAWAGUCHI, 1932) and high temperature treat-
61
ment of newly laid eggs (H.'\.SIMOTO, 1934). Polyploids as well as mosaics and
gynandromorphs arc also easily induced by the action of a recessive mosaic
gene rno ElLU) (V-13\.
In tripi'oid and tetraploid silkworms, the cell size is larger but the num-
ber of cells is smaller than diploid. Consequently the silk productivity is not
expected to be increased (KAWAGUCHI, 1937).
3. Practical application of induced chromosome mutation
The powerful s2x-determining function of the W chromosome was appUed
for practical purposes by TAZIMA. As in Japan an the farmers rear PI hybrid,
the discrimination of sexes is a very important procedure in the commercial
egg production. Usually the sexes are discriminated by the tiny spots on the
ventral side of the growt1 larva. T AZIMA obtained a strain in which a frag-
ment of chromosome II, which carries dominant! genes for noticeable character
is attached to the W chromosome. In this strain, all marked larvae are female,
while all non-marked larvae are male; thus the sexes are easily separated.
The females of this strain, however, were not hardy enough owing to the
)lyperploidy of the translocated part. TAZIMA attempted to remove this defect
V -l-J.. OrigimLl sable transll1. (W+I1.p8<') ,
62 T. Y O[\0YUlll<l
V-V1. Practical applicaliull of lral1s1ucalioll in the silkwurm (1)
1" +(111
IV __
z_ ......

<1>'1

z.".,""""
1-';[
\\r,Ptl'f] (\v. ; 11 i r 'F:" ! .'
X'LI\' "

rt:"I.,(,1.

.-. ..,..,. "_..,.",,
M-strain

II
,I, , ,
\\1\'\ 1 I"liJli:lrily tYIll:

IV_ ....... z=_
-l"zEZ:'tCiI-
\\1. : j1 i J' i ",1 ',I jJ i J'oJ ,Il
111 I I'rld ','
Pre:,ulIlcll ell ['dlil',J.>' I illL' (011,,1 rud i. III
W....".....;".- IV""""";;_
z= ....... '1.= .......
w--
Z= .......

I-;f. of \Vl' :;[ r.dn
(prilllHi1.v ilJJprtJH:d lYJlll)
:-1

..1
711:
r.1 hI 1/"", ndY:,

-'---
jJ! ill1i!lT f;tJl'oflriaql r1f1f!Il;d
I Yl1eH t
CllG
Improvement finished
Trilnsferred heronftcr to
brt:cd pnH:tical by
to tJu(JulHf
Y. TAZIMA
Silkworm Genetics 1llustrated
63
so as to be used for commercial purposes. After repeated irradiation of X-
rays, he succeeded in eliminating the superfluous part of chromosome II, leav-
ing only a small fraction of the chromosome where marking gene located.
Thus the strain became to be utilized commercially (V-14l.
The eggs laid
T. Yokoyama
TAZIMA wa;; not content with this, but took another forward. He
intended to make it pOl;sible to separate female alld male egg;; by the same
principle, because the male worm cOlllparati vdy larger amount of
silk than the female. By utili;.:ing same procedure as hy HASIMOTO (IV-
6) he could induce translocation in which W chrumosome was tagged with a
fraction of X-chromosome involving the dominant gene to white egg color
IV-I4).
In this strain females come out of the Illack eggs while ll1ales hatch from
the white eggs. lIe alRo in cooperation with an engineer, an auto-
untie machine for eggs of strains with photoelectric tubes, which
enables to discard less useful [emale eggs bc[un.' halchillg' IV-IS, V-IG).
I
V 15 Practical application of tran:;locatiol1 in the silkworm (2)
uf eggs hy tlw color
,
\\' c::=J ....

, ,

,. r::::=:.:. 1::::=1

WIll
WL":..:._.=",_/c:::=I_

',,1. , ,
WC_;--'l :'_' I Ic::::::.J
?l. -;,1/'-_',1---' ,', . '},::.:-.-l
I.) "'/ (I I
r.Jllp1'r
(.'I"",II",I"IJI.!
[-._"1
i:f-::-:-:-l r_--
'--'
w c=::rda i
2c::...J c=:J?c::=:It::=:l
lV" .) 11',
\\,11111
6.7
U
Y. TAZIMA

65
V-16 ea) Automatic sexing machine for the eggs
V-16 (b) Reverse side view of_the_machine
Y. TAZIMA
66
T. Yokoyama
VI. Mosaies
TOYAMA 119(0) 111"8t descrihed a mosaic silkworm in detail. Aflerward a
recessive gene 1/1(1 was founc! by GOJ.I>SCIlMIDT & KATsuru (137) in the pre-
sence of which mosaics easily obtained. Recently the mechanism by
which various kinds of mosaics and ll1()ttlings come nut has been explained
and the method of ohtaining mosaics has been improved to a rcmarkable ex-
tent contributing much to the advancement of silkwol"ln genetics.
For this exhibition, the following arrangements arc made to show:
1. Mosaic clue to 11U1 gcne.
Mosaicism and
KATSUKf anel AKIYAMA Wl37) [ound a heritable mosaic strain in which a
number of mosaics lor hoLh :--lOmatic and sexual characters appeared as they
were mixed with the normal im;ed" in Olle batch, l\ATSUKI worked further
with GOl,nc;cIIMlllT (1 i'l on a mock of occurrence of the mosaicism
genetically and cytologically. The is causc.d by one recessive gene
!}lO possessed by mother insed. In cytological observations of the strain
homozygous for mo, it was founel that 1 wo egg' Iluclei appear in egg plasma
and each of them was fel"tili;wd with a sperm nucleus. Therefore, it was
assumed that each fertilyzec1 nucleus might develop into each side of insect
body. Somatic mosaics:are expected to OCCUI" ill one of the crosses: J. mo,
A/a><a/({; 2.1110, il/Ic<!l/({; ::. 1J!u,a/a:: .1/a (el alld (( represent some allelic
genic symbols), Further, mo, is a gene I>y Ilat Ufe, hut acts as domi-
nant in mUltiplied conditioll, as by \l\),11), in tetraploid
VI-l Mo"ail:imn and p;ynandroJl1urphism
nil!' tn a Gl'ne tU{)
.... " , .
L Tllldl'll', ,n\1] Plll;11 1111111' _. Hd.ltl'r:d 111"'"li .. " f(,,. p.v
C(Hljlq';lli'd u;,ll 'il'I'fm rl"'I','(II\,._-I\ :\1111 fl . h\' mo
in o( "1(1 :.\ rcllll
,1. Bilil\(,'ral for 1. ,Ifill +
K. KATSUKI & R GULDSCHMIDT
VI--2 Mosaics of markings
+Pcil-P' C/.
+r:h ell
pZe_p+Z, I-I. CI!IKUSlil
VI-3 f Mosaic of silkgland color
I T
0
11W,P"+Y'PY,C p+'IP+r, C
Gl
mottling
mOl mosaic gene
pll, moricaud pattern
p,
]" yellow blood
+ Y, cDIorles3 blood
C, deep yellow cocoon
"'. @.Osho\\'ingbltJod color
67
M. HARIZUKA
Though the blood color genes arc clearly identified from those of the cocoon colors
as shown in other figs., the Yl!llow blood gene, Y, which enable to be permeable the
intestinal mucosa to carotenoids into blood circulation, was proved to be also concerned
in permeability of silkgland wall to the pigments from blood into silkgland,
fiR
T. Yokoyama
fcmale JIIo/mo/1/l() I which produced mosaic offspring in spite of posscssing a
dominant gene +.
VI-2 Mosaics of some larval markings.
VHl Mosaic of silkglancl color.
y, yellow blood, and I 1', eolorless, eontrol lhe permeability of the intes-
tinal mllcosa to caroLenoic1s and are in linkage relation with JJ
M
, llloricaud pat-
tern. Mosaics in the silkgland obtained in the mo,
all of which showed nlOsaieisl1l also in larval markings, JIM and j), and were
pale yellow in blood color, i.e., illlernlt'diale betweell 1
r
and +. In mosaic
silkglands there were two kinds of c('tls, yellow and colorless. This indicates
that Yand _IX, which control the' permeallility of intestinal mucosa to
tenoicls, arc also possible to control that of tile silkgland (llAIHZtJI(A 1940, 1948).
Mottling due to the chromosolllal fragllll'ntaLiol1.
strains of \wn' ohtail1(!cl by treating with x-
rays Llw pupae' heicroxygolls for normal line! iranslUCl'nt genes. As the chro-
mosomal fragment carrying a normal g('I1(' located on the Z chromosome is
deleted in certain cells of epidermis and otbel' organs, so tho allelic recessive
translL1cent gene exprc'sscs its action, while other cells when' no such deletion
occurs show normal appearance, making the pattern" mottling"
(VI-8).
VI-8 Pleiotropic function of od genc'
The od-mot[lec1-transluclmt (OIl"'I) larva charactt'rizl'c1 hy the mosaicism of
normal and translu(,ent skin is due to the eliminatilln in sO!l1atogenesis of the
fragnwnl carrying 1".1 gene. 1n this larva vitally stained
with neutral reel, all of the ()rgans andt of ('C(oclel'llllll origin C'pidennls,
Vl-H I'lvintrupic fUllc! ion !lr lid gVIU'
del1wm;tratvd liy vit:li
nUll t t ril
SIUII!efi whh
11I'utl';11 rt:d
prothof{\clf.' uland
H. AkUOA
silk glands, Malpighian tubules, prothorasic gland, hypo-stigmatic glandules,
sali vary glands, ocnocytes, and others--were stained in mosaic pattern, where-
as those of mesodermal and enclodermal origins were stained homogeneously,
This fact shows that ad gene expresses its action in the cells of almost all
VI-5 (a) S-mottled due to the elimination
of the fragment carrying P' gene

sC;'l1,reg:il.:d (rom
muttled-'tripod
II CHuwsm
VI-ri (b) MOltling due to elimination of chromosome
!jtralnli, A,
U. Jnll!fnJel.lilllc;
C, D. liilutcral mosaic
A B D

In\egumllnt of Smottled
wllitL' scanty
Hdl(l\'hHlT III :iupl)rnUmerary chrom()son1(! (S)
during- till' milluration division
li, !.;upernuIl1Qrary chrommwmt!
A c
t, TAKAsAKI
70
T. Yll!(Oyalllll
organs and tissues of cl'ioclerl11al origin of the silkworm larvn, and the cells
of organs and tissues originated [rom ectoderm have lhe same cytoplasmic
characterisLics necessary for tlw of ()d-oily gene (AIWGA, H142) (VI-B).
Vl-ri Tile nnd of
in t.he distl"iiJull'ci
in examil1l'd an'll fWO
(While
Frl.'lllltmcy (If the (dimillHI ion o( the
:;upl:nILlllll'rary cl!ronwsomc in
I.(L'I'1\1 ('('lls lIf rliffl!)"cnt sexes
(White
5U
.iI
11\1
ih'I'I',I/t{l' \'-'.lhll"" 01 the
I.:limln.u lUll
....... \;t 1 t U
,.../ .............. , - ,', l"i.i.t,II'J:i '
...............
' .
.....
II .... '_""";:""'l'-
Vl-!i Elimilwlin\l Ill' fragnlt,'111
Fruqllul]cil'!{ (If l'lirnilltlli(l1l of tilt' SlIIWl'l1l1IIH.'l'1II'Y dll'OlllO:ionw,
jJN fraf\llH'IlI, in lilt! dil'fl'l'l'lll ('UIlHtiluliolH;.
(Sm) and (]'), ,("l'I'I':1111 t'!II'Otlln:;ullws Ill'oclll('('(] Ity hrt'nkilll{.off
or till' 2nd l'1l I I, till! wlio!(' rhrOllHtS(ll1It',
l'111'0 1) 1(\:;( Illlal ('ow;L i t (( t inn
(Sm) / t I / I I
-
(Sm)/(Sm)/ I 1/1 I
-
(Sm)/( n/ I I
-
-
-
(Sm)/(Sm)/( nl I I
-
-
(Sm)/ I I
-
(Sm)/(Y)/(l')
-
ttV(lcaR(' vnlu('s of
t ('I i nti Ilal inn
11l'1'('1'111 :t!!;l'
'I'
.U'i.O."()
!I.D
1).0
Il,O :t .[
T. TAKASAI\l
A few kinds of striped-mottled were obtained by treating with X-rays
the pupae heteroqgolls for striped marking (ll) anc! normal. This kind
of mutant is characterized by the appearance of numerous white patches in
the black marking and is clue to the elimination of the fragment carrying
the dominant gene jJ" on the .second chromosome in the epidermis, making the
appearance of allelle recessive gene /_]I. The freq Llency of elimination of the
chromosomal fragment is different according to the struclme and behavior
of chromosome. It is high when the chromosome does not exist in pair, as
in trisomic zygote (Vl-G, VI-G, VI-7),
VI-7 Elimination of chromosome in trisomic zygote
Elimation of the whule 21ld chromosolllc during
gamcl.ogellush; in trisomic ;lygule!:l, jJ'/ Y / I-''"!
sex
cl!mllHlliLJll percentage of each
of 8 chroinosllllws
total
p'
y f"fJa!
2.11% 4. !J. 8.9%
7. u 8. 1 7.1 22.2
Trii:iomic mmmics and their
11 ..

3. Mottling due to a mutable gene.
P'
32
o
progenies
Y ;.oal
132 144
142 142
T. TAltASAI<!
This mottling is clue lo the mutabilily 11'0111 nor111al to oily of oat gene
located on the second chromosome, and characterized, in appearance, by the
irregular distribution of normal and oily parts of mixed in one
individual (VI-9, VI-i0) (TAKASAKI 1940).
T. Yokoyama
Mottling ell\(! to mutable
Hereditary in mutability lIr
oat gelw and its direction
",,/ (II
('lIlhl
lollo,t)
V-ll1 Maturnal of inhiilitor, Ie
(Ill till) lIlutabilily lit' fI(lt gene
I'
.
F
I
73
VII. Genetics
TOYAMA was a pioneer in embryological study, as well as in the field of
genetical study of silkworm. His paper (1902) is even now, referred to as
one of the most valuable reports on slll{worm embryology.
The non-c1iapausing egg of the silkworm hatches about 10 days after the
deposition at 25C. showing no pigmentation of the serosa (VII-I), while the
diapausing egg begins to show the characteristic pigmentation of the serosa
in about one day after the deposition except for the case of white egg mu-
tants. The serosal pigmentation is completed in 3-'1 clays, and the egg color
is determined by this pigmentation incombil1utiol1 with colors of the chorion
and of the yolk.
VIl-l Embryonic development of the
non-hibernating, egg lit 25C.
/
The silkworm egg is deposited at the metaphase of the first maturation
division (reductional), and the ferlilization takes place in the egg in about
two hours after the deposition. Unfertilized eggs deposited by non-mated
females or dissected out from the ovarian tubes of virgin female moths can
be induced to develop parthenogenetically by high temperature treatments
which are known to cause a union of two haploid cleavage nuclei or forma-
tion of the restitution nucleus in the eggs (SATO 1934, '42) (VII-G), The high
74
T. Yokoyama
temperature treatment of the eggs newly laiel by ordinarily mated moths also
causes the androgenesis (so-called" merogony") (IIASIMOTO H)29) (VII-G, VII-
7, VII-8J,
The formation of egg color mosaics which is often observed in the eggs
treated with high temperature within one hour arter the deposition is due to
the participation of cleavage nuclei derived from a union of two sperm nuclei
in the embryonic development. The mmmic paUern of eye color, when it
appears in the silkworm llwlh, is always antero-pmltcrior stripes parallel to
the body axis, in contrast to the mottling lype which is usual in Drosoplzila
(TAZIMA 1942, KOYA.MA l!)!'i()) (VlI 51.
The germ cells clilIerentiate t!Jemsl!lves 1'1'0111 the blastoderm cells in a
definite region on the ventral side of t.lw egg. The periphl'ral plasm which
is specially destined for the place uf germ cdl differentiation, sLlch as the
poslerior llolar plasm in Dipteran and Culeopteran inst!cls, can not be distin-
guished morphologically in Lhe silkworm, thoug-h a region of the
germ cell differc.ntiali(l\1 hm; nmplled tm ventral side o( the
egg by cauteritmtioll experiments V 1I. ,1). For I he formation of the
gonad, the germ eel I:> thus clifi"erenliated, lllllst he receivt!cl illLo the genital
ridges which have origin, independent of the gm'lll cells, in the coelomic
sacs at the (i-9th segments Ill' tlle embryo IVII :1).
In the silkworm, many lUlll atiol1s wb iell cause 111 a ]formatiull in various
organs or have lc1hal have hl!en discovere(l, :supplying' L1sdul materials
for the developmenlal studies. E-alleJes arc a group which draws Ollr special
attention and on whit-ll many studies have been nmclc g'iving' very interesting
results. f\ remarkable characteristic of H alleles is the Yael that lhey show
pleiotrupic actioJl on variolls t isslles am! organs (Jf edudD1'11utl origin, 'Inc!
cause morpholugical as well as physiological c1is(l1"(krs in c\evdopmcnl. This
pleiotropic al'lioll of the genes sLlggests that their ad ion is com:cl'llcd in the
cell differentiatioll at an early stage of Llw embryonic devululllllcnL It seems
that each 01' these alleles ads with its own [lotc'IlLial in a definite c!irection
(TAK1\.5J\Kl HHD) Vll-17). A dcfcd or thl'. ).!;onad rormutiun in the
homozygous UN embryo may iJe cited all example of these Slllclk!-l (ITIKAWA
19M, MIYA 1955). At the germ banel !-ltag'c, the dit"[ercutiaLiOll of germ cells
proceeds normally in the EN I EN embryo as in the l1ol"malulle. The EN gene,
however, begins befol"G long to affect clctenninatlull of tlw body segments,
and it makes the abdominal segmcnts develop into the ones resembling tho-
racic segments (VIl-,lJ, VII-H). It also inhibits f()rmation or Lhc genital
ridges which appear at tbe (l-9th segments in the normal umbryo receiving
the germ cells. Owing to the detect of the genital ridge formation, in the
EN I EN embryo, the gonad formation can not be initiated, and ge.rm cells
remain free amidst the mesoclerm cells and finally degenerate (VII--15).
The kidney egg is a maternally inherited recessive character. That is,
a female homozygous for ki gene lays only kidney-shaped eggs which die
without hatching because ot a lack of the mesoderm formation (VII-12).
In the silkworm there are many characters, the expressions of which are
affected by environniental factors during embryonic development as seen in
the change of volt in ism by the effects of temperature and light (see the sec-
tion of Physiological Genetics). Here cited are the l11ultilunar marking and
the mons tar larva (VII-9, VII-lO).
VII-ga
A multilunar marking appears in the presence of gene L, but the num-
ber and localities of the lunar patterns are influenced by other genes and
environmental conditions.
111 the lowest extremity (2L), the patterns appear in the fifth and the
eighth segments where there are markings in the normal pattern worm (2L).
In the highest (7L), the lunar pattern appears in every abdominal segment,
from the fourth to the tenth.
There are several intermediates between the two extremities ..
VII-9a The variety of m1l1tilunar
0ct
+
*
+
*
Black and red -I- ligures show ordinary types of the multilunar. These types
can be obtained as almost completcly stabilized oncs. The blue tigured ones * are
very rare types, and it seems probable that 5 and G are genetically comparable
with G (red) and 7 re!'lpectively. T. HmoBE
VII-9b
Therc are two centers as the lunar pattern occurs, the fifth and the eighth
segment.
The lunar pattern appears primarily in two 'center segments, next comes
that of the sixth, the seventh, the fourth, the ninth and the tcnth segment in
order. However, the order between the fourth and the ninth-tenth segments.
is sometimes altered.
The patterns in the segments aecorciing to the above mentioned order are
highly heritable, while those on the segments in different order are scarcely
carried over to the offsprings.
76
T, Yokoyama
Order of manifestation
10 1\ 12 13
Crit ic:;a I period for determining manifestHtion
9 r
L. ___ _ J.
10
I
15 dll),' (al 15"(;) 30
I ; ----,.. l
I .. __ :_ ... ____ ____
egg Jllylnll .
A II c
H1nwtukinC;ih. hatch
U 1, l'
Crilh."nl in dtHMmlnlng t. is H .. II", hcrilnbl(!
nHJllstar II nnJ the \'ullif1h.nl lIr!! A .. C .111<1 ... H,
T,
The \$ the ml.lltihmar paUm:n i\:\ inHu(mccd by the incuba-
tion temperature, 'rhe lower tlw temperature (within the range between 15
and 28C,) do t.he more llaltcrn\:\ appear (VIH)(:.\,
The critical period when the 1ll1111ber of' paU(ll'llH is lIlm;C affecLed is be-
tween the sixlh ane! tlte fifteenlh day of incubatioll al I fi"C,
VlI-$\.: II [ lIlflni [t'slaLiun
by inculmt.itllllell\Ill'.I'alUrt:
L,llUllIif
....
1 HIROB
For 1!llt'il'cll'(1 IIi\U1'l: refer to
!)n, wltilL' whilv ligures iii) and ((i) cor
to lil) lind (Ii) ill VII-911,
+ +
J\l'l'lIWS hl'l WI'l'lJ cil'l'l,"!\ ,,\lllW the
cl!allge of t. lit! Hum her of lhe lunar
IHIII.L'rn according to I.he gCllcticl\l
comilil.lItilllllln wlJlI <If, to liw illcuba
t ion tl)llI[ll!ratuJ:e,
VII-9c
When the eggs heterozygous in L gene are incubated under a low tern
perature (15C), the manifestation of lunar pattern is nearly the same as that
of homozygous ones incubated under a high temperature (25C).
There are, however, two exceptional cases in which the influence of the
incubation temperature is not observed, i. e. 5L, and 7L in VII-9c.
VII-IO
The manifestation of the monster, mse-II, is different according to the
treatment of eggs. The appearance is very rare in the eggs treated with
hydrochloric acid soon after laying (VII-lOa). In the hibernating eggs there
is a high frequency of appearance (VII-lOg).
Besides, many experiments were carried out on the occurrence of mse-II
using the eggs treated after the hyclrochlorizatiol1-refrigeration method or
the refrigeration-hydrochlorization method as well as eggs treated after other
methods, obtaining the following c(:mclttsion: if the treatment in which the
hatching is more difficult, the more frequent beco111es the appearance of ab-
normal worms. However, it was found that refrigeration has no direct in
tluence upon the occurrence of the monsters.
VII-IO Changes in manifestation of the monllter (mse-II) gene
dLle to hatching method
eRg laying hatch a.bpnpoe',mread"
a
treated With acid ISh t ..
'0011 nftc" lilYlng 0.;; 'K)>e-------12 days--lncub.t'Qn (25'C)----M almo'l none
b "{ter clays%'J,w(.(!j)/)////(.I'lffff?1J;W...-il days.... very few
aCI .. mcnt
c alr(c
t
ac!d a.-40h days..... few
er gcratlOn ...,-
" 00--72h __",wff/fo,w)/;/plm'i/:21 days-tO' more abundant
f refrigerated
g kepl Ut:dO
t
d" tcmf" re(rig.ratlon(SC)
naturn "m ILIon (25'C) many
of this monsler are different in grade among different
strains, so the pcrcentagc of abnormal:; can not be decided, But i.t is
noticeable that the abnormals can hardly be obtained by the ordinary hat
ching l11ethod (a)
Physiological analysis of lethal actions will be an interesting problem in
future as. hinted by a study 011 the lethal non-hibernating egg (VII-ll).
VII-ll
The lethal non-hibernating, !-n, is a recessive single Mendelian gene, seg-
regating 3: 1 in one batch in F
2
The eggs homozygous in l-n die around
the blastokinesis stage of the embryo.
78 T. Yokuyama
Ii.'
Ii!
"
'" g
'"

m
::;.
!

0.
U
cl'
VII-ll Difference in 02tlplake anel development between
normal and lethal l1(1nhilJeratillg' (I-II) eggs
200
100
VII-4 Maps of the i I'q.r,iOll of tlw
egg at tlw cJl!(lVage stage.
1. ventral view
a. view
...

. ".:.:" _._ .'

"
""n,II.1 ,.,1.
2. lut (.!ral vil'w
.1. cnH,ti
x /1.5 genital region
00'
0",;
-:;:::-;'1(100 rt:gi(lll
tlU'
T. TAltAMI & I';.
79
When eggs are treated with hydrochloric acid, normal ones hatch in about
10 days after the treatment (A), while I-It eggs die after about one month's
slow development showing less 02-uptake (B). When eggs are kept at 25C.
without the acid treatment, normal ones enter the cliapause showing little O
2
consumption (D), while l-n eggs whose O
2
uptake is somewhat higher than
normal, show slow development before death, reaching the blastokinesis stage
in about 70 clays.
The yolk segmentation begins in about one day after the deposition, and
the yolk material is divided into small spherules, the yolk cells, each includ-
ing one or more yolk nuclei. As all the yolk nuclei are derived from the
same cleavage nucleus from which the nuclei of embryonic cells come, the
embryo has an intimate relationship to the yolk cells, but not to the yolk
material itself which is of maternal origin. The interaction between" the
embryo and the yolk is one of our problems to be solved, and the androgenesis
(" merogony") and in vitro culture of embryos are used, in addition to the
formerly used reciprocal cross, as the new techniques in this field.
VII-2 Differentiation of the geml cells
, .
" ,:
3. Cross section of nn 18 hours
old embryo
L Cross section oC a 16 hours old egg,
showin!( th" ,ogrogation of the germ
cdls [rom the ((crm b:lnci
-"" ',,, 4. Cross section DC a 21 hours
,,'-';1\ old embryo
2. Longitudinal scclion DC a
15 hours old egg
5 .. Cross section ;'24"'hours old
cmbr),o, the germ cell. separate
completely from the germ baud,
and, .rc by the somatic
ccl Is- presumptive mesodermal
cells
1<. M,YA
so
T. Yokoyama
Finally, it is worth saying a few words about the results obtained in the
field of experimental embryology. The si Ik worm egg belongs to so"called
mosaic (non-regulative) type, and a defect due to partial killing of its embryo-
nic area can hardly be fLlled up by regulation, giving rise a partial embryo.
The presumptive regions of organ pl'irnodia have roughly mapped on
the surface of the egg (VII 4).
VII-2 DIfferentiation of germ cells (K. MIY/I).
In the sill<wol'tn, as in the Dipteran and Coleopteran insects, determination
of the germ cell seems t.o be brought aboul by entrance of some cleavage
nuclei into a dcftniLe region of the periplmnll, only difference bdng that this
region is not located at. or near the posterior pole of lhe egg but in a definite
part on the ventral sick o[ the gel'lll cell lakes place shortly
after the formation or the germ band, and a group of cells which have been
differentiated into germ invaginate inward from tlH! germ band.
VlI-a F(1rmulinn tlf the gonad
1. seL'lion lhrullllh lhe Hlh
of a nu hUll I'H old C:lllhryo
4. LUllgit udillal lIf
uf a 72 holt n, old
l'mbryo !-\howinn the gl'nitul forti
2. uC a GO !wnnl uld
embryo lhruugh llll' Uth (1\) and
the Htll (Il) showing'
lhat the f(cnltnl I!xtl,tld,
Inward and till! Hl'l'lll celb
fi. L.ongitudinal scdillJl of the
7-Sth Se!(ments "f it 78 hout'u "Id
l'mbn'o showing lH,tdllal
eontraclion of gl'nil,tL coru
3. Cross GG hours old
embryo through lhe (A) :lnd the
9th (B) showing sL'paratlon
of the genital ridge from the coelomic
sac and formation o( the gonad llrimodium
K. MIl'A
81
VII-3 Formation of the gonad (K MIYA).
The gonad formation begins with the differentiation of genital ridges in
the coelomic sacs in the 3rsI-6th abdominal segments. These genital ridges
extend inside toward the germ cells, roll back to enclose the cells, and then
segregate from the coelomic sacs to form the gonad anlage separately in each
segment. Next, these gonad anlages connect successively to form a pilir of
cord-like tissues (genital cord) which later contract toward the 5th abdominal
segment, forming the gonads.
VII-5 Differentiation of the compound eye.
There is a well-known fact that the mosaic pattern of eye color, when it
appears in the silkworm meith, is always antero-posterior stripes parallel to
the body axis, in contrast to the mottling type in Drosophila. This can be
explained by the direction of the differentiation observed in the imaginal bud
(N. KOYAMA).
VII-5 The pattern of eye color appears as
al1teroposterior stripes ia the silkworm moth
N. KoYAMA
VII-6 Doubling of the chroll'/-osome number in the egg developed by arti-
ti.cial parthenogenesis ..
In the unfertilized egg artificially activated after laying, the reduction
division proceeds normally, but conjugation of the two haploid daughter nuclei
82
T. Yokuyama
resulted frol11 the of len place giving riHe to a single cleav-
age lludeus which seems to dc'velo]) into 11 diploic! individual. On the con-
trary, when unferlili7.cd eggs dissected out from the mother's ovarian tube
are artificially activated, doubling or the chromosome llmnber is accomplished
by formation of the restitution nucleus Ill. SA.'I'OI.
VII-G Duplication of number in art ilielal partheJlogenesis
u( 11 tdp!o'ld In'brill hCl{wccr'I a
Il'! r;1ploll1 1(,!I1II1L' ami iI -tUpluid rtw.le
1. CunJUI1Rlhlt\ nf till: l\\'11
frmn thl! third '" lin ilf!llrdnllr
udival(!11 (l vlndn

2. A rc;\tltutlun nul'iclith
In tin l'l-!l!: lIut
IWnI O\'Url'lll tulle
Iwiort.' nud
lu;tirutlJu Wilh hiUh
nJflt'.tlninij'
ilfld
J lJlvi.llclll
3 r':quIltluII41 dlvluioll
01 IIII' p".,litntillu
nw'll'n,.. IlIfl1lllliol
1I111y ulle IHllur
5. 'II
ill tl IlIiuHlrr
"'IIP r ITIII 1 I.IL' ... , ,
VIf. 7 Ar(ilicial ill. I {.<\SI MtI'J'(1l.
lj, H,I 1 HI <I
'1I'l,'rIlUJiwqtl!
l'arllwl1llgenesh, h, Illml to I)('l'llr s[HlIllalH'lllmly ill till' while it
is rather easy III Ill' in<illl'l'cl by artilkial In'aln}('llt or with hot water.
The silkwor!11H ckvdo[ll'cl by arl.ilkial art' (til' salllL' in gC110-
V((-t i\rlili<-i:ti parlh,ulI.l';t'IIV::i!;
I. loid by
virgin mothHj not
:!. Ullfl!l'lili"':cll (.'lUpi 1!lId hy
virgin moth\! in uaym
nnd If(mlcd wllh hUl Wi.1tcr
('I.o.t\'1 1;i!iJ hy
mUlad funmillb irl II h'w hllurli
ll. llNl'MOT<I
83
type to the female parent, that is, when a mother is heterozygollS for some
genes, the parthenogenetically obtained progeny are also heterozygolls for the
same gencs, and are mostly diploid females with some exceptional tetraploid
and mixoploid i11divicluals. The body of the mixoplont consists of both diploid
and tetraploid parts. As far as artificial parthenogenesis by heat treatment
is cone l1l"l1ed, eggs dissected out of the ovary at the end of pupal stage or
eggs of emergecl moths, and those cleposited by virgin moths are the same
in potential in parthenogenesis.
VIl,-8 Androgenesis (union of two sperm nuclei in the egg) (H. SATO).
A union of two sperm nuclei in a silkworm egg exposed to a high tem-
pt:,'ature soon after the deposition. The resulted nucleus develops into a
diploid individual without normal fertilization (so-called /I merogony").
VII-8 Merogony
Union of the two sperm nllclei in the Rilkworm egg
eXJlosed to high temperature at the time of oviposition.
The I tcd nucleus develops withoL! l fertilization of
lhl' egg nucleus. I-I. SATO
VII-12 Development of Iddney egg.
A genetical character, kidney, is a recessive mutant affecting the shape
of the ep,"g and causing death of the embryo formed within. In the kidney-
shaped egg laid by homozygous hi females. the mesoderm formation can not
be observed, while differentiation of the ectodermal organs proceeds inhar-
moniously (K. SUZUKI and M. IcrIIMARu).
VII-13, 14 E-gene group (N. I-rIKAWA).
Generally speaking, the occurrence of extra markings and su pernumerary
legs in the silkworm larva is related to the E-gene group. Of this group, 15
genes are known at present, all of which having the locus (0.0) on the VI-
chromosome. These genes are all dominant to the normal.
EN and EO(/" among the rest, are very suitable genes for study of devel-
opmental genetics. Embryos homozygous for EN or Ec
a
die at the later stage
84 T, Yokoyama
Embryonic dl'vl!iopnwnl of Llw kidney l)gg
I. + nnd iii ORR"
am; amnion
doc: den.nor.tlng .011
for: lorm.llon coli
Ip: loglIke prot
mt: main lrachanl lnmk
"": oral cell
pn i portion
Mer: aerol"\.
lit: IIllama
SUI):
t: n
Ihp: lI,orock port Ion
ye: yolk coli
;' ,n
uo,' It' ""
... ",
r'
\
2. A (ully J'
Ill' cmhn'o
r-"'-'--
38. Crml" iiCl'licm
,hrouRh Ihe
procophdli<' parl
511. Longitudlnnl
Qr the
prucclllml k llart
LongitudInal
action uf a G dnys
old Iti
.. "--....

'''"
(D. Cro.. ..cllon
lhrouuh lh. mlddl.
I1lU\ l'( the Ilbdohll)O
GA.
ut II Y
uld
flC. Crmm 5D,
1Il'('t!<'n through In.val-tlnnthHI
till! uhdolllCI1 ur !.iliMma ill
r,E. Furn"'t lun
{II thl' mllin
trnl.'hcul trunk
ahd.ll'ncn
4C. CrOtlll "",,11011
,hrouRh lh.
pro ph nile pori
7. Longitudinal ..eetlan
01. 12 doys old
ki embryc>
.110; .. ' ..
..
K !-iUlllKl & M.
of embryonic ckvel()jlll1cnt without any l-\iHIl of hlal-\(oldnesis, TIle
appendages, hristles, all(1 t nll'lw;tt, ()f llH' 1 st"ll! h ll()(ly sl'gments in
the EN/EN embryo are of forms Il(' llw thoracic segment
in the normal embryo, while and t r<lcllt!Ut' are rudimentary, while
no abdominal appendages arc seen in the l,:r',,/ RI'" embryo, Moreover, gonads
can not be observed at all in either EN/EN or E"" / Eri" embryo. According
to MIYA (1955), the EN gene does not affect normal differentiation of the germ
cells, but development of the genital ridge.
85
Embryos homozygous for Nc (located at 1.4 on the VI-chromosome) lack
some thoracic segments, and die at the later part of embryonic stage.
All of the EN, EGa and Nc genes in homozygous condition cause abnormal
differentiation of the thoracic and abdominal segments.
Genes of the E group have not a true allelic relation one another. ITl-
KAWA has found that recombination occurs between EGd and EN", obtaining
some EGa EN"/+EN individuals in the offspring of EGd EN"/++ 9 XEN/+(O).
These individuals had many supernumerary appendages on abdominal seg-
ments. It seems that these individuals have abnormalities in the chromosomes
as a result of crossing over occurred in the E locu$.
VII-l3 Embryonic abnormalities in Emutants
After ahout 3-4
day" InCUbatlon,\
the embryos can
be distlngldghQd
I;!ach other by
Lholr characteristic;
nlmormalities
Fully grown
embryos
+(+
Nurmal
"(EN
Thorllx1 ike
ab<lomlnul5cgmenu,
gonad absent, no

.....
U
I
Abdomlnol leg.
and gonad
nlH;lcnt, 00
lJllIstokinl!sls
VII-14 Extra legs in the pupa and moth
(EGrl ENe lEN
1. pUpD. 2. Plath
.,'
Individuals, obtillncd 1]$ a
recombiontion of genes,
havt! rllllny or cxlra legs
00 the ubdomen N, i'l'IKAW'"
N. ITIKAWA
86 T. Y"kllyanw
VII.15a J\bs<"l1cc of the gonad in 1 he 1,'N j eml>ryo m:. MIYA).
Tile gonad formation in tile divided into twu the Ol1e
is c1i1Tenmtiatiol1 of germ cells at the germ band slage, and the other is for-
mation or the gcnita I ridge at II\(' formation The absence
or the gonad, which is one of the eiwraclerh;lics or the ENjJ.,'N embryo, is
explained before.
1. Cl!J:,'; "('1'111/11 lhl'IIIIIdl :J. r",t,:, ./'1110" Ilulilluh ,II,! /JIll
lith ',"V,IILI'lit .-1 11'1'; 11"'11", \l1')!JlLt'1i1 01 .I 11(, hULII",tid
ult! JIIIJ'1H1l "/uh,'v'l Ii 'rill /1111,1 .... , TIll' I', IllIt
1:,11, ,\It:{'H,h''oI,d II\' III,' ,'1'1[. jolllll'li .1I1t1W'IIHII'II, l"I'lIl.dll
'" 11h' allil,11t'
pl1!JllldlLlln ", '''11111'11.
I lid/II ill ,,','lltlll "I til,'
):II;llIJ:1J ';"}:I/11'II1 1)/ ;( /:'
hl'III" .1\,1 IIPlilial 1'lItil1'\'11
(;"1111 ,.'11" \"1'.[ 111'1' .1II1"UI'
,I ... 1IH"'\I,d"III1,d .. "II,
I. I.II"V,L\HlIiUi,j ',1'1 (jnll lie
lit,' .. I ,I lUlurti
lilt! 1-.'.\1""'\ "111111\'11, l"I'l'llI tl'lh
,", I', 'I,,'I\' :unnUH" III!'
!l/I" II!!'" 11/," ,db .I' III liv. :!.
Studies Oil tfie' ill till' silkworm (M. TS1IJl'J'Al.
Crossi I1g (lve'r 1>('t I,: l/' a n<1 1','Jd> was r(llLnd iTSllJ IT \ '!i:l, '!iil I, When
male heterozygou8 for 1,,'1/ and /,;1.'" rrum tIll' allt'lk llH'lIt iOlll
'
(\ ahove
lllated with norlllal f('malt-s, a ve'r!, small llUllllll'r oj' individuals with
the ciscoJlflguratioll p;ellotope Hfl H"J'/ I +. "'(I1'(! ol)taifH'd. TIll'c;" larvae have a
new phenotype which was only t'xpt'l'tt'd alllOlli; 1lll' ()r FII nncl RK1',
where there occurred J'(!l'oillhi nati()n. The J'l'I'o1lliJinal iOll illd i viduals wefe
characterized hy the or ('xlra-alldol1linal IV.I'S (lr til(' first and second
ahdominal segments well as Ill' eXlra-semilullar paUl'rllS Oil the ab-
dominal segment. They arc tllLW ea"ily clistingllishC'C1 from tlw Hli I j I E
1
(P
larvae.
A few larvae with genotype Hli and KjJ/ I larvae always appear in the
cross + x II KP;'-I- + obviously by recombination,
Silkwotm Genetics Illustrated
87
Furthermore, from the cross between Ell ane! EN", Ell and EAt", Ell and E'u"
and between Ell and EN, a very small number of recombinants can be obtained.
From the facts presented above, it is clear that genes belonging to the
E-series behave as complex loci. This E complex loci may be called by the
term, "E region". It consists of several genes with similar effects arranged
in close sequence at the end of chromosome VI.
VII-15b The E p;;eudoallelic genes in the silkworm
+/+
M. TSUJITA
VIT--IG The role of E-allclic genes in the embryonic development (T. TAKA-
SAKI).
The most rcmarkahle characteristics of E-allelic genes are that they show
pleiotropic action 011 several tissues and organs of ectodermal origin, and also
t.hey manifest morphological as well as physiological disorders in the develop-
ment. Studies on the pleiotropic aciioll show that these allelic genes are
possihly concerned in early cell differentiation of embryos. As far external
arc concerned, the disorder is most remarkable in the crescent
markings ane! in the prolcgs. What is noticeable is not the abnormalities
themselves manifestcd in specific body segments of an individual, but the
mode of manifeslation of gene actions which has special connection with the
body axis. When morphological characters are comparee! between E-mutants
and normals, it is evident that the corresponding character appears with some
shirting- of the place of expression along the body axis in these mutants.
Each character has an expression center in a segment where the expression
is the 1110St frequent in occurrence becoming lower toward the anterior. or
poster lor direction.
88
T. 'Yokoyamd
It is assumed by TAKARAl<I that each of the R-genes has a cel'tttiil actiOil
potential to make the expression part shift toward the anterior, or
clirediol1. For example, the action of begins to appear at the first ab-"
dominal segment and spreads posteriorly, I1Htking crescent markings appear
011 the third segment in addition to the sc!concl segment which is a normal
area of the expressioll, and also making the character which appears on the
fourth segment shift to the [tfth. Thus the star spots normally appearing
011 the fifth segment often duplicale in Lhe sixth and seventh segments, The
direction of action is definitc! [or each gene, and cetLain genes act in the
posterior direction (Eli" whi Ie others in the reverse direction Morc"
over, there are some genes which act inlhe poslerior direction on the dorsal
side and in the anterior direction Ull the ventral side lEKJl,:::!.), and vice versa
... ).
The action potential is not the same at different levels along the major
axis of the embryo, Thc potential of pill is the highest in the fHth ahdomi-
nal segment, becoming lower in the next segm.cnt, so t.he absence of star-spots
on the 11fih segment. is a stablo charader, whereas the extra-crescents un
the third segment is variable in expression.
The direction and power of action are different in . ,
[i;D and Elip not only among thelll but also till! dorsal
al1(1 ventral sides of the same individual. The curves show
the power of action in the thorax and the abdomen.
T.
89
VII-17 Disturbance in segment formation by EGa and ED gene (T. TAKASAKI).
The EGa gene causes abnormal fusion of the fourth abdominal segment
with the fifth, or the fifth with the sixth, the fifth abdominal segment being
VII-I7 Deformities in the segments by EGa and Ell genes
lind mechanism of their formation
head .. " ....... A
thora. de {I :::
II'" ..
seginents III'" -
1 ....
2
3
abdominal 4 ..
segments 5
B C D E F G H
A-B: ECa/+, C-D: ED/-I-, E-G: EGa/ED, H; EGa/EKp
I
1: Ventral deformity rarely observed in Eca/EEp and EDIEKP
Changes in manifcstioll of deformity due to genotype
deformed segments
genotype
direction
of action
I
I
thoracic abdomiml1
i percentage
of
deformities
Eca/+
Eli/+-
E1(1'/ -I-
EU" /R"
EcalEJ(Pi
I
I
I
EIJ/EJ.:l'
1/+
---" ...)
(d)

Cd)

(v)
.= (d)
----->-
(cI)
--+
(v)
(--;
(d)

(v)
(d)
::::::; (v)
..
I
I
I

2 3 (v)
III '" 1 '" 3 5 '" (j Ccl)
5",G(d)
3 (v)
III 1 2 (d)
3 (v)
III - 1,..... 2 Cd)
total
110.
5895
2969
The in appearance of deformities is affected to a great extent by
coexistence with any other allelic gene. Cd) : dorsal, (v) : ventral sides, arrows
directed to the right indicate tendency of posterior shifting, and vice versa.
T. TAKASAKI
90 T, Yokoyama
the center of the abnormality, while the HI} gene causes the similar fusio11
between the third thoracic segmcnt and the fin.;t abdominal segment, or be-
tween the first and the second abcltlminal segmenis, having the center of tht:!
abnormality in the r1rst abdominal segment. In cases, the abnormal
fusion is limited to the dorsal or the larvH, and docs not extend to the
ventral side, Each gene, tillIS, has a cxpression center where the
gene action is the sl rtm,!{esl.
One of the 1110s1 of tlles(.' almurmalities is lhat the
expression is influenced to Ll large hy (,()!llhinat ion with 01 her
genes, For inslance, Ilw alilwnnal in percenlage io fi,1396
in RO"(FD, anel to D,Ofi?il ill Hn"/HIII', in cOlllparison witll O,;ll?6 ill
HOIl/HI, whilD it iH in HIl/I,'I, !O,riW'(i in },'Il/}.;r"', and in HlI/HI\I' it is
llcarly tile same as in F I,
The body wall of silkworm larva has itc\ OrIglll in a dorsal L'xlention of
the lateral margins or Ihe gl'rlll lmnd, 1 hal thc frolll either side
of the germ hand mCl'e and along 111(' lilw lIll Ilw clorsal siele,
forming a continuullS wall. Jr, thlTt'i'ore, titl' l'Xh'llSiullS are asynllndrical, it
will expected that th(' fusion Ull Ihe dorsal becollles iI regular, a part
of the c:xtentioll n'l11ainiIJ,t:: fret' ill a sl'gillc'ni, or with the extension
in the neighhoring Sl'g'llICnt. I\'-allelic .f!,'<'lll'S SCVlll to ad in the course of wal!
formatioll,
When twu aIle-lie gl.'IlCS in c()!llllinalilin arl' (lppw;ilv ill t'lwh direction of
cell as in ,,"""//<.'II (,'.1, it is pm,siille tllat the uSYllll1lclry in
extension, irrl',l(ularily l)l'l'llllll'S lal'gl'r ill l'rl'qul!lll'Y, rCHlllting ill
forlllal ion III' a largL'J' llul1l1wr (If abnllrmal individual,;, Un Ihl' ('(mLrary, if
Iwo gcncs in C()mliitiatioll al'! ill tfll', ,;anl<'. dirl'l'lillll ill 1,;('''//<.'11'1' ( :), [uwcr
percentage ur alJIIUl'llla[ individuals will lie eXIll'l'!t'cl.
The rad thaI there is !10 J'l'llwrkali1l' diJll'rL'nl'c ill th(' of ab-
betWl'l'll H"/HI anrl (' _.1 l'an lil' wilh the as-
sllmption thaL E"I' h:lrdly inJlUL'Ill'l'S Ull' lin;! aiHloJ]]inal Sq,:'Il11'l]t ill which
formatiun of tlw alJ1lo}'J1laiity is dUlllin<lil'(1 by "'II,
The shifting in 1)(I:.;itill11 and {'Xll'llsioll ill area (lr ahnoJ'malities, as
in may be hy tll(' asslllllptioll that the Sllm or ndioll putcn-
tials of two allelic which ad in oJlPo:.;ilv (tireci illns is enough to
suppress completely th(' llurmal formatiul1,
Silkworm Genetics 11111strated
91
VIII. Physiological
Under this heading, voltinism, i. e, the characteristics of the silkworm
related with the number of generations repeated in a year, molting and the
hormonal control of both are mentioned. These characters have practical
significance in sericultun' and research interest and results in this field of
silkworm physiology characterizes one phase of Japanese biology.
1. Voltinisl11.
If genetically bivoltine eggs are incubated at 15"C. after the blastokinesis
stage of embryo, the moths developed from the eggs lay non-hibernating eggs.
If the eggs, however, incuhated above 2'1 DC., the moths lay hibenrnating eggs
(WATANABE EnS). The light also inOuenccs this characteristic, i. e., the long-
day effect of more than Hj hours is comparable to a high temperature, the
daily illumination shorter than 12 hours to a low temperature (KOGURE and
KOBA.YASHI 1928).
2. Molting.
There arc in silkworm varieties tri- (lltJ3) and penta- (Mr.) molting ones
besides ordinary tetra- (_i_N) molting ones, and they make multiple alleles in
sLlch relalion as llfl>-i .11>M" on the VI-chromosome. The manifestation of
the characteristic is affected by the rearing temperature and the nutrition;
that is, a high temperature in the first and the second instars and good nutri-
tion tend Lo diminish tIw number of molting.
S. Hormonal control of molting and voltinism.
BOUNIUOL (1 B3n) and independently showed that the silkworm.
whose corpora albta are extirpated enters pupal stage without undergoing
furLlwr larval molting.
FUKllnA (HJLiO) extended lhe study and conclucledlhat whether the silkworm
undergoes larval molting or pupation is delermined by the relative strength
of the hormones of corpus allatum and prothuracic gland, where the corpus
allatul11 hormone restraining and the proliloracic hormone promoting meta-
morphosis.
IhsE(iAWA (E1Gl) and FUKUDA (H1Gl) independently proved that the secre-
tion of the sub-oesophageal ganglion is the controlling factor in hibernation
of eggs. The hiilernating character of egg is also affected by certain kind
of chemicals sLlch as uranium nitrate (lLil.5F:GAWA 1943).
VIII-1 Change in cliapal1se chantc1.er of eggs laid by the moth which de-
veloped under different day-lengths and temperatures during egg
and larval stages.
Motlls of bivoltine race are, as a rule, to produce 110n-diapause eggs at
the first generation and diapause eggs at the second generation under natural
conditions. The nature of eggs are changeable with different day-length and
92 T. Yokoyama
temperature applied Lo the moth at various Hlages of life.
Both factors are most effective in lhe embryonic stage after blastokinesis
in the course of incubation. DurinK the effective period, treated with com-
plete dark of short-day illumination uncler J2 hOlIrs a clay, anc! a low tem-
perature at 15C., the female lllothH developed fnnll the eggs may produce
nOll-diapuuse eggs. While trealed with a long-day illumination above 16-17
hours, and a high temperature of 25"C., they may produce diapause eggs.
In larval sLage, illumination and temperature are also effective in the
same direction but wiLlt less degree than embryonic stage. In late larval and
in pup,!l stage", on the contrary, a lower temperature below 20"C. may give
a higher rate 01 c1iapau:>e egg producer ()f l1lolhH.
The threshold of percl:ption of light is O.OD r. c., Violet light, 350-510 m,LL
wave length, has the same cJ'fect as white light, while a light over 550 m,LL
wave length of orangt'.-ycllow has no for lhe production of the dia-
pause eggs (Temperature, WATANAun: 1DlD; light, [\OGUlm and KOBAYASHI 1928,
K()GURE 1D:-33).
VIII-l Chang() ill lliupause charadur uf L'ggS
Through lht! effect of day.length and temp. UpOIl mullwrs (lJombyx nlfJl'i)
1. aiJove 21i"C.
illl1minated [or I(i hl"s.
illulll. [ur Hi
illulll. for 12 hrs.
.[. IJduw Iri"C.
i1lulll. fur 12 hrs.
larval i"ila).\l! I l"iwradc,l' or the
'I jlU pall molh treated as
Isl.-:II"<1 inslal'! .1I11-lilh stagl' Ildl.
I I
, ,
I all lllOlhs lay
regardk's:; of daylellgth <llld l('III/). I t1iilpamw l'gg
: hakltes
illul1l. [ur Iii ill'S.
bduw
aliuvl' 2H"C.
illulll. j"m hni.
regardlt.::,;:; of daykngllt ami Illllll.
1
111m,t Illol.hs lay
, diapausl' l'gg
I haldwH
IllUsl. lllolhH lay
Illll1(1 i apllUHU egg
! haldwH
all l1l0[J1S lay
lIulldiaptlUHU ()gg
IllltdlL!!;
M. KOGURg
VIlI-2 DiscQvery of suboesophagcal ganglioll the! tliapuusc center of the
silkworm.
According to II. Kna;:AwA m)l1;l), it is easy to line! out according to Ehr-
lich's diazo reaction whether t he eggs arc desLined 10 grow 10 cl iapaLlse 01'
llon-dlapause ones as early as they are within a pupal body. The reactioll
was applied by HASEGAWA to his studies to find out the organs responsible
for the determination of voltinisl11.
Silkworm C-enetics Illustrated 93
Firstly, immediately after pupation female pupa which had been deter-
mined to produce diapaLlse eggs was ligated at the level between the 6th and
7th abdominal segment as ovaries were located in the anterior part and then
the posterior part was cut off. The eggs developed in the anterior part were
of hibernating nature, showing that the segments posterior to the 6th abdomi-
nal segment had no relation to the determination of voltinism. Secondly,
female pupae were ligatured immediately after pupation at the level between
the 3rd and Lith abdominal segment as ovaries were located in the posterior
part (Fig. 1). Eggs developed in the ovaries located in the posterior part
were not influenced towards diapause. But if the ligation is not complete
and there is a slight communication between the both parts, eggs may be
" "
'
I r'
S: III.
.
. ..
,..;
bll

",i
bll

oj
<l)
.!::
...,
u
<l)
C
C
o
u

<l)
b.Il
.a
p.
o
Ul
<l)
o
u
o
cr5 ..: .n <.0
on
bll Ilil Ilil
ri:;
T, Yokoynmll
affected towards diupausc, The results show thal lhe organ or organs to
produce c1iapallse factor, iJ' \vollid not be localized in the posterior
but in the anterior portion of silkworm pupa, Thirdly, decapitation"experi-
ment was carried oul. Decapitalioll of newly pupated insects forced them to
lay 110lNliapause eggs regardless uf wlwtller silkworlYls had been determined
10 yield cliapaL1sc or non-cliapaulle eggs by incubation l'OIH.litions, From this
it wal:) inferred that the diapaLlsc (cuLer ,vollid be localml in or about the
head,
/\. pupal ahdonwn isolated fro1ll lhoracic sl'gllwnts illll1wdialely after pupa-
tion' was clllllluyc:'cl to Lest a ruudion (If llt'gptl til" organs in question, It
is call1'd "le!1L alxlunlCll" illig, I I, Eggs in t(':-it alldorllcn shall be neutral
flll' diapausc. nature. TIll' Il':;L l".t1'l'it-cl out hy illlplalllat ion of organs or
a sLl!1[ledccl org'<Ln togl'llll'r wilh pI'o[hOl'al'il' ,ldnnd \'vllicIt promote the dif-
ferent iaLiol1 or lIw It'st alldollH.'Il,
Implantation of each organ sllch as bruin. corpus allalull1, salivary gland
and sulJot'sophageal gland aneI of various c()mhinal iOJIH of these organs gave
no satisfadol'Y resulL. Only t 11<: implanl of gallglion, however,
exertl'c] It certain erfed Oll lite gruwin.1.!; l'gp;S ill I he t(!st ahdolllen und made
thCll! diallllUSL' egg (Figs. :: and ,[ S(;), Till' ohtained wilh test abc1o-
IlH.!llS [llen !1uhstan1iakcl hy l ).!;<lng;lia into
the silkworm of Ilttlul'l', In JUri[), it. was l'l'vt'alcd lhat the gan-
glion plays an illljJ()J'lalll part in dl'kl'lllillllin,L\ diapltUHt' llalmc, liberating the
diap<tusc factor,
Tlw larval gallg'lia inlo l'l'l'ipiunts \vill dl!vdop and will take a
furm like thaL of ganglia (Fig', :l).
\.Y<m"jlhLl11 \ll' \\m)'t lan'(\ ',', <\1:H.\
and molh (i'ip:, Ii, BSCI (L!lll'lillllS ill alll'rill,L', 1l1111-diapam;ill,l( l'ggS inlo diapctu-
sing Jiur[llt'l'Illll1'1', it h; [hal I Ill' sulHll'sullhagcal of
Tussal' silkwo),lll, .llll/w/'ll('([ j>('I'Il,1'i (Fip;, [iI, whit-Ii ('Ilk}'" inlu diaJlClUsL: in pupal
forlll also (lOSSl'SSC'S [Ill' ahility lo l11al((: l'ggS ill' /lrllllhyx silkworm Idilernate.
In lD[i(), lL",sl':(;,\W,\. dist'uvC'rl'd Ihat tltl' I'al'lll1' origitmks in lhe
suiluesllilhageal p;allglillll, The I'at'( was indl'lll'lIt Iy r!lulld hy :1, FUKUll.'\. and
he has l'sLaillishucl a hyp()[ltl'sis that Imtill is (If primary ililpUrlanl'l: t.o conlrol
the vulUnisl11, activating or SU[Jlll'l'!1sillg 1I1L' fLllll'1iOli of sul)(ll!!1opilagl'l.d gan-
glion by way uf cin:ul1l-ocsupitap,'l'al l'UllllL'C[ i ves (Fig. ::. DC I.
In all CaHC!1, it is jusLiJiahle to say Uwl 111(' sulJocsophageal gan-
glion liberates the diallau!1c fad or. Furthermore, Ilw transplantation of the
gang'lion l1Hecl tn ojlLain (IhllHlLlSI' or polyv()itine enabling iO
preserve the race wit hOL1 L repeal ing several gem.'rat iOlls \ I IJlSI':(;AW A 1 DfiO),
VlII-3 The neurosecrct ion in the siI kworm,
It was established by FUKUDA (1940--'44) thaL the larval molting <lnc1mcta-
95
morphosis of the silkworm are controlled by the balance between the hormones
liberated fro111 the corpora allata and frol11 the prothoracic glands. But it has
not yet been known that brain of the silkworm plays any role for molting and
metamorphosis in this insect as well as others, for instance in Rhodnius pro-
lixus Hl40-) and Piat)'samia cecropia (WILLIAMS, 1946-) ..
It is believed that the phenomena of the molting and the metamorphosis
in various insects have been correlated with the neurosecretion of the lJrain
(SCHA.RRER, 1 (54).
It has recently been reported by HASEGAWA (1951-) and FUKUDA (1951-)
that the diapause of silkworm is controlled by the diapause factor secreted
from the suboesophageal ganglion.
KOBAYASHI IW8 performed physiological and histological investigations on
the brain, the subo;)sophageal ganglion and the corpus allatum during the
metamorphosiH of ]iombyx 11/ori.
1. The hormunal function of the brain.
When a pupa is extirpated its brain immediately after pupation, 110 im-
aginal differentiation is obServed in the pupa. As it is in a cliapause-like
state, the pupa is called" Dauerpupa". If a fresh brain obtained from a
normal pupa is implanted into the head of the" Dauer-pupa ", the latter be-
comes imago in 14-1 (l days after implantation.
The above results that brain affects on the imaginal differentiation
probably by secreting a hormone. It seems that the hormone secreted from
brain initiates an imaginal differentiation affecting the prothoracic glands in
the silkworm (VIlI--J, Fig. 1).
2. The cytnhistolDgy of the neurosecretory cells in the brain.
Several giant called neurosecretory ce1ls arc observed in both the
intercerebral is and the ]lrutocerebt1l111 in the hrain of the silkworm. A secre-
tion liberated ['1'0111 the cells flows along two pathways; the
Ol1e if; that [rom the l'ci [8 into corpus carc1iacul11 through the nervi corporis
carclaci and I lIen corpus allaLulll through the nervi corporis allati, and
the other is thai from 1 ht' brain to the h[ood circulation through the cortex
in the brain. The latter HCCI11S to be either a brain hormone or its precursor
(VIII<l, Fig.
3. The nenrmwcl'ctory cells in the fluboesollhageal ganglion.
Suboesollhageal contains also it number of neurosecretory cells,
larger than in diameter, which differ in number according to incubation
temperature, high temperature incubation giving larger number of neuro-
secretory cell than low temperature incubation.
The stainability of the neurosecretory cells in the ganglion during the
period from 1he spinnig time to 24-48 hours after pupation is weaker than
those im.t11ediately after the 4th larval stage. B\.lt the cells from the 72nd
!:l6 T. Yokoyama
VIII-3 ion in tlw (1)
l.]I,'Iolh uhlair1l'd b\' rl}imrdaIlL.ltion ,1,l.\rillll,\l 3nJ tillY ilfll..'l IJllpalion
O. The numher of neurosecretory
c(dl:-; in ganglion
of brain Intu the hCiltL tlf 1111
cl(!rnul pupa.

...
1l.lj;,o

o
.... -:-: .
t Ilai/Il : r. .. I
'';; TU:illl'd III "
'\ TIl'rtllid 10 liJY
/) lIumrlap,llHil.'
/
2,BHtin ut i!!ntJ hour (If IVth illsuu' 5, Hru.ill ill uller
NI:!2 &
xl:Jl:1 : :.'(). 1-- _ . .,..;..'
Utl)lItgtlllGI:l ..,r Il\()lh
Iu
lit) 711
NIIUIlIP(- III Cull')
Each pniltl the llllm
bel' (II' \l('UI'llsNTetory cells in
(11'1[' gungliol1.
3.Bruin at 90th huur or Vlh imtlur
0: Tlw 111('al1 value of the number
or cells.
M. KOBAYASHI
Fig. 1.
Fig. :!.
Fig .
. )
".
(Tlw l1l'lIrm,('cTd illn in Ille silkwOI'll1) [':xplnn:ll iOIl of Figure'".
A molh o]Jl:lillL'd by r('illlplanial ion or brain inlo IIH' IIl':ld or a "Dmw,' pupa ".
Tlw I>r:lill lix('d al 111(' i:!nd hOllr or the 'lih in;;l:lr. An <lxh, from the neuro
t;eLTL'tory ct'll Itl Iwrvi t'orlH1rh; ('arioc] i,; Ilh;;('rvt'(j ill tIll) brain.
TIll' brain lix('d nl 1 Ill' !llllh Iluur of tlw Gtll inslar. J\ vanwie about
!ill,/I ill dianwll'I' is ill till' cyloplasllI hi stailll'd tlt'(.P with add fUl'hHin
and tl\(' sPrl'l'lory ll!aI('rial>, IIow inlll nn :IXOII.
Fig .. l, TIll' Ilraill IiXl'c] on Ill" :lnI day ai"ll'r pupal ioll. Till' llt'llrtlSC'(TL'lmy Cl!l1;; arc
lar,[(('r than thc' ('()nlllliln lIt'rvl' I'db (abollt 111," in dianll!II'r). A large vucuolc
is S(,('11 ill I hI' cyloplasm.
Fig. G. An adult hrai 11 lixl'd inHllt!di:llely aftt!r A val'uoll' contain
ing agglutinate sulHitallt'l' i,; "I>,;ervaldt!.
Fig. Ii. The lIumher ur cdls ill sUhlH'sllpha.I.\I':d )';:lllgliol1 is compared
bctwucn silkworm larvae Irl'ated so as tn lay c.hapallsl' l!!!,),(S and lhmw to lay
At the last larval i11;;tar. l.he I'orlller had nwl'l! cells than
the ]aLl_(!!'. In Ihis nCl1l'llSl'l'l'dlll'Y smallL,r Ihan in dia
meter not cOllnted.
hour after pupation to the previous day of the emergence are again deeply
stained. The stainability of the cells in the moth again becomes weak with lapse
of time. These facts suggest that the rise and fall of the stainahility of the
cells has been cOllnected with a quantity of secretory material in the cells.
97
VIII-3 Neurosecretion in the silkworm (2)
1. Sul>(I(l8ophngen.l'ganglion 30. Corpus rlllptum nl 96th hour
at splnnln,g stage, of Vth \nstar.
8, SUbocflophngenl gnnsllnn l1. allMum "15th day
immediately .(Ior 4th molting. niter pupation.
9, Corpus cnrdlpcum al 48th J2, Corpu$ tlJlallJm of moth
J10l,lr aftor 3rd moltIng. lmmedlately aCLcr cmergcmce.
M. 1{OBAVASlll
Fig. 7. The suboesophageal ganglion fixed in SUSA'S fixing mixture and stained with
Mi\LLORY'S at the spinning Blage. The neurosecretory cells in the ganglion
arc stained cleep reel.
Fig. 8. The Hubocsophageal ganglion fixed in BOVIN's and stained with GOMORI'S
chrome-alum hacmatoxylin-phloxin immediately after the 4th molting. The
large cells stained with huematoxylin are the neurosecretory cells.
Fig. 9. The corpus cardiacLll11 fixed at the ,18th hour after the 3rd molting. The
neurosecretory materials is in ,it (a phase contrast microscope).
Fig. 10. The corpus allatullI ilxed at tJle 96th hour Clf the 5th instaL The black
globules in the nervi corporis nllati arc the neurosecretory material. The
globu les arc also seen in a corpus allatum.
Fig. 11. The corpus allalull1 fixed on the 5th day after pupation. There are numerous
vacuoles in cytoplasm of the corpus allatuJ11. It shows that secretory materil
is elaborated in the cytoplasm of the corpus allatunl and secreted into the
blood stream.
Fig. 12. The corpus allatum fixed immediately after emergence. It has many large
vacuoles showing u degenerate state.
From these results, it seems that the diapause factor is secreted from the
neurosecretory cells in the suboesophageal ganglion (VIII-3, Fig. 6-8).
4. The histo-physiology of the corpora allqta and the corpora cardiaca
during the metamorphosis.
The secretory material originating in the brain is seen both in corpus
caI'diacztm and in corpus a/tatum, as reported by BOUNHIOJ." et al. (1953). But
T. '( uiwyum<l
the phYHiological signillcance of this secrc10ry material Hccreted during the
pl'riocl from lhe last instal" to the IlU pal stage, has nol yet beel1 known in the
silkworm (Fig. (KOJIAYASIII 1!l5R).
VIlI- 11 Diagram showing the hormol1al mechanism of the determination of
voltinism in the silkworm.
The diapause fad or which controls [he rlinpallse nature of silkworm eggs
is secreted from the suboesophagC'al ganglion during the pupal Btage. The
secretion is controlled hy the brain. The brain of the pupa destined to lay
c1iapausc eggs has a function by which lIw subocsophagcal ganglion is stimu-
latt'c] to release the diall<lllSC fa('tor, and on the cOl1tntry lhe brain destined to
lav non c1iapallse eggs inhil1ils to rc'lease it.
VIII-Ii Diagram Ilw 11IlnnollllI 1l1l'l'ilaniNI11 ,,(' the
(If v()llillism ill Ihl' >iilkwonn
SI;mul,lIi""
tlLld,,!' 1111' illl\llI'WI' Ill'
lilt' Llt'llIl'
t 1"'\II\lhaj!I'al
l'Utllll'l I i
'VilliI'll! lilt' Illlhll'U,'" ot'
Ihl' ,11.1P;lII',I' f.H 1111
vrrf-[j On the nllc of pl'otliol'ilcic ,u,"lnnd and corpora ;dliltil on metamor
pbosis in llw silkworm.
Salivary glancb;, protllOrack .t.;Hnglia alld fat bud ie's lakt'll (lui [rom (i-day
o1d larvae of the last instal' were transplant cd J't'Slll!('tivl,ly in[o u hody of a
old larva of the last instar at tlll! \llh l)ody sl'gment. After about G days
from implantation, it was ligalul"cd at two levels: 011C betwl!en the meso
thoracic and the postthoracic segment and the other between the gonadal and
99
postgonadal segments including the 9th body segment. 'the result was that
the post gonadal segments implanted the prothoraclc gl!ijqd pupated, while
those implanted another organs did not. The results show .. that prothoracic
gland has the function to make the silkworm pupate. The :(pnction is exerted
by me1jlns of some diffusible substance. An attempt to test the existence of
the substance was made as follows; a matured larva was divided into two
portions at a level between the 4th and the 5th body segment and both were
connected again with a glass tube. The experiment resulted in pupation of
the both parts at the same time, These facts show that the communication
of the substance from the prothoracic gland for pupation 'is not propagated
through the nervous system but by means of diffusible
Corpus allatum is also another important endocrine gland for metamor-
phosis of the silkworm. If corpora allata are taken out from body before
the critical period for hormonal secretion.in each instar, the silkworm does
not repeat larval molt but pupates, emerges lays eggs despite of, its
iustar age. The experiments were carried' out llsh1g third, fourth and fifth
instar larva.
VlII-5 On the role (If prothoracic gland and corpora
allata 011 metamorphosis in the silkworm
:'1 ",":t,'" ' I"" '1:'
, 1 .... ' .. ti
2. Puplltion 1)( pUlOteriur rHln
1. lndul'lion uf pupaliuliin pmitiHiur of ffilltUrt; Inrva connected with
uf 1I}(l:llureci Illrvne ' anttr\rJf purl U)' !I tube.
by transplunlutioll 01 p!uthoradc
gland
3. PUP;;I!! (If which
wruorll .dlalS! JUIV!! "
hcen ill
:lrd
1/1 .lIh imitlJf
(11111..1). In 5lh (Jr
(right).
. Cocoon 5 spu n
by larvae openucd
.as mentioned in
fig, 3
5. MIJlha frorp
lhe three pupae shown
in h,Q:, 3
U. Eggs l.:.lIlI by lhe :;howIJ in II!:!. 5
S. F'UKUDA
VIII-6 Parabiotic experiments on the hormonal mechanism in determina-
tion of embryonic and pupal cliapause.
100
T. Yukoyama
When two Bombyx females were connected at their heads, they emerged
parabiotically and each of them could lle crossed with male and laid eggs. If
two pupae different in nature (or c1iapallse, c1iapallse and non-diapause, wett'e
chosen as each of the parahiotically uniting bodies, non-diapallse-egg-.prbctucing
nature or one individual was chani!;ed to diapmlse-egg-producillg-nalure. This
shows the transport of the c1iap[luse facto)' from one to the other.
J>/1i1osamia c:"I1.lhia, whose nature is to diapaLlse with pupal form, and P.
ricini, which does not: c1iapause in any (orm are united parabiotically in pupal
slage. This procedure results in simultaneous emergence of both sides. It
may he explained frolll this that the llrcJthoracic gland of cynthia is influenced
to be activated by the brain hormone of ricini and release the hormone Lo
emerge. Two pupae of J>lIi!osmnitt cynt Ilia, one of which had been kept for a
year in a cold storage to acl i vale I he brai n for hormone release and the other
of which pupated newly and had not yL'i iwell activated, were united back to
back parabiotically. Both inclivicil1als cI1wrged sitnulianeollsly.
Vlll-0 Parabiotic l'xperilll(,l1t:l on the hormonal
in (kterminatioll of ('mbryonk [mel pupal diapuusc
1. "BHIIlII\ \ 111111 II. UlJikd III'"d III hl',ul
pafilhiuli. ;dl\' Il'IlIlI pllllai ,11\'
(In, 'v i! . .;, lli,tPOIII';t q.:J!
\llilll1l1"l'l :Iud lill' it
iI 1'1'111111, lilt, latlt"
Ill' ILl by II IliaIJall!,t' ")U!
b,iCl II
'" :1 d",i"I'1i 1101111 11(11;111111111 1111!' II
ot HI
fh',,11 ,Ind ;Ii/II,tlHrlU,.l
I, ')'1"",,1. . ..
i Hllmhp.: flHlth. Ifliitl
unill'lI JI,II',lhllllit'lIl1r hi.tl'h. III hack
ill'l'l'IIIHtiatlllJ.{.
Till' h tlw !iHIIW lI'i til{, 1.
l"lilulation hi 1'111' 'f [II
Jar I'm!; I, hU'1 lUI l'lfl,,'t 011 ':HIJ:
diapalL',"
,I. !\II-dill; JWlll paralll"lk 1111111/1
pi ,I ]Jupa kl'JlI ,ll lllw II'mp. 1.lr
tllW Vi'aI' and IlI'Y.'ly 1IL1j1iltl'd pupa
(If P'Ii'osamia ('y'lf"'11 I
S. l-'UK1.1UA
VIII-7 Influence of the unbalance between hormones by prothracic
gland and corpora allata upon the development of the silkglancl.
When a silkworm of the fifth stage was implanted with a prothoracic gland
Sill{worm Genetics Illustrated
101
of the young fifth stage larva, it did not pupate but repeated further larval
ecdysis except only a part of the mouth which pupated. 'Such a phenomenon
is called prothetely. The fact shows that the larval ecdysis depends not only
upon the hormone from the corpora allata but also upon the hormone from the
prothoracic gland. It seems that the antennae are the most susceptible organ
to the hormonal unbalance.
Sometimes there are cases in which the silkworm cannot spin cocoon due
to some unknown cause which cannot be attributed to any known disease.
Dissection revealed that the silkglands have some abnormalities. The ab-
normalities are also induced by a high temperature shock at 32C. given at
a certain period of the third or the fourth instal' which would coinside with
the release of hormone for ecdysis. FUKUDA (1956) assumed that an abnormal
high temperature condition during the rearing coqrse causes one of the major
factors responsible for non-spinning larva' frequently seen at farmer's house.
VIlI-7 Influence of of hormones secreted
by prothoracic gland corpora al1ata on
the development of the silkglanc1
1. Nurmul 5th intitnr larvn (Idl) prolhetcllt:
6th imslM larvn (riglll) prudu.'cd by transplantAtion
01 1 he prullwnH:!c glands.
1, Non.spinni'ng larvl! among
normally spinning unu:].
appear in
percl'lltanl's In a

5. Vnr!ous types of abnormal
silkglands from nOIl.spinning
one h normal),
ddormations of the glands
urc rutl\l.'r 'illglll tlwn those of
prothcwlk ones.
S. FUKUDA
VIIJ-8 Gene-controned mechanism of the internal secretions concerned with
the growth and development of the silkworm (S. MOROIIOsm).
(A) Traits of the genes controlling two kinds of secretions.
102
T. Yokoyama
A growth-promoting hormone (G) is secreted (1'0111 the corpora allata (CA)
and converts the duration of larval slagc, body weigh!, secretion of silk-matter
and 11101tinhHll (molt-numbers) into the direction to decrease but voltinism
(generation numbers) into the direction to increase.
A growthinhibitory hormone (II is secreted from the suboesophageal gan
glion I SG) and converts larval c1 Lll'atioll, body weight. silkmatter and molti-
nislll into the direction to increase !Jut vo,ltinism into the reverse direction.
"Maturing modifying gene" is located at about 1,.0011 lhe Z chromosome
and controls the maturity of insects. According to our investigations, this
gene is likely to conlrol the function or tbe brain and modifies the activities
of the corpora nllata anc1 Huhne::;ophageal ganglion through the nerve cOJ'dmis
sures from the brain. There arC' live 1 YlJeH; maLuring ILm'), inter-
nwcliaLe maLuring {I
I
"'I"I 1.11(1, 1.
1
.>11', 1 rII/'I} and late maturing (Lm). The
dominancy relation is L1JI>II."".;: ... !J."" 'II1II'.>Lm'.
"Major gene controlling molLinisl11" is located ett 8.0 Oil the VIchromosome
and likely to modify the function of the corpora aIIata controlling the growth
VIII-H Gl'!lcconl:lollerl ltH'chnnhilll (If the ilHC'l'llal spcnllinns COnCl!J'neo
wiLll I he gruwl h ancl (Ilwclo])ntcnt of lIlt' silkworm
(irur
rliff('wnlilllhm
of ',Irlilm flf IH'IW
l\f:lin g-erlC cnntmllifllf ......_.....

({)CiUkA, '31; SHEMUtlAllIA,
lInl',)
JrU(,f"{IClion of j{cmc
t'lJrpU9 ullut um
(N, e. Ul1t),)
Scx"linlwti mudif)'lng ..... Brain
(T,"'>-tf,m>/.m
t
)
{Smm. '2fi: N"OATOMO, ':!Ii: ([\111110110')111.
MllIlOml!Uu. I,m: TAKAfiAI{I, '!'i(,) \Inn.)
Main Rein'S mnl rollins'
vll,ltlnism r.-\")
(nrllwth :}
(J!OWUll(ll,. KIM, '3\)
Nl'rV(' }

C""i<' !llIlaner Ilil(. :11
I Mfl.lII)
'rim
fil'llk lnj{. Ijl
Il'/IJ,,)
'fili)
}
(MOROIIO'lHI !:1m! ':'iG)
Un\I,)
Hrllwlh
luhlhitlll'Y
rFlntllUA, ';,1:
IIAllluiAwA,
Bruhl hnrTllllrll' ....,.
"I'mlllll';Il'il-glallIl ......
illlrllHlIW
(FUKUlIA, "In: MUIHI(.A,
'111,
o.,,olopmOnlal 1",lann, (fig. 9)
Lm+Af I (iruwlh fUlwthm
V IUIU.:tlun
(MonUIlUl:illl, unp.)
1
IJllrmopal antagonistic
iJalance (fig. r),6) .. ,
I(;f(/l)
(,I\1UllOlltHUII. unp.)
Al'IlVUllon {If llrulhoradc
1:11\11(1
!WII,I,IAM!I, 'li: InIlKAwI\
,,(Jlht'r\:i, '[il!
-I- Variou. lypo. of ... Mullini,,", 7.HJ _ Vollinl"" Ifill. 7.8)
molting' cycle lIormo"al 1",lnncc 1(;/(;/) lI'lflTlOIUII balance (iii/G)
GenIc [)Ulnn,c (M/V/I.m) Genic hulan.c (VIM/I.",)
unp.) (MORilHOiUII. unp.)
(MOROIiOtlollh unp.)
S. MOROHOSm
103
promotion (parallel to the function molt-numbers). There are three
types; trimolting (M3), tetramoltil'llg' (+M) and pentamolting (l11). The domi-
nancy relation is
"Major gene controlling voltinisni" is locatecl at about 4.0 (perhaps +4.0)
on the VI-chromosome and is likely to modify the function of the suboeso-
phageal ganglion functioning to inhibit growth (parallel to the function con-
generation numbers). There are three types, univoltine (VI), bivoltine
(+V) and l11ultivoltine (va). The dominancy relation is VI>+Y> va.
(B) Interactions among three kinds of genes (Figs.
Fig. 1. Principle of developmental balance.
Left half, loci of the genes, Lm, IvI and V; right half the relationships between
the function of the brain (Br) moclifyl:n'g the activities of the corpora allata (CA)
and suboesoplUigeal ganglion (SG) aI1t[:the function of the corresponding genes.
The early maturing gene (Lme) sligll:tly accelerates the function of the corpora
allala and slightly inhibits the of the suboesophageal ganglion through
the nerve comtnissures. The late gene (Lm) weakly stimulates the
activity of the suboesophageal ganglion' and slightly inhibits the activity of the
corpora <Illata through the nerVe COl1'l'1l1isSLll'e. The function of the corpora allata
is strongest in the l'vID gene among three genes, .M.", +M, IvIs, and that of the
suboesoplutgcal ganglioIl is strongest in the VI gene among three genes, VI,
+v, V".
Fig. 2. Diagram illustrating the function of the brain (Br) controlling thc corpora
allata (CA) and suboesophageal ganglion (SG).
p, promotion; i, inhibition; t, relative secretory power of hormone; @,
@, relative quantity of corpus al1atlll11 hormone; . @, @, relative quantity
of ganglion hormone; ;=, antagonistic action of two hormones.
The functions of CA and SG organs arc different according to moltinism anc!
voltinisllI. There arc live types iu the sexlinked maturing gene controlling the
htnction of the brain: The early maturing gene (Lme) weakly stimulates the
secretioll of CA <lllll weakly inhihi ts the secretion of SG through the nerve com-
missure:.>, while the late mal Llring gent) (Lm) weakly promotes the secretion of
SG and weakly the secretion of CA through the nerve c0111missures. The
growl,fl.pl'omoting hormolle G and growt.h inhibitory hormone I act to each other
'aIHagl;nistically .. An antagonistic balance in the earlier stages determines tnolti-
nislll ;IIHI that in the later, vollinism. Moltinism is genetically controlled by a
genic balanc!:! (liJ/V/Lm) and vollinisll1 by a genic balance (V/kI/Lm). These two
arc important self.regulating characteristics throughout the life cycle.
Fig. 3. Diagram illustrating a genic balance between the major genes controlling
llloitinism and the sex-linked maturing genes .
::!. direction in change of mol tillisll1; --, -, , stable, intermediate,
unstable relation in chang!,! of moltinism; H. L. high or low incubating tempera-
tures. The early maLUring gene (Lme) diverts the major gene controlling molti
l1ism 'towards a dominant direction, and the Jate maturing gene
(Lm) towards a reverse direction, 1\11"-,,+ High incubating temperature
(II) COllverts into a recessive direction, MtJ_.'-Ii'J_.1\I1", and low temper-
ature (L), int(l a dominant direction, 1v1
9
<- + M<-1\I15. The modification of inoltinisll1
by ihcubating temperature may be clue to a change of the fun_cti(ln of the young
brain.
104 T. Yokoyama
VIII-8 (B) (C) Interactions among three kinch, of and antagonistic
balance btHwecn t wu kind!; of hormones
Fig. 3
--+ II --1-
_1.
:,
A. Organ,
(@)
r;(&)
11
r@ @
n. FUI1I'liom
bl. 2nd. :JnJ. ,Ith. [jth.
L:\f .... al (iura lion
Fin l. Ptindvlc IIr dcv(!loll[Ueolu.t balance

rM10
VIr", 1_ Y I
:I.il
-----_._----
(./IIf, ,I,m, tnt

! ,ell
L---------J
Alllag(llli::;tlr.
hnlaIH7c in IIII! hal;ltlt'(! in
eurllrr adVi!lIl'(.,d
J J
(M/F/{.>II)
(lellit' haianl'f'
IV/MIT.",)
i
1
1v!liltinbm (i/l)
IlpVcl0\ll1lttHal 11alann!
tlI itrllwl II L
[J('I,1('ll11Illwntal t.
FiR. ,I
--11--
<:"--- L <E--
Fig. fJ A. l-:.ICh Inslar
(!'iliffl'
,;-F1"'''.;;---..

Rl r::1 =-: __ J
III III 111 III
:1
II, AI\ \",'.,,1 stages
VrgrtatAv{' IIhiHi(' ncpro
1
\ucliv(' pha!ie
Mollillimn
'C
VOl lin I".
S. MCmOHOSlU
Fig. 4. Diagram illustrating a ,L,,'cnic balance between the major geneH controlling
vlllLiniHl11 and the Hexlinked mnl.urillg g()llCH.
<=, direction in change of voll:inislll; -_, __ , ,HWble, intermediate,
um .. t'able relation::; in dl<lngu of voitinislllj H. L. high or low incuiJating tempcra-
turcs. The early maturing gene eLm") divertH t.he majo!." genes controlling
voltinhnn [:owarc1H a recessivl.' direction, V'J,,", + J' -<- VI, and the late maturing
gene (Lm) towards a dominanL direction, V"-> II' -> VI. High inwbaling temper
ature (H) 'converts voltinism iilto a dominant direction, Vll_)o -I V --> VI, anel low
105
temperature (L), into a recessive ditection, V<-+ V <- VI. The modification of
voltillism as well as that of moltinisl11 by incubating temperature may be due to
a change of lEg'r{lllction of the young brain.
(C) Antagonistic balance between two kinds of hormones.
Fig.' S:' A. Diagnlm illustrating relative functions between corpora allata and sub-
,,' oesophageal ganglion.
The balance between the function of corpora allata and suboesophageal gan-
glion is affected by age; in the very earlier insLars the function of corpora allata
which secrete a growth-promoting hormone is little affected by the of
suboesophageal ganglion which secretes a growth-inhibitory hormone, while in
the later illstars the fUllction of corpora allata may be largely affected by that of
su]JoesoplHlgeal ganglion. Thus a critical point in the larval stage exists in the
third imltar, as seen in Fig. 5 B.
Moltinis11l as well"as voltinism are too complicated to explain by only a genic
balance as above mentioned_ ThuH an antagonistic balance theory in relation to
mollillh;nl and v(Jltinism proposed by the writer.
B. Diagram i1luOltrating a relutive function of two antagonistic hormones (G
and I) in the larvul lifc.
(C), the critical point at the third instar; 1. 2, 4, 5, each- instar. The larval
tllll'aLion of each instar is determined by the physiological balance of two hor-
mones and the wdght of lmv'll boclieH. The greater the difference of the func-
tions of the two hormones and tlw increase in weight of larval bodies are, the
longer becomes tIll! larval duration of each insta!"_ Thlls the larval duration of
each instar in tlw silkworm is shorter in the followiug order, as a rule; 2nd,
1st, :lrd, 1t11 and 5th instaL
Fig. G. A. Diagral1l illustrating the effect of age Oil the relative functions of the G
and I hormones ill each insta!".
1,2,3, 11,5, each in;;lar; Jl1, molting; p, pupation; white areas, growthpromoting
function; dotted areas, growth-inhibitory function. The rise and fall of the two
fUl1CtiollS in e:lch instal" is rl!sembling that of the entire larval life, though there
are some variatiolls. In the earlier stage of each iUHtar the growth-promoting
function is Huperior to the growLh-inhibitory function and in the later stage the
reverHe is the case. J\ cri tical stage of each instar exists in which the two fUllC
tions arc uljldvalenl.
13._ ])iagram illlistraLillg the relative strength of growth-promoting and
-inhibitory futlctiom, during the entire larval life. A, vegetative phase (lst-3rd
in;;tar8); 13, rCllJ'otluctive phase (4th 5th il1star;;); C, the critical stage in which
the two functions arc L>aluncetl; wliile area, growth-promoting function; dotted
area, growth-inhihit\Jry funcli\ll1. The earlier instal'S are the vegetative phase in
which the growth-promol.i ng function is superior to the growth-inhibitory func-
tlOn and till! latur instur;; arc the reproductive phase ill whi ch the growth-
inhibitory function is superior lo lhe growth-promoting functiou_
Moltinism and voltinisl11 are two important self-regulating characters in
the life cycle of the silkworm. The former is determined in the earlier
developmental stages and mainly controls the molt-numbers. The latter is
determined in the later stages and mainly controls the generation-numbers.
106
T. Yokoyama
Although both moltinisIll and voltinism are determined by the' of
the two antagonistic hormones secreted from corpora aUata and
ganglion, moltinisI1l may be c1elenninecl predominantly by the quantity of l1he.
growth promoting hormone secreted from the corpora allata, and voltinism
may be determined predominantly by the quantity or the
hormone secreted ftom the suhoesophageal ganglion. The brain controls the
secretive activities of these two hormones through t1W nerve commisSL1res.
VIII-9
There is the recessive trimolting, rl, besides ordinary dominant trimolting.
The former is on VII chrolllosome, D,96 distant from q. The duration of
feeding period of rl is different that of ordinary lrimolting 1\1\ the former
develops at the same rate as the telral1lollings, eliminat.ing the last stage
(VIII-iO).
VIII-I0 Comparis()n of lilll'al ion:-{ C'xpn'sl!ci by Hcvural
t.lw moll ill!,'; cJutnll'lcl' in Hombyx mori
day 1 2 3 4 5 G 7 R H 10 11121314 Hi 16 111.819 20 24 25 rocoon
weight
tetramoult.
.MI
I
I
1=-,=
[
o
recessive [
I
I I I
other J'Qce!lsi\,i?
I I I 1 :I
lrimoull.
t!'lmoult. Lr
iIf' . I I
'I
11lm.Jult.
11-// ___ .. I ____ J
lIt I
hatch lIloUlt. in!-ilar ripe
The worms homozygous ill 1'1 lWCOlllC tdralllll11ings or trilllolLings c1el.51m.d
ing on the nature (If environmental conditions. il is possible to
seled different strains of trinlOlLings or lelramoltings.
If the F
J
hybrids the two strains mentioned above are incubated
and reared in a lower or tI higher t.empcrntul'(:!, lhe longer the period of a
higher temperature, the less trimolting::l appear \Vm-Dal.
There are many devc[opmentally and genetically interestinp; behaviors of
rt when ;it is cOlllbined with dominant tritl1011ings, lIl:J or usual tCiramoltings
Ml (lX,
Sil k worm Genetics Illustrated
VIII-ga Change in manifestation of rt in the crosses of two
stnlins, A and B, inc,lbatioll and rearing temperature
black while lines show reciprocal crosses
between both strains
rtfl "'rl
pCTt:cnl.'!lf.' 01 rCCCS!;iVl' I ri,mo/ting
" ....... _ .. ___ .. ___ ... ____ _____ Jo
' .. 28
::3
28 E =s
"1':
.. 2" ... 23 ", 2H
'\. .... C... sHain A and 13 do nol l;h.mge
'. their rWlUfC hl nny londilion
VIII-Db Changes In 11101tiniSI11 by the combined
action of two genes, I't and M3
phenotype
gene combination
1'1 hal> a llaLure ttl cuL oJT the last instal'. III the case of NIHrt, how
ever, 1l10St. of larvae do llot bil1lo1t but become lila. In the CElse
of M I rt almost all larvae becol11e socalled "recessive trimol t ", but
sOllwt.imes othel' twu type>; uC larvae appear as follows: 1, normal tetra
mol t as same as in the case uf 1I1"l't, 2, uther type of trimoulting which
is similar lo M:I but recessive onc. T. EmoBE
107
108 T. Yokoyama
IX. Biochemical Genetics
The early stage of biochemical genetics oJ silkworm is marked by a
brilliant work of MATSUMURA. (HJ34) Oil the amylase activities. A remarkable
work which followed was a study of KIl,I<AWA on the tryptophan
metabolism. There have been numerous stud ies made on the chemistry of
cocoon coloring mat tel' and the pig men t in the integ umcnt. .l nteresLing results
are eXrJectecl in near future concerning the relation between a certain pig
ment and the viability or disease resistance of the silkworm.
1. Amylase activity in the digestive juice and in the body fluid.
As as U134, MATiWMURA investigated the gClletical behavior of the
amylase activities in the digestive juice and in the body fluid of the silkworm
and succeeded in Jincling out genes, the linkage relation between them,
distribution of Llw genes among various varieties of silkworm and the rela-
tion hetween the HlHylase activities and the vitality.
:2. Tryptophall mctabol iBI11.
In the cm;e of silkworm, there are many lllutants which are different in
the color of eyes ane! in that of eggs, normally, LIte moth laying
black eggs, while the wbitt(!yecl one laying while eggs. The black coloring
malLer is synllH:tizcc1 in t ht' body of sillnv()l'm through seventl steps from
tryptophan in the presence of genes, each corresponding tll a differenl step
(Kn;.KAwA ID:17, etc.). Tlw main route is shown. as:
Tryptophan-) kYllurenine :HlycJroxykYllUrenine -) Pigment
In a lllutant WI' kYllurenine is syntlwli:wd hut not 3-hydroxykynurcnine
sillce the 'lllutant has IlO cm:ynw concerning that: chemical reactioil. III another
lllutant l(l,!, :)hydroxykYl1urenilW is made hut not the pigment. 1t waH [oune!
recently by INA(;AMI (HlGfi) t hat in a mutallt r/l, there is no ellzymeH which
are concerned in the chemical processes to COllvert kynurenine into <lnlhra
ny Igl yci no and :1 hydroxy kynurcnine into hycll'OxY'Lln thrany lic acid, sLlch
cll;r.ymes exist in normal strains.
3. Melanin.
Melanin is a blackbh pignicnt commonly fUlind in the integument of
silkworm. There are several gei1cs related to the melanin formation. The
interaction among ane! bdwedll them and the genes of transparency of the
skin was sLudied by lhuw\ (19,1;1) and hiB collcagues. The cllvironmental
condition which affects the melanin formation was studied by HAIm:uKA (19
1
12)
In a mutant black pupa, /1)1.
11. Pterin, riboflavin and uric add.
In the integument of a mutant silkworm lemon, [em, there is a pigment
pLerin, water soluble and beautifully fluorescent. XanlhopterinB, leucopterin
109
and Ieucopterin-B are proved to exist in the skin (KIKKAWA 1950, ARUGA 1951,
et al.). Several investigators Suppose that pterin is derived from riboflavin,
a great amount of the latter substance is found i)1 the Malpighian tube of
the silkworm (KOYANAGI 1938) and the amount of which is closely related to
the translucency of the integument (KIKI(AWA 1944). The translncent integu,
ment of the silkworm is brought about by the lack or weakness in the
accumulation of uric acid in the epithelial cells of the skin, which is control-
led by several genes (luccr 1932, SHIMIZU 19'13); The uric .acid formation is
closely related to ri boflavin (HATAMURA 19
1
19) .
. In a certain instance the presence of pterin is related to the hardening
of the cuticle of the skin (TSLJJITA 1955) or to the virus susceptibility (ARl:JGA
1955).
lX-l Tryptophan metabolism in Bomby.'!: mori.
The work in this field was started by KIKKAWA in 1941 in connection
with the problem of eye or eggcoror formation, and then extended by
BU:I'ENANTlT, .lNAGAMl, etc. The figure shows the pathways of tryptophan
metabolism in Bombyx mari, especially during pupal and egg stage, The
conclusion has been obtained fr0111 biochemical studies using several mutants
such as white-l and white-2 elc. concerning tryptophan metabolism.
IX-l Tryptophan metabolism in Bombyx mori
Cll,-yH-COOH I
If Nil, -Vn.!J Oil
J II Indole lactic acid
l"rynlophon 011
ro Kyull,ln
I. 011
I'o,mylkynureulno or I Ai
t / _/ 9-COOH
O
eo.cll -ell-CDOII :-.. N
Nil, """ Kynurenic X
Nil, OCOOII
I(ynurcn!nc rcd blood I Nil, """"'0
1
CO-NII-CH,-COOn
..J...,wlu/o"l
Anlhronllk all Nil

CO.CII,-CII-COOIl "':Idm '
Nil V I Anlhrnnllylgiycinc
Nil, '_____ ,
Oil 6rr' N 4, S.Olhydro"yqulnolin'
'\
S:OOH ,cd blood I' ,,,JCOOlI
Yll-NII. r COOII 0' N "..-/ y
<;.g. 110 I Xonlhurcnlc
N" "N -NII-CIl.-COOa
:-... 0 :-.. I, NIl,
o Nlcoll"I.. 0 I 0 I
XnnthommOlln Id 3.H)'droxy.nlhronllyl.
(pigment) ac
--_Actu.l pathwQY H. K'KKAWA
----Unlikely Jl.lhwny A,
1(, INAGA"'!
110 T. Yokoyama
IX-2 Mutant .eggs concern Lng' tryptophan metabolism.
1. White-l (tot). In this mutant, tryptophan is converted into kynurenine
but kynurenine is 110 furlher converted into 3-hydroxykynurenine. Therefore,
kynurenine is accumulated in this mutant, and h1 a later stage of pupae or
just before hatching, kynurenine is converted into various substances, such
as kynurenic add and qL1lhranilic acid etc. as shown in the figure.
2. White-2 In this mutanl, lryptophan is converted into 3-hydroxy-
kynurenine through kynurenine, bul 3-hyclroxykynurenine is not converted
into pigments. 'l'11ere1ore, the accllmulated ahyciroxykynurenine is converted
into xanthurenic add and 3-hycll'oxyanthranilic acid etc., in a certain stage
ot pu[>ae and C!!'ms. Bul, ni.colinic add does not se'em to be produced from
ahydroxyanthntnilic acid in Homby ,
;1. Reel (I'l'i. /\. mULant of fuscanil1c pigment:;, KnG<AWA et af. (1954)
hllve shown thal the mutant lacks a purplish pigment in the compound eyes
o{ llloth and in Lhe serosa of eggs .
i. Normal ( f' I. kind:; or fuscctninc pigments are assllmed to
exist.
5. Arti ([cially pigment.ed eggs produced hy the injection of a-hydroxy-
kynurenine int.o tile pUJla of /I't l11utant. The compound eyes of the 1110ther
moth are also piglllented. These facts indicale thal the metabolic palhways
after :1-hydroxykYlluJ'lllinc. formal ion llormal in this mutant (11. KIKKAWA).
lX-!! MuLant ctlllt:cl'lIing tryptophan
No.1 white 1 ("'I)
No. white 2 (u'J)

,'.
,"
n
Nil. t nurm;\1 (+)
NII.:i Artifkiilily
cgll'!
I1Tfldurcd h\' the 111-
kl'( lUll (If 3lwtlruxy-
kynurt'niili' in II) lhl'
IHlllil IIr 11'-1 IIl'ulant
IX--3, IX-I!, IX.-G Tlwse three exhibiliolls show II,Ie studies hy INAGAMI
(19Gfi) Oil the tryptophan metaholism in the silkworm.
The body-fluid in normal strain changes its color into black when exposed
to air, while in a mutant strain l'b into red in stead of black. The difference
111
in constituents of blood between normal and rb is that in the latter there is
enormous accumulation of 3hydroxylcynurenine which is concluded by the
author to be converted into 3hydroxyanthranilic acid, which is again changed
into 3 yet unidentifIed substances F, I and L. (IX-1 X, Y, IX-3 (2), IX-5 (1)).
The red pigment which is a characteristic feature of rb is formed by
the oxidation of 3hydroxykynurenine, while the 11orma1 black pigment is
formed by the oxidation of tyrosine like sUbstance. If the both substances
are in different proportions the color fluctuates bet.ween red and black, for
instance, in vitro experiment, the mixture of 2 mol dopa, 1 mol 3.hydroxy.
kynurenine and t.yrosinase changes into blackish hrown, while the mixture
of 0.5 mol dopa, 1 mol 3hydroxyl<Ylll.lrenitie and tyrosinase changes into ,red.
The blood of individuals of the cross -I- X rb shtlws various shades of
color when exposed to alr (fX-4). The author added another example to the
IX-3 Manifestatioll of red blood (rb), concerni;lg tryptophan met'lboJism
exposed in the air
1. Normal body fluid --------, black-brown
rb hody fluid --.--".---_> red
2. Formation of the red pigment
1) Abnormal accumulation of 3hydroxykynurenine
3hydroxykynul'onine content ill the body fluid of some
lllutants nt instar measured by diazooxide method
normal
-I- 4O/lff/ml
white
till
white
U'2 52
reelblood rb 782
2) Formation of the red pigmellt llitro
Normal body lluid -.--------) black-brown
Normal body Jiu i<1 -I- 3hyclroxykYl1urenine -> red
dihyclroxy.
phenylalaninc
-I-
2 mol
1 II 'I
0.5 "
O. 25 "
3-hydr oxy-
kynureninc
1 mol
1. "
1 "
"
-I-
ty rosinase
(potato)
+
+
+-
+
+
+
color precipitation
black
blackbrown -H-
redbrown
+
red

red
yellow
T. Yokoyama
one genc--onc enzyme theory by showing that there is no enzyme in rb which
converts 3-hydroxykynmenine into anthranilic acid (IX-I, IX-5 (2)).
spot
!\
Il
C
[)
E
II
G
II
I
.r
K
L
IX-4 Variolls shades of blood color of rb when exposed to air
F, of (,1':' rb) 'pot"
of body nuld
,II luf'.a,
,b
tlO 1"I1ia I normnl rll
type lrpO
hnrir body n\l
nliid
I,
3hrdr"I
kynurenine

+ 2mol 1 mol -
3.h)'<lro"r _ L 101 2 mol +
krnur-unlnc n
Lyrol!linosc ,j' + ...
ahrLil'oxykynu 1'I!IIInl)
ubtnincd from tit!.:
(MdV Iluill of ,b
K JNAGAMt
IX-fi (1) Tryptophan metabolism of [family:>.: mori
l'aJll'1' chromatogram of tryptophan meLUholit it'" in urine of
sonw lllutants. Solvl'llt, butanlll: acetin' add:
I norma
IH lluort'scelll'l,! i
! 1
glul:ul'Onide of :l-CHIkynlL. II. IIH ! yellow i
:!:
sulphate of :HH!kynu.
!
II. Hi
I

:lhydra xykynllrt'ni IH' II. :lH
I
1
kYllul'L!llillu
II. hllw grelm
I
1
xnnllllll'Cnic adll 0. ,IH
yL'llow grl!en i
F 0. [i[i yellow iJlue
I
II
an t 11 ra ni lylglycilll' O. iH Jlurple
II
/1.Hdihydrllxyq llinolinc 0. HO yellow
O.H2 pale blue
II
ahydroxyanthranilk add 0.88 blue pllrJlll!
1
untltrani lie acid
10. \)2
jlul'ple
:J:
L I O. [is pale blue
kynurtininr -t : UJ1t_hrnniliC add
-I' tv
3hydroxy. 3hydroxy.
kynurenine rb anthranilic acid
kynureninase
1
1
111
II -1+1

-I
+
-I
II
m 11-
-I-
1+
-I-
-I
IX-5 (2) One gene (l'b) to one enzyme (kynureninase)
>.
'j 0.3 I
'C
' 0.2 D.kynuren!ne
.- 0.1
without D, L-kynurenine
1
30 60 90
time In minutes
EnzYIl1<:1tic degradation of kynurenine
51111. of 0.002 M n, Lkynurcninc, 5 mi. of 0.1 M phosphate buffer.
pH H.l, 211101. of Cl1%YlllC SOhlLioD, H
2
0 to 15 mI.,
temperaLure 37C . wavelength 360 mfl K. IN.A,GAll'll
113
IX-6 Histochemical on tyrosinase concerning melanin formation
of s-l11oltlec1 IS. KAWASU).
In an aUempt to demoJ)strate the mechanism of the melanin formation
in 1 he silkworm larva, hislochemical studies were carried out on tyrosinase
of black and while parts o( the integument of a mutanl "mottled Striped".
Tn general, the melanin pigmenL 1n t.he integumenl of the silkworm larva
deposits in ex()culicle, buL not in endocuticle nor in epidermis. And tht
melanin pigment is renewed in each molt, that is, at the molting period the
old cut icle casi 0[[ and new cuticle coni alning melanin pigment is produced
by the epiderm IL been reported that the tyrosinase &cti vHy in the
integ-ument of :iOI1W larval marking' mutanls measured by manometric method
is than thal of a normal lype, and the difference dudng the molting
period clue In that of ti1C tyrosinase activily in the epidermis with new
cuticle. However, m icroanalYBis 0 f the enzyme distribution in the integ-ument
1 observation on Lyrosinase
concerning melanin [ormation of s11lottlcc1
Integument C)C
Smoltlcd
Q
Schematic figurr of Integument
'''lih whit(: 'Iml blnck purts
of Sonoltlcd
Cross section of integument showing
changes from 4th to 6th Instar
114 T. Yokoyama
can not he done unless histochemical method is used. For this purpose, it
is convenient to use X-ray mutant" mottled Striped ", because in this mutant
there are while patches among the black melanin part and, therefore, both
wh ite and black parts can be investigated in Ol1e preparation (IX-6, a, b, c).
Tyrosinase can be demonstrated histochemically by observing the formation
of brown and dark produced in the tissl1e sections which have been
incubated in substraLe solutions. For the present investigation, freezing-
drying' method was adopted, and as substrates, tyrosine, catechol and dopa
were used.
Tyrosimlsc occurs in the exoculiclc, cndocut.iclc and epidermis as we}! as
in the muscle and adipose tissue (lX-O, h, c). In a [ceding stage, there is no
diffel'cllce on the tyrosinase action ilcLween white and black parts. At the
early stage of tlw molting period, the C'pickrnml cell--upper part or all part
of the cell--of t.he pigmented part was stained in slightly darker color than
Fn!e%ing drying section of integument
incuhatcrl wilh calechol Ht early Htngn
of 4th moulting
IX-(k with ('al('l'I101 at: middle
Hlage of 11th Ill()ulting ( :lfiOO)
innthatccl with clopa at middle stage
of 4th moulting (>:1500)
ext' ...
em: ...
nc ...... lWW
l ......
ty ...... lyt'oHinase
IllllH ... IllU"c!c
h, h' ... of 1Jluck and whitc
jlurtl-l in old and new
culklcH
S. KAWASE
115
that of the normal one. 1\t the middle stage of the moli when new cuticle
was produced by the epidermis, the tyrosinase action of the new cLlticle in
which melanin pigments would be produced was stronger than
that of nonpigmented part (lX"G, b). However, in the other part especially
in the basement part of the epidC'nnis, the difference of tyrosinase action
between the both parts could llot be observed. 1,'rol1l these results, it can be
said that the melanin formation of the silkworm larva is closely related to
the tyrosinase activity in the integull1ent.
JX-7 Influences of the dominant chocolale gene (I-a) Oll the manifestation
of some larval markings, egg color and skin color (S. SASAKI).
The dominant chocolate gene (f- (I) is located on the IX chromosome,
which always inhibits the formation of melanin in the larval exocutic1e
controlled by the genes belonging to pmultiple Ellleles, such as norm"l rmrk.
IX-7a [-Cl inhibit" tnc'lanin formalion in a
few larval marki I1gs
I hI/I, I-a/p,' I-a/I, PIp"
IX-71l [-If dol'S 1101 inhibil melanin formiltiol1
IIf b'hr:! (,7,{?) and Ursa (U)
"
116
T, Yokuyama
jng ()-1'), Striped (PI, Black 1/1'\ J\l[oricaud 1/1"'",,1r\ In the of lX--7a,
the inhibition of and J)" by r (l j,; illustrated.
The I-a gene dnes not inhibit t11Q blackish brown or yellowish brown
pigments produced by sLlch ac1ditionalmarking gem's m; Zebra (Zo), Multilunar
(L), Ursa (VI. When I-a is combined with om' of these marking genes, the
lal'vl:ll marldng becomes (lilllte in color. This is clu(' to the facl that I-a
inhibitH the melanin fOl'lllation by t1w ilIad: color g(:I1(, of the newly hatched
larva (I
e
/,) and tile fUJ1danlC'l1tal markil1g I 1
1
'1. In the pll, Ze, I-a or
pll, [1, ra individual, melnnin produced by }l'1 is inhihited, while black pig-
menl formed by Z(' OJ' U remains uninfluelll'c'cl (IX 'ib\'
In mating a hlack l\I.Ui; Clnd it white L the infllH'I1l'C' tlf the black egg
on the white onc l'lIntimws to cxbt in till' ('0101' or the nt'wly hatched larva,
and it is m-lLlul that, when ils lllot\l('l' I /n'l l'01H;titutiol1 till' young larva
with wt/w{ looks like a (ll1l'. 011 tl1l.' olill'1' hand, the colors of the
dominant chocolate YOLlng larva just aftt,!, Iwkhill,l; in tlte (TOSS of I-a/I-a
;<-Il't/U'j or in ll10 back (If F
I
'!lI'I/,I'1 of tW() ldncls of shades,
1. e., dark and pale. The' ronnel' 1)('['0111(''; n mot h and the latter
a white eyed. I\l10tl1('1' nol ict'alill' f,wi. is that the color uf hybrid F!
(UII/W! ',1 r ((IT ([ \)) is alnlOst l'olll1'iloss a!HI t lIt, Cll1'nl11ll,l.(l'll 1'('111<1in8 as kynure
nine ill the egg (lX-i l', (\1.
IX-Ie qf' 1"1 IIYllI'jd (/I',', /-rr.',1 ar<'
1111,] I'! 11'11 11 1 "-";< '11 l','m;lill!; :1:; !;"1111I"'llilll' ill l') ';.1 ';
If'l 1/'1
vVhcn l11dallill is (lII'111l'd fly i ", Jr", e
'
, /,1t, I "ZI', 1"rT etc, in
the larval eX(lcuLit:ic, X<l11111lIpt('1'in-l\ P1'Ot!IIi,(,(j I,y !('/Il ,!';C'lll' h SC<ll'cl'Iy recog
nized in the hYLJoderlll]s (Ii' th" l'l',l('i (Jil , 11<1\\,('\'('1', Wij('ll I If ,U;('Il(, inhibits the
formation of mL'lanin in tIl(> l'Xl)l'ul il'il' 01 tIll' 1'I',!(i<lIl, x<l111 hll[JIl'ril1B comes to
exist (IX-7 el),
IX-7d, lOri. In kll' of t 11(' larva wit 11 !';l'lll'l icalcll]1st itution,
+I-rt/_I_I-"', jJ"/ I 1', II/mil, Illere ('xisl it ('()w:iricra/JII: IlIIlJJ/)('I' uf I)iack pigment
granules (Pg), while in the lIypodcl'mi,; ur Llll.: l'i',l.dUIl 11ll' hlack pigment
granules are scarcely recognixed, as S('L'n in Fig-. ] 02, ill'cause in t his indi-
vidual the development of ,I black ", j)ll, is not inhibi1l.'d <Inc! black pigments
117
to form the pattern are accumulated in exocllticle as if at the cost of the
hypodermis pigments. In the picture the bJack granules are seen in the
hypodermis which are not of ll1elanin in nature.
IX-7c1, 102-1. In the exocuticle of the silkworm with the genetical con-
stitution of I-al+, pnj-p, +/lem black pigment granules (pg) in the larval
exocutic1e are few. In <ll111ost all cells of the hypodermis the brown granules
such as bpp: are observed, but their number differs according to each. cell.
It is concluded that these granules are not of xanthopterin-B in nature but
of tryptophan derivative.
IX-7c1, 100-], There exist a considerable number of the black pigment
granules (j)/;') in the larval exocL1ticle. A small number of the granules of
xanthoplerin-B are scattered in the hypodermis.
IX-7cl, 103-1. At the basement of the nodules (nod) few black pigment
granules (pg) are found. In the hypodermis there exist a considerable nutn-
ber of transparent and small granules which do not' exist in the case of
Fig. 102-1. These are considered to be xanthopterln-B granules. When the
production of the black pigment In the exocuticle is inhibited by the func-
tion of I-a gene, many granules of xClnthopterin-B come to be found in the
region.
IX-7c1 Interaction of pn and I-a genes on
the formation of xanthopterin-B
(0
f
1
-"/+T-tl,PIl/IP, +Item
+1-" /1-1-", pH !-I_ P, lemlfem
nod. nodules
cxo. exocuticlc
encl. endocuticle
hypo hypodermis
1-"/1_, pH / -I'P, +Iiem
l-a/+,pD /-I-P,lem/lem
x-g. granules of xanthopterln-B
bpg. brown pigment granules
pg. black melanin pigment
S. SASAKI
118
IX-8 Relation t he eli! ute h lack (lid) and several markings,
The elil ute black elmt rolled by a rC('l'ssi vc gene fnl iH characterized by
dilute black color 0:1 dorsal ::;idc of larva, with darker color on the thorax:
and caudal portions, '['his characteristic is manifested in homo7.ygous incH-
vidual of bll, but uncleI' lIClc"ozyg(lLlS c()ndition IJr/1llodil1es the mallifestatiol)S
of such markings as 111)rnuti I f P), Sll'ipccl (Jh, Zebra (;(0), Multilullar (L),
Unm \U), etc" by darlwning' the culor of these For
brownish lllultilunar 111arking' is ctlnlm;l perfectly changed into blackish Olle
in bd/bd, L/L inclivicltlal, hut is partially changed in In/II, D/L one.
I X-H [xC' lal iOll 1)[.'1w('('11 t hl' tlilu( l' hlack (hd)
and SI'VI,I':Jl lIIarld ngf\
I"
7,'
T, 11
lJl'liu
llutmnl Slrilwc\ MlIl'i,lIl1d 1"hl".1
Multi"

IUIIIII llUlll'rn
I'
}I,l ).jl',d /" /Iii 1/ "d
lId
I. I,d UM
II. (\1/111,>\ ,I\( N,
(',\'::')'a Transpi'atllafll)n Ill' HIl' (arV;[( lllf('gUl1ll'llf I)':' i'{\(;\SllIMAI,
The flf l<li'val in(\'!(lllli('\11s wnl' ('arri('d out lly using
sl'vl'ral [arvid markings, it.'llIlln y,'IIIIW and utl-t r<lldul'(,llt ill thvir :lrc1 larval
inslal's, ant! p!tYSill,LKI'IWI it'al Wl'I'l' lIIil(]i- (In tltt, transplanted parts
wlwl1 tIll' host elltl'rl'c[ till' Itll hrv;il in::tar. In tltt, intl'.l\ul1wI11fl of
stripl'd (Ir') , ,'_"lllUUIL'd Is'''') and IIILtltilllll,tr ({,I with-II I'llI'm llll'lanin in their
cuticles Wl're t I'ansplalltl'd til t plain 1 III OJ' 1Ii1l'l1Iitl I I "I, 11ll'lanin pigments
were not ohservl'tJ ill III(' donor's l'utiek::. llrk al'iet ill tlw of
the j>" in natural POSilion (nut Ireal('(\1 was vcry liltll! in IllwnLit.y and the
hypodermis was nll( t:lailH'rl wi lit lll'lItral 1'('d, Illil ill tlti' (';\S\' Ill' (he tran.s-
planted piecl! uric acid in 1111' jr" illl'l'i'ilS(,(\ Illuch and
with neutral reel. In this \:aSl', Wl' l'ollle! j'('('().f!:lIize t!tat Ihl' of uric
acid were transl11i (teet frol1l t Ill' host's Ii I () tIle' clOllor's OllC,
However, in the Citse lemo!] 111'111) w11 ich cont ains xanthop.
teril1-B in the was transplanted to tilt' normal, xanthoplerinB
was not affected by the transplantation, i\nd when oil-translucent (od) in
Silkworm Genetics IIILlstratcd
119
which uric acid is very little in quantity in the hypodermis and the integu-
ment is transparent, was used as clonal', the hypodermis of clonor became a
little whitish or transparent.
When the transplanted individuals had grown to the 5th instar, melanin
which is due to pH gene was formed in the majority of the transplanted
pieces, but no apparent differences between the 4th and 5th ins tars.
From these experiments of integument transplantation we can consider
that the functions of b?th gene and cytoplasm have important effects on the
manifestation of characters:
hUBt:
dunor:
hOBl:
donor:
IX-9a '[nlllsplantation of the larval intogument
lem lem +1'
!emK [emK {emK
-( 1J
lamK
lem lem +P
tP J( p [( ps K
Smt
SlItt]i:
Smt Smtlem + P I-a I-a
pSl{
I-Ct
p' [( SlIttf( S'JLtleml{ Sl/I1lemJ( Smtj(
t
P
psl{
P
LI{
pi ch P
+1Jl{ odK
dark is stained with neutral red
IX-Db (upper) Transplantation of knobbed (k) (E. NAGASHIMA).
When the protuberance which is formed clue to the gene function of
Knobbed (1\) ill the !JIll segment was t ransplunted to the normal (+k), the
clonor's integument was not lJuric.c1 into the codol1l of the hosi owing to the
protuberance. So, a Humber of pieces fr0111 the 5th segment with Knobs
were used as donors in all these experiments of the integument transplan-
tation.
IX-9b (lower) Transplantation of striped (P') and Smottled (smt) (E. NAGA-
SHIMA).
The larval integuments of the normal ( !-P) or S-motiled (smt) were trans-
planted to striped (PS) and S'!11ottled (S"'t), and the melanin formation in the
host portion neighboring the donor was observed.
120 T. Y olwyamu
IX-9b Transplantation of the larval integulllent
1.ran::ijJlanLaUu!l u[ knobbed (in
hosL : jl dUllor: II' Ii: host :Il'J( ciOllOl' :IPK
transplul1tation of striped (P') and S molt led
host: I'" I JI K
NAGASHIMA
Tbe format ion oj' ml'lanin whkh is (lLw t() thv gl'J.W function of P' is
wry lit tk or nOI1(', and ill cast' tltl' , .... n" is llsed. ,tS the liost and donor the
lllottlings arc scarcely IVlmcovur, (he hypoderlllis white
anel rail hc stained wilh 1H.'utrul n'd.
pitcllullll'IlCl were frOIll t.he ca:,;l' uf /i" integullwni whkh
was transplanted to /),1 individual and SI) It is consic1ered that these
differences occurred hy the dfl'L'is or the donor ii/I I .
lXDc Transplantatiun of !-il)oLy (Sri) IE. N!$i!\SJIIM.$.
In order tu kllUW Lhe of Llw melanin formation in thE! pupal
integument ill case or Buoty (so I which forms black melanin
in its pupal cuticle was used.
When the normal larval ill [l'g'1I1l1en!. with J\nobH ( I l'K) waH transplanted
to in the 1th-fit 11 i11sta1's, the color [one of dOIlur'H cuticle in the pupal
stage was brown, showing no cUrfen'ncc from 1.11(.\ normal one.
On the contrary, in case the sooty was tranSplanted to the normal, the
donor's cuticle showed brown color and not black, and the melanin which is
due to the gene function of sooty was not recognized.
121
IX-9c Tran!;plantatiol1 of sooty (so)
host: so donor: + P J( host: + P donor: $0
L'llrly stage
middle !stage
last stage
formation of epidermis
122 T. YtI!wyaIlltl
IX \Ie[ Sl'cliol1 of I he lransplanted portion (E.
The sUl;LiullS or the part beLween the host and the donor in
the integument transplantation were observcd under microscope.
In t11e early sLag", the syndtiulll which was (ormed from blood corpuscles
COll1Wtted both tissLles, and th:.' S,'cl)[Hlary layers from both sides extended
t hemsd ves ancl lmike\. Tlwl1, Of ;\ Iluclt-i or the merged with
each other and (Drtlled I he mlc[ci (If the !lew hypodermal cells, These cells
lied in a Ii nc L1lldl!r I he ncwly [urnH.:d se,'onc]ary layer anc1 the connection
IlC'cumc complete.
IX10 Ilbck pupa If,/) and temperature I.M. IIAHIZUI(A, Hl47).
(;V1W /II), w 11 kll I:; n':;]JIIIl,;ihl(' i'ur tll(' dVPllsit illll or dark pigments in
pllpal illll';';UllIt'IlI, i:; located Oil IT,I positiOIl or XlI-chrolllosome, The gene
is wieldy ciistrilwtl'd <;ilI\WIII"lII ill the Chinese uni-
vollilll' strains. 'i'u;l wilh til(' wild Hllm"J'x lItaJUfarinrt shows
I !lal /'1' ulso rL's[JllllsiLdc for t Ill) black [lupa of n. 1Il({llriarinlt, which is
IU(lhd II 11111 I liS wild IYlll' (Ii' clOlllv;:tit-al.uc\ silkworm.
Tile pi,l(IIH'nlat iUll !InletS,: ill lilt' 1>1at'k JlU [J<L is highly sLlsceptible to
Iclll\ll'I'atu\'(' l'llllcliliullS, Th,' lCllllll'ralurv dfC'l'livc p('riod I.T, E, ['.) is rather
narrowly limiLed, rallgill,l': 1'1'11111 I ltc' l'lul Ill' 10 Hl'vcral hours before
[>II]1al i()ll; l'. g" :.!() iI(lIll'S al dural illn or 1 illl(' lWCl'SSaI'Y Jur il1-
ltillitillg tIll' rlJl'llI:tlilill l'll<1l1,P:i',; witll llIi! according to
Ill,l;arillllllk CllrVl', 11ll' lligllll' lllL' Il'III]ll'rat Iltl' slwrl.l'l' I Ill' liuw, as shown
in the tallIl' Iwlow.
llllr:lli'lil til lilli" 1I"I"'",;:Ii'Y IIII' inhiililillg 1111' pi,l';lIlt'111
[mIlI:ll i"ll :iI diffl'i'I'lll
11'111[11'1':11111'1'
111
HI
!I!ill
,11111
11111
(ill
;:0
1'-
"
7
IlJi II ,
ill wall'!'
I (1011

,Ill
13
The Lemllcraturc-lillll' both ill it dry dl<Lllll)l'r and in llOL water,
follow t a fkr j\ rrlwll ius' i'lI1'JllU la, in w h kh t he It value for each
b IIll a \{'vl'l Ilf :iO,()(I() ral. which is dlw neither 10 the gT()w!h rate
llor to t]Ie IIxidatioll lIf ('l1ZYl1lt' hut to heal death of living !Jtdngs or protein
coagulation.
123
Tyrosinase activities of the body fluid showed difference between bP a.nd
-/- during the T. E. P. There is no difference, however, in the enzyme acti-
vities or in chromogen content of the blood during the period from the
ripening to pupation between bpinsects which are kept at 20 and 40'C.
during T. E. p" though the former becomes dark-colored and the latter not.
IX-10 Black pupa (bp) and temperature
!!. lctiviLy in blooU
during fJrcpupul
,
>

+
,
@
bP I,P
:WC dLlrillf(I'l';;P dO C TEr'
3. 50lublu prolt.:ill in Will,:{ l"Ulidc uf hp pupa,
/I) jllt'l hdorl..' pUllatiull
20'C Juring 30'C during
'tEl' TI,I'
4. DL'nuturullon HYPOll1C'shi" as to the of(cet 01 lcrnpcrillurc
UPOLl lhe cutic.:ular o( insect
WP. wtltcr solubh.l proleIn,
high lemp.
,
@)-
WIP, willer prolein
Water soluble protei n, which is connected with
is changed in nature by a high tempera
ture, and it fails to have a to be combined
with the l11uking the formation of
llwlanin impossible. M. HARIZUKA
Water soluble protein content in the integument just after pupation varies
according' to various temperature treatments applied during the T. E. P.
Treated over 28C" the quantity of the water soluble protein decreases. In
hot water extract, however, it is not reduced.
From the fneis mentioned above, it is assumed that water soluble protein
plays an imporlant role in the pigmentation mechanism giving a site where
chromogen sllch as phenols is oxidized by enzyme. A slight change occurs in
nature of the protein caused by high temperature, resulting in failure of the
pigment formation and loss of a chance of combining the reactioning system
which involves chromogen, enzyme and the water solubler protein, on which
the pigmentation reaction may proceed (I-IARlzuKA 1947),
124 T. Yokoyama
IX-.r 1 Differences in pterins in epidermis of several mutants (n. ARUGA
and N. YOSHITAKE'J.
Three kinds of p1.erin recognixablc in tlte epidermis of the silkworm
larva: leLlcopteril1, isoxanthopterin and xanthopterin BIT. In addition to these,
xanthopterin HI called xanthoplerin .(3) is also formed in the epi-
dermis of lemon lTlutant by the recessive lern gene. Much isox<tnthopterin
and lellcopterin is contained in the epidermis of Striped (P') larva characteriz-
ccl by hlackish marking, and xanthopterin Bj. and 131l is hardly formed in the
od-iram;lucent (oel) in which the uric add content ill epiclcnni::l is less than
normul.
IXII or ptl'rin,; in epitll'rllli,; of several mutants
p [em
P'
od i)' tem ad lem
Rf valLie mutant.
x:tlllholllvri II III yl'll(lw O. ,[[i
().
II
I
lid jJ" [em od (om
I
II I
plerin
colur ur , ,
JIuoresCt'IIl'C i sudium i
I
tll:el)t' i :I,'('lal('
I !
J) [em
I
X<lllll1Upl,l'ill BTl ydlnw \I.,IG \1.11 , I
I
i su,a III hupl" rill purple) 1I.:I:C
II
It'lI"[)ptl'ri II palll bIll, O. I:!.
II
II. AIW(iJ\ & N. YOSllITAKI,
IX Rd1l1i(J1l lJdwCCll ptcrin f()rmation and nil and (II. J\IWGA).
The ptcrin [ormati(}11 Ill' tlw silkworm larva was investigated
hy LU;ing (od'/l
l
) and (8
'111
) mutants ob-
tained IJY X-ray treatments. The lllOttkcI-lranslucl'111 is cllaraclerizecl by thc
mosaicism between l10rnml and translucent part.s of the and is
due to the occasional elimination of the fragment or chromosome carrying
.\.,,,1 gene during' somal.ogenesis, and mottlcrl-stripccl is characterized by the
appearance of numerous white patches on the black marking clue to the eli-
mination of the chromosomal fragmont carrying p" gOile. The xanthopterin-
Silkworm GenetiC's Illustraled
125
B [ormation in the epidermis controlled by lem gene was investigated by us-
ing ad
mt
lem and Sott lem larvae,
The [ormation of xanthopierin-B was markedly inhibited in the epidermis
where od gene expresses its action by the elimination of the fragment carry-
ing -1-(1 gene in od"" lem larva, and the same phenomenon was observable in
the epidermis in which P' gene ads to form melanin pigments, whereas xan-
thoplerin-B is fonned in the while patches where t.here are no melanin pig-
From these result.s it. may be said that. t.he xanthopterin-B formation
by lem gene is inhibited by the action of od translucent or p'.
When od,nt anel smt larvae are vitally stained with neutral red, phenomena
resembling the formation of xant.hopt.erin-B described above are observable
as to the stainability of the inlegument. The epidermal cells in which +oc!
and p genes express their actions are stained in deep color, but those in which
oct and P' genes express their actions are not or very lightly stained with
nculral reel.
IX-12 RclaLioll between plcrin formation and od, p genes
! motll.d I,"olu,.nl (odm'l
-_ .... _-_.;;{.
-----""'.'"

" ..... '
01
1
n, moltled .trlped (S.')
p
.... ---

..

adm' 10m odfI'Lt (8t1I1Tl(!d wlth nLiutrai red)
Sm, {em Slftl (fj:taincd wIth neutrnl ,"cd)
I
;
ept-M cuticle

culicJc
epJ- ''''' <
dermis ,.
II'
I ..i.'
I ,
)
I

H ARUGA
IX-13 Relalions among pterin, uric acid and 11lelallin pigments (H. ARUGA).
There arc somc close relationships among pterin, uric acid and melanin
pigments in larval integuments in certain silkworm mutants, St.riped mark-
ing controlled by P' gene is imperfect dominant to plain (P), showing darker
hlack color in homollygous (P'IP') than in heterozygolls (P'/Pl, On the con-
trary, the uric acid content in epidermis among three types, P'P', P'P and pp,
displays an inverse relation with the case of melanin pigment, i. e. PP>pj)'>
P'P'.
lsoxanthopl erin and leucopterin arc contained in the epidermis of plain
(P), and in addition to these pterins, xunLhopterin-B is contained in the larva.
of plain and lemon (p lem), The relationship between the melanin and pterin
126 T. Yokoyama
IX-13 Relations among pterin, uric acid and melanin pigments (Integument)
1. Relationship between melanin ancl uric acid
substance
quantity of melanin
quantity of uric acid
uric acid mg/tissue g
type_.
pp
pps
II
'fit
2. between melanin anc! plerin
5.8
3.6
substance I
quantity of
quantity of pterins
melanin
xanthoplerin B
I
isoxanthopterin

p
+1-
p lem 1f\. 1-
ps
-Ht
lem
"r
+1-
3. Relationship between pterin ancl uric ,It:id
.. ... .. _ ,-,
- " quantilY of uric quantity 01 plerins
P
oil
lem
lem od
------_ ..
I acit! uric acid I
mg/tissue g
8. 8
G
8.
,1
x'lnthopterin B iouxanthoptcrin
-/1
-1-
-111- -I

I eucopteri n
+
+
+
+
Icucopterin
+
+

formations has been investigated using p !em and Ii (eJil. In jJ' {em iudi vidual,
an antagcinisLic relation for the formation of melanin and xanthoplerin-B in
the manifestations of Striped and lemon was observable. Not only melanin
formation was suppressed to a certain degree, bLlt the Cjuantity of xanthopterin-
B also markedly decreased, whereas no quantitative changes in is()xanthopterin
and Ieucopterin were recognizable.
The relationship between plerin and uric acid has been clarified by using
lemon and ad-translucent ([em oil). The uric acid content in [em od is almost
the same as in ad-translucent, but the manifestation of lemon character is
markedly iniluenced by od translucent, the amount of xanthoplerin-B decreas-
ing remarkably.
Silkworm Genetics III llstrated
127
IX-14 Studies on pteridine metabolism in insects (S. NBS
AWA, AKAGUCl-n,
T. T.II IRA and M. TSUJITA).
In regard io the mechanism of pierin metabolism in insects such as Dro-
sophila and silkworm, the followIng scheme bas been proposed by the authors:
Xanthopterin B->2 amino-Ll-hyclroxypteridine 6-carboxylic
acid->2 amino-4-hydroxypteric1ine (AHP)->Isoxanthopterin.
In both Drosophila and silkworm, an enzyme oxidizing AHP to isoxan-
thopierin was found and its nature investigated.
PteridinC' metabolism dedllced from the studies
of Drosophila and Bomb),x
Cormnol1 all N
Pre(lIr<,o, N
J
)COOH
--t-C I __...,aoiJo .... Red
tv ) b' N se, ct Pigment
v, en, etC' t Lahyl
Brown Pir;mrnL Yellow Pic;menl

AHP Xanthir,e
Dehydrogenase __.... t j'Y ___,.t
cnPN Acoopto,) HN>
0
Ii H
Isoi:antlloptcrin Uric j\cic]
lVI, TS1:JJTJ\
IX-l S Studies on the maternal inheritance of the yellow-lethal silkworm
(M. TSUJITA).
Maternal inheritance of yellow lethal silkworm is shown by c1iagram-
maHcal figure,
IX-IG Excretion of uric acid in translucent skin mutant (S. SHIMIZU).
The crystalline substance precipitated in dermal cells of silkworm iaryae
is, in general, uric acid, which is cont.ained much more in the skin of normal
larvae than in that of translucent mutants, ill roughly proportional to trans-
parency of the skin containing it
In both types of the larvae, the crystals of uric acid in dermal cells be-
gin to dissolve and Dow out of the cells toward maturation for spinning,
128 T. Yolwyama
IX-15 Maternal inheritance of lethal lemon silkworm
Fcnmic moth
,
.
,
.
1
N"rm,'
Iintching
Ydtuw larva (Dies)
---Yst1ll0It
M. TSDJITA
IX-lG Excretion of uric acid in skin ml1tant
Uric ncid content in thl:' excrements of matulc larvae
lIinl1ll
normill (210 rng/g) ll'ansluccnt (5 lng/g)
TJrie acid (ontl'nt in dermal cclls
nrJrrnal (1(1., mg/rr)
(3 lng/g)
Uric content in the llrinc or muths
I)ormal nIsic-)
(5 lUff/g)
S. SI-IIMIZU .
129
finally disappearing at ahout the end of spinning. This change can evidently
be shown by chemical analysis of the skin, and it can also be observed directly
under microscope.
The difference between the two types of larvae, in uric acid excretion of
the Malpighian tube, ill not yet BO large at the voracious period of the fifth
age. But it becomes evident at the end of maturing when the crystals in
the Malpighian tube (" absteigender Schenkel ") and the acid in lhe excre-
111enl are much morE' in normal than in translucent, in parallel with the trans-
parency of skin. The same is said as to the acid content in the urine ex-
creted during the pupal stage'.
Amylase activity
A brief description of the studies carriee out on the amylase in digestive
juice and body fluid of silkworm is given in the following with figures from
IX-17 to IX-23.
Explanation of Lhe figures
lX-17 Simple test of amylase activity of the digestive JUIce and body fluid.
When electric current of high voltage, about 3000 volts, (the amperage
being small) is applied to the silkworm, it instantly vomits digestive juice.
Thus the digestive secretions can easily be collected. In most cases, two
drops of digestive juice were added to 2 c. c. of 0.05% soluble starch solution.
The mixLure stood at room temperature for 1 hour. For measuring Ule
amyloc1exlrinizing power, 2 drops of iodine solution (1/20 N Lugol's solution)
were added lo the above mixture, and the colO! thus produced was tested.
When the color lurns yellowish brown, the amylase activity is strong or
lX-17 Simple tc..,i of amylase activity of the
digco.tive juice ::Ind body flUld

high \olt
n1l111-
Ilmpt.rc
1
amylase
activity
(+J

amylase
activity
(-)
- nfter 1 hour
at room temp
S MATSUMURA
130
T. Yokoyama
positive (+), but if it turns blue, the enzyme action is weak at negative (-).
The body fluid can easily be obtained by making a small incision in one
of thoracic or abdominal legs, and getting the body fluid which drops out.
The amylase activity of the body fluid were tested in the similar way to
that of the digestive juice.
The author succeeded in selecting four strains which showed marked
difference in amylase activities iIi body fluid and digestive juice. Two strains
showed strong amylase activities in digestive juice and body fluid, while the
other two very weak or almost negative in both activities.
The results of his experiments which have been carried out since 1924
are shown in the following figures (IX-IS to 23).
IX-IS Genetics of amylase activities of digestive juice (A) and body fluid
(B) in Bmnbyx mori 1.
a. Two strains which differ in amylase activities in digestive juice, one
very strong (+), the other very weak (-), can be observed within the same
race or among different races.
b. Two different strains in amylase activities in body fluid, as well as
in digestive juice, are found within the same race or different races.
c. The strength and the weakness in amylase actions in digestive juice
behave as an a11e1011101"ph, according to mono-Mendelian heredity.
The strong amylase action (+) always dominates over the weak (-) as
shown (A).
d. The strong and the weak actions of amylase in body fluid also behaves
as an allelomorph like the case of digestive juice (TIl.
e. The two pairs of characters, can be represented each by a pair of
different genetic factors (genes), that is, the strong ancI the weak amylase
fX-1H Genetics uf amylase activities o[ digestive' juice (A)
and body fluid (8) in BO/1Z{IYx mori
.
, I I
l1I'jal'

t !m'
-1/(11' t /ar tIt
(Ill
k_',:::;;"t;l)
'I'

t/he
I /h/' -1/"(1

bt'/lie
((e. a gene of negative amylase activity ill digcsl'ive juice.
be. a gene of negative amylase activity in body i1uicl.
The enzyme activity of heterozygous individual Ulkes an
intermediate value of the parents. S. MATSUMURA
Silkworm Genetics Illuslraled
131
actions in digestive juice each by +n
c
and ae, and those in body fluid each
by +br and be.
IX-19 Linkage of the amylase genes of digestive juice and body fluid in
Bombyx mod L.
There is a complete linkage in female, and partial linkage in male. The
crossing-over value in male ca. 1%.
I',
IX-19 Lil1kage of the amylase genes of cligestive
juice a11d body fluicl in Bombyx mOl'i
r ..
x
I /It'/ ! bit
af! t /ttfJ t
l-__ ,-__ -'
I,

...... -- .
, _ __ _1 ___
ae b"/(le be
r,
(/11 1111 1 I tliJ bll/I bv (It' ('l",I(, I at' (l1/' II'
\N t)9
Till' l. rtll unr vdluc of hilI ham) ilbl gcm,:'l, ac and be. 1'!J C(l I
IX-20 Distribution or Lhe <l.mylase types in variOlls races of Bombyx mori L.
From tire facts as mentioned above, the silkworm can be classified into
the following four types as far as the amylase genes of the digestive juice
and body Ilu ic1 arc concc'rned:
1) I_"r I_Ill type, 21 I '''be type, 3) ae+
b8
type, 4) aebe type.
IX-20 of the amylase types ill various
of Bumliyx mori L.
UllI\oltlll(! UII11I. ...L UIIl\o!!inc
\ lutl Llll.{)Oll 1.ILl \\hIH lll(II{)I\ I
. __ J
I I
1lype
"",11 ....ail!!! ....
I- un (10 I no be
IT Iype ITlIIM IV type
o
Indl,m nluln\olune IIbht
r.lu.
132 T. Yokoyama
The amylase test was carried out with 67 Japanese, 107 Chinese, 5 Indian
and 28 European races, the results of which are shown in the following table:
Groupe Race n umber of each amylase type
I Japanese univoltine white cocoon (14 races)
II Chinese univoltinc whitc and yellow cocoon (45 l'aces)
III Japancse & Chinese bivoltille white cocoon (115 races)
IV European llnivoltinc white cocoon (13 races)
V European univoltinc yellow cocoon
VI Indian multivoltine light grecn cocoon (5 races)
+ ne be> -I- 'lr -I-b">ae+ '''aebe
+ae +''', ae+'''>+a'be, aebe
--I- '" -I' be, -I ne be, (fa I "'j>aebe
aejI"+""be, liebe, +ae +b"
ae -I 'Ir, aebe,
+ar +'"
By the app1ication of the amylase test, one can easily differentiate allied
races or strains in the same race, which are difficult to discriminate by any
other morphological or physiological characters.
IX-21 Distribution of amylase types among the indigenous races in China,
France, Italy, India and Japan.
As we arrange the amylase types of 207 races tested in relation to the
districts where those races are collected, the distribution of amylase types is
shown in the following table.
Districts Italy & France India China Japan
% % % % %
Amylase types
+(11 +IJI'
'1. 7 10O 30.G 32.8
+(1r' be
25.0 3G.I
ae-"!/lt' 55.5 39. '1 28.8
{Ie be 30.G 5. 0 1.7
IX-21 Distribution of amylase types among Lhe incligenous raccs
in China, France, Italy, 11lllia and Japan (207 races te-steel)
& France
Japiln
.......a .......a
S. MATSUMURA
'H +b.
av+
ac ha
Silkworm Gcnelils Illustrated
133
IX-22 Functional differencc of the digestive amylase among dWerent strains
of Bombyx mari L.
The activity of the digestive amylase gene +ae is variable with different
races as shown in the following.
Daizo> Ryulmku>Chi nese No. 101 > Koishimaru > Hei >Ch usu>Kojiki ..
Ihaku.
On the cont rary, the activity of {(e which concerns with the weak amy-
lase action (almost negative activity) is nearly the same for each race.
The amylase activity of the digestive juice of +ueoe individuals is half
as strong as that of 1-'" I ne ones.
Fj hybrid hetwecn 1_"" between tli[[erelll strains shows intermediate Of
the parents in amylase action.
The variable activity of the digestive amylase in +ae strains may be in-
terpreted by the quantitative theory of gene action of R. GOLDSCHMIDT. If the
intensity of the amylase action is controIIed by the quanLity of amylase, and
the degrec of activity of the amylase production depends upon th2 +a.
action, the variability of amylase activity may be attributed to the quantita-
tive difference of the +'" action.
IX-22 FUl1ctiol1dl cliffcl'Cl1CC of the amylase
among c1 iffercnl sll<tins of Bombyx mon
(
8
7
-780 (I1nl,o) p
I If I 0'/0.
am) lase Dcth Ity 'i 0 aml'lllse act I 0
a F,
" 10.
.. Ij am) laso nctlvlty 2" 2
1/+
'HuyJu c

/'1'> (R) IIk""u)
1 60 I( 103)
1 3'> (L 101)
05 (hul IlIflluru)
-... fJ(l (IILI)
'-. j'1i hu,u)
"J 50 (1\IIJdo Ih,11,u)
I(C 103)
107)
(f 7)
'- (C n)
p
f/J I +i"
aLtl\.ll} 7 1 amvb">c ncll\Hy.4 4
"
liilbC' r11.1lvlly 6 3
5 MATSUMURA
[X:!3 Amylase activitIes in digestive juice and body fluid in the 5th instar.
a) Di,l',c"live amylase.
1. Enzyme action of I a" larvae is feeble soon after the 4th moli, then
gr,\dually increases in its action until the full growth in the 5th stage
and decreases when nearing the ripening.
2. Male surpasses female in amylase action.
3. Through the 5th age, amylase action of oc gene larvae is totally negative,
134
T. Yokoyama
b) Body fluid amylase.
1. Enzyme action of _/_be larvae is feeble at an earlier stage, it increases
gradually in proportion to the growth at the 5th age, but declines
again near the ripening stage.
2. Female slightly surpasses male in amyJase action.
3. Through the 5th age, amylase action is totally negative in the larvae
of be gene.
IX-23 Amylase in digestive juice
and body lluid in Bambyx mm'i
--
t bl'/ '/It'

1-11
, ........ '--" 0
f, Ii .,
in 5th
1. ...ity (If digL':-.livc ilm ....
1.11

.:!
O.M '
0.7
' Illi
"
O,!j
0,,(
lit 1/1/1' j
-9
....... .:..
l'"!!-J-T-i;-"G7-
in ;jtll in\;IH
'J .. hi ivit r o( hody 1111111 ,l1nrJ,!,';('
S. MATSlIMl1l1A
IX-24 Genetics of Cal'OicllOid cocoon color
}I
'I
. '
\\ ,;
mirl 11q.;lafll!
Schematic figure of pathway of
carotenoid permeation
A: carotene %011C C,-carotene)
B: x<ll1thopilylls zone (lutein,
violaxanthin, taraxnnthiLl
etc.)
M. HARIZUKA
J - ([I-du.),
II cr X-L'lu.) .
Yl'IiOW blood
'iuol)' plnin wliite 1>100[1,
i ILitilJi{s }'
J II ), yellow blood inhibitor,
lTIay bl' alJtolic lo (/
C (XII-ehr.), g;olc1l,n YL"low cocoon
()/ ( II ), pale YL'IIow cocoon
c
s
/( II ). slraw L'olurt,tl ['OL'COL1
Ci ( /I ), yellow intwr layer of
CllC[)[)ll
Above four art' multiplc alleles to
white iuner layer I" and mainly COllcern
in permcability of :,;ilk gland to xantho
phylls.
F (VI-chr.), ilesh colured,
amI Cli, pink and cinnamon-huff,
when cocxio;t with 1;:
Aboye three COLlct,rrl in ".carotene
permc:ation.
135
IX-34 Genetics of carotenoid cocoon color (M. BA1UZUKA).
Two main pigments are known in cocoon shell of the silkworm; one is
ethersoluble, carotenoid, and the other is water-soluble, flavone. The caro-
tenoid is said to be derived fr0111 mulberry leaves upon which the larvae
feed, because there is exact similarity between the carotenoid in blood, egg,
fat bodies or cocoon shell and those extracted from the leaves (DIm 1933,
MANUNTA, 1033, '35, '37). Really, !1-carotene, neo-/1-carotene B, lutein, taraxan-
thine and violaxanthine, all of which occm also in mulberry leaves, were
detected in cocoon shell (OKU 1933, MANUNTA 1937, HARIZUKA 1956).
There arc kinds of COCOl1 color fundmnente;Ily. Chemical analysis shows
that each color has its own cOhstitl.tonts as shown below (Vid. IX-25).
COCOOll layer
I
t lrou- I
gllllUt ,
outer'lTIost
ollter-, mid- or inner-
I flavone
CUCOOl}
color
blood I cocoon 1------- '.
gUile I gene I
while in
a, 1;
blood or +1'
while in
any
yellow +"-I-lY: +1"_1 CJ
blood
outer
yelluw
straw-
culoreu
inner
yellow
lk"h-
colored
pink:
greell
"
"
"
"
"
G/,
C
C't
c
t
Flu
PI,F+u
Ott
abun-
dant
I
++
++
neo-fi- .
carotene BI lutein
I
++++
trace
+ +-1'++
trace
++++
trace
-I-
+ ++
tara viola-
xanthine xanthine
++ ++
++ +++
++ ++
trace
+
trace
+
N.B. 1,0.1-1.0; ++.1.1-5.0; +++,5.1-10.0; ++++,10.1-20.0;
-I- -I- -I -I- a!Juvl) :!O.l 11115/100 g cucoon reared at 23'C during the
lasl larval imilar (M. HARIZUKA UJ5(J).
It h; reasonably assumed that genes responsible for the cocoon pigmenta-
tion with carotenoid control the permeability of intestinal mucosa and silk-
gland for the pigment concerned (J uccr 1949). Following this concept,' caro-
tenoid genes of the silkworm are classified into two groups, blood and cocoon,
according to the sites where corresponding genes exert their action. -Those
of blood group control the permeability of intestinal mucosa to both c!'rQtenes
136
T. Yokoyama
and xanthophylls. Those of cocoon group are classified further into two
groups according to the kind of carotenoid to which they control the per-
meability of silkgland cells; carotene group sllL:h as and F and xanthophyll
group such as C multiple alleles (viel, VI<-l). Main portion in t.he silkglanel
at which each pigment enters is different. Almost all carotenes enter the
gland at the posterior division. of the middle silkglanc1 called carotene zone
and xanthophyll mainly at the middle division of that called xanthophyll
zone. As carotenes enter the silkgland at. earlier time of the larval in-
star, they advance forward and reach the top of the silkglancl at the ripening
stage and are spun at the beginning of the cocooning. Accordingly, they
always come to the outer layer of a cocoon. While the time of entrance of
xanthophylls is not so simple as carotenes which hereditarily explained
by IL".RIZUI(A (1956).
IX-25 Effect of rearing temperature condition during the 5th ins1.ar on
carotenoid pigments of COCOon.
WATANABE (1918) repor1.ecl that a depth of yellow cocoon color
was variable with change in rearing tempc;'rat me during the last larval in-
star. Ill' concluded that yellow cocoon color bccan1l' deeper with increasing
temperature. Since then the effect of tempcrature conditions upon the process
of development of cocoon color has long been clisl'u:ssed among of
Bomb),x in Japan.
According to UnA (1\)22), ill Eur()pean yellow COCOOll, flesh color or cream
yellow at the of cocoon at a lower temperature and yellow
at a higher temperature. Ill' insistecl that F Ulesh colon against
Y:I' (yellow COCOOll) at a luwer temperature and vic(! I1I'J'SIt at a high tempera-
ture. This was, however, denied hy (H):n) with the explanation that
the fact attributed tll incol1lplet.e ckvel0pl1wllt 01' llesh color by F at a
high iem peratu reo
I-L'\.Rlz1.IKA (lD[i(i) has estimated fraction of carotenoid occurred in dif-
ferent cocoon layer at different lire and fOllnd that a comparatively
larger W110UIlt of the !Jigmenl was acculllulated ill cocuon shell ilt J:i"C. than
in that at 17.5"C. or at 2S"C. The rate ul' lluctuat iun with lliffcrent Lempera-
ture conditions was larger in carotenes of tile ()utcrllHJst layer than in xan-
thophylls of outer or inner layer. In olller words, the develuping
of carotenes are marc to tcmperature conditions than those of
xanthophylls. He was able to explain the whole problems nwntionecl above
from the stand-point of the difference of SllSL:l'ptihi Ii Ly ]Jc! wucn both pigments
to temperature and the location of each pigment in (liffel't'.nt cocoon layer.
" At the Lime, Y was not C\ symbol of yellow bloot) gUile as present time, but
yellow cocoon.
IX-25 Effect of rearing temperature during 5th jnstar
on carotenoid pigments of cocoon
17 S"C JO'C
c
I?il II
, I. p"cQroteno
10 I ::;. U 1
'" IV. tnraxanthln
'"
? ___ n __ ,,-.. [L._ CL ___ V. vIol anthin
" IDWlVV lomwV JOINV
IUC Z,'C 30"(
PkF"'lltEJ tWill BfEI
137
138 T. Yokoyama
X. ,Breeding (Including Heterosis)
It is said that silkworm was introduced from China into Japan more than
a thousand years ago (X-I). It is not certain whether there had been any
aboriginal Japanese race of silkworm before the introduction from China.
However, the old Japanese race spins a peanut-shaped cocoon which is differ-
ent from the usual Chinese one which is oval or spindle in shape.
Most Japanese varieties were white in cocoon color, a few races
yellow. They were usually uni-voltine, a few being bi- or tetra-voltine.
were usually tetra-molting ones.
For breeding purposes, the egg producers reared the stock strains.
times they tried cross-breeding between Japanese and Chinese (X-I).
in the 19th Century European races were also used lor cros::;-brecding.
being
They
Some-
Later
K. TOYAMA studied hybrid vigor and empha::;izecl the benefits of utiliza-
tion of it in 1906 (X-I).
X-I History of breeding
Old Ja[JanC5C silkworm mainly from
aborigin.1I and Chinc:ic race!:> Int ruduced abOlll I
years L,ltcr in Hllh Cl.!lltlln' EurujJt:all !'ace!; Wl:n;
<lisa micd for
1845
1894
1D06

C-! 'j1 hivoilinc hy 1. NAICAYAMi\ &
( "_. .1 lInivfJllilW j. FUJIlIfI1T(}
- --I ]npilllcsC! by T.

K.'TuYAMA hyhrirl illltl
(If utilbr.atiun 111' it
Tht: Scrh:ultural Expcrill1lmL M.A. F.
ill 1!J11 <11111 ilk 1U1:L\UU\ of
hrceuinlr W<I:, acl:qltcd fur L1H! of
Vilrioll!; 1)[ il:. wdl .n.
hyhricJ!:I lhem
1014 C:J A new of Fr-hybl'itl \ltarh!ti
by the guvernment. \Vithln !il!vcral year!; 1!1L7-1!I:!:!
nearly all 5ilkworm adopted FrilyiJrid.
1926 improved with \;Ir!:n ;:UWHlIIt (If
!jilk
ID34. Rat:e!. illJapll'd I'llr !liv!'! \\'r:ll: ill't:d: I'.
WJfld lur nlW :,ill .. uf !.lIfH!riul' tJualLiy, 1'1lr
flli.1PIc :,ilk, fur for fl:,hillg and raCCn
welt ueJ.lplt.!u Cor rearing in I:l1\'ironllTt'nl.,
and til' whirl! al'l! rkh ill V, Ih
HH5 ill q\l;llltitr a:i qU;llil), ha:i
been and utili t.:uJllinuing advLlnt:ing.
Scientific silkworm breeding began in 1, when the Government estab-
lished the Sericultural Experiment Station. K. TOYAMA was in charge of the
breeding project. Practically all the varieties in the country were collected
and the characteristics of each variety were examined. Then,
139
between any two varieties were tested in their' qualities. The results of the
experiments were published in 1917, in which 156 cases of FI-hybrids were
recorded. FI-hybrid is superior to the parent pure races in the rate of de-
velopment, viahllity and silk production as shown in Figs. X-10 and 11.
In 1914, the government decided to encourage the farmers to reaf hybrid
worms by distributing hybrid eggs through the specially organized route.
pltlnvay of silkworm egg at present from breeders to farmers is shown
in Fig. X-1G.
X-15 Pathway of silkworm egg from breeders
to Bilkworm rearers

EXllCriml'nL
,station, M. A. F.
C)-c::::I
0-0
00
(hreL!d)
--1.---'1 riln ..'
IJrecdw:l (I cd I
\'mt f)l'udul"l.l':i

commercial egg
producers
c:J
0-0 ,0
(rour) (I"OOU'")
silkworm rcarcrs
c::>-eJ - O-CJ
0-0 0-0
(h,-.,,<I) (roar) (produ"")
I
Jfii2) grand 31 rc
p
rOdt1Ct.ive,.l)gg
neaC parent egg
o 0 ocommerclal cgil
Chinesc ' {F"hybrld, cgll)
The hybrid silkworm gained popularity among farmers so quickly that
it prevailed all over the country whithin about five years since the first dist
ribution, increasing the yield of cocoon per gram of eggs (Fig. X-H).
X-14 Diagram shuwing the relation of the propagation .
of hybrid eggs and cocoon.yield
(while circle, spring: bla.ck dn:le, 5ummor-Autuml1)
140 T. Yokoyamll.
The breeders have tried to improve the reproductive races in various
characteristics useful in practical sericulture, as well as to and the best com-
bination between them for producing hybrid. Various races adapted for clif-
[ere11t uses, e. g., good for raw silk of superior quality, for staple silk, for
flossy silk, for fishing line, and races well adapted Jor rearing in unsuitable
environment, and races pupae of which arc rich in vitamin were bred
around 193'1 (X-I).
;;
7ri
c
8
8
'0
E
"
''ij
"
1

g
40
30
20
10
X-3 Increaso in cocoon shell weight
".
".
PrilCtka] hyhrid
- -i7i'f.:fi:.;"" -1;;t Ho:i7,-:o:, 17,1 -:;"":",,,,"""'7
1
!:':':'l::"'n

X-7 Progre:-;s of the raw silk [Jl'H'(.'rtl<lgc of l:(ICllOI1
(Sprillg
____ years
141
Every effort was concentrated on increasing the silk production by a
single worm, sLlcceeding in raising the gain from about 20 cg. to about 50 cg.
during the period from uno to 1950 (Figs. X-3 and 7).
Usually high productive worms are not vigoroL1s. A compromise of the
two antagonistic characteristics was attained chiefly by the impr9vement of
ih::! rearing method. Through such improvement the increase in efficiency
for production of raw silk was resulted (Figs. X-3, 7 and 14).
The effort of silkworm breeding is at present directed to the. improvement
of the quality of' fa w silk sLlch as size deviation, neatness, exfoliation and
boiloff, decreasing neither quantity of silk nOr viUility (X-Z).. For instance
the result of improvement in neatness of raw silk 1s illustrated in Fig. X-g.
X-2 Aims of breeding
1. hluher vleld
of l'IlCOfln
2. belll'f rcclnbilHy
o{
3. Jess deviation hl
1101Ivicr weight of
sIngle COCOO/l and
richer lllHWlil}' rl[
sJJk
hCIlIthy !iilkworm
unfr.wot'llble one
and grctltcr h:mgth of favorable ono
cocoon filamunt II1II ----
uelll'f
of
[j, h.'!'1s
V, 10'. bollar{la'S
(IQSS quantity of seridn)
Several examples of practical breeding of silkworm are shown in Figs.
X-1. 5, 6 and 8.
SW:UIU, HASIMO'l:O, later TSUJITA, TAZIMA and TAKASAKI in the KUMAMOTO
Branch of the Sericultural Experiment Station started the breeding of a
European and a Chinese race to increase the silk production in 1927 according
142 T. Yokoyama
1
X-9 Progress in neatness of raw silk
(National avtJragc of ca.:h year)
93
92
91

0

90 0
<;

if
"tl

1
88
1937 '3S '3[.1 1,10 '17 I,m '50 'Gl 1!j2 '5;; '51 '55 '[16
X-S Selection for length of C()CllOI1 lilanwllt
in Chinese racL ... MK
(IY:l8)
gCIlr.!raLions
(1950)
N.llA'fJl.
Silk worm Genetics Illustrated 143
to the plan of Y. TANAKA, This work has continually been carried out,
suceeding in increasing the silk production of a single worm from 33,8 cg. (in
average) to 70.2 cg. in the European and from 25.3 cg. to 56,8 cg. in the Chinese
(Fig, X-5, 5a and Fig. X-6, Ga).
Hybrid silkworms are vigorous and high-productive in silk, but it is dif-
ficult for the egg producers to rear the pure races to produce. hybrid eggs.
Double cross or half-double cross eggs are more convenient in this point,
because the parents for the production of commercial double cross eggs are
.<l
1<

..
:;:
.5i
<
0:
rJ
<Il
j
]
l<:
Ul
(:!
0
0
u
0
U
'+<
0
v
bJJ
ro
......
(:!
v
u
'""'
Q)
Po
Q)
.<=:
'-'
'""'
0
'+<
(:!
0
'';:;
u
OJ
OJ
Ul
V

OJ
'""'
:;j
Po
<J.)
.<=:
f-<
i
K
144
T. Yokoyama
already hybrid and easy to rear and the number of eggs laid by one moth
is also large. Half-doublc cross eggs were first distributed by the govern-
mcnt in 1925 and double cross eggs in 1911G (X-141.
:!tll
Hill
X-5 ProgrcCls in cocoon qualities by [lL1re line: sclcLtioll
A. ran', n]o

1911
1t:11
/I
II
:'
.. ....
!, ,:\ /
............. "4 ". ,

......... "/"'" \ :: \ / '.. r

,'" '-........ I ," .,"
',.' r: ",-="" "
.
r-/ .. . '
"
.' .
. . . ...
....
.... \:"
:/ \
... .' '.
" 'I" I I
._'.1 \:'
...... ,
" .'
I
,
,< ,
.. " 1
.
.. ,_/ \. ,
".,'
l'iLllflllr 11ll' wl'il:hl ofa l'fll'()(lll from 100 Cfj,',lq',) til \1,,1'.') 111 Olll
. \1'11111 tllll rg) 1(1 (lit (i ill tilU
"f tlH' (',h:(IP1\ :11(,11 rail,,' (ruin 1(111 ll:l.-;" .... ) III lliH in 010
{nml 1110 (If I. :1.",,) to 1m III 1ilU
(If Ilw wdEdl! III pupa from lOOp. III tIl) (!!. I:! /1) ill 010
IllIlil 11111 (1 Plt'.) (I 1I161U
X-5a Specimen showing the progress in silk
amount of cocoon, a EuropeaI) race 010
145
146
T. Yokoyoma
X-
6
a Specimen showing the progress in silk
amount of cocoon, a Chinese race 610
HeterQsis in silkworm'
If we plot the value of anyone characteristic of FJ hybrid against that
of mid-parent vaille, we get a linear relation between the two, e. g. in the
cocoon weight lhe relation is expressed according to the formula:
'y=O.826x+O.609
.y is the cocoon weight of FJ hybrid and x is that ot mid-parent (X-IO).
X-IO Cocoon weight of Fl hybrid and its midparent value (spring 1944)
.....
g __..,
230
E
::; ;:i ::; n
Y
j
'" ./ j. l:: ., .:;:.. ii
Observed F,hybrld Valu .. (x)
_/.:)/ ... : :T . H
1.70 / 11.P' ":::,: ,
.---+ct:hctcrosis,F\-M 1. 7 r::.;0.83 3
t..:- /' __ M:mldparenlv"lue
..... 1',: F,.hl'brld
j 40 nx 1 1133 6810BI5 $,1.l0131l4 43 3122
I,U J 70 2.00
g
mld.parent value
C. HARADA
X-ll Hybrid vigor expressed 111 various charactetil:ltics
Ion
80
fiU
4U
20
(The figure In parenthesis shows the absolute
% or Yield of cocoon wt.!ight of w(,light of
monality fram 10.000 hatched c.Oi,oon shell sIngle cocoon
lilf'-,I!!
21 7%) (l7.3 kg) (405Lg) (U8g)
(31 5 cg (! 18 g
(13 51<g)
(122%)
o pnrQnlsF, hybrId P 1',
p r,
P F,
C. "ARAPA
148
T. Yukoyanta
In X-ll the absolute values of heterosis in variOllS characteristics in
economical races, mortality, yield of cocoon from 10,000 ants, weight of cocoon
shell and weight of single cocoon are shown to be \12.2% (PI): 21.7% (P)),
(17.il kg.: 13.5 kg.), 140.S cg.: ;11.5 cg.) and (2.0S g.: 1.78 g.) respectively (X-ll).
Very interesting two examples are shown in which the silkworm with
particular gene shows <l remarkable degree of heterosis. The one is K gene
(X-12) and the other is 1'1 gene
X-1:! lIt't(']"osis l'Hl1:>I.'(\ by gent.'
(K anel m/ action
knobbed normal
i,'!II:OC!1l wt.
C-J
11)0
r!'tl:;:i (ru!'>s
.11 "''' I
LJli:Nl 11.-
C ___ ) C_)
111) J LU

(:)
120
T.
K I kno\)h('c\ \ is :t ill n.() posit ion !In Xi-chromosome. There
is ano[her ,[;,cne JIl/' <I[ :!.l.() Oil I he' S;lI1H.' ch rtJllHlSUI11 e. IImUlm made comparison
Iwlween ,I comilinationS!lf li](' I\V() a!)()v(' menli()ned gene!"> and their
alleles, and found Ihal (11(' ('()l'IHlIl w('re arnll1,l(,ecl jusl in the order
of degree or or gelll' l:()11lIJinali()1l, i.
1 / + //IN 111/' J 00
-1- /I\_ Nil>! ))Ij.> I 1.1 ()
1/1 mN I
I IK 111/)/1
Recessive trillloiting rt i!"> a on Vll chl"oJ1)o!">omt'. If 1'/ is concerned
ill Fj-hyhrid, the heterosis in !">Udl cJwractcrislic as cocoon weight
or COi._'oon shell weight, especially ill the lattvr, is remarkably hig-h, i. e. 17()-
190?;; and respectively, in contrast to the ordinary F( cases which
103-1:25 anc! 105-125 in the respective characteristics (X-U\),
SJll(worm GCJ1CliLS rated
X 13 Heterosis caused by rt-gene
P!gurcs are ShOW11 in index for 100 of the mid parent value
F,-hybrld of
1'/ x +
Ordinary cases
01 F:L"J1_lbricJ
cQcoOn woighl

cocoon shell weight
(;)
125
In [0\ progeny of l't)( +, heterosis of oocoon shell{ mote
remarkable than that of pupa-weight. The fact may be ex-
plained follows:
In tt, the recessive Lrimolting', silkworm eliminates the 5th
instar of normal 11.h moller, the silk production is pOQr all
compared with lhe l1orm,ti. F, individuals of rtx+ beconie
'1lh moIler regaining the last in$lat, during which eill{.gland
gl"OW remal leabl y.
T. }Imom;;
COCOOllS Exbibited
Before the Mdji era (1792-1867)
149
150
X-17 Kinko
X--19 Koishimal"ll
The Mciji era (18()S. H118)
X-21 Holdren
X-23 Nihonllishiki
T. Yokoyama
X-l8 Matamukashi
X-20 Scihaku
X-22 Kuniic)1i
X--24 Yamatonishiki
Silkworm Genetics lIlustrated 151
Early period of the Sh5wa era (1927-1950)
152
T Yokoyama
x :jG ]!) CIO:i X-:Hi Tailwi >' Chaan
Presenf time
Varieties for spring rearing t w() arc parents, lower F,)

,
':11>-
,

,p,

, ..
r\!!
1

iii
-.....,
i\_
. .,

,

\
\ ,
..
r

X-37 ]122;<
SilkWOl"Dl Genetics Illustrated
i5S
X-39 Hoko X Shingyoku
154
T. Yokoyama
Varieties for summer and autumn rearing
X-tlO Sh i1ka x Giti1"ei
X-41 J122 x ell5
Silkworm Genetics Ilhlstrated
Varieties especially improved in lousiness-free
.. ,:1
,
X-43 J501 x C502
156
T. Yokoyama
X-,14 Scnsh u)< Bunko
Silkworm Gelletics Illustraled
XI. Uelation between the Sill{worm-races and
the Pathogenicity of Muscardines
157
There are many diseases in the silkworms. However the recent advance
in the study 011 the muscardines, especially such topics as their species speci-
ficity and the difference in pathogenicity of differcnt varieties, including dif-
ference in resistance oj' the silkworm races, arc 'rnost interesting. About 150
species of l11uscardine which arc panisitic all various species of insects are
known in Japan, and 20 of which are (;ound attackiL1,g silkw0rm. They
are 110t only different in the pathogenicity on different species of insects but
also different in the time lag between the infection and the appearance of
symptoms. In the of muscardine which has a long time lag,
treatment whe:1 the are observed is little effective. lL is, therefore
necessary Lo clhinfcct 2.S early as poss:ble. This principle in rnuscardine dIs"
infection contributed very much to stabilize the cocoon crops in the district
where sllch a C]Llcasc grec;h-lTIuscardine with long latent pEriod LS prevailing.
XI-l Mllscarclines <lttacking .silkwo1"rrLs
Whitt' ll111';,'"rdine
(/it'(/I/l'('}'irl i'llssiall'l (BAl..) VUJI.I..)
GreGll I1ltlscal'ciine
(Spicaria /)raSi12a (MAUBL$.) AOi{l)
Yellow
(lsi/ria /arillosIJ (PICKS.) FR.)
Satllma fungus
,l1irsutella satumaensis AOlU)
158 T. Yokoyama
XI-1 Muscardines of silkworm and other various insects (1936-1950)
XI-2 Relation between the pathogenicity of l11uscardines alld the kind of
insects
j nocllllllll
-cOncel1trat16ri oC"
i noculating-spure-
sllspem,ion
nurilber of spores
in 0.01 c. c. uf inuclI-
lating:spore-
suspension
,-_ .. -----_,_.--
Ilumber of insect
inoculated
diseased
insects
si lkworm

pyral iLl
mulh
Bealll'cria bassi{[J1({ Ilsaria jllmosorosea I Isaria /arillosa I "check
:-1(1: 1 :100}0 1:1 I 1 :100 :11 1 :1 :101 1 I-
I! I'
: I- 1
1
- -I -I 27t1 i I- -I
I '
1 0 i 10 1 (J 10 ! 10 H) 10 lO lU
HI 10 10 10 ! 1I 0
() (} [i
2 0 0 o
0 0 0 0 to 10 III 10 7 5 1
.,
,J
2 o I
:1 o i,
o I
IO JO 10 10
I
o
Remark>;: (1) lnscds teste',ll: silkworm (-1111 slage larva), kyoso (pupa), pyralitl-
mull! (5lh stage larva)
(2) and humidity during the experiment period:
27C.,
(3) Inoculation: O.ll1 c. c. of RIJore suspension was inoculalecl Oil indi-
viclual insect
The pathogenic activity of white- red- and yellow-muscardine (Beauvel'ia
bassiana, [saria fumoso-l'osea and [saria farinosa respectively) is the highest
Silkworm Genetic.;; Illustrated
159
in the host on which they are the most frequently found in nature, namely in
the domesticated silkworm (Bombyx mori L.), Kyoso-:fiy-pupa (Stu,1'mia smicariae
C.) and pyralicl moth (Mal'garonia pyloalis W.) respectively (XI-I, AOKt et
1951 a, XI-2, AOKI et al. 1951 b).
The p:tthogenicity of white-mllscardine is different according to the races
of silkworm (XI-3, AOKI et al. 1951 b.)
XI-3 Relation between the pathogenicity of white-muscardine and silkworm-
races
Table a.
silkworm race Mus. 1 NU5/C108 N112xC 110
conccnt"ratiolJ of 1 .
i11oculat!ngspore 1 :111 :100
1 i 1 :200 I 1 I
--........... I ----.. --.. - --
I
number of spores
i 11 0.0] C.c. of spore f
sllspel1Hion
number of 1
10
silkworm tested
5 daysli --
G 10
I
diseased
silkworms
after the
inoculation 10

I
11.
I
10
total 110 j 1()
..-" ... -.- _ .. -
6
<1
1U
2
4
4
10
286 164 \
10 10 I
+
10 ! 10 i
I
5
286 164
10 I
10
Remarks: (1) Silkworms tested: 5th stage larva immediately aiter ecdysis
(2) Temperature and humidity during the experiment period:
2SC., 95-100%
Table b.
10
o
silkworm-race . N 115 . C 108 I N 115: C 108
concentration .of 1:1 ..1 :1. 1..-=-
sporesuspenslOn . . __ '. ) i. 1 ..... ___ __. I _. .._ .-- -- 1 .. ___
number ol--'spores in U.Ol c.c.
1
I . I i I ! I 'I
I I + 259! -- ! + I =-.1 I I =-.
:::':::: I ::: I I:: '::
"---... - ---- ---_.- --- ....... -." ..... ..
Remarks: Temperature and humidity during the experiment period: 26
Prv
213
P
C.,
75,...,100%
IGli T. Yokoyama
The pathogenicity of white-, red- and yellow-muscarc1ine to the various
races of . siH::worm, Kyoso-fly-pupa and pyralid moth respectively is directly
proportional to the germination rate of conidia. the growth rate of germ-tubes
and the speed of cylindrical-spore formation of the causal fungi in the host's
blood in vitro (XT-4, AOI\:[ ct ai. 1951 b).
XI--4 Relation between the germination of conidia in the insect blood and
the species of 111uscarclines or the kind of insects
llU;cct bloocl
I silkworm
I
..
I
dines
<C llSJN 112 C 1101
kyosll-lly
Mus. 1 N 115
.. l';crminat ion
I
length of
while 21 l<i
I
13 2
germ luhcs red
II 2
.,
2 32
"
(1)
Ylllnw K I)
I
G 13 L
I
1
white \18 \lG \12 16
p;l'l'l1linati n,l(
rL'" .'?is 11
],I
12 87
rate (:l)
yellow (j\)
!iii 73
furmatiull of
while
HI-
-H-
11-
cylindrical reel
tIf
c::)
rd!u\\' I I -I- -Iff
XI .. !) of of
illld a((:lt:king variuus ;11;;(,,:1;;
A Il c
(A: I, .':; NlI:i 'Cllti,
C: NIl:! "ClIll:
hr:,. incuhation) (c)
(Al (0)
llir:;llli!lla AnKt
1.11\ (1\)
(B) "
T ...ytlifmlrhl/rmlitla lC)
(,\) (11)
it"
(El
A B
IIHt!iCitnlilH!
UII wurllW {\
NuctuitlilC B
Yellu\V IIlU:,CiHdilll: 011 (t\) (A) (B) I _
l.h'l/lifO/U}jUIil !lj'l'{/,thili,'j Oil
J\l:Ll1lidilt.' (C)
Ct'ramIIYI'illal! (1)
Pnalid Illoth (E)
[sllr;tl Aot';'I
1111 (A)
lIlad. /Jijlriml nij)prmicll (B)
(01l"/!1I1"11 di".,/rw/rJr METCH.) K. AOlU
Pyralid-
moth
3
32
17
24
77
-I-

XU. Wild Silkworm
Here are presented following wild or semi-wild silkworms found in Japan.
Bombycicleac
XII--l Bombyx mandd1'i11(t MOORE (Kuwa-ko in JaI)anese).
Food plants: Morus. MosLly three generationS a year, hlbertlatioll {;\$
eggs. No practical use of cocoons at presel1t in Japan. Iudoor-breeding of
larvae is difficult. '
Saturniidae
XII-2 Ant/zetaea yamamai GUERIN (Japanese Tussah Silkworm. Yamamai
in ,Japanese).
Food plants: QU(:'rCltS acutis-sima, QUernts_sel"rata, GC!J!taneq c,r;e'l,Zcr;ta, . ..
On,Q gen,Qrattan. a YQ.at, ht\)C!tn.atlcm, ae, Q.gg'O.. CG<:G0.1J.S. l'lt8.c.lkJ.!J.ty
as materials [or expensive silk cloths. Moslly field breeding On QztfM:US
acutissima, in Nagano prefecture: '
XII-3 Anlizeraea pemyi Guj1iluN (Qhinese Tussah Silk:worrn. Saj{INan in
Japanese).
Food plants: .il!!.qrczts fil, acr4fc4
Castanea crenata, Sqlix ll.imitzalis
b
generations a year,) nlbefnai.iQn as
Cocoons are used as materials of sllk clc!)tl1s. Mostly no practical
breeding in Japan now.
XII-t! Phi!osamia cynthia pryeri BUTLER (Sinzyu-sat1 in Japanese).
Foods plants: Ailanthus glandulosa, Picrasma qzlassioides, Pheltodendl'on
mnurense. Two generations a year, hibernation as ptlpae. Cocoons are used
for spun silk.
XII-5 Philosamia cynthia ricini BOISDUVAL (Eri-Silkworm. Hima-san in
Japanese).
Food plants: Ricinus communis, Ailanthus gtanciulosa, Ilex integm.
Polyvoltine. Cocoons are used for spun silk.
XII-6 Dictyop!oca japonica MOORE (Kuri-ml1si in Japanese).
Fooel plants: Castanea c1'enata, Ginkgo biloba, Ci1'lnantollum camp/lOra.
One generation a year, hibernation as eggs. Cocoons are materials for
spun silk.
162 T. Yokoyama
Larvae of wild silkworms
XII-l Bombyx mandarina MOORE
XH-3 Antherrrea Pern.1'i GUERIN
XII-5 Philosamia cynthia ricini
. BOISDUVAL
XII-2 Antizeraea yamamai GUERIN
XII-I! Plzilosamia cynthia PI)'I?I"i BUTLER
XII-6 Dictyop/oca japonica MOORE
168
COCoons of wild sllkworI\1s
.x II 1 PlnlosamlCl XII-5 Phllosamlct cynthza XIt-6 Dztyopl(!cCl Japol1ica,
/n I'erl BUTLrR rZClI1t B01SDUVAL MOORE
Silk and cocoons of Vi lid
IG4
.w
XII-l BOn!hyx
mandarina
Moorm
XII-4 Phi/osamia
cynthia pryeri
BUTLER
T. Yokoyama
Moths of wild silkworms
XII-2 Antlzeraea
),amamai

XII -5 Plli/osamia
cYlltflia ricini
BorSIJUVAL
XII 3 Antlieraea
pe1"12yi
GUERIN
XII 6 Dictyop/oca
japollica
MOORE
Silkworm Genetics
XIII. Genetics of Mulberry (incllldhig food plants
of the Silkworm otlier tlian Mulberry)
165
OSAWA (1916) found that triploid mulberries prevailqllite widely in Japan
among practically useful varieties.
Afterward he obtained tetraploid (XIII-5, 6) plajl.ts [rom the hybrid between
the triploid (XIII-2, 4) and the diploid Clm-1},
SEKI (1951) discovered that MOl'Zts titiaejolia MAKINO is hexaploid
XlII-I, 2, 5 Chromosome type at mi!.taplUlI:lC or
cell division in ovary tissue of nlll)berry
XIII-l diploic!
XIII: .. 2 triploid tetraploid
XIII3, 4, 6 Microscopical preparation of root Lip cell of mUlberry
I
XIII 3 heJ{aploid
XUI-4 triploid
XIU-6 tetraplOId
Thel ear.:: about 1000 mulberry in JapaJ1, about 200 of them being
practically cult.ivated. The Sericultural Experiment Station M. A. F.
in breeding 3 varieties of high productivity (1949): Kokuso No. 20, K, No.
166
T, Yokoyama
and K, No, 27, and bas been distributing the scions thereof for grafting to the
farmers,
As for the distribution of mulberry species in Japan; Morus bombycis is
found in the northern districts, 1'v1. multicaulis in the southern districts and
M, alba in the intermediate region.
In the exhibition, there are found polyploid mulberry trees, various mu-
tants, mulberries under breeding, popular mulbeny varieties, other food
plants of the silbvorm including Cudrallia jrll'allensis, C. triloba, Broltssonetia
Kajinol?i, Ulmus sp" Rosa sp., Lartuca Smriola, L. r/!mticulata, L. dmroglossa, L.
laciniata, 7'amxaclt1n pIa tJ'C({l'jmm , 11denoplwrrf I'Pr/idlfata, CryP/o/ aenia canadensis.
Another interesting demonstration to be shown is a technique by which
artificial crossing of different mulherry trees can be carried out easily. When
XIII 7 Usual method of artificial
crossing in mulberry tree
1. Branches with
[Japer llag priur to
r) are t l.) Ill..!
Xl1r Sa Posl-harvest-jlollination
TIle procedure is as fo!loll's:
1. The cuttings (If l-year-olt! an' stured at cole!
temperature of U"c.
Tlwy are. 1 hen, Lntl1sferrl'd to a Warm trawl' ill
fur growlh,
3. Tlwy arc able to blo;;solJ1 amI k'rril'H.
4, Thus the artindal cross pollination 011 the cutling, re.
suIting in production, is possihk t llrollgllUul the
year.
Silkworm Genetics Illusttated
XIII8b Cllttings arc ready [or blooming at 25C.
left; Wit]1 female IJoWI,'!TI, rigllt: wJth male flower.s
X III 8e Shoot blooms and QellrS 1)l!!tries at
any season by the procedure
lG7
168 T. Yokoyama
a branch or shoot of mulberry is made cutting in a pot at 25C. after a cold
of about 30 days, it blooms and bears berries at any season of the
year (XIII-8) (SUGIYAMA 1951).
In the exhibition there were pictures of practical crossing techniques,
pollen-gun, isolating envelope and propagating nursery (XIII-7), as well as
the pictures of male and female flowers of mulberry (XIII-g).
1)
2)
3)
1)
2)
3)
4)
XIII-9b Floral diagram of mulberry
o o

..
The exhibition also included live plant with pot as follows:
211 Pot No. 1
Pot No. 2
Pot No. 3
Pot No. 4
3n Pot No. 5
Pot No. 6
Pot No. 7
6n Pot No. S
311 Pot No.
g
Pot No. 10
4n Pot No. 11
Pot No. 12
Pot No. 13
Fot No. 14
Pot No. 15
Pot No. 16
5n Pot No. 17
Mixoploicl
Pot No. 18
Polyploid mulberry
Natural diPloid and polyPloid
]ushima (Monts bombycis K.)
Ichinose (Mortls alba L.)
Kairyajumonji (Moms alba L.)
Rosa (Morus latifolia P.)
Akagi (Monts bombycis K.)
Shimanouchi (llIo1'lts alba L.)
Fukushimaaha (Moms allia L.)
Keguwa (Mo}'zts tiliae/olia M.)
Induced Po(vploid
No. 3604 (48) (411? >,211 0)
c (Lla ',1 :< 211 0 )
Tok Ll 42 (311 ,;' y, 2n 0 )
23CN4, originated from2n seedling treated with colchicine.
23Ck3, originated from 2n seedling lrealec1 with colchicine.
27CE, originated from 2n seedling treated with colchicine.
b (6n q ;( 2n 0 )
S (twin from 211 seedling)
a (4n<;Jx611o)
23CRy4, originated from 2n seedling treated with colchi-
cinco
IB9
170 T. Yokoyama
Pot No.8 Kcguwa (ill P()i No.!) No. 3(iOl (-1S) :3;1 Put Nu. 10 C 3n
Pot No. 11 Toku 43 411 Pot No. 12 23 CN
1
lIn Pot Nu. 13 23 CKa '111
171
Pot No. 17 It 511.
172 T. Yokoyama
Improved new varieties
Pot No. 19 Kokuso No. 20 (Naganumax Garyii), high-yield, suitable for KantO
district.
Pot No. 20 Kokuso No. 21 (NaganumaxTsukasaso), high-yield, suitable for
the southern district of Japan.
Pot No. 21 Kokuso No. 27 (Naganumax Kairy6nezumigaeshi), high-yield, sui-
table for the middle district of Japan.
Pot No. 22 No. 18, cold resistant.
Pot No. 23 No. 14, resistant to blight disease.
Pot No. 24 Tani No. 10, resistant to blight disease.
POi No. 25 Tani No. 2775, resistant to blight disease.
Pot No. 2G Tani No. 2830 (2), resistant to blight disease.
Pot No. 27 No. 19, an early budding variety.
Pot No. 28 Hata Fl (Okinawaguwax Shiwasuguwa), suitable for subtropical
region.
PoL No. Kairyiiichinosc suitable for Kumamoto
Prefe.cture anel t1w regioll of similar climate.
Put No. 19 Kukuso No. :.W Pot No. 20 Kokuso No. 21
Pot No. 23 No. 14
174 T. Yokoyama
Pot 1\'0. Tani
No. 2830 (2)
Pot No. 28 Hata F1
Pot No.:2.1 1':0. 1\1
Pot No. 29 Kairyaichinose
Silkworm Genetics 11lustraictl
Popular varieties
Pot No. 30 Kairyonezumigaeshi, a widely distributed variety in Japan.
Pot No. 31 Ichinose. a widely distrilm1.ecl variety in Japan.
Pot No. 32 ShUkakuichi.
Pot No. 33 Tomieso.
Pot No. 311 Kairyoroso, suitable [or western Japan.
Pot No. 35 Kairyoakameroso, suitable for western Japan.
Poi No. 36 Kenmochi, suiLable for snowy region.
Pot No. 37 ]ushima, suitable for snowy region.
Pot No. 38 Nekoyatakasl.lke, suitable for snowy region.
Pot No. 39 Ichihei, an carly budding variety.
Pot No. 31 Ichinose
p,)t No. 32 Shukakuichi
175
176
T. Yokoyama
Pot No. 3:-) Po! No. 3-1 Kairyoroso
Pot No.;J(i Kenmochi
Pot. No. :{S Kairyoakameroso Pot No.:-37 Jnshima Pol No. 38 Nekoyntalwstlke
Pot. No. 30 Kairyanel.tuuigaeshi Pot No. 39 . Ichihei
Mutants and malformation
Pot No. 10 Kibajiimonji, yellow leaf.
Pot No. 41 Shimpaku, chlorophyll-free in shoot tip.
Pot No. 42 Garyii, branch winding.
Pot No. 43 Shiclareguwa, drooping mulberry.
Pot No. 44 Marubaichinose, an entire-leafed mutant from a lobed variety.
Pot No. (15 Chijimiguwa, shrinked leaf.
Pot No. 46 Mozaikuguwa, resembling mosaic-diseased plant.
Pot No. 47 Itoguwa, deeply-lobed leaf.
177
178
Pot No. 40 Kiln.
ju lllunji
]>01 No. ,.W Shiclarl'guwa
1'. Yokoyama
I'ut Nu. 11 Shillpaku
PoL No, 112 Garyil
Pol No, 4-1 Maru]JakhinoHu
Chijimiguwa
Other food plants of silkworm
Trees
Pot No. 48 Cudrania javanensis TREG.
(.Moraceae)
Pot No. 49 Cudrania 11'iloba TIBe.
(Moraceae)
Pot No. 50 BroZ{ssonetia Kajinoki 5mB.
(Moraceae)
Pot No. 51 Ulmus 8p. (Ulmaceae)
Pot No. 52 Rosa sp. (Rosaceae)
POL No. 48 Clldrania jaitJrmons
i
'6 r
180
T. Yokoyama
Pot No. 49 Cudrania triloba I-I.
Pot No. 51 Ulms sp.
Pot No. 50 Bl'ouSSol1etia
Kajinoki S.
: . \ ;1 ..
".
Pot No. 52 Rosa <lp.
Herbs
Pot No. 53 Laduca Scariola L. var. saliva
BrSC}IOFF (Compositae) ,
Pot No. 54 Lactzeca clenticulata
(Compositae)
Pot No. 55 Laciztca clracogtossa
MAKINO', (Compositae)
Pot No. 56 Laclzeca laciniata
MAKINO (Compositae)
Pot No. 57 TaJ'axacu!n plalycarjJwn
DAHLST (Compositae)
Pot No. 58 flclenoplwra verticil/ata
FISCH. (Campanulaceae)
Pot No. 59 Cryptotaenia canadensis D. C. var.
jaj)onica MAKINO (Umbelliferae)
181
182
T. Yokoyama
-
p".'\" to
/" I, J iJ'
l.f,AfjIo, iffJ
c:.;.\of,;H
PuL No. R!l Lat'luca M.
t\' N \' i
T.I ,'," .1,' II 1/1
pk'i/ ,:I'{1,Uli 1
1
,...,\:,'
.: tor

\
:to ..
Pot No. 57 Tara.tacztin jJ/atycarpum D.
\
\
/ \.
,
I'u! Nu. JUIJJl(}j'/WYlI ,'eylidllata F.
Pot No. CrYi'totacmia cClnadensis D. C.
Bibliography
AOKI, K, e( aI., 1951a J. seric. Sci. Japan 20(5)
---,1956b Ibid. 20(G)
ARUGA, II., 1939 Bul!. serie. Exp. Sla. Japan 9(G)
---, 1942 J. seric. Sci. Japan 13(5)
---, 1944 Ibid. 15(1-4)
----, 1951 Ibid. 20(1)
ARUGA, II. and S. WADA, 1954 Agric. and Hortie. (Japan) 29(10)
BOUNIIIOL, J. J., 193G C. R. Acac1. Sci. 203(5)
183
---, 1953 Transactions of the IX international COngress of Entomology
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FUKUDA, S., 1940 Proe. Imp. Ac. Tokyo 16(8)
----, 1951 Ibid. 27(10)
---, 1955 Ziklwn Ii:eitaigaku Sago Toronkai KocnyClshi (Fukuoka)
FUKUDA, T., J. KIRIMURA, M. MATSUDA and T. SUZUKI, 1955 J. Bioeh. 42(3)
FUKUDA, T., 195G NatLlre 177
---, 1!l57 Ibid. 180
GOLDSCIIMID'l', R. and K. KATSUIG, 1927 Bio!. Zbl., 47
---, 1928 Ibid. '18
---, 1931 Ibid. 51
HARlZUKA, M., 1940 J. sorie. Sci. Japan 11(4)
---, 1942 Ibid., 13(1)
---, 194.7 Bull. seric. Exp. Sla. Japan 12(5)
---, 1948 J. serie. Sci. Japan 17(1-2)
---, 1956a Ibid. 25(2)
---,195Gb Jap. J. Genei. 31(10-11)
llA,\EGAWA, K, 1943 J. seric. Sci. Japan 14(1-2)
---, 1950 GizyulsuShiryo, Ministry Agric. and Forest. Japan (29)
----, 1951a J. serie. Sci. Japan 20(1)
---, 1951b Froc. Jap. Acad. 27(10)
---, 1952 J. Facul. Agric. ToHori Ulliv. 1
HAsIMoro, B., 1929 Jap. J. Genet. 11(3)
---, 1933 BulL serk. Exp. Sta. Japan 8(7)
---, 1934 Ibid. 8(9)
---, 1941 lbid. 10(5)
M., 1943 Ibid. 11(3)
---, 1!l49 J. selic. Sci. Japan 18(5)
IlIROBE, T., 1939 lap. J. Genet. 15(2)
IT!KAWA, N, 1944a Jap. J. Genet. 20(1)
---, 1944.b Ibid. 20(3)
INAGAMI, K, 1955 J. agric. Chemist. Japan 29(12)
JUCCI, C., 1932 Boll. di Zool. 3(1)
---, 1949 Proe. 8lh. into Congr. Genet.
KATSUKI, K. and T. AKIYAMA, 1927 Sangyoshimpo 35(406)
KAWAGUCHI, E., 1928 Z. Zellforsch. 7
---, 1!l33 Cytologia 4(4)
---, 1934 J. seric. Sci. Japan 5(2)
---, 1935 Sapporo Noringakttkaiho 26(126)
184 T. Yokoyama
---, 1935 Kagalm 5(8)
---, 1936 Fac. Agric. Hokkaido Imp. Univ. 38(2)
---, 1937 J. serie. Sci. Japan 8(2)
---, 1938a Cytologia 9(1)
---, 1938b Ibid. 9(2)
KAWASE, S., 1956 Jap. J. Genet. 31(10-11)
KIKKAWA, I-I., 1937 ZooL Mag. Tokyo 49(10)
---, 1941a Ibid. 53(10)
---, 1941b Genelics 26(6)
---, 1943 Bull. seric. Exp. Sta. Japan 11(3)
---, W44 Ibid. 11(6)
---, 1950 Gakujutsugcppo 3(2)
---, 1951 J. serie. Sci. Japan 20(1)
KlKKAWA, II., Z. OGITA and S. FUJITo, 1954 Kagaku 2'1(10)
KIM, ].. 193!:J J. serie Sci. Japan 10(2)
KOBAYASHI, M., 1957 Bull. serie. Exp. Sta. Japan 15(3)
KOGURE, M. and N. KOBAYASHI U)28 Bull. Naganoken serie. Exp. Sta. Japan (2)
KOGURE, M. 19:13 J. Dept. Agric. KYLlShu Imp. Univ. 4(1)
KOYAMA, N. and S. TANAKA, U)5G J. Facul1.. Text. and Sericult. Shi118hu Univ. (6)
KOYANAGI, T., 1938 Bull. Seric. Exp. Sta. Japan !:J(4)
MACHlDA, J., 1929 Z. Zellforsch. 9(3)
MANuNTA, c., 1933 Boll. Suc. Ital. BioI. spero 8
---, 1935 Atti Soc. Nat. Mat. Mudena (6G)
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MATSUMURA, S. 1934 BulL Naganoken seric. Exp. S[a. Japan (28)
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MIYA, K., U)55 J. Serk. Sci. Japan 25(3)
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Y., HJ52 Jap. J. Genet. 27(5-G)
NAKAO, Y, TAZIMA, Y. and SliGIMlIRA, '1'., 1954 Ibid. 2D(4)
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OIW, M., H)2f) Bull. 1\gric. Chem. Suc. Japan 5
Hl30 Ibid. G
1932' Ibid. 8
---, nm Ibid. \)
OMURJ\., S., Hl38 .r. Fac. 1\gric. HokkaicHl Imp. Uni v. 4() (4)
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---, Hl,l:l Ibiel. (i(4-G)
----, 1912 14(1)
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TAI{ASAIU, T., HMO Bull. Serie. Exp. Sta.
---, 1943 Jap. J. Genet. 19(1)
TAKASAKI, T. and Y. TAZIMA, 1944 Jap. J. Genet. 20(3)
TAKA,SA,I<I, T., 1949 Ibid. 24(3-4)
TANAKA, Y., 1916 J. ColI. Agric. Tohoku Imp. Univ. 7(3)
---, 1927 GakujulsttKyokaiH6koku (3)
TANAKA, Y. and E. KAWAGUCHI, 1932 Jap. J. Genet. 7(4)
TAZIMA, Y., 1941 J. seric. Sci. Japan 12(3)
---, 1942 Jap. J. Genet. 18(1)
---, 1944 Bull. seric. Exp. 5ta. Japan 12(2)
---, 1951 J. seric. Sci. Japan 20(1)
---" 1953 Ibid. 22(3)
TAZIMA, Y. and M. MIK!, 1955 JaP. J. Genet. 30(4)
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TSUJITA, M. and B. SAKAGUCIlI, 1952 Ann. Rep. National Inst. Genet, Jap!ln 2
---, ---,1953 Ibid. 3
--- ---, 1955 Ibid. 5
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---, 1906 Ibid, 7 (2)
WATANABE, K, 1918 Bull. seric. Exp. Sta. Japan 3(8)
---, 1919 Ibid. 4(2)
WWGLESWORTrr, V. B., 1934 Q. J. M. S. 77(2)
WILlJIAMS, C. M., 1946 BioI. Bull. 93
Supplemen.tary Bibliography
AIWGA, II., 1939 BllII. seric. Exp. Sta. Japan 9(6)
---, 1940 Ibiel. 9(9)
194.3 1biel. 11(4)
CIIlKUSJII, H" 1938 J, seric. Sci. Japan 9(2)
KAWAGUCllI, E., 1936 J. sedc. Sci. Japan 7(2)
OSAWA,1., 1916 Bull. seric. Exp. Sta. Japan 1(4)
SAKATA, T., '194.3 J. serle. Sci. Japan 14(3/4)
SEKl, Il, 1951 J. seric. Sci. Japan 20(1)
SUZlJIO, K, 1929 Jap. J. Genet. 4(3)
T AJ(ASAlO, T., 1947 Silkw. Inform. Servo (2)
TANAKA, Y., 1943 Sangaku
T AZIMA, Y., 1938 lap. J. Genet. 14(3)
---, 1943 Bull. seric. Exp. Sta. Japan 11(5)
---, 1943 ]. seric. Sci. Japan 14(1/2)
--- "uel OTA, N., 1952 Jap. J. Genet. 27(5-6)
Errata
page
for
--.1 -----______ .. ________ _______
1, Preface 24
12 19
on
p'
14 Plate II
15
27
29
29
29
29

36
36
36
36
37
38
38
39
39
39
39
39
39
39
39
39
40
8
II-110
19
20
21
22
27
19
20
21
22
13
12
28
19
23
between 24 and 25, to
be added
26
30
34
between 34 and 35, to
II-32 no-glue
(Ng XII-D. 0)
(!-sp)
+'
1+'
,
+'
+'
+'
! +P'
+P'
+P'
.+P'
ze
f
Ys, straw
(body cQlor)
U
B
'
(serosa color)
be added i
36 I (body color)
38 NNs, new no-sta.rs
(::jkin marking)
21 (1938)
of
pS (The same correctibn y/ill be
made in all p' hereafter.),
left fig. of the lowest rqw is
unfertilized eggs bearing no
number of exhtbition
II-32 no-glue (Ng XII-D.O) (no
picture)
(l-sp)
+0
+0
+0
+0
+0
pI

pS
P'
Ze
f
Yr
(skin marking}
unr
(color of chorion)
XVI-chromosome
lu, lustrous (eye color)
cts, cheek and tail spots-
(skin marking)
XVII-chromosome
Em, black moth (body
color of moth)
bt s, brown head and tail
spots (skin marking)
OW, waxy translucent
(skin character)
(to be omitted)
"
elp, ellipsoid (egg shape)
(skin marking)
or, or translucelft (skin
character)
(1936)
2
page
40
42
47
47
50
52
53
53
55
55
55
55
56
56
56
57
.58
59
eo
61
62
66
68
72
72
75
75
75
75
77
78
79
S7
91
92
92
93
93
98
98
100
----- -_ --- T'
-- ... -_--_._-
... --!.-
for
27
Plate III
5
6
8
Plate V-l, upper fig.
Plate V-3, title
"
Table V-6
10
I (1928)
I III-9
I Z+Qd
I
i Z+Od
j
, were
I
X-ray dcse
, does
! 84.4 1
: pMpSY
i dil.psy
conc.pSY
pMY
Suzuki '39
21 . jP'
27 ' Tazima '52
16 pY
3 cryptal
I Plate V-ll (a), title Alianthus
, 1 V-ll
(1927)
III-6
I 2+
od
, 2+O(l
was
X-ray close
dosis
84.4 r
pMpSY
I dil.pSY
I cnnc.p8Y
I
pMY
, Suzuki '29
pSa,-2
read
, Tazima & Ota '52
;py
I cryptol
I
'. Ailllnthus
i V-ll(c)
13 character character,
Plate V-I4, upper part W-PSa(W'-I-PtY'PS"t
Y
, W-PS'a(W.+l'+Y,psa+Y)
22 Further, mo, Further, 1110
1 I1I0/mo/mo;- mo/mo/mo/ +
Plate VI-9, title Hereditary changes Changes
Plate VI-9 embryo lethal embryonic lethal
8 , monstar monster
10 A The
Plate VII-ga, explana- Black and red + Blac1{ and recl marked with +
tion
"
fl (recl)
15 if the treatment
Plate VII-ll, end of. before death.
explanation
7
! Plate VII-I5b
i
6
12
Table VIII-I, right
column, 1st row
Plate
Plate VII-2, right
fig. 1
3

70 days.
[<;11/+
'll/EII
characterizes
(LOS f. C.,
. lelt

of ESC: brain
(KOBAYASIIl 1958)
nondia pause
with
6 +
in the treatment
before death (C).
70 days (C),
. Ell/+
: or Ell/Ell
I
(I.OS f. c .
left
VIII-2
ESC: brain-
(KOBAYASIII 1957)
l1on-c1iapause
in
3
vni'?s:_ -- WA,
106 i 11 I rl is I rt larvae is
106 I 21 ! trimQltings I trimolting
106 I 21 tetramoltings: tetramolting
107 i2nd line from the last trimoulting I trimolting
108 28 acid, such I acid, Such
109 ! Plate IX-I, last line by a mutant by a mutant gene
112 ! Plate IX-4, explana- (+Xrb) spots : (+Xrb); spots
upper row ,
II tion of left fig. in the !
112 Table rX-5 (1) I C I 3-hydraxykynu- I C I 3-hydroxykynurenine
! Tenine I
113
113
115
118
U8
118
119
119
119
120
120
123
125
I 2 ' s-mottled , S-mottled
Plate IX-Ga, title s-mottled 'S-mottled
1 normal one I white one
9 ' individual individ uals
9
15
I 10
4=112
i 17
,lone
I (sm,)
lk
I Plate IX-9b, title of ' (smt)
lower row
5 p"
Plate IX-lO, explana- I which is connected
, tton with
I Plate IX-12, upper delation
part
ones
(sm,)
K
I +K
r sm'
(sm,)
+p
: which is to be connected with
i
deletion
126 Table IX-13, 3. 2nd quantity of uric acid i quantity of uric acid mg/tissue
column uric acid mg/tissue g ! g
127 9 ' yellow-lethal ' lemon lethal
127 11 , yellow lethal , lemon lethal
130 6 ' four strains four kinds of strain
131 Plate IX-20, explalla- i oivoltine bivoltine
tiOI1 of 2nd fig. from
right in the 3rd row
132 upper table, V ! European univoltine European univoltine yellow
yellow cocoon cocoon (15 races)
133 11 FJ hybrid between FI hybrid between
133
134
134
134
last line
Plate IX-24, left fig.
Plate IX-24, explana-
tion of left fig.
+ae between
ae gene larvae
I }.
xanthophylls zone
Plate IX-:?4" explalla- + 0
tion of right fig.
ae larvae
y
xanthophyll zone
____ - __ ___ O, _ __ ...... .
line I for
_- 1
148
148
149
158
158
160
168
175
175
176
178
179
181
: mentally
3
Plate X-12,
I
I single coocon
left fig. micropteral ...
of 1st row
Table X-13, 4th
of explanation
)ine I In rt, the recessive
I trlmolting, silkworm
: white- red- and
I yellow-muscardine
last l'me :. jumoso-rostdt
1
Plate XI-5, explana-! Dendmlimum
tion of 2nd fig. from :
left in the upper row ;
23 23Ck3

ill)
Pot No. 36
Pot No. 41
Pot No. 46
Pot No. 53, explana-
tion
Kairyoakamcroso
Kenmochi
Kenmochi
Shinpaku
Mozaic-mullJerry
sCCll'iola
read
fundamental cocoon color
single cocoon, are
micropterous
In rt, recessive trimolting, thE
silkworm .
white-, red- and yellow-
muscardine
fumosorosea
Drmdrolimus
i 23CK3
: Kairyoakameros6
I Kemmochi
I Remmochi
Shimpaku
I Mosaic-mulberry
Sea rio/a

Bicho Seda Takayana

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