i nheritance of chlorophyll deficiencies beta vulga...i nheritance of chlorophyll deficiencies in...
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I nheritance of Chlorophyll Deficiencies In
Beta Vulga l V F SVlTSKY
Introduction
The illhcriLtllce oj dcficicIlCic ill sugar Ct~ls
inheriLIl(c h~l attracted lhe and
Hoi 0(
DiscuSiolb 10
the localilltion
attelltioll of scielltists because mall oj sucrose content anei root em
tilt rooL
and tilt 01 sucrose ill (~I j()) ano also the
rest Oil the Ilew bus leaves IJUt in lilc where radioactivemiddotmarked it was indicated tllll
the mobile ugar ill the vascular Systelll
lhe 01 t he LOpS or the roOlS In
lor nutrition lile radiotcllve ca rmiddot
lorlllatioll 01 sucrOse in the leal blades ollsider SlICI he L mobile sugar buill
ill leans but llot ill the fOOts oi SIIar beeb
UTel 11h
unknowll in beets (eJ I)
dd(
dilkrclll stnlClUlCs 01 01 tlte
interllal [isles s()m~ colorless Tin I UllCfioll 01 III
and OIJlT crops showed [hal lile size color aud ltUlulJer
sucil g~lIn also hI illlCll-
III COllllCClioll with Ihis a 01 certain citarallcrs ill
IJieu
III genes wb iell [lclerosis ill the
DiHClclll (
I) ( lrLaili III the
obsencd III Ilallh alter CPOSlllC to tWllIi r~Hliati()n
of the inheritallce oj
san lor lhe gcnct ie and work lor Ihe breeding 01 irradiated beels
lalerial and Ielhods
The Llces smdicd were isolated ill ari~Ld popushylatiolls of sugar alld [udder beCh inbred lines Iud 111 this til lIatur1i mlllII]IIS but not mutati()lls stimulated radiation ric
F secd the inmiddot
lkptrfmefll oj
I01RM OF TllE C S S IS 1
hcritance 01 the slIlall dillcrcnces ill the 01 lhe parshypropashy
to obla ill F 1 a uti F seed
genes causing ~lnd other gtllcs known in 1Jra L will not be discllssed ill this paper
Results
The inher-itau(e 01 albinism The of allJillislll in beets was laned 188) and IBtl Proskowcl sl miin] scyeilleell chimeric with altern ale grccn and whilc sC(lors He camc lO conclusi()ll albinism was inhcrited
on-Mendelian inll(rirancc of chlorophyll deficieTHY status aifJomashy
cuiatus III IlI l Stehlik loune a sCld bect which olle green alld one whilC branch The growl from lhe seni balls
from thc white branch were lIe concluded Da diese von dell in dell ZehlclJ yorhandellell Farbstollc del
was ctwcd ) could not chlon)
and perished at all stage derived from seed I)alls of tIl( grccll Imlllch gne liOO llormal green plants without allY trace of dhinislll Stehlik VI a the ilrsl to stalC thal albitlism III heels was inherited through the as StOWIl from studies ill Milllbilis
alboll1acuaa Corrells in IJOl) Stehliks onclusiom that albinism in beets is inherited through tht
cylllplaSllJ was confirmed Iullerali in J~)28 and J~lI2 (2 2()_ Yhmcrati indicated that chimcri( Ileus OCTulTcd ill lions at the rate of one per flOnOn or HlOOO green 10 inheritatlcc or albinism YIlItlcrati collCcLCd aboUl 300 chimeric C1I in commental varieties
The prog(ny lOllsbted of while when the lowcrs the white branches or grcet) branches
Seed collected from the I)ranches as m live
from white or green im ill beets ill inherited III
limbil lIlt ti) PrilJlllia sinewis (II and in Zell lIIa)s (g)
In 1940 Owen alld abo dcscribed one case of inherilance- of albinism from studi 01 the progcn from white and green branches 01 albino
I have studied mall in sligar awl fodder beet OllC
Illhenlic casc was (slablishcd the nuclear In all other 01 ill the
to note lhat among be for recessivc I have I1c(r obsclvltd chimeric
IX lo middotRY I
III bCd populations lllOrc oltcil a a ugtllcquellcc or hcrilallJc ( In the rather than 1 a resliit 01 ll1utation 01 lluclear gents This typ of albinislll becausc of its jrcqucl1t appearance attracted the attelltioll o( the first G
In a review of the inileriLalln ul ddomphyll ddicicncies ill which a reurlle of all cOllduned ill all crops to 1~)U was (iled H dc Ham (Ii) IllClItiollCd for one Ii pc or dcmiddot jl( I ypl
lIendeJian inheritance of albinism-the beets of Yellow Eckcndorf
green char of sugar and lodder bCCLS as l11 as the prccll(c or aiJsencc ill seedling
III F hyiJrib derived from pollinatioll pollen collected frolll whi te branches
2)0 and 79 albino to the mailC
derived from collected froIll the grecll branche of the families lor albino albino III this way the green
lor the reteivc
Eckendor[ plant was
r denoted as IV This gene produced The (himcrit nature or the
of llormal grcen ugar iJeet of 1 variegated
The color of theoe albino
to one
for the
gene which the hOlllozygOte Eckcllltiuri plalH
was caused the lallation of the dOll1inallt gene 111 into a I((essin gClle w
Recessivc gellcs causing complctc albinism or arc COlhldclshyrarer in beets (han the other rtC(IVC g-clles for chloroph)lI defieinides
b for the g(lllS middotirbCllIs ehlorin1 elL
Tahle l-kmidian Inheritancc of Albini ill Beta VulgllS [
-lumber of Seedling
Whitlt x
In
Observed Cal(ulltt(cd Ratio ])
Observed 77~
Calculated (Rat io l [ ) 7)o ij
appearance oj albino hethe dOlllillalH gelle mea or
III F somnimes rat io of Olle to I) normal green w[th true of gene crops
IW S S BI
b Ollie
I( I iOIl
ill the to deal
i II other
Aurea found OIll
IIJ
coloI 11111
type-the dominant plaJlt which
Color
dnminalll gellc III wilh sCldlings (rc all)ino similar gClle laThe
derihcd In ILwcr in middotnlndiwlill and ill and iJ
(olTem (J) ill IIira IInIIS This gene knowlI ill olher crops too Table 2 holls the dau of (I I he i 1 I I ~J)l wi I Fi)
(Tossed to 01 a ginlI lype 01 was ltol llllilorln green and
I idied green plallls
oiTspring iJUl the F promiddot for green alld alhillo
Tahk 2-1nlumiddotritanrc of U1TJ ryp ill Heelgt
(eneratioH Cal
utnher of
lint- Grnl
Si(d Ji n~
1llomiddot lhino
01raquocnd (akuatcd
(rat io I I
I -)~
(1117
Ohscn __ c llculatcd
11)
I q~)l ~II
2-120
II
lJ 1I ~ 1
Ob-cncd III 1111
(rnio I 1 ) 107 ()o)
()bs(f cd
11cuatcd (rat in 1 I )
~l_i Imiddot J I b
J09
I 12~)
II IH
)~) I
V~) if)
Ohscrnd CalclIlatcd
latin 12 I l
F 191 I H I I Hl
171 Otgtl
Ohscn cd CalcUi1Cd
(ratio I I)
I))) I K I
I(ll
101
Ill
point or prohahillt for r gClILl
POl1lt of prnbabilJt Inl D~l
LUlesccns type-gene III The rcccssilT gClIe (II Wh
between sligar and fodder beets Red Eckclldori 1 6219 h) I) 3) In
and sometimes the pair 01 leaves 11so showed llorma I leaves bccallle mol( and morc a
planb dietL In was redUlcd and did Jot excecd SIX penellt of the
normal gTCCll of the -ltlme The F planls llormal
of these rHCS arc Slalltlan) hecanse
accumulatcs later grccll hut lw following leaves become gncner alld cOlHjiliollS 111(1 ()Il
not of Ilormal grcen hects gClles or the arc Ihe of of
wilh Ihis in Ilw rallc or
uluiHr of Plans dth SC((Hld and Thinl Pair of LCltlfs Colored as follows
Gf(~(n
2iO
Calculated 04 IIi
point of prohabillt)
point or problhilit~ X (iJj-l
Tahle -I-Segregation in F~ After Crosses of Two Raus of Vircs((~ns Type (rit Vil 112 and tlJ 1 V) of chlorophyH dc(icicntits in Reta Fulgas L
umbtr of F~ Plants Suryh iug
Green Yellow x
Observed 2()O
Ctlculated (ral in 9 t) )01
point of proiJahil it U)middotj
poinl of prohahiiiq X)
A II these Vhell two diflerelll vircs(cns raccs arc norma I green
(41) The double heClllse ()f
III gene Cfe also oj
harwl the grcgllinll sllOwcd nille green to four I)
J0LIZgtL OF Till ~~ S S B~ T
The percellt of
root ill
weight of root of the normal
when it the ratio of three
there was 110 cOlltillual the sallle oiJscrnd bv I
In isolated at which showed
11 hanesl was middotUO green in the same F
is sejltLlted [rom other middotiresccils gelles to one III such
wit h Iccesi( genes in hces~
pClh
the Salt Lake Cit) station allother t till stronger deby in chlornshy
tlescril)ed g-enes and Ttlble illustrates the illhcritalllC of the Observatiolls (rc (ontiuClcd for three years lI1( 1 HTC normal tlCCIl III the F mOllomiddot
was ohserved
umber of ~umhcr Seedlings
G(fura~ Poptlla~
Year lion rions Gn(u Ydlow
cd I (F h
Ohscrnd F
Calclated (ratio i I 10
Obsertd I~Jtd F
Calcnlat(d ratio I ) 0729
ObscrHd IlJI F Hil
Calculated (rlio I)
()hscly(x I Hl
Calculated (ratio i I) 71)7)
OINned 1~~~)~1 19 Calculated (ntlio 11 ) j)1ckrrols 22 22 OR181
x = lHI
X lUi-
mlltn chlorophyll dcfkiencies--the deficiencies arc based 011 a
gene W1S obsened ill beets this gene
(
shows h1e [iowered and produced s(cd it the IncHs
The dala for and
F a lid i II hal klTOSS of plan is
hI Owen to normal
x
belg (I) found allother gcne in i)(cts the gelle which green to lIormal grecn SCt lors
was indicated bcgg (I) Yellow to senors The inlwriullce of lhi gCllc was not hOWll
The deficicilcies which dcvc]op ill the
VOL IX io I J-IARY I
of a 111 u ta ble gCllc ha c beell descrihed ill erai (l g 19 SimilJr genes llOt found in lJia
Chlorina type-genes (h and defens of ill beels
redu(cd
-rahle (L-inherlrance of Cldorina rype Deficiency in JJeia ~JUg(H L
UtUh(T of StcdJings (~cncramiddot ~unlb(r of
tion lamiH~ Green Yellow
Gene cl1 isolated in CS 32 Ohsentd
CainTatcd (ral io I)
h tl~ Ii
t()) i)OI]
Obscl cd
(llntlated ratio I
11 1 F tnl
lO-)7t)
10
ILW)~ll
OhsltrCi[
ClicuiarClt1 (rat io I)
lFI) Fc Ii 2) ~f 1111
100 Olilil
Obf-(led
Calculated (ratio I )
1~ )~) -) Sit 112
IIIgt O ~19L~)
G(llC ell isolated in Great fesitrn hetmiddott
Ol)srycd
Ohgterved
Calculated (ratio I I) 114 117 [0]
Gene (1 isola(d in
Obsenclti
Kleinwanziebn b~t
Calculated I) 5525 IR 07511
()hS(T cd
Calculated (ratio ~~l) JlL
point of probahiliq
point of prolJahili~ X~ 6fH
X
gCllC differ ill t he oj rcdllClioll of the Ihe gene ( which was isolated ill tile
reduced the 0 roob about 0 pcrccllt Another earlier (11) dccrcased the rool much more
which wcre isolaUd in the inbred lille SLC F hiJri(b normal
hI ilrids eTC l1onnj grecn III
Ilormal lk H E
Sugar which crossed S7R
crossed Ilormal green Fe hybrids howed mOl
Vhcll Iwcts of the chlorina 11]( beels SLC color of cotduiolls and obtained from these
rhus til( beCIS il h 10
oj the gene oi 1(11 liS the
ltlificnlIl sugar hlCI yarietics showed 19gt1 and I ~Fj fi)
K oj lhl Creat VVcsterll light
i II Longmont Colorado alld to clllorillli tYI)l hcets SLC
which I rcccIIcd hom Longmont F plants green and
[n 19) T studicd ilso the rc([i( gene 01 Ihe isoblcd III Ihc iniJred lil1( SIC I02~ lun which from KkinwlI1zlebcns beets Ii)
Interaction In i t t~
shown Ih11
l the iahle
were combined wilh til( llOmogou gellts cnhini( antHhcr type o[
for CXllllpIe Ihe gene of chlorill[ type thell two via illc under illlnhousc
between certain genes for of the Circsccm tvpc 0 the combination of (110 different
nOIl-
the
Hld
genes of Ihe ircs[cIlS
conditions which carried the above-mcn t iOlled and I he F (lrogcln
nine grcen a 11(1 ((n
Tahe 7-Intcllcion Uetwccn inSHIll and Chlorina GHtS for Chlo1ophyl1 fhgtfirknc i Il F~~ G(n(r~Hion
I-Jvbrids
Sredlill
Ydlow x
Ob-crcd
CdkllIalnl (raio ~17) jll(j
I pOlllt of prohlbili ~UH
poinl uf probahi[ X = (dH
VOl IX C()i R I ~)7
Tahil also show) that it Ias piblc 10 distinguish by [h( color of and mltllrc leaves lhe of the genotype from the chrh
genotype ill thest
When the gem VI yirCSCCllS comhined with the gene A II
cgrcgatioll complicated wcre observed F could he because tbesc plallls carried the genc Au pro dneeo after F which [he ratioll l green
(4 Vi1lilIl(lU ViViAu(lu) I I Vtlillllllll) I alhillo (I II
The threc viable classls gIT F families which (Ollshy
d assifica Lion
The effect 01 emironment on chlorophyll formalion III different Culthatioll 01 beets the main
lIchas urea differenl (nvironmcilts showed that
vcry much the lormation of
DiflprclHCs fIl thc lspecially obmiddotiots dlCIl these or under field rondi t ions D i ffercnt
a nel fall
and action of the gencs 111 (h
of ClIirolllllcnt on
could I) illmtratcd
Lethal (hlolophyll mutants-A HAil Ull died ullder field (onditiolb tllrce
Semimiddotlethal mutant~
a Imost norma 11 y i 11
under gnulhnmc cnnditiom kaves and pro ltlured a fair amount of seed The mutant
field and had
to accumulatc
bccomes nmdishyThe raccs which
wcre denotcd a irltS(C11 under conditions classified as xtnllu in held The races with which green wcre dll1otcd as vircsccns llIore viable than otiter races with ccdling
Table- 8 Interaction BeOHen Vircs(cns and Aurea Genes for Chlorophyil Uttirienlty in I Generation
Seedlings
Hyrids frc(~n Yellow Gold Albino
Ohscn ed
CalClllated (ratio
o hlcrvc I
Caku]arcd (ratio 36 1lt1)
J9gt1
IIl
qj
II
ltIR
17~)
IRI
(l
~17
1)[
ll
18
J2]
121
iii
LlGI
ltUR9i
7 point of prnblbi1it)- X2 7tZ
point of prohability
obtain seed even ill thebefore and from although their
of vironment whicb
The differences 111 the the in(]uencc of enshy
ill beets were also descrihed for olher crops Akerman (2) in oats Smith and Holm (I tIl
The of dnmi Hallce of the genes also varied undu different conditionslt of chlorina type were as viable ill the chell looked much JigJ1tcr in The Chell did not differ in the green The gelle dl was reccssive ulldel Another kind of behavior was observed in the field gnw for the were light and could be
the to
eft were normal grcen while the Tn the latc summer the
In sncll a way the of dominance changed peTiod as he seen from the data or segregation in (Table 9)
the
Tahle 9-Segregation in F Chlorin Hybrids (Ficld Obsenatiol1s in 13)
-0 of plants ith folor of Icn(t as follows
Reading in Grecn Ligh (middotgrcell Total
July 11
October 17
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
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ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
I01RM OF TllE C S S IS 1
hcritance 01 the slIlall dillcrcnces ill the 01 lhe parshypropashy
to obla ill F 1 a uti F seed
genes causing ~lnd other gtllcs known in 1Jra L will not be discllssed ill this paper
Results
The inher-itau(e 01 albinism The of allJillislll in beets was laned 188) and IBtl Proskowcl sl miin] scyeilleell chimeric with altern ale grccn and whilc sC(lors He camc lO conclusi()ll albinism was inhcrited
on-Mendelian inll(rirancc of chlorophyll deficieTHY status aifJomashy
cuiatus III IlI l Stehlik loune a sCld bect which olle green alld one whilC branch The growl from lhe seni balls
from thc white branch were lIe concluded Da diese von dell in dell ZehlclJ yorhandellell Farbstollc del
was ctwcd ) could not chlon)
and perished at all stage derived from seed I)alls of tIl( grccll Imlllch gne liOO llormal green plants without allY trace of dhinislll Stehlik VI a the ilrsl to stalC thal albitlism III heels was inherited through the as StOWIl from studies ill Milllbilis
alboll1acuaa Corrells in IJOl) Stehliks onclusiom that albinism in beets is inherited through tht
cylllplaSllJ was confirmed Iullerali in J~)28 and J~lI2 (2 2()_ Yhmcrati indicated that chimcri( Ileus OCTulTcd ill lions at the rate of one per flOnOn or HlOOO green 10 inheritatlcc or albinism YIlItlcrati collCcLCd aboUl 300 chimeric C1I in commental varieties
The prog(ny lOllsbted of while when the lowcrs the white branches or grcet) branches
Seed collected from the I)ranches as m live
from white or green im ill beets ill inherited III
limbil lIlt ti) PrilJlllia sinewis (II and in Zell lIIa)s (g)
In 1940 Owen alld abo dcscribed one case of inherilance- of albinism from studi 01 the progcn from white and green branches 01 albino
I have studied mall in sligar awl fodder beet OllC
Illhenlic casc was (slablishcd the nuclear In all other 01 ill the
to note lhat among be for recessivc I have I1c(r obsclvltd chimeric
IX lo middotRY I
III bCd populations lllOrc oltcil a a ugtllcquellcc or hcrilallJc ( In the rather than 1 a resliit 01 ll1utation 01 lluclear gents This typ of albinislll becausc of its jrcqucl1t appearance attracted the attelltioll o( the first G
In a review of the inileriLalln ul ddomphyll ddicicncies ill which a reurlle of all cOllduned ill all crops to 1~)U was (iled H dc Ham (Ii) IllClItiollCd for one Ii pc or dcmiddot jl( I ypl
lIendeJian inheritance of albinism-the beets of Yellow Eckcndorf
green char of sugar and lodder bCCLS as l11 as the prccll(c or aiJsencc ill seedling
III F hyiJrib derived from pollinatioll pollen collected frolll whi te branches
2)0 and 79 albino to the mailC
derived from collected froIll the grecll branche of the families lor albino albino III this way the green
lor the reteivc
Eckendor[ plant was
r denoted as IV This gene produced The (himcrit nature or the
of llormal grcen ugar iJeet of 1 variegated
The color of theoe albino
to one
for the
gene which the hOlllozygOte Eckcllltiuri plalH
was caused the lallation of the dOll1inallt gene 111 into a I((essin gClle w
Recessivc gellcs causing complctc albinism or arc COlhldclshyrarer in beets (han the other rtC(IVC g-clles for chloroph)lI defieinides
b for the g(lllS middotirbCllIs ehlorin1 elL
Tahle l-kmidian Inheritancc of Albini ill Beta VulgllS [
-lumber of Seedling
Whitlt x
In
Observed Cal(ulltt(cd Ratio ])
Observed 77~
Calculated (Rat io l [ ) 7)o ij
appearance oj albino hethe dOlllillalH gelle mea or
III F somnimes rat io of Olle to I) normal green w[th true of gene crops
IW S S BI
b Ollie
I( I iOIl
ill the to deal
i II other
Aurea found OIll
IIJ
coloI 11111
type-the dominant plaJlt which
Color
dnminalll gellc III wilh sCldlings (rc all)ino similar gClle laThe
derihcd In ILwcr in middotnlndiwlill and ill and iJ
(olTem (J) ill IIira IInIIS This gene knowlI ill olher crops too Table 2 holls the dau of (I I he i 1 I I ~J)l wi I Fi)
(Tossed to 01 a ginlI lype 01 was ltol llllilorln green and
I idied green plallls
oiTspring iJUl the F promiddot for green alld alhillo
Tahk 2-1nlumiddotritanrc of U1TJ ryp ill Heelgt
(eneratioH Cal
utnher of
lint- Grnl
Si(d Ji n~
1llomiddot lhino
01raquocnd (akuatcd
(rat io I I
I -)~
(1117
Ohscn __ c llculatcd
11)
I q~)l ~II
2-120
II
lJ 1I ~ 1
Ob-cncd III 1111
(rnio I 1 ) 107 ()o)
()bs(f cd
11cuatcd (rat in 1 I )
~l_i Imiddot J I b
J09
I 12~)
II IH
)~) I
V~) if)
Ohscrnd CalclIlatcd
latin 12 I l
F 191 I H I I Hl
171 Otgtl
Ohscn cd CalcUi1Cd
(ratio I I)
I))) I K I
I(ll
101
Ill
point or prohahillt for r gClILl
POl1lt of prnbabilJt Inl D~l
LUlesccns type-gene III The rcccssilT gClIe (II Wh
between sligar and fodder beets Red Eckclldori 1 6219 h) I) 3) In
and sometimes the pair 01 leaves 11so showed llorma I leaves bccallle mol( and morc a
planb dietL In was redUlcd and did Jot excecd SIX penellt of the
normal gTCCll of the -ltlme The F planls llormal
of these rHCS arc Slalltlan) hecanse
accumulatcs later grccll hut lw following leaves become gncner alld cOlHjiliollS 111(1 ()Il
not of Ilormal grcen hects gClles or the arc Ihe of of
wilh Ihis in Ilw rallc or
uluiHr of Plans dth SC((Hld and Thinl Pair of LCltlfs Colored as follows
Gf(~(n
2iO
Calculated 04 IIi
point of prohabillt)
point or problhilit~ X (iJj-l
Tahle -I-Segregation in F~ After Crosses of Two Raus of Vircs((~ns Type (rit Vil 112 and tlJ 1 V) of chlorophyH dc(icicntits in Reta Fulgas L
umbtr of F~ Plants Suryh iug
Green Yellow x
Observed 2()O
Ctlculated (ral in 9 t) )01
point of proiJahil it U)middotj
poinl of prohahiiiq X)
A II these Vhell two diflerelll vircs(cns raccs arc norma I green
(41) The double heClllse ()f
III gene Cfe also oj
harwl the grcgllinll sllOwcd nille green to four I)
J0LIZgtL OF Till ~~ S S B~ T
The percellt of
root ill
weight of root of the normal
when it the ratio of three
there was 110 cOlltillual the sallle oiJscrnd bv I
In isolated at which showed
11 hanesl was middotUO green in the same F
is sejltLlted [rom other middotiresccils gelles to one III such
wit h Iccesi( genes in hces~
pClh
the Salt Lake Cit) station allother t till stronger deby in chlornshy
tlescril)ed g-enes and Ttlble illustrates the illhcritalllC of the Observatiolls (rc (ontiuClcd for three years lI1( 1 HTC normal tlCCIl III the F mOllomiddot
was ohserved
umber of ~umhcr Seedlings
G(fura~ Poptlla~
Year lion rions Gn(u Ydlow
cd I (F h
Ohscrnd F
Calclated (ratio i I 10
Obsertd I~Jtd F
Calcnlat(d ratio I ) 0729
ObscrHd IlJI F Hil
Calculated (rlio I)
()hscly(x I Hl
Calculated (ratio i I) 71)7)
OINned 1~~~)~1 19 Calculated (ntlio 11 ) j)1ckrrols 22 22 OR181
x = lHI
X lUi-
mlltn chlorophyll dcfkiencies--the deficiencies arc based 011 a
gene W1S obsened ill beets this gene
(
shows h1e [iowered and produced s(cd it the IncHs
The dala for and
F a lid i II hal klTOSS of plan is
hI Owen to normal
x
belg (I) found allother gcne in i)(cts the gelle which green to lIormal grecn SCt lors
was indicated bcgg (I) Yellow to senors The inlwriullce of lhi gCllc was not hOWll
The deficicilcies which dcvc]op ill the
VOL IX io I J-IARY I
of a 111 u ta ble gCllc ha c beell descrihed ill erai (l g 19 SimilJr genes llOt found in lJia
Chlorina type-genes (h and defens of ill beels
redu(cd
-rahle (L-inherlrance of Cldorina rype Deficiency in JJeia ~JUg(H L
UtUh(T of StcdJings (~cncramiddot ~unlb(r of
tion lamiH~ Green Yellow
Gene cl1 isolated in CS 32 Ohsentd
CainTatcd (ral io I)
h tl~ Ii
t()) i)OI]
Obscl cd
(llntlated ratio I
11 1 F tnl
lO-)7t)
10
ILW)~ll
OhsltrCi[
ClicuiarClt1 (rat io I)
lFI) Fc Ii 2) ~f 1111
100 Olilil
Obf-(led
Calculated (ratio I )
1~ )~) -) Sit 112
IIIgt O ~19L~)
G(llC ell isolated in Great fesitrn hetmiddott
Ol)srycd
Ohgterved
Calculated (ratio I I) 114 117 [0]
Gene (1 isola(d in
Obsenclti
Kleinwanziebn b~t
Calculated I) 5525 IR 07511
()hS(T cd
Calculated (ratio ~~l) JlL
point of probahiliq
point of prolJahili~ X~ 6fH
X
gCllC differ ill t he oj rcdllClioll of the Ihe gene ( which was isolated ill tile
reduced the 0 roob about 0 pcrccllt Another earlier (11) dccrcased the rool much more
which wcre isolaUd in the inbred lille SLC F hiJri(b normal
hI ilrids eTC l1onnj grecn III
Ilormal lk H E
Sugar which crossed S7R
crossed Ilormal green Fe hybrids howed mOl
Vhcll Iwcts of the chlorina 11]( beels SLC color of cotduiolls and obtained from these
rhus til( beCIS il h 10
oj the gene oi 1(11 liS the
ltlificnlIl sugar hlCI yarietics showed 19gt1 and I ~Fj fi)
K oj lhl Creat VVcsterll light
i II Longmont Colorado alld to clllorillli tYI)l hcets SLC
which I rcccIIcd hom Longmont F plants green and
[n 19) T studicd ilso the rc([i( gene 01 Ihe isoblcd III Ihc iniJred lil1( SIC I02~ lun which from KkinwlI1zlebcns beets Ii)
Interaction In i t t~
shown Ih11
l the iahle
were combined wilh til( llOmogou gellts cnhini( antHhcr type o[
for CXllllpIe Ihe gene of chlorill[ type thell two via illc under illlnhousc
between certain genes for of the Circsccm tvpc 0 the combination of (110 different
nOIl-
the
Hld
genes of Ihe ircs[cIlS
conditions which carried the above-mcn t iOlled and I he F (lrogcln
nine grcen a 11(1 ((n
Tahe 7-Intcllcion Uetwccn inSHIll and Chlorina GHtS for Chlo1ophyl1 fhgtfirknc i Il F~~ G(n(r~Hion
I-Jvbrids
Sredlill
Ydlow x
Ob-crcd
CdkllIalnl (raio ~17) jll(j
I pOlllt of prohlbili ~UH
poinl uf probahi[ X = (dH
VOl IX C()i R I ~)7
Tahil also show) that it Ias piblc 10 distinguish by [h( color of and mltllrc leaves lhe of the genotype from the chrh
genotype ill thest
When the gem VI yirCSCCllS comhined with the gene A II
cgrcgatioll complicated wcre observed F could he because tbesc plallls carried the genc Au pro dneeo after F which [he ratioll l green
(4 Vi1lilIl(lU ViViAu(lu) I I Vtlillllllll) I alhillo (I II
The threc viable classls gIT F families which (Ollshy
d assifica Lion
The effect 01 emironment on chlorophyll formalion III different Culthatioll 01 beets the main
lIchas urea differenl (nvironmcilts showed that
vcry much the lormation of
DiflprclHCs fIl thc lspecially obmiddotiots dlCIl these or under field rondi t ions D i ffercnt
a nel fall
and action of the gencs 111 (h
of ClIirolllllcnt on
could I) illmtratcd
Lethal (hlolophyll mutants-A HAil Ull died ullder field (onditiolb tllrce
Semimiddotlethal mutant~
a Imost norma 11 y i 11
under gnulhnmc cnnditiom kaves and pro ltlured a fair amount of seed The mutant
field and had
to accumulatc
bccomes nmdishyThe raccs which
wcre denotcd a irltS(C11 under conditions classified as xtnllu in held The races with which green wcre dll1otcd as vircsccns llIore viable than otiter races with ccdling
Table- 8 Interaction BeOHen Vircs(cns and Aurea Genes for Chlorophyil Uttirienlty in I Generation
Seedlings
Hyrids frc(~n Yellow Gold Albino
Ohscn ed
CalClllated (ratio
o hlcrvc I
Caku]arcd (ratio 36 1lt1)
J9gt1
IIl
qj
II
ltIR
17~)
IRI
(l
~17
1)[
ll
18
J2]
121
iii
LlGI
ltUR9i
7 point of prnblbi1it)- X2 7tZ
point of prohability
obtain seed even ill thebefore and from although their
of vironment whicb
The differences 111 the the in(]uencc of enshy
ill beets were also descrihed for olher crops Akerman (2) in oats Smith and Holm (I tIl
The of dnmi Hallce of the genes also varied undu different conditionslt of chlorina type were as viable ill the chell looked much JigJ1tcr in The Chell did not differ in the green The gelle dl was reccssive ulldel Another kind of behavior was observed in the field gnw for the were light and could be
the to
eft were normal grcen while the Tn the latc summer the
In sncll a way the of dominance changed peTiod as he seen from the data or segregation in (Table 9)
the
Tahle 9-Segregation in F Chlorin Hybrids (Ficld Obsenatiol1s in 13)
-0 of plants ith folor of Icn(t as follows
Reading in Grecn Ligh (middotgrcell Total
July 11
October 17
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
IX lo middotRY I
III bCd populations lllOrc oltcil a a ugtllcquellcc or hcrilallJc ( In the rather than 1 a resliit 01 ll1utation 01 lluclear gents This typ of albinislll becausc of its jrcqucl1t appearance attracted the attelltioll o( the first G
In a review of the inileriLalln ul ddomphyll ddicicncies ill which a reurlle of all cOllduned ill all crops to 1~)U was (iled H dc Ham (Ii) IllClItiollCd for one Ii pc or dcmiddot jl( I ypl
lIendeJian inheritance of albinism-the beets of Yellow Eckcndorf
green char of sugar and lodder bCCLS as l11 as the prccll(c or aiJsencc ill seedling
III F hyiJrib derived from pollinatioll pollen collected frolll whi te branches
2)0 and 79 albino to the mailC
derived from collected froIll the grecll branche of the families lor albino albino III this way the green
lor the reteivc
Eckendor[ plant was
r denoted as IV This gene produced The (himcrit nature or the
of llormal grcen ugar iJeet of 1 variegated
The color of theoe albino
to one
for the
gene which the hOlllozygOte Eckcllltiuri plalH
was caused the lallation of the dOll1inallt gene 111 into a I((essin gClle w
Recessivc gellcs causing complctc albinism or arc COlhldclshyrarer in beets (han the other rtC(IVC g-clles for chloroph)lI defieinides
b for the g(lllS middotirbCllIs ehlorin1 elL
Tahle l-kmidian Inheritancc of Albini ill Beta VulgllS [
-lumber of Seedling
Whitlt x
In
Observed Cal(ulltt(cd Ratio ])
Observed 77~
Calculated (Rat io l [ ) 7)o ij
appearance oj albino hethe dOlllillalH gelle mea or
III F somnimes rat io of Olle to I) normal green w[th true of gene crops
IW S S BI
b Ollie
I( I iOIl
ill the to deal
i II other
Aurea found OIll
IIJ
coloI 11111
type-the dominant plaJlt which
Color
dnminalll gellc III wilh sCldlings (rc all)ino similar gClle laThe
derihcd In ILwcr in middotnlndiwlill and ill and iJ
(olTem (J) ill IIira IInIIS This gene knowlI ill olher crops too Table 2 holls the dau of (I I he i 1 I I ~J)l wi I Fi)
(Tossed to 01 a ginlI lype 01 was ltol llllilorln green and
I idied green plallls
oiTspring iJUl the F promiddot for green alld alhillo
Tahk 2-1nlumiddotritanrc of U1TJ ryp ill Heelgt
(eneratioH Cal
utnher of
lint- Grnl
Si(d Ji n~
1llomiddot lhino
01raquocnd (akuatcd
(rat io I I
I -)~
(1117
Ohscn __ c llculatcd
11)
I q~)l ~II
2-120
II
lJ 1I ~ 1
Ob-cncd III 1111
(rnio I 1 ) 107 ()o)
()bs(f cd
11cuatcd (rat in 1 I )
~l_i Imiddot J I b
J09
I 12~)
II IH
)~) I
V~) if)
Ohscrnd CalclIlatcd
latin 12 I l
F 191 I H I I Hl
171 Otgtl
Ohscn cd CalcUi1Cd
(ratio I I)
I))) I K I
I(ll
101
Ill
point or prohahillt for r gClILl
POl1lt of prnbabilJt Inl D~l
LUlesccns type-gene III The rcccssilT gClIe (II Wh
between sligar and fodder beets Red Eckclldori 1 6219 h) I) 3) In
and sometimes the pair 01 leaves 11so showed llorma I leaves bccallle mol( and morc a
planb dietL In was redUlcd and did Jot excecd SIX penellt of the
normal gTCCll of the -ltlme The F planls llormal
of these rHCS arc Slalltlan) hecanse
accumulatcs later grccll hut lw following leaves become gncner alld cOlHjiliollS 111(1 ()Il
not of Ilormal grcen hects gClles or the arc Ihe of of
wilh Ihis in Ilw rallc or
uluiHr of Plans dth SC((Hld and Thinl Pair of LCltlfs Colored as follows
Gf(~(n
2iO
Calculated 04 IIi
point of prohabillt)
point or problhilit~ X (iJj-l
Tahle -I-Segregation in F~ After Crosses of Two Raus of Vircs((~ns Type (rit Vil 112 and tlJ 1 V) of chlorophyH dc(icicntits in Reta Fulgas L
umbtr of F~ Plants Suryh iug
Green Yellow x
Observed 2()O
Ctlculated (ral in 9 t) )01
point of proiJahil it U)middotj
poinl of prohahiiiq X)
A II these Vhell two diflerelll vircs(cns raccs arc norma I green
(41) The double heClllse ()f
III gene Cfe also oj
harwl the grcgllinll sllOwcd nille green to four I)
J0LIZgtL OF Till ~~ S S B~ T
The percellt of
root ill
weight of root of the normal
when it the ratio of three
there was 110 cOlltillual the sallle oiJscrnd bv I
In isolated at which showed
11 hanesl was middotUO green in the same F
is sejltLlted [rom other middotiresccils gelles to one III such
wit h Iccesi( genes in hces~
pClh
the Salt Lake Cit) station allother t till stronger deby in chlornshy
tlescril)ed g-enes and Ttlble illustrates the illhcritalllC of the Observatiolls (rc (ontiuClcd for three years lI1( 1 HTC normal tlCCIl III the F mOllomiddot
was ohserved
umber of ~umhcr Seedlings
G(fura~ Poptlla~
Year lion rions Gn(u Ydlow
cd I (F h
Ohscrnd F
Calclated (ratio i I 10
Obsertd I~Jtd F
Calcnlat(d ratio I ) 0729
ObscrHd IlJI F Hil
Calculated (rlio I)
()hscly(x I Hl
Calculated (ratio i I) 71)7)
OINned 1~~~)~1 19 Calculated (ntlio 11 ) j)1ckrrols 22 22 OR181
x = lHI
X lUi-
mlltn chlorophyll dcfkiencies--the deficiencies arc based 011 a
gene W1S obsened ill beets this gene
(
shows h1e [iowered and produced s(cd it the IncHs
The dala for and
F a lid i II hal klTOSS of plan is
hI Owen to normal
x
belg (I) found allother gcne in i)(cts the gelle which green to lIormal grecn SCt lors
was indicated bcgg (I) Yellow to senors The inlwriullce of lhi gCllc was not hOWll
The deficicilcies which dcvc]op ill the
VOL IX io I J-IARY I
of a 111 u ta ble gCllc ha c beell descrihed ill erai (l g 19 SimilJr genes llOt found in lJia
Chlorina type-genes (h and defens of ill beels
redu(cd
-rahle (L-inherlrance of Cldorina rype Deficiency in JJeia ~JUg(H L
UtUh(T of StcdJings (~cncramiddot ~unlb(r of
tion lamiH~ Green Yellow
Gene cl1 isolated in CS 32 Ohsentd
CainTatcd (ral io I)
h tl~ Ii
t()) i)OI]
Obscl cd
(llntlated ratio I
11 1 F tnl
lO-)7t)
10
ILW)~ll
OhsltrCi[
ClicuiarClt1 (rat io I)
lFI) Fc Ii 2) ~f 1111
100 Olilil
Obf-(led
Calculated (ratio I )
1~ )~) -) Sit 112
IIIgt O ~19L~)
G(llC ell isolated in Great fesitrn hetmiddott
Ol)srycd
Ohgterved
Calculated (ratio I I) 114 117 [0]
Gene (1 isola(d in
Obsenclti
Kleinwanziebn b~t
Calculated I) 5525 IR 07511
()hS(T cd
Calculated (ratio ~~l) JlL
point of probahiliq
point of prolJahili~ X~ 6fH
X
gCllC differ ill t he oj rcdllClioll of the Ihe gene ( which was isolated ill tile
reduced the 0 roob about 0 pcrccllt Another earlier (11) dccrcased the rool much more
which wcre isolaUd in the inbred lille SLC F hiJri(b normal
hI ilrids eTC l1onnj grecn III
Ilormal lk H E
Sugar which crossed S7R
crossed Ilormal green Fe hybrids howed mOl
Vhcll Iwcts of the chlorina 11]( beels SLC color of cotduiolls and obtained from these
rhus til( beCIS il h 10
oj the gene oi 1(11 liS the
ltlificnlIl sugar hlCI yarietics showed 19gt1 and I ~Fj fi)
K oj lhl Creat VVcsterll light
i II Longmont Colorado alld to clllorillli tYI)l hcets SLC
which I rcccIIcd hom Longmont F plants green and
[n 19) T studicd ilso the rc([i( gene 01 Ihe isoblcd III Ihc iniJred lil1( SIC I02~ lun which from KkinwlI1zlebcns beets Ii)
Interaction In i t t~
shown Ih11
l the iahle
were combined wilh til( llOmogou gellts cnhini( antHhcr type o[
for CXllllpIe Ihe gene of chlorill[ type thell two via illc under illlnhousc
between certain genes for of the Circsccm tvpc 0 the combination of (110 different
nOIl-
the
Hld
genes of Ihe ircs[cIlS
conditions which carried the above-mcn t iOlled and I he F (lrogcln
nine grcen a 11(1 ((n
Tahe 7-Intcllcion Uetwccn inSHIll and Chlorina GHtS for Chlo1ophyl1 fhgtfirknc i Il F~~ G(n(r~Hion
I-Jvbrids
Sredlill
Ydlow x
Ob-crcd
CdkllIalnl (raio ~17) jll(j
I pOlllt of prohlbili ~UH
poinl uf probahi[ X = (dH
VOl IX C()i R I ~)7
Tahil also show) that it Ias piblc 10 distinguish by [h( color of and mltllrc leaves lhe of the genotype from the chrh
genotype ill thest
When the gem VI yirCSCCllS comhined with the gene A II
cgrcgatioll complicated wcre observed F could he because tbesc plallls carried the genc Au pro dneeo after F which [he ratioll l green
(4 Vi1lilIl(lU ViViAu(lu) I I Vtlillllllll) I alhillo (I II
The threc viable classls gIT F families which (Ollshy
d assifica Lion
The effect 01 emironment on chlorophyll formalion III different Culthatioll 01 beets the main
lIchas urea differenl (nvironmcilts showed that
vcry much the lormation of
DiflprclHCs fIl thc lspecially obmiddotiots dlCIl these or under field rondi t ions D i ffercnt
a nel fall
and action of the gencs 111 (h
of ClIirolllllcnt on
could I) illmtratcd
Lethal (hlolophyll mutants-A HAil Ull died ullder field (onditiolb tllrce
Semimiddotlethal mutant~
a Imost norma 11 y i 11
under gnulhnmc cnnditiom kaves and pro ltlured a fair amount of seed The mutant
field and had
to accumulatc
bccomes nmdishyThe raccs which
wcre denotcd a irltS(C11 under conditions classified as xtnllu in held The races with which green wcre dll1otcd as vircsccns llIore viable than otiter races with ccdling
Table- 8 Interaction BeOHen Vircs(cns and Aurea Genes for Chlorophyil Uttirienlty in I Generation
Seedlings
Hyrids frc(~n Yellow Gold Albino
Ohscn ed
CalClllated (ratio
o hlcrvc I
Caku]arcd (ratio 36 1lt1)
J9gt1
IIl
qj
II
ltIR
17~)
IRI
(l
~17
1)[
ll
18
J2]
121
iii
LlGI
ltUR9i
7 point of prnblbi1it)- X2 7tZ
point of prohability
obtain seed even ill thebefore and from although their
of vironment whicb
The differences 111 the the in(]uencc of enshy
ill beets were also descrihed for olher crops Akerman (2) in oats Smith and Holm (I tIl
The of dnmi Hallce of the genes also varied undu different conditionslt of chlorina type were as viable ill the chell looked much JigJ1tcr in The Chell did not differ in the green The gelle dl was reccssive ulldel Another kind of behavior was observed in the field gnw for the were light and could be
the to
eft were normal grcen while the Tn the latc summer the
In sncll a way the of dominance changed peTiod as he seen from the data or segregation in (Table 9)
the
Tahle 9-Segregation in F Chlorin Hybrids (Ficld Obsenatiol1s in 13)
-0 of plants ith folor of Icn(t as follows
Reading in Grecn Ligh (middotgrcell Total
July 11
October 17
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
appearance oj albino hethe dOlllillalH gelle mea or
III F somnimes rat io of Olle to I) normal green w[th true of gene crops
IW S S BI
b Ollie
I( I iOIl
ill the to deal
i II other
Aurea found OIll
IIJ
coloI 11111
type-the dominant plaJlt which
Color
dnminalll gellc III wilh sCldlings (rc all)ino similar gClle laThe
derihcd In ILwcr in middotnlndiwlill and ill and iJ
(olTem (J) ill IIira IInIIS This gene knowlI ill olher crops too Table 2 holls the dau of (I I he i 1 I I ~J)l wi I Fi)
(Tossed to 01 a ginlI lype 01 was ltol llllilorln green and
I idied green plallls
oiTspring iJUl the F promiddot for green alld alhillo
Tahk 2-1nlumiddotritanrc of U1TJ ryp ill Heelgt
(eneratioH Cal
utnher of
lint- Grnl
Si(d Ji n~
1llomiddot lhino
01raquocnd (akuatcd
(rat io I I
I -)~
(1117
Ohscn __ c llculatcd
11)
I q~)l ~II
2-120
II
lJ 1I ~ 1
Ob-cncd III 1111
(rnio I 1 ) 107 ()o)
()bs(f cd
11cuatcd (rat in 1 I )
~l_i Imiddot J I b
J09
I 12~)
II IH
)~) I
V~) if)
Ohscrnd CalclIlatcd
latin 12 I l
F 191 I H I I Hl
171 Otgtl
Ohscn cd CalcUi1Cd
(ratio I I)
I))) I K I
I(ll
101
Ill
point or prohahillt for r gClILl
POl1lt of prnbabilJt Inl D~l
LUlesccns type-gene III The rcccssilT gClIe (II Wh
between sligar and fodder beets Red Eckclldori 1 6219 h) I) 3) In
and sometimes the pair 01 leaves 11so showed llorma I leaves bccallle mol( and morc a
planb dietL In was redUlcd and did Jot excecd SIX penellt of the
normal gTCCll of the -ltlme The F planls llormal
of these rHCS arc Slalltlan) hecanse
accumulatcs later grccll hut lw following leaves become gncner alld cOlHjiliollS 111(1 ()Il
not of Ilormal grcen hects gClles or the arc Ihe of of
wilh Ihis in Ilw rallc or
uluiHr of Plans dth SC((Hld and Thinl Pair of LCltlfs Colored as follows
Gf(~(n
2iO
Calculated 04 IIi
point of prohabillt)
point or problhilit~ X (iJj-l
Tahle -I-Segregation in F~ After Crosses of Two Raus of Vircs((~ns Type (rit Vil 112 and tlJ 1 V) of chlorophyH dc(icicntits in Reta Fulgas L
umbtr of F~ Plants Suryh iug
Green Yellow x
Observed 2()O
Ctlculated (ral in 9 t) )01
point of proiJahil it U)middotj
poinl of prohahiiiq X)
A II these Vhell two diflerelll vircs(cns raccs arc norma I green
(41) The double heClllse ()f
III gene Cfe also oj
harwl the grcgllinll sllOwcd nille green to four I)
J0LIZgtL OF Till ~~ S S B~ T
The percellt of
root ill
weight of root of the normal
when it the ratio of three
there was 110 cOlltillual the sallle oiJscrnd bv I
In isolated at which showed
11 hanesl was middotUO green in the same F
is sejltLlted [rom other middotiresccils gelles to one III such
wit h Iccesi( genes in hces~
pClh
the Salt Lake Cit) station allother t till stronger deby in chlornshy
tlescril)ed g-enes and Ttlble illustrates the illhcritalllC of the Observatiolls (rc (ontiuClcd for three years lI1( 1 HTC normal tlCCIl III the F mOllomiddot
was ohserved
umber of ~umhcr Seedlings
G(fura~ Poptlla~
Year lion rions Gn(u Ydlow
cd I (F h
Ohscrnd F
Calclated (ratio i I 10
Obsertd I~Jtd F
Calcnlat(d ratio I ) 0729
ObscrHd IlJI F Hil
Calculated (rlio I)
()hscly(x I Hl
Calculated (ratio i I) 71)7)
OINned 1~~~)~1 19 Calculated (ntlio 11 ) j)1ckrrols 22 22 OR181
x = lHI
X lUi-
mlltn chlorophyll dcfkiencies--the deficiencies arc based 011 a
gene W1S obsened ill beets this gene
(
shows h1e [iowered and produced s(cd it the IncHs
The dala for and
F a lid i II hal klTOSS of plan is
hI Owen to normal
x
belg (I) found allother gcne in i)(cts the gelle which green to lIormal grecn SCt lors
was indicated bcgg (I) Yellow to senors The inlwriullce of lhi gCllc was not hOWll
The deficicilcies which dcvc]op ill the
VOL IX io I J-IARY I
of a 111 u ta ble gCllc ha c beell descrihed ill erai (l g 19 SimilJr genes llOt found in lJia
Chlorina type-genes (h and defens of ill beels
redu(cd
-rahle (L-inherlrance of Cldorina rype Deficiency in JJeia ~JUg(H L
UtUh(T of StcdJings (~cncramiddot ~unlb(r of
tion lamiH~ Green Yellow
Gene cl1 isolated in CS 32 Ohsentd
CainTatcd (ral io I)
h tl~ Ii
t()) i)OI]
Obscl cd
(llntlated ratio I
11 1 F tnl
lO-)7t)
10
ILW)~ll
OhsltrCi[
ClicuiarClt1 (rat io I)
lFI) Fc Ii 2) ~f 1111
100 Olilil
Obf-(led
Calculated (ratio I )
1~ )~) -) Sit 112
IIIgt O ~19L~)
G(llC ell isolated in Great fesitrn hetmiddott
Ol)srycd
Ohgterved
Calculated (ratio I I) 114 117 [0]
Gene (1 isola(d in
Obsenclti
Kleinwanziebn b~t
Calculated I) 5525 IR 07511
()hS(T cd
Calculated (ratio ~~l) JlL
point of probahiliq
point of prolJahili~ X~ 6fH
X
gCllC differ ill t he oj rcdllClioll of the Ihe gene ( which was isolated ill tile
reduced the 0 roob about 0 pcrccllt Another earlier (11) dccrcased the rool much more
which wcre isolaUd in the inbred lille SLC F hiJri(b normal
hI ilrids eTC l1onnj grecn III
Ilormal lk H E
Sugar which crossed S7R
crossed Ilormal green Fe hybrids howed mOl
Vhcll Iwcts of the chlorina 11]( beels SLC color of cotduiolls and obtained from these
rhus til( beCIS il h 10
oj the gene oi 1(11 liS the
ltlificnlIl sugar hlCI yarietics showed 19gt1 and I ~Fj fi)
K oj lhl Creat VVcsterll light
i II Longmont Colorado alld to clllorillli tYI)l hcets SLC
which I rcccIIcd hom Longmont F plants green and
[n 19) T studicd ilso the rc([i( gene 01 Ihe isoblcd III Ihc iniJred lil1( SIC I02~ lun which from KkinwlI1zlebcns beets Ii)
Interaction In i t t~
shown Ih11
l the iahle
were combined wilh til( llOmogou gellts cnhini( antHhcr type o[
for CXllllpIe Ihe gene of chlorill[ type thell two via illc under illlnhousc
between certain genes for of the Circsccm tvpc 0 the combination of (110 different
nOIl-
the
Hld
genes of Ihe ircs[cIlS
conditions which carried the above-mcn t iOlled and I he F (lrogcln
nine grcen a 11(1 ((n
Tahe 7-Intcllcion Uetwccn inSHIll and Chlorina GHtS for Chlo1ophyl1 fhgtfirknc i Il F~~ G(n(r~Hion
I-Jvbrids
Sredlill
Ydlow x
Ob-crcd
CdkllIalnl (raio ~17) jll(j
I pOlllt of prohlbili ~UH
poinl uf probahi[ X = (dH
VOl IX C()i R I ~)7
Tahil also show) that it Ias piblc 10 distinguish by [h( color of and mltllrc leaves lhe of the genotype from the chrh
genotype ill thest
When the gem VI yirCSCCllS comhined with the gene A II
cgrcgatioll complicated wcre observed F could he because tbesc plallls carried the genc Au pro dneeo after F which [he ratioll l green
(4 Vi1lilIl(lU ViViAu(lu) I I Vtlillllllll) I alhillo (I II
The threc viable classls gIT F families which (Ollshy
d assifica Lion
The effect 01 emironment on chlorophyll formalion III different Culthatioll 01 beets the main
lIchas urea differenl (nvironmcilts showed that
vcry much the lormation of
DiflprclHCs fIl thc lspecially obmiddotiots dlCIl these or under field rondi t ions D i ffercnt
a nel fall
and action of the gencs 111 (h
of ClIirolllllcnt on
could I) illmtratcd
Lethal (hlolophyll mutants-A HAil Ull died ullder field (onditiolb tllrce
Semimiddotlethal mutant~
a Imost norma 11 y i 11
under gnulhnmc cnnditiom kaves and pro ltlured a fair amount of seed The mutant
field and had
to accumulatc
bccomes nmdishyThe raccs which
wcre denotcd a irltS(C11 under conditions classified as xtnllu in held The races with which green wcre dll1otcd as vircsccns llIore viable than otiter races with ccdling
Table- 8 Interaction BeOHen Vircs(cns and Aurea Genes for Chlorophyil Uttirienlty in I Generation
Seedlings
Hyrids frc(~n Yellow Gold Albino
Ohscn ed
CalClllated (ratio
o hlcrvc I
Caku]arcd (ratio 36 1lt1)
J9gt1
IIl
qj
II
ltIR
17~)
IRI
(l
~17
1)[
ll
18
J2]
121
iii
LlGI
ltUR9i
7 point of prnblbi1it)- X2 7tZ
point of prohability
obtain seed even ill thebefore and from although their
of vironment whicb
The differences 111 the the in(]uencc of enshy
ill beets were also descrihed for olher crops Akerman (2) in oats Smith and Holm (I tIl
The of dnmi Hallce of the genes also varied undu different conditionslt of chlorina type were as viable ill the chell looked much JigJ1tcr in The Chell did not differ in the green The gelle dl was reccssive ulldel Another kind of behavior was observed in the field gnw for the were light and could be
the to
eft were normal grcen while the Tn the latc summer the
In sncll a way the of dominance changed peTiod as he seen from the data or segregation in (Table 9)
the
Tahle 9-Segregation in F Chlorin Hybrids (Ficld Obsenatiol1s in 13)
-0 of plants ith folor of Icn(t as follows
Reading in Grecn Ligh (middotgrcell Total
July 11
October 17
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
LUlesccns type-gene III The rcccssilT gClIe (II Wh
between sligar and fodder beets Red Eckclldori 1 6219 h) I) 3) In
and sometimes the pair 01 leaves 11so showed llorma I leaves bccallle mol( and morc a
planb dietL In was redUlcd and did Jot excecd SIX penellt of the
normal gTCCll of the -ltlme The F planls llormal
of these rHCS arc Slalltlan) hecanse
accumulatcs later grccll hut lw following leaves become gncner alld cOlHjiliollS 111(1 ()Il
not of Ilormal grcen hects gClles or the arc Ihe of of
wilh Ihis in Ilw rallc or
uluiHr of Plans dth SC((Hld and Thinl Pair of LCltlfs Colored as follows
Gf(~(n
2iO
Calculated 04 IIi
point of prohabillt)
point or problhilit~ X (iJj-l
Tahle -I-Segregation in F~ After Crosses of Two Raus of Vircs((~ns Type (rit Vil 112 and tlJ 1 V) of chlorophyH dc(icicntits in Reta Fulgas L
umbtr of F~ Plants Suryh iug
Green Yellow x
Observed 2()O
Ctlculated (ral in 9 t) )01
point of proiJahil it U)middotj
poinl of prohahiiiq X)
A II these Vhell two diflerelll vircs(cns raccs arc norma I green
(41) The double heClllse ()f
III gene Cfe also oj
harwl the grcgllinll sllOwcd nille green to four I)
J0LIZgtL OF Till ~~ S S B~ T
The percellt of
root ill
weight of root of the normal
when it the ratio of three
there was 110 cOlltillual the sallle oiJscrnd bv I
In isolated at which showed
11 hanesl was middotUO green in the same F
is sejltLlted [rom other middotiresccils gelles to one III such
wit h Iccesi( genes in hces~
pClh
the Salt Lake Cit) station allother t till stronger deby in chlornshy
tlescril)ed g-enes and Ttlble illustrates the illhcritalllC of the Observatiolls (rc (ontiuClcd for three years lI1( 1 HTC normal tlCCIl III the F mOllomiddot
was ohserved
umber of ~umhcr Seedlings
G(fura~ Poptlla~
Year lion rions Gn(u Ydlow
cd I (F h
Ohscrnd F
Calclated (ratio i I 10
Obsertd I~Jtd F
Calcnlat(d ratio I ) 0729
ObscrHd IlJI F Hil
Calculated (rlio I)
()hscly(x I Hl
Calculated (ratio i I) 71)7)
OINned 1~~~)~1 19 Calculated (ntlio 11 ) j)1ckrrols 22 22 OR181
x = lHI
X lUi-
mlltn chlorophyll dcfkiencies--the deficiencies arc based 011 a
gene W1S obsened ill beets this gene
(
shows h1e [iowered and produced s(cd it the IncHs
The dala for and
F a lid i II hal klTOSS of plan is
hI Owen to normal
x
belg (I) found allother gcne in i)(cts the gelle which green to lIormal grecn SCt lors
was indicated bcgg (I) Yellow to senors The inlwriullce of lhi gCllc was not hOWll
The deficicilcies which dcvc]op ill the
VOL IX io I J-IARY I
of a 111 u ta ble gCllc ha c beell descrihed ill erai (l g 19 SimilJr genes llOt found in lJia
Chlorina type-genes (h and defens of ill beels
redu(cd
-rahle (L-inherlrance of Cldorina rype Deficiency in JJeia ~JUg(H L
UtUh(T of StcdJings (~cncramiddot ~unlb(r of
tion lamiH~ Green Yellow
Gene cl1 isolated in CS 32 Ohsentd
CainTatcd (ral io I)
h tl~ Ii
t()) i)OI]
Obscl cd
(llntlated ratio I
11 1 F tnl
lO-)7t)
10
ILW)~ll
OhsltrCi[
ClicuiarClt1 (rat io I)
lFI) Fc Ii 2) ~f 1111
100 Olilil
Obf-(led
Calculated (ratio I )
1~ )~) -) Sit 112
IIIgt O ~19L~)
G(llC ell isolated in Great fesitrn hetmiddott
Ol)srycd
Ohgterved
Calculated (ratio I I) 114 117 [0]
Gene (1 isola(d in
Obsenclti
Kleinwanziebn b~t
Calculated I) 5525 IR 07511
()hS(T cd
Calculated (ratio ~~l) JlL
point of probahiliq
point of prolJahili~ X~ 6fH
X
gCllC differ ill t he oj rcdllClioll of the Ihe gene ( which was isolated ill tile
reduced the 0 roob about 0 pcrccllt Another earlier (11) dccrcased the rool much more
which wcre isolaUd in the inbred lille SLC F hiJri(b normal
hI ilrids eTC l1onnj grecn III
Ilormal lk H E
Sugar which crossed S7R
crossed Ilormal green Fe hybrids howed mOl
Vhcll Iwcts of the chlorina 11]( beels SLC color of cotduiolls and obtained from these
rhus til( beCIS il h 10
oj the gene oi 1(11 liS the
ltlificnlIl sugar hlCI yarietics showed 19gt1 and I ~Fj fi)
K oj lhl Creat VVcsterll light
i II Longmont Colorado alld to clllorillli tYI)l hcets SLC
which I rcccIIcd hom Longmont F plants green and
[n 19) T studicd ilso the rc([i( gene 01 Ihe isoblcd III Ihc iniJred lil1( SIC I02~ lun which from KkinwlI1zlebcns beets Ii)
Interaction In i t t~
shown Ih11
l the iahle
were combined wilh til( llOmogou gellts cnhini( antHhcr type o[
for CXllllpIe Ihe gene of chlorill[ type thell two via illc under illlnhousc
between certain genes for of the Circsccm tvpc 0 the combination of (110 different
nOIl-
the
Hld
genes of Ihe ircs[cIlS
conditions which carried the above-mcn t iOlled and I he F (lrogcln
nine grcen a 11(1 ((n
Tahe 7-Intcllcion Uetwccn inSHIll and Chlorina GHtS for Chlo1ophyl1 fhgtfirknc i Il F~~ G(n(r~Hion
I-Jvbrids
Sredlill
Ydlow x
Ob-crcd
CdkllIalnl (raio ~17) jll(j
I pOlllt of prohlbili ~UH
poinl uf probahi[ X = (dH
VOl IX C()i R I ~)7
Tahil also show) that it Ias piblc 10 distinguish by [h( color of and mltllrc leaves lhe of the genotype from the chrh
genotype ill thest
When the gem VI yirCSCCllS comhined with the gene A II
cgrcgatioll complicated wcre observed F could he because tbesc plallls carried the genc Au pro dneeo after F which [he ratioll l green
(4 Vi1lilIl(lU ViViAu(lu) I I Vtlillllllll) I alhillo (I II
The threc viable classls gIT F families which (Ollshy
d assifica Lion
The effect 01 emironment on chlorophyll formalion III different Culthatioll 01 beets the main
lIchas urea differenl (nvironmcilts showed that
vcry much the lormation of
DiflprclHCs fIl thc lspecially obmiddotiots dlCIl these or under field rondi t ions D i ffercnt
a nel fall
and action of the gencs 111 (h
of ClIirolllllcnt on
could I) illmtratcd
Lethal (hlolophyll mutants-A HAil Ull died ullder field (onditiolb tllrce
Semimiddotlethal mutant~
a Imost norma 11 y i 11
under gnulhnmc cnnditiom kaves and pro ltlured a fair amount of seed The mutant
field and had
to accumulatc
bccomes nmdishyThe raccs which
wcre denotcd a irltS(C11 under conditions classified as xtnllu in held The races with which green wcre dll1otcd as vircsccns llIore viable than otiter races with ccdling
Table- 8 Interaction BeOHen Vircs(cns and Aurea Genes for Chlorophyil Uttirienlty in I Generation
Seedlings
Hyrids frc(~n Yellow Gold Albino
Ohscn ed
CalClllated (ratio
o hlcrvc I
Caku]arcd (ratio 36 1lt1)
J9gt1
IIl
qj
II
ltIR
17~)
IRI
(l
~17
1)[
ll
18
J2]
121
iii
LlGI
ltUR9i
7 point of prnblbi1it)- X2 7tZ
point of prohability
obtain seed even ill thebefore and from although their
of vironment whicb
The differences 111 the the in(]uencc of enshy
ill beets were also descrihed for olher crops Akerman (2) in oats Smith and Holm (I tIl
The of dnmi Hallce of the genes also varied undu different conditionslt of chlorina type were as viable ill the chell looked much JigJ1tcr in The Chell did not differ in the green The gelle dl was reccssive ulldel Another kind of behavior was observed in the field gnw for the were light and could be
the to
eft were normal grcen while the Tn the latc summer the
In sncll a way the of dominance changed peTiod as he seen from the data or segregation in (Table 9)
the
Tahle 9-Segregation in F Chlorin Hybrids (Ficld Obsenatiol1s in 13)
-0 of plants ith folor of Icn(t as follows
Reading in Grecn Ligh (middotgrcell Total
July 11
October 17
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
J0LIZgtL OF Till ~~ S S B~ T
The percellt of
root ill
weight of root of the normal
when it the ratio of three
there was 110 cOlltillual the sallle oiJscrnd bv I
In isolated at which showed
11 hanesl was middotUO green in the same F
is sejltLlted [rom other middotiresccils gelles to one III such
wit h Iccesi( genes in hces~
pClh
the Salt Lake Cit) station allother t till stronger deby in chlornshy
tlescril)ed g-enes and Ttlble illustrates the illhcritalllC of the Observatiolls (rc (ontiuClcd for three years lI1( 1 HTC normal tlCCIl III the F mOllomiddot
was ohserved
umber of ~umhcr Seedlings
G(fura~ Poptlla~
Year lion rions Gn(u Ydlow
cd I (F h
Ohscrnd F
Calclated (ratio i I 10
Obsertd I~Jtd F
Calcnlat(d ratio I ) 0729
ObscrHd IlJI F Hil
Calculated (rlio I)
()hscly(x I Hl
Calculated (ratio i I) 71)7)
OINned 1~~~)~1 19 Calculated (ntlio 11 ) j)1ckrrols 22 22 OR181
x = lHI
X lUi-
mlltn chlorophyll dcfkiencies--the deficiencies arc based 011 a
gene W1S obsened ill beets this gene
(
shows h1e [iowered and produced s(cd it the IncHs
The dala for and
F a lid i II hal klTOSS of plan is
hI Owen to normal
x
belg (I) found allother gcne in i)(cts the gelle which green to lIormal grecn SCt lors
was indicated bcgg (I) Yellow to senors The inlwriullce of lhi gCllc was not hOWll
The deficicilcies which dcvc]op ill the
VOL IX io I J-IARY I
of a 111 u ta ble gCllc ha c beell descrihed ill erai (l g 19 SimilJr genes llOt found in lJia
Chlorina type-genes (h and defens of ill beels
redu(cd
-rahle (L-inherlrance of Cldorina rype Deficiency in JJeia ~JUg(H L
UtUh(T of StcdJings (~cncramiddot ~unlb(r of
tion lamiH~ Green Yellow
Gene cl1 isolated in CS 32 Ohsentd
CainTatcd (ral io I)
h tl~ Ii
t()) i)OI]
Obscl cd
(llntlated ratio I
11 1 F tnl
lO-)7t)
10
ILW)~ll
OhsltrCi[
ClicuiarClt1 (rat io I)
lFI) Fc Ii 2) ~f 1111
100 Olilil
Obf-(led
Calculated (ratio I )
1~ )~) -) Sit 112
IIIgt O ~19L~)
G(llC ell isolated in Great fesitrn hetmiddott
Ol)srycd
Ohgterved
Calculated (ratio I I) 114 117 [0]
Gene (1 isola(d in
Obsenclti
Kleinwanziebn b~t
Calculated I) 5525 IR 07511
()hS(T cd
Calculated (ratio ~~l) JlL
point of probahiliq
point of prolJahili~ X~ 6fH
X
gCllC differ ill t he oj rcdllClioll of the Ihe gene ( which was isolated ill tile
reduced the 0 roob about 0 pcrccllt Another earlier (11) dccrcased the rool much more
which wcre isolaUd in the inbred lille SLC F hiJri(b normal
hI ilrids eTC l1onnj grecn III
Ilormal lk H E
Sugar which crossed S7R
crossed Ilormal green Fe hybrids howed mOl
Vhcll Iwcts of the chlorina 11]( beels SLC color of cotduiolls and obtained from these
rhus til( beCIS il h 10
oj the gene oi 1(11 liS the
ltlificnlIl sugar hlCI yarietics showed 19gt1 and I ~Fj fi)
K oj lhl Creat VVcsterll light
i II Longmont Colorado alld to clllorillli tYI)l hcets SLC
which I rcccIIcd hom Longmont F plants green and
[n 19) T studicd ilso the rc([i( gene 01 Ihe isoblcd III Ihc iniJred lil1( SIC I02~ lun which from KkinwlI1zlebcns beets Ii)
Interaction In i t t~
shown Ih11
l the iahle
were combined wilh til( llOmogou gellts cnhini( antHhcr type o[
for CXllllpIe Ihe gene of chlorill[ type thell two via illc under illlnhousc
between certain genes for of the Circsccm tvpc 0 the combination of (110 different
nOIl-
the
Hld
genes of Ihe ircs[cIlS
conditions which carried the above-mcn t iOlled and I he F (lrogcln
nine grcen a 11(1 ((n
Tahe 7-Intcllcion Uetwccn inSHIll and Chlorina GHtS for Chlo1ophyl1 fhgtfirknc i Il F~~ G(n(r~Hion
I-Jvbrids
Sredlill
Ydlow x
Ob-crcd
CdkllIalnl (raio ~17) jll(j
I pOlllt of prohlbili ~UH
poinl uf probahi[ X = (dH
VOl IX C()i R I ~)7
Tahil also show) that it Ias piblc 10 distinguish by [h( color of and mltllrc leaves lhe of the genotype from the chrh
genotype ill thest
When the gem VI yirCSCCllS comhined with the gene A II
cgrcgatioll complicated wcre observed F could he because tbesc plallls carried the genc Au pro dneeo after F which [he ratioll l green
(4 Vi1lilIl(lU ViViAu(lu) I I Vtlillllllll) I alhillo (I II
The threc viable classls gIT F families which (Ollshy
d assifica Lion
The effect 01 emironment on chlorophyll formalion III different Culthatioll 01 beets the main
lIchas urea differenl (nvironmcilts showed that
vcry much the lormation of
DiflprclHCs fIl thc lspecially obmiddotiots dlCIl these or under field rondi t ions D i ffercnt
a nel fall
and action of the gencs 111 (h
of ClIirolllllcnt on
could I) illmtratcd
Lethal (hlolophyll mutants-A HAil Ull died ullder field (onditiolb tllrce
Semimiddotlethal mutant~
a Imost norma 11 y i 11
under gnulhnmc cnnditiom kaves and pro ltlured a fair amount of seed The mutant
field and had
to accumulatc
bccomes nmdishyThe raccs which
wcre denotcd a irltS(C11 under conditions classified as xtnllu in held The races with which green wcre dll1otcd as vircsccns llIore viable than otiter races with ccdling
Table- 8 Interaction BeOHen Vircs(cns and Aurea Genes for Chlorophyil Uttirienlty in I Generation
Seedlings
Hyrids frc(~n Yellow Gold Albino
Ohscn ed
CalClllated (ratio
o hlcrvc I
Caku]arcd (ratio 36 1lt1)
J9gt1
IIl
qj
II
ltIR
17~)
IRI
(l
~17
1)[
ll
18
J2]
121
iii
LlGI
ltUR9i
7 point of prnblbi1it)- X2 7tZ
point of prohability
obtain seed even ill thebefore and from although their
of vironment whicb
The differences 111 the the in(]uencc of enshy
ill beets were also descrihed for olher crops Akerman (2) in oats Smith and Holm (I tIl
The of dnmi Hallce of the genes also varied undu different conditionslt of chlorina type were as viable ill the chell looked much JigJ1tcr in The Chell did not differ in the green The gelle dl was reccssive ulldel Another kind of behavior was observed in the field gnw for the were light and could be
the to
eft were normal grcen while the Tn the latc summer the
In sncll a way the of dominance changed peTiod as he seen from the data or segregation in (Table 9)
the
Tahle 9-Segregation in F Chlorin Hybrids (Ficld Obsenatiol1s in 13)
-0 of plants ith folor of Icn(t as follows
Reading in Grecn Ligh (middotgrcell Total
July 11
October 17
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
belg (I) found allother gcne in i)(cts the gelle which green to lIormal grecn SCt lors
was indicated bcgg (I) Yellow to senors The inlwriullce of lhi gCllc was not hOWll
The deficicilcies which dcvc]op ill the
VOL IX io I J-IARY I
of a 111 u ta ble gCllc ha c beell descrihed ill erai (l g 19 SimilJr genes llOt found in lJia
Chlorina type-genes (h and defens of ill beels
redu(cd
-rahle (L-inherlrance of Cldorina rype Deficiency in JJeia ~JUg(H L
UtUh(T of StcdJings (~cncramiddot ~unlb(r of
tion lamiH~ Green Yellow
Gene cl1 isolated in CS 32 Ohsentd
CainTatcd (ral io I)
h tl~ Ii
t()) i)OI]
Obscl cd
(llntlated ratio I
11 1 F tnl
lO-)7t)
10
ILW)~ll
OhsltrCi[
ClicuiarClt1 (rat io I)
lFI) Fc Ii 2) ~f 1111
100 Olilil
Obf-(led
Calculated (ratio I )
1~ )~) -) Sit 112
IIIgt O ~19L~)
G(llC ell isolated in Great fesitrn hetmiddott
Ol)srycd
Ohgterved
Calculated (ratio I I) 114 117 [0]
Gene (1 isola(d in
Obsenclti
Kleinwanziebn b~t
Calculated I) 5525 IR 07511
()hS(T cd
Calculated (ratio ~~l) JlL
point of probahiliq
point of prolJahili~ X~ 6fH
X
gCllC differ ill t he oj rcdllClioll of the Ihe gene ( which was isolated ill tile
reduced the 0 roob about 0 pcrccllt Another earlier (11) dccrcased the rool much more
which wcre isolaUd in the inbred lille SLC F hiJri(b normal
hI ilrids eTC l1onnj grecn III
Ilormal lk H E
Sugar which crossed S7R
crossed Ilormal green Fe hybrids howed mOl
Vhcll Iwcts of the chlorina 11]( beels SLC color of cotduiolls and obtained from these
rhus til( beCIS il h 10
oj the gene oi 1(11 liS the
ltlificnlIl sugar hlCI yarietics showed 19gt1 and I ~Fj fi)
K oj lhl Creat VVcsterll light
i II Longmont Colorado alld to clllorillli tYI)l hcets SLC
which I rcccIIcd hom Longmont F plants green and
[n 19) T studicd ilso the rc([i( gene 01 Ihe isoblcd III Ihc iniJred lil1( SIC I02~ lun which from KkinwlI1zlebcns beets Ii)
Interaction In i t t~
shown Ih11
l the iahle
were combined wilh til( llOmogou gellts cnhini( antHhcr type o[
for CXllllpIe Ihe gene of chlorill[ type thell two via illc under illlnhousc
between certain genes for of the Circsccm tvpc 0 the combination of (110 different
nOIl-
the
Hld
genes of Ihe ircs[cIlS
conditions which carried the above-mcn t iOlled and I he F (lrogcln
nine grcen a 11(1 ((n
Tahe 7-Intcllcion Uetwccn inSHIll and Chlorina GHtS for Chlo1ophyl1 fhgtfirknc i Il F~~ G(n(r~Hion
I-Jvbrids
Sredlill
Ydlow x
Ob-crcd
CdkllIalnl (raio ~17) jll(j
I pOlllt of prohlbili ~UH
poinl uf probahi[ X = (dH
VOl IX C()i R I ~)7
Tahil also show) that it Ias piblc 10 distinguish by [h( color of and mltllrc leaves lhe of the genotype from the chrh
genotype ill thest
When the gem VI yirCSCCllS comhined with the gene A II
cgrcgatioll complicated wcre observed F could he because tbesc plallls carried the genc Au pro dneeo after F which [he ratioll l green
(4 Vi1lilIl(lU ViViAu(lu) I I Vtlillllllll) I alhillo (I II
The threc viable classls gIT F families which (Ollshy
d assifica Lion
The effect 01 emironment on chlorophyll formalion III different Culthatioll 01 beets the main
lIchas urea differenl (nvironmcilts showed that
vcry much the lormation of
DiflprclHCs fIl thc lspecially obmiddotiots dlCIl these or under field rondi t ions D i ffercnt
a nel fall
and action of the gencs 111 (h
of ClIirolllllcnt on
could I) illmtratcd
Lethal (hlolophyll mutants-A HAil Ull died ullder field (onditiolb tllrce
Semimiddotlethal mutant~
a Imost norma 11 y i 11
under gnulhnmc cnnditiom kaves and pro ltlured a fair amount of seed The mutant
field and had
to accumulatc
bccomes nmdishyThe raccs which
wcre denotcd a irltS(C11 under conditions classified as xtnllu in held The races with which green wcre dll1otcd as vircsccns llIore viable than otiter races with ccdling
Table- 8 Interaction BeOHen Vircs(cns and Aurea Genes for Chlorophyil Uttirienlty in I Generation
Seedlings
Hyrids frc(~n Yellow Gold Albino
Ohscn ed
CalClllated (ratio
o hlcrvc I
Caku]arcd (ratio 36 1lt1)
J9gt1
IIl
qj
II
ltIR
17~)
IRI
(l
~17
1)[
ll
18
J2]
121
iii
LlGI
ltUR9i
7 point of prnblbi1it)- X2 7tZ
point of prohability
obtain seed even ill thebefore and from although their
of vironment whicb
The differences 111 the the in(]uencc of enshy
ill beets were also descrihed for olher crops Akerman (2) in oats Smith and Holm (I tIl
The of dnmi Hallce of the genes also varied undu different conditionslt of chlorina type were as viable ill the chell looked much JigJ1tcr in The Chell did not differ in the green The gelle dl was reccssive ulldel Another kind of behavior was observed in the field gnw for the were light and could be
the to
eft were normal grcen while the Tn the latc summer the
In sncll a way the of dominance changed peTiod as he seen from the data or segregation in (Table 9)
the
Tahle 9-Segregation in F Chlorin Hybrids (Ficld Obsenatiol1s in 13)
-0 of plants ith folor of Icn(t as follows
Reading in Grecn Ligh (middotgrcell Total
July 11
October 17
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
gCllC differ ill t he oj rcdllClioll of the Ihe gene ( which was isolated ill tile
reduced the 0 roob about 0 pcrccllt Another earlier (11) dccrcased the rool much more
which wcre isolaUd in the inbred lille SLC F hiJri(b normal
hI ilrids eTC l1onnj grecn III
Ilormal lk H E
Sugar which crossed S7R
crossed Ilormal green Fe hybrids howed mOl
Vhcll Iwcts of the chlorina 11]( beels SLC color of cotduiolls and obtained from these
rhus til( beCIS il h 10
oj the gene oi 1(11 liS the
ltlificnlIl sugar hlCI yarietics showed 19gt1 and I ~Fj fi)
K oj lhl Creat VVcsterll light
i II Longmont Colorado alld to clllorillli tYI)l hcets SLC
which I rcccIIcd hom Longmont F plants green and
[n 19) T studicd ilso the rc([i( gene 01 Ihe isoblcd III Ihc iniJred lil1( SIC I02~ lun which from KkinwlI1zlebcns beets Ii)
Interaction In i t t~
shown Ih11
l the iahle
were combined wilh til( llOmogou gellts cnhini( antHhcr type o[
for CXllllpIe Ihe gene of chlorill[ type thell two via illc under illlnhousc
between certain genes for of the Circsccm tvpc 0 the combination of (110 different
nOIl-
the
Hld
genes of Ihe ircs[cIlS
conditions which carried the above-mcn t iOlled and I he F (lrogcln
nine grcen a 11(1 ((n
Tahe 7-Intcllcion Uetwccn inSHIll and Chlorina GHtS for Chlo1ophyl1 fhgtfirknc i Il F~~ G(n(r~Hion
I-Jvbrids
Sredlill
Ydlow x
Ob-crcd
CdkllIalnl (raio ~17) jll(j
I pOlllt of prohlbili ~UH
poinl uf probahi[ X = (dH
VOl IX C()i R I ~)7
Tahil also show) that it Ias piblc 10 distinguish by [h( color of and mltllrc leaves lhe of the genotype from the chrh
genotype ill thest
When the gem VI yirCSCCllS comhined with the gene A II
cgrcgatioll complicated wcre observed F could he because tbesc plallls carried the genc Au pro dneeo after F which [he ratioll l green
(4 Vi1lilIl(lU ViViAu(lu) I I Vtlillllllll) I alhillo (I II
The threc viable classls gIT F families which (Ollshy
d assifica Lion
The effect 01 emironment on chlorophyll formalion III different Culthatioll 01 beets the main
lIchas urea differenl (nvironmcilts showed that
vcry much the lormation of
DiflprclHCs fIl thc lspecially obmiddotiots dlCIl these or under field rondi t ions D i ffercnt
a nel fall
and action of the gencs 111 (h
of ClIirolllllcnt on
could I) illmtratcd
Lethal (hlolophyll mutants-A HAil Ull died ullder field (onditiolb tllrce
Semimiddotlethal mutant~
a Imost norma 11 y i 11
under gnulhnmc cnnditiom kaves and pro ltlured a fair amount of seed The mutant
field and had
to accumulatc
bccomes nmdishyThe raccs which
wcre denotcd a irltS(C11 under conditions classified as xtnllu in held The races with which green wcre dll1otcd as vircsccns llIore viable than otiter races with ccdling
Table- 8 Interaction BeOHen Vircs(cns and Aurea Genes for Chlorophyil Uttirienlty in I Generation
Seedlings
Hyrids frc(~n Yellow Gold Albino
Ohscn ed
CalClllated (ratio
o hlcrvc I
Caku]arcd (ratio 36 1lt1)
J9gt1
IIl
qj
II
ltIR
17~)
IRI
(l
~17
1)[
ll
18
J2]
121
iii
LlGI
ltUR9i
7 point of prnblbi1it)- X2 7tZ
point of prohability
obtain seed even ill thebefore and from although their
of vironment whicb
The differences 111 the the in(]uencc of enshy
ill beets were also descrihed for olher crops Akerman (2) in oats Smith and Holm (I tIl
The of dnmi Hallce of the genes also varied undu different conditionslt of chlorina type were as viable ill the chell looked much JigJ1tcr in The Chell did not differ in the green The gelle dl was reccssive ulldel Another kind of behavior was observed in the field gnw for the were light and could be
the to
eft were normal grcen while the Tn the latc summer the
In sncll a way the of dominance changed peTiod as he seen from the data or segregation in (Table 9)
the
Tahle 9-Segregation in F Chlorin Hybrids (Ficld Obsenatiol1s in 13)
-0 of plants ith folor of Icn(t as follows
Reading in Grecn Ligh (middotgrcell Total
July 11
October 17
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
VOl IX C()i R I ~)7
Tahil also show) that it Ias piblc 10 distinguish by [h( color of and mltllrc leaves lhe of the genotype from the chrh
genotype ill thest
When the gem VI yirCSCCllS comhined with the gene A II
cgrcgatioll complicated wcre observed F could he because tbesc plallls carried the genc Au pro dneeo after F which [he ratioll l green
(4 Vi1lilIl(lU ViViAu(lu) I I Vtlillllllll) I alhillo (I II
The threc viable classls gIT F families which (Ollshy
d assifica Lion
The effect 01 emironment on chlorophyll formalion III different Culthatioll 01 beets the main
lIchas urea differenl (nvironmcilts showed that
vcry much the lormation of
DiflprclHCs fIl thc lspecially obmiddotiots dlCIl these or under field rondi t ions D i ffercnt
a nel fall
and action of the gencs 111 (h
of ClIirolllllcnt on
could I) illmtratcd
Lethal (hlolophyll mutants-A HAil Ull died ullder field (onditiolb tllrce
Semimiddotlethal mutant~
a Imost norma 11 y i 11
under gnulhnmc cnnditiom kaves and pro ltlured a fair amount of seed The mutant
field and had
to accumulatc
bccomes nmdishyThe raccs which
wcre denotcd a irltS(C11 under conditions classified as xtnllu in held The races with which green wcre dll1otcd as vircsccns llIore viable than otiter races with ccdling
Table- 8 Interaction BeOHen Vircs(cns and Aurea Genes for Chlorophyil Uttirienlty in I Generation
Seedlings
Hyrids frc(~n Yellow Gold Albino
Ohscn ed
CalClllated (ratio
o hlcrvc I
Caku]arcd (ratio 36 1lt1)
J9gt1
IIl
qj
II
ltIR
17~)
IRI
(l
~17
1)[
ll
18
J2]
121
iii
LlGI
ltUR9i
7 point of prnblbi1it)- X2 7tZ
point of prohability
obtain seed even ill thebefore and from although their
of vironment whicb
The differences 111 the the in(]uencc of enshy
ill beets were also descrihed for olher crops Akerman (2) in oats Smith and Holm (I tIl
The of dnmi Hallce of the genes also varied undu different conditionslt of chlorina type were as viable ill the chell looked much JigJ1tcr in The Chell did not differ in the green The gelle dl was reccssive ulldel Another kind of behavior was observed in the field gnw for the were light and could be
the to
eft were normal grcen while the Tn the latc summer the
In sncll a way the of dominance changed peTiod as he seen from the data or segregation in (Table 9)
the
Tahle 9-Segregation in F Chlorin Hybrids (Ficld Obsenatiol1s in 13)
-0 of plants ith folor of Icn(t as follows
Reading in Grecn Ligh (middotgrcell Total
July 11
October 17
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
Table- 8 Interaction BeOHen Vircs(cns and Aurea Genes for Chlorophyil Uttirienlty in I Generation
Seedlings
Hyrids frc(~n Yellow Gold Albino
Ohscn ed
CalClllated (ratio
o hlcrvc I
Caku]arcd (ratio 36 1lt1)
J9gt1
IIl
qj
II
ltIR
17~)
IRI
(l
~17
1)[
ll
18
J2]
121
iii
LlGI
ltUR9i
7 point of prnblbi1it)- X2 7tZ
point of prohability
obtain seed even ill thebefore and from although their
of vironment whicb
The differences 111 the the in(]uencc of enshy
ill beets were also descrihed for olher crops Akerman (2) in oats Smith and Holm (I tIl
The of dnmi Hallce of the genes also varied undu different conditionslt of chlorina type were as viable ill the chell looked much JigJ1tcr in The Chell did not differ in the green The gelle dl was reccssive ulldel Another kind of behavior was observed in the field gnw for the were light and could be
the to
eft were normal grcen while the Tn the latc summer the
In sncll a way the of dominance changed peTiod as he seen from the data or segregation in (Table 9)
the
Tahle 9-Segregation in F Chlorin Hybrids (Ficld Obsenatiol1s in 13)
-0 of plants ith folor of Icn(t as follows
Reading in Grecn Ligh (middotgrcell Total
July 11
October 17
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
VOl IX 0
of green color of ithin commtT(ial pOjlulalioth in the leaves The diflennces ill the represents the main factor causing lin grcclI colnr of leaYb in individual beets 11) showed a
the melcr ill the color Therefore for
green color of leaves colol slandards
ma) Cllssifictl iOll of 1ltllll[wrs of III a short lime
To ucterlllinc of grnl color of irans the ( slandard
of E F Votehall was This color suuHlard I Ii daes for the or grcc color green 10 dark green Each ebbS is ltlidded
into IOllr submiddotdelsscs which do llot diilnclltiatc ill the phyll but shades o[ beet leaves was of the color stalldards the
(bsscs of this standard equaled per one squtre U11 of the leaf blade
heel races rich poor ill foi obtaincd lftcr selectioll lor the of leaves C OkOIlCllko the first to of chlorophyll He showcd tbat plants selected green color and isolatiolls produccd ditT(jCllCCS 111 the imcllsi ty of green color of the
I studied the inheritancc of intensity of green color hy mtnA sugar lwll dom 3~)11) 10 shows the the dOlHS and of Iheir F V and iJlcklTllSS hyhrids the of green color was done s(cral for each The dJt1
showed the
I ltThen cloll C to the 1 dol1t [61 I were allllost as the i r pa rents
When Ii I was crossed to dark grccil clone F as the paren ts
3 fhCH ( lone was ([(cd to dark green clone (i(j the were Interlllediate ill color in both
crosses () f lOll
i middotVhCH back t()
green clone the F j and vicc nrsa whcll back to the dark gTCCIt dOlle F III bnth
the ba(kltTo~s wcre intermediate in clear (olnr betwcC1l the F and recurrent
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
Table JO-Vadation in the Intensity of Grc(11 Color of Leavc~ in Sugar nCe[ Clones and Their fl f~ and Bu-kcross Hybrids Determined c cordillg 10 Color Scale of Prof E Votchal
Coeff Number of Plants With the Grark of Green of
)Cj I4natiuli Color as Iollows 1ulnher Vari~
of Parents of Standard ability (encratinn and Hybrids 9 10 II 12 l~ 14 15 16 Plams lean U(viatinn Percent
Clone with light gTecn Ica n~ gt~ ~ OHIi
1 gr(CH lca(~s ~l iK24 01] j(i
F 1-17 x Hi 11
Pmiddot()ti dil leave 1middot1000
CIOI1( Pmiddot20) rith dark Jeaes Ii ] ~J) 120
PmiddottWi X 2ns 21 It) 97 19 107
Ii x p-nfi If q1 ~I
F p(iIj X P ~)7 l 1 I 109
Backcn)ss p )7 X (P37 X Zl 214 24 26 II LfF 2~Leuro1
Backcross X 117 X 1(16) 28 If JI III PI 11D J7 210
F Plt17 X pmiddot(it 11 24 B 21 II K I 17 ~K7
Pmiddot66 X P 20 2( 37 (i 10 19 17 II Ii) 260
Illt(n~it STC(l1 color ith the increase Yo
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
VOL IX -0 1 JA-l Ry 19)
G F and lc hybrid obtained lrom (H)SSeS between light glTell done 37 and dark grcen donc GG ( This was an indication that the gemiddotnetic diHennee of green (olor which were observed ill the bcct populations were caused mainly hI the luclear gene but not clelllenu of ltytomiddot plasm The backeIm (ollfinllCd this ultldmiol1
7 The genetic of the F and F as well as of the (ormiddot responding backcross 11OW(ltI lhal the intcllsity oC grccll color ill bects is a quantitative character which b caused the actioll of polygenes Thereforc the inheritance of the oj in sugar hcet races must he studied Oil the basis 1)[ method for the or the However 11w analysis of
01 intellSl y of the grcCll of th paper and will Iw elucidated
Genetic beets
Aurea it) lutecellS (gene Ill) yirccclls ( gcncs-TJ1 1t and chlorinltl (11 and
1 he 01 interaction between genes vi[ and and Ii and All are dis(llsscd
01 cllironmcnt on somc indicated gClltS is showll
caye (othisting of Illosaic white and green tilI( senor llIore lten as a result of chang-cs ill the heritable ul)tltlnccs 01 than ill thc result of lluclear mutation
study of differcllt hyhrids showed that the inherited diflermiddot (lletS in the ltjuantity of ( which arc peculiar to different withill commcnial arieties arc determined the nuciear inheritance that is the ael iOIl of Thesc ill color of leavegt and ill or Thcrclore
a of
b
c calise the (hloromiddot
References
(I) IlEG( F A 19middot10 List of charactCrs and gcne the Blta ndtlri L 110( ArneI Soc 2109middot111
KFRfAr- l~l22 UmerslHhungeli uiJer cine in direktcHl Sonnenmiddot lichte nicht Icbensfahigc Sippe Oil Anella Eatiltl Hereditc 3 147middot]77
111
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
onL 01- TilL S S B J
(3) -IJERSOE G 1lt)23lalltmal inilerilauee 01 III malle
Botanical Gazette 7li41lmiddotmiddot1l8
Ihuow C vo I ~I)) Die Bea Rubel Zuchter 2gt 101middot122
(5) 1J07 nlirrililllllll
(ti) lhCR E 1010middot1 j Y01l Chroma tomiddot
bd Mcialldriulll Amirrhi1l1lJli lIud ZeilSchr L lndukrive LHImiddot102
CLAUSl R E and CA~lERO D ellloroshy
(H)
1291-329
CORRESbull C 1918 Zur Kelllllui eiulachcr IlIllIddner Ilaslarde Pnus Akad der Wis SitlIJeL 11221-26(1
(10) GOIOLlAKO S E 1954 Plastids and slarcil ill root 01 su)ar uecl (In Russian) kad auk SSSR (1) (181middot881
(II) CREGORY R 1 191) On ill Prilllula sinemi 01 Geneticsi 305middot321
(12) CrsrAFSSOi IH2 The in arjuus types 01 chlorophyll l111tatiob Ifcrediath 211HlH92
(13) GusrAFSSOi 1946 The eJeel o[ OIl aribi[ and igour Hcredi las l2 2fi)-28(i
(H) GUSIAFSSOi A JI17 The advantageous cHen oj deleterious llIUlatiOlls
Heredi las )3 573middot575 (Iii) GUSTAFSO~ bull iYIOI md 0 VEllSlEIi L 190
iactors and heterosis ill barley Hcrcltiits l6IHimiddotllt)2
(16) HAAN H DE 19)L Inheritance oj ell Gellet JO357middot 16
( 17) HOIL G 19)1 Chlorophyll mnlaliollS m
1gt157-471
(IH hlAI Y 1936 Gellomiddotand of )
Genetics )3 169-195
(19) hHL Y 1916 Chlorophyll and ids Zeits(r fInd Ahst
KARPER R J9)0 The ellell in the in sorg-hulll
KlmiddotRSAiOV A L and PAVLl~OVA O A sugar bcn
K URSAgtOV and TURK M R J954 About the fonn of mobile sugar in the ular sYLel1l of sugar cHad auk SSSR I)) SS)-HSS
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
VOL IX 01 JALRY 19
J AllRHT II IUi l bel rhlorophyllmulanlcn bei Pi~llll Hlld die Vercrbullg etntr lIlmiddotIHmiddotll Mutante Hort Genet ]0 1-18
lATllER K 1H9 Biometrical Jew York Dover Publicatiom rlC 1)15 p
1Iu-ERATI O I J28 L nternltltl fur
of Zcitschr 2 1137-1 112
IUERAlI O 1~112 Die bei Bela L Zuchtll 11214-21fgt
OKOImiddotNKO 1 C IlJ21 Zm des dn Zuckershyrube lind seiner Sakharotrcl Moscow Sortovodn()shy
of the PBSCD of the Sugar
OKON~KO A C I J28 The colour of leaves and the of in them a characters in sugar beets (In Rus
COlllributions trolll Sciclltific Plaut Institute Kiev 219middot80_
OKOzElKO A 1938 About ariations of the ( in sugar beet Jeans Kid Acad Sci InsL Bot J 26 III-12i
OKONElKO A S 1 Ill Assimilation of carboll Formation aIld transshyler of sugars in heelS (In Russian) Vcsoiuznyi Nauchnoisskdovatcl skii Inlilut Sakharlloi Promyshlenosti Sveklovoustvo (All-C Ilion Scientific Rcsearch lmtilutc of the Sugar Industry Reel Culshyture) 12n-28R
S TOIAClIOV J 11 alld KI-KLCH A M 1928 Inshyin ugar btCb with difIerelit quantity of
Con lri bmions Scientific Kiemiddot21947
(h F V K 19c12 Somc M endd ian characters In
Bela obsennl III lile Y -R-B group Jour 01
(3~l) PAL H 1 alld RA~LllJA S 1941 new type of variegation III
rise Indian Sci II 170-17G
POWERS IlRoy LOCKE I F alld CARRlmiddotn T C 1950 Panitioning method of appli(d to characters of
gric Tech BulL 998 I-JG
Tber die Vcnrblichkict der hei Zuckerruben O l Zcitschr l Zucherind 11
Lll1dwirtsch 20 149-156
RHOADES M 1 19middot1fi Plastid Hlulati()m Cold N Y BioI Lab Cold f-brhor HioL
II 202-207
(17) RWGVAY R 1912 Color LIldanh and color lWIIlCnclalUlc
tOil D C Ui page alld plate
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
(41) STElllK V 1921 Zum Studium del iJllOllnaiitatClI bei dcl Zuckcrrulx Zeitschr ZuckerilHllbtlic (zccloslovakidlul 45 tOJ11L
STIlllH II and IIRsClILJo K IH I libel cinCH IllOJl(lgell
Fall VOIl Helerois bel nlirrhilllllll D Bot (CS1H-lHi )58
TABESKY t structure of the indicaor or dlC Jill or the lell (Ill
~l all
Nauk_ SSSR IZHstia SCI BioI JG09-GI
VVINGI O I )19 Oil tlte loll-Mendeliall inlicrital]Cl III COlllpt Rend Tra Lab
VRIGIIT SEWALL IDi2 GL Brit Res COllllcil
held at the Institute of oj the
In plpcrs rea(]
Genetics Research Coull(il
ov 1m S S B T
SAVITSKY V F 19~O Oil oj the 01 sYllIJlwms of Bea ita B
SAVITSKY
ill II da
tics alld
1 (Ill lk]aYil Ccrko Plant Breeding
To the the
SAVITSKY V F 1932 vlalcrials to the 01 sugar bcet raccs ~ludlllOissJcdOatd ski IllS1itul Sakmiddot
J p
(fl) SAVITSKY V F 19middot10 Gelletics 01 the Nallchlloisskdmatcl skii I
losti Svcklovodstmiddoto ScielltillC Rcsearch
Kiev tklaillskii
beet (Ill Sakhallloi
I ~19middot(j8J
SgtIITII J C H 1919 lwccscs Jorlllalioll
In Franck J and Loomis E cds IhoOsl1thesis til plants eh 9 loa Slate Col
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