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The Use of the Monoploid Method for the Production of Homozygous

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University of Nebraska - Lincoln
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Open-Access* Master's Theses from the University
of Nebraska-Lincoln
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1953
The Use of the Monoploid Method for the
Production of Homozygous Diploid Lines of Corn
Melvin Dale Rumbaugh
University of Nebraska - Lincoln
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Part of the Agronomy and Crop Sciences Commons
Rumbaugh, Melvin Dale, "The Use of the Monoploid Method for the Production of Homozygous Diploid Lines of Corn" (1953).
Open-Access* Master's Theses from the University of Nebraska-Lincoln. 2.
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THE USE OF THE
MONOPLOID
METHOD FOR
THE PRODUCTION OF
HOMOZYGOUS DIPLOID LINES OF CORN
A THESIS
Presented to the Faculty ot
The Graduate College in the University ot Nebraska
In Partial Fulfillment ot Requirements
For the
Degree ot Master ot Science
Department ot Agronomy
The writer
wishes to express his
appreciation to Doctor -RKQ H.
Lonnquist for the suggestion RI the problem and IRUthe valuable
assistance given
throughout
Sherret S. Chase, Research
Department, Iowa
the course ot this study; to Doctor
Associate Professor, Botany and
State College,
for supplying some
Pathology
ot the material
used in this study and for his suggestions as to experimental procedure; and to the Department of
for providing the field,
Agronomy,
greenhouse
made this work possible.
f.t
University ot Nebraska,
and laboratory facilities which
"
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bsul_ ......................... " " " ...... ..
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111
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•
•
41
4•
~oplold
lD41Ylduals baYe baea obearYed to occur 1. sWferal .peol.s
ot pl. .,. at y-mAC. thous'b. lleus11,. low, t'nqueaol.s.
art1t101811)" lJlduee4 doubUaa ot the
SpOII'. .011S or
oln"oIaoaome OOII'pleaeDt or oella t •
...tore ot the pi.tl11&_ and '-'lute aertateru ot the IIlOBOpl.old pl...,s
8ftatly 1........ the eha. . . tor ......fu.1 eelt·tarUliaat1oD.
'ftle
hoIIoaycoua diploid 1 ..... eo pr04uoed aN the • •etto equlftleate ot
adYaa0e4 geMn.t1oa 1.bft4 It..s.
Ttlis prooe4Ul"e haa oerta1J1 a4TeJltq_ O'f'er the
OU.~1a'fT
preotloe
"1' ot ....1'11tl0•• to pJ'Oduoe lnllNd 11....
ot .81t-poll1.'10. tor a __
T1w treque. .,. of oo.rna..
ot .780.... homoS)'lOU8 tor eertatl1 d••lrab1e
lIfte•• III the ,ro..,- ot aD. lad1y14u8.1 beteroSJ'lOue tor 'he ..... 111
queation, will .. the aquere ot the tJ.'eCluenct:r ot oOQU:rftDee ot ,be
_ _a
.~«
,he 4aa1red . . . . .
geneti. aye'- ot indiyidtuU.
" . --.op10i4
_'hocl 1'1" ,_
.,..t•• la teata'l.. nprodueUu. tOn.
'1'beH8ul tad aporoph'" are 8OIIplet81y homoS1lOUS u4 a1loul4 pro••
a4yaataa-ou. tor the ."'4'1 ot eo-nata .Mdl. . probl....
"the••
homo.,.••
would ottar an
oppo~t,.
tor the study ot
het.rost. 1. a . . li.llt l'UUlttDl 1• • olearer eXplanation of Jqbrid
Yl.r t.rOlI the ....tleal po...t of Yin.
!'be GCu'Pan.oa ot releted
popul..tlou ot 111M. 4erift4 .." taa doubl1a& ot lIOaoplotds 8Jl4 11._
obtalM4 by ial»re.alq 8Bd ••1",108 WOUld be
toftlla'ttoa •• to tt»
dp~ ..,
11a14
ta-
.tte.t1....... of' . '••11. . _tIaod. 1n exoludlq
deletenoue . , .••
1t1e dlplold8 derlyed by this Ihtbod OOUld also lila utll1M4 tor
,be etudyof 81:p&r1aea'tal 4eal• • la the ne14 aad tor detctJ."m1Jl1q
the residual Yarlal»111ty due to
.UY1~tal
f.4Itors.
'festlag ot t_ IIOROplold. thase1• • tor aerw.h aaronomla ohare...
te1"1.t108, wOh es d1.8ftH real.tea•• , tollowed 'by baokoro•• lng tbe
plllDt. &0 • •ted
'0 the .ouree stook should
hetOl!'O!Q'1OUD stoOk. wi th
the 4.....10pllHBlt
ot
Pftftilt h
4eY.lo.....' ot
hiSh frequno1e. ot de.lred 8 ••••
a praot,loal
_~
ot bree41Dg oom tllroush
,be utl11at1oD ot llosIoa1FWI 11.8 obta1Jled troa the dou'bl1q
ot
lIOaoplo148 wUl depend. ullOa the obtM.1Jl1D8 of a ree..oaabl,. 111gb. raM
ot monoploid.
tor
001'11
trOIIl a.ay panloul8t'
;produotlon, aad
OD.
stook, NPreseat1Jll d.s1mble a. .te.
tile . . . .sstv.l cloulUiDa ot .ala lIOaoplolda.
Thia atud1 Is .. 1....tlgat1oD. ot vano.. upeo. ot tUM prob1....
TlI8 11 wnl1;UI"e OD ptlrihellOQI8us18. hapl01d,.. 8lld oolohlo1ne teohniques 10
~.
ra~.r
exteatTe end
pre8.t dud,..
1'1
oomplete l"eYlew 18 be,oad the 800pe ot
!J.1.eretore, only 'tbat portion of the l1teratU1'e 0011-
1....8tl.'.4 111 the i1lb.er plallts baa boa sel.oW tor NY18W.
11ttle of the literature de.liD« with the
orto~tl0
Alao,
aspect. ot the
pl"O'blem hea been 1llo1uded hereln.
'Dle tel'm1aology or the particular author kaa. ln ttll OtlaH. beea
apecle.,
_on
a8
those ot tbAt
fjeJIWJ
)flcot,I!!&, will preseDt 4ata en4
0'b.8"8.'101&8 o<SMralnc -'pl014 pl_ta 'l&h 1. are Aot IIODOplo1da.
.,.,.1"8 haTe be_ 1nolude4
to%' Hf1_ 'beea.uee
'il"bM. .
at the al1dlcnt,. or the
.al0 pro08. . . ad p:robleJ18 W tho•• or ape01•• wha1'O th. baplot4
plata ere 'bue llOaoplo14 ••
Roplo1d7 - . tiftt oytologloally prewed and ftported b7 Blalceal...
BelUnl£. J'amh_ 8tld Dersner (1) 1A the pau. Dature.
1
"ith lnoreee1.
lmo..184fJG aad :rettaemeat ot 1ieohD:1quea 1. the tteldG or -p1Mtlce _4
plOint breed1n&.
D\UI!lWOUG
iutazt...
hi_er pluta baTe be_ :reported.
tbe tollow1ac " , f t , ttlIIOJa8 othor.:
or
the ooourftn08 of haplo1d. lD tlle
Their
'lrlt1oum•
In all
~_1'8,
and
haa be_ noted 11l
Cfl281OW3. Ont:e18. )')atura,
00887plum, B0I'48!!!. Ll;t..l• • LxggaertllOU11,
~reum.
OOovr1'el:lC8
E!:!erla •
.IE" ... Oenothera. aolaaum.
1i!!.-
the 4eteot1on of the pal'thenofJuaott 0 haploida 1s
po881ble In hybr1d propBiea ot lntlm- or interspeclfio nnd InterpD8rl0
oroeses prlnser117 by' _eir nUMfablaaoe to the mete:rael
to the pa.terml '1PO where clOmin8Dce 1s IJrfOlve4.
~
ln oon'tl'tta"
'lb. opposite altaatloa
exist. Where eadrogenetlc haploids ere distinguished by their reaeroblaaoe
to the paternal type in oontrn8t to the maternal type.
'lith few cn:oepUou the f:nqUlllnoy 01' ocouneaoe 01' haplolcl8 hAs Hell
Yff'r1
low.
Beneea 1931 and 193'. Satlnca, meDel.. , _
lated '3 Dature
h~plo1d.
Aver"/ (40)
180-
trom a population ot 410.000 lnd1Ylduela, e
t'requenoy 01' about 1:5.600.
lohftnMA (26) reponed an UIlususl t'l"e<luenoy
ot 1:100 f'Joa one eou.neta1 lot 01' barley_ HAl"lftl'u'l (21) tound a rate 01'
1 haploid amona 3,000 to ',000 111ftnta ot sM-ls1and cotton.
Jon.. aM
Loagley lSI} dhoove:l."ftt1 11 haploid p18nto smoBS .p,Proximately 300,000
1"148 plut..
JCatll78llft (!'1) ha. obaerve4 thom to ocour at
tbe 8tuIMt epeet8. but touad a t'Nq\umo,. of 0.1
tbe p8.1'tbao....t10 de"'eloPlMll'
ot
&alth (41), ho....er. worked with
0.48 per cent tA lXitlet"!! 1'1101100000..
Obae'f"f'at1o". by RaJldolpb (39)
8 ~u_G7
~r
.ent or 1:1.000.
he". lDdl08ted thRt
ot tbe
ea Btloleu.
in
_1_
the
tMque.,. ot
UDder tbe
.t1Jlull 01' polltaatiOD 8.Dd D01'I.Ifll • •aapem development 1_ of tbe ordor
ot
OAe
In ..., thous.at,
01"
0.05 per
cellt.
More reoent in",eat18atiou, 11. . . .r. baYe 8h0WJ.t thet perhaps ,be
trequeae1 may
reponed
1:900.
1IOl"8
o~a1nl_
oloselyapproXSMte . . til oae tiboWlQll4.
(lhqe (ft)
43 . . .oplolda fro. 38.6M ..8411... , " treqUAmo7ot
lUuet (12) tound that progenie. troll tl'1.Qldo matze plant.
pollinated b7 JlO%Wll diploids ss.e a f'Hqueao, ot aoaoplolda of 1:958.
J"requenoie. ha" varied, howe.er. trOll 1:100 In a diploid JllU.1tlple re-
o•••l.e ' ••tel' used by stadler (uapub11abad) aad reported by Oh&_ (4)
to 0:1<4•• &68 obta1Jaed 1)1 F'ontmO (1S)
pollinuted malae
.~Jrl.ti
aJm"
pro_i•• of 1'1_ open-
...
The euploid reducUon of Obl.'OllOa.,. BUllhers that »104 .... laplo14
apo1'Op,w. lu1a been abae1'ftd to be aoo0llp8Jl1e4 bJ' oeJ1ain chaaaea In
llOrpholoQ the' ftre oha1"lloterl.tl0 ot the aonop1014 001141 tloa.
ette.'.,
U
The.
\"lOud be expecned, 8 . a reYeftal ot thos. _loll MY. be_
w:14e1y obaened W ooau.1' •• a _SQ.lt ot euploid il101"eaae. ot Ohromo.OIIIe
1I.W1Sbere.
'!'h. 8441'loa
ot .. eotaple'.
sao.
baa bee lIOted to bftBl
.'bout .. lacre... in ad. .. ot auo1.1, eell., 884 so_ or.... .. -11 ..
,l"Oloqed _p.t1.,. growth period. end oansln pkpl010sloel 81""'10118.
Th... ettaet. haft
-.11. repofta4
(IB) • •01lJ&le (36), aad
The lIOat 41reot
plo14 ooadltlon 1.
0"1'••
.~
~..t
ll)' E~.r aa4 Ruttle (13). Konort
r..ul. tbe mil the 1Jd \t at 10D ot the lIOao-
ot a ftli4uotloa 1. the abe ot ..118 aat1 auol.l.
leostott U.8) .tudled the rat10 of both Ita.....ad YOl. . 4 _ _10u ot
oe1l.e ot hap1014 _4 diploid toru ot three .,.01•• ot 1I100tl_ and
Crepl. ft211larta.
•e.
Root cella.
no-tal eella, end pollen _t_r ..118
weN .a.'Ur84 turi_ tlle aour.e ot tJuI atld,.. 'ftle least
8lIOW'lt ot
oocurred 1. 'he 11aear 41__1011 ot po1lea JIOther cella ot
the M1la ot the 4tplold toa
oal1. ot the _»1014 tbftt..
-.:re
1IItre
I.
oha_
811.,...tri8•
1.31 per _., 18.l'aer thlm 'the
'!'be 4ate Oll tbe
t~
apeot.. ot nco'l_
ttlnher w.'bataAtlat_ 'b7 the . . . author 1A a latter paper «89' ..
Qat•• _4
GoodWia (18) IlOte4 that tbe eelloot • haploid Oenotben
ru.!!:1oa1s _re a.11er thall tho .. ot tbe 4lplold tom..
ad Ae.ae (1.,) toad the, haploid polln
S1II11ul,.. CIlda.
lID"er ..lla of t£U10U11l ooapaot.
hWlbo14'U. were 8II8l1er tba. 'boae or ,he 411'1014 plat..
'IMnon (1',
ftportect that tke _all. 41_nera of nuole1 or po11a _'her _110 1. the
diploid ead haploid torma
or
o..otb!r~ tlna,I!!!!a were 12.9 aad 9.'
moroM. reapeotlwl,..
1'I. .aahla (M). who worked with a polyploid aerie. ot C;N218
oa!i1lar1a. found a atroas poettl•• oor..1atloa between the okromattD
. 8 . illS _allUred b7 tbe 182:l8th
ot the 011210____ 111. the _rlat_ cella
of root t1p. as th.Y' B:ppee.red 18 _tapM•• GIld the .ell 'fOluaea.
He
noted that ,be oell areaa 8Jld . .11 yol18e. ot tbe haploid. toR of thi.
apeel.a were Dot quit. bal1' .. large 88 tho.. of the dlpl01d form.
Tbe .ui( or ttett."'" (45) aad
JCostot't (29) baa d8llOutJIBte4 that
the llOl'PlIoloc1oal alkrat101l8 01' _plold. aa4 8xp$riJllt1lW autopQIJPlolda
are ulti_tel,. due to alt...tloa. III '011 all18.
WI
more dell••• aaaa-tol'lU
'9'8.I"ioua
OJ'",
or
the 41pleld. with a re4uotton lit at_ of
aM plat parta.
Sndtll (.:1.) 0baeJ'Ye4 that the 18_••
pl. .,.
Haploid. la_all,. appear
or
we" M1"JIOWI' 8Ild .0ne1' than .....
Haploid Cmt. el1la1'1e pl. .:t.
_l"8
haploid T£1tlewa
1I!pIlOOO!!!J!
le. . . or tM 41pl014 pleata.
ehowa to
f t _ _le
reduee4 dtplolda
b1 Boll1...heed (!S) but the,. differed 80!11wt&at 1a lea1' 8hepe.
Ohue (t\) note4 that a morpl\olopoal el1.arUterl8tl0 01' _1_ moao-
pl01ds Gould be ued to • •t111&\11_ tl'I.em f'rttta 411'1014 pl_'a.
'But t1m
••e4l1a, leat ot IaOD.Oplotda we.e fouad to be 1 ... tJum OM-halt aa 10DB
tile oompereble l . t of dlpl014 ...aU.... ot tb.e IIOUMe
all luteneee.
Be
noOk III allloa'
Fonuao (lG) Me also aote4 aa tU tared l.at appeareaoe
1J1 ha,lold _1.. plata.
lltta.'ta a1milar to the •• l1aTe be_ reported in .,e:r10\1.8 apool....,.
Blak•• lea, BellJal, II'8ftharzl aut Jar_1' (1). Browa (2). Chrl.teJUID aa4
Damto1'd (e), hr.A (14'. Qa,•• _4 Ooodwta (18). LamIa (30). Llad.1:i1'Oll
ea4 Kooa (31). MoOr.,. (31). Rol'4euktold (3", aa4 Toatol'p (43).
A IJ'M.1:i aaQ ot 'h. heplotdaobee1"f'8d ia pl.t...... art ... 8IIOIlI
IlJ'brtd pro...... ot 11l1:ierepeclt1o 01' la'e'J',aerto
t_
OJ'lO.....
J'or thl.
reuon. 1t hea be_ beld 1.'tJ 'f'&rtw. M>1"ken. a. tor axmaple, Kostoff (D) •
. .t b7'bndlza't1oa 1.
ot .plotu.
ODe
ot
DOre .tt.ott.... _thode tar . . J)ftdtlotloa
,
By orooll1. . Bt'lOtlana ",,'." . , SOl!!!! alE!!. Fe"".la vl01. .a,
tI!gpra1nm epleatua. A!.!'2e Bel;la4p88, _d DetU1'8. atraaloalua. IVaaoT
( ~4) attempted to study tbe etfectS.v..... of hT8ri4i ..t,oa la the
duotioa
ot baploldo. Oa17
partbeaos&&atto diploid
0 ••
naltl. ned
Ll!0p!ra~oum
In ~ tlowu monoooe"m, Sm1 th
})1'0-
wa. obttl.1aed !IIld It • • a
.!O!l!l!!l.
(41) obeerre4 49 h.e:plold. &.II!lOAI
2,4'18 h,-brld. (2 per oeat) aner oro•• l .. with
1.
aeRloeld~•• a4 ,
amoag e,'fOO lJldlvl4ua1. (0.08 per oat) trca "~17 .elte« prolm•••
Haploidy
1m.
o'bael'ftd 1:a Oeaothora 1Iy DaYl. set JCul1carnl (9) wi tb.-
out emplo11Di oroaa1ag, wb1le
maar.OIl
(14" Oates tmd Oood-s.a (18) t en4
Cate.alde US) touad haplo:Ula 1a the . . . IJ8IW8 Ilf'ter IBterapeolt1o
DtrI1a (8) !'eUed to 1a4t.lCe
h.ybr141zatloJl.
tu
4eYelop&l8llt ot baplo14.
1n 002l0tJ!!ra L!!!!£Hl!!! by polU. .tto. IJ&'f01V1l\C wide oro.....
Sbdlarly, sattaa. Blakula., aIId AYerr
(40) have reIJ01"W tJaat
tbe t'raQ.UIl87 of l1eplolda 11l Detura 1'IoJa ,..1te4 pm,al•• 40•• .,t 41fter
tJooa tha' 1a Pl"Opal•• t:rcm later- aa4 1Btnapeolft'l OM. . . . .
.1'....
'1be tiret
!'arsd'l.Ga, 8Jld
t~
Da!!!£a _plola. obtalaei 'by make.lee, Belllag.
(l) ,.... tJOII a group ot altDoraally allpearl. . Sati-
Y14uala _0u.re4 ill .. attempt to blkoe .1'Omoaoaal Ift'&8Ular1tl••
the flppl1 oatt oa of 0014
.a ..
attaalua.
R.dolph (38) touad a haploid 1.
dur1nC the course ot
.rteota ot heat
au
~
111 _terlal t1'08te4 by heat
eX]Mt1"l!lellt dfUd.pe4 prbael'11y to
t1"G8. _ _t.
em
lrT
~
ten
tbe
t ......Ul. .tloll plt')oe88.
Aooard1n13 to Kostoff (1!g) t tbe exper1laeat8 with tempeJ'tltuN led
Hebel'la:adt (el) to bel1ne tllat HOl'OAO:NO.S t1l'OIl dyiJl« GeUs 1n tM
nelpborllood or the au _ned to ettElate 1ta partbeaose_tl0
d WYel.GplIIB at.
HoweYor. Owttatsaoll (10) rejeoted the 14e. or
~ st1lMlua
8
hOm dying "11a 111 010.. prox1m1 ty 110
1'.
lfeOhanltu!tl atlmw.atlOll ot _1la _4 IN..lel ta th41 owl.. ldtbout
tenlUzattoa waa att;eapte4 by DaY18 (8) 18 u etton to laduoe parthe-
.ogeae"I..
SOIa
aoo
on!'!.. ot
~a!!!!tra
1....l'Cld.aaa
_1'"8
}')l"loltec1 aa4
a1aaJw4 w1 thou. tau......
Cllem10al tftatJleata to lnduoe pl.lMheaop..ala
Van overlM8t, CoRk11n. end Ble• •l •• (4').
-'1'0
ett.pt.ed by
A lIOIl-Yla'bl. paeud~
ooulaU D8 ot ..._Al hua4red eell. cterlYfJ4 froft tIl8 1Jmer 1.,..r ot ••
1JttellJ1:l*lt
we PI'Octu.eed '7
o't'8J'1. . ot
D.!V~
the lnjeotlon
ot _phthal_. . .tl. .14 law
nl'8mOJllwa. SMeral other sub8t . . . . lnjeoW 111 com-
'bia.ti_ .UIl awWt taUed to lDl'lu. . the tormatloJl ot nabl. 8Mb.
the pel" cent
ttt haplolda bl
$h.. pro",.l.. • • t .....e.eli 1I'lftl'ke41,..
By per-
tU tt1J\s: 3. 6, tt."\d 9 dqa to 1 • • bet,.,... the openlag 01' tlae t1nt floftt.
Acoord1D! to Sltith (41) t SWl. (UlllJU'l1sU4) had aot t.nereaae4 tlMt
frequency or haploid. ta mat_ ,,. delay1Jlg polU.ae.tlon.
J'ol"tuno (16) touad that 'the mqueno,. ot the
III _1_ was aot lJlfb'eflae4 "
OGCa1l"1'e1lCfl
01' haplol4a
de18184 poll1Mt1c:ms tor perl04. of elpt
or
:rr.-
CIleee (4,5) o,.tlled atmtlar Rld.nee that dslaJ1't4 'POllination
ltsel1' 1. DOt a autf'1ot.at .xplarultice to. the wide Yf'.rtetlou 01'
Qwm07 01' l:atq)101dy record... 111 _1_.
h
tNqueoy of lIOIloplold d..... loJlll'l&nt ill . t H _S found
l»T Ohaae
(',5) to ba 1"&e11 ••peden u,on the __ atook MU!! 'POlls. PU'ellt wae4.
ae"'aln .toolea ....n uaed ae aeed pereats teade4 to produce .1&uttoantl,.
hlper t1''8q\llJ.101ea ot JIIODOplo148 thAn other stooks, lrreapootl.,.e of the
pollen peren' used, end aUd.l8,rly oertel» oOOD W'lan used
1'18
poll. .
pereat. tended to produoe higher frequenoles of moaoplotde 'ho othera,
lr1'8apeotl.... ot tbe me_ pareata 1nvol....a.
ot aollOplo148 olrtalud In the
pro~.,.
ot ay part! ouls.r oroa.
ot
to be 1Jltlueaoe4 by both parente in spite
ptI1'e.1It
'!'hU8. thft aotu@l Incla..c.
tl:le tact tbat the pollea
contri'buted nm. ot Ita sene. to tbe mmsoplold plant.
derl....d troa IIOnoplold.
DOgeDSt!O
Wft :round
\m. toUD4
Dlplol48
to be partioula.rly t8vore1tle PanAe-
stoot••
1'he 8l"tlttelal product'ion or baplo14a 1t1 X-rfty tftPtaent ot tho
female org_. or pollen cella hna beeJ1 praottoed
reth~r
enea1ftl,..
but the 'Haul'S. obtatnM 'by tbp Yftr10us \1IOrtce1"8 do Dot appear to be
unitorm nor conoluaSye.
The 1"1 rat haploldlJ deri...ed b"!, ;(-"1'
treame' or pollea mitre 01»-
taiJMKl by sta410r ('2) In corn (~!!!I!.).
Kata7fmUl (Sf) looated ! Ilnpl01d.
ruaoua m
lndlvldunla (5.16 per
cent) pl'04uoed by spike. ot TrltlOU1R 1ItOJlOoocoum ·[-ra,ed aurial meloal ••
•
0
_\1)"
POlleD WU lrra41aHd and applied
IlOH haploid.
(1'.158 per 08.)
_1"8
were poll1D.ate« wIth aormeJ. pollen.
der1Yed.
DO
to uc'Z'll8l plst11a _ y
..'D.
b."radiated pinUa
haplOid. were obtel.e«.
la~
OODolwted that the apeftl au.olei f'l!cm In'841atttd pollen lIOthel" ..Us
ItO_thee degaR.teet an.. s._riag the OYUl., wtthout tu.l _ _ th the
eM. but thetr 9""80e atlaulnted 'the
en tQ
d ....olop pertbetlo• •'toal17.
xoatott (29) Ifttf.)l'prttted Katayama-. obeel'ffttlo1'l8 ae deNdt. proot
tbat 1'0111. .t10u 'fd 1h X-ra,ed pollen areatl,. laOl'&e8.. the
haploids 'but .8.1.HII1Id tlUl' tbe treq.ueae1 would
specie. wte4.
'I'8.ry
t'JIequeJlO7
wi th the .train
of
ot the
10
Acool'dlac ~ lol"tWm (16) Yete1ka ea4 Va.l1., .... ,4'1) aJ.80 studted
-
... .ff"t8 Of Xo-l"9y trea __ ' of poll•• OD haploid trequaao7 1a 8 't.ri-
t1!\B spe01 8S an4 ",afte'tl...
(is.,
o~
0".1" a I_
Aa
~,l!ooe
a~Jdo
t1:t8 11810
0"'a18
ee.>.
pel"
OH
They t'r.IlUl4 I 11ftp1014. . . . . S5 8N4.
a I._ mate
N!,11&O!!!.
I!Ul4
the
ver. N.t!;I.
"'»1014 exb.I.blttag all the ace••lye Oha:noters of
pal'ftt • •
o".1aed b7CerulliaoYa (19)
flow" t1"e8.1Ie4 with
!,eo~!!!
"'ft.
X-l"fl)'II
trom the pro_t•• of
_4 JI0111JMne4 1'4'h 1101"II8l.
pollea.
:Pollol'ltBl X-ray t:ten_at of po11_. IyQ.O't' (M) 100eted tour
51!!!'!!!! !"!!!toe lulploldth
0_
",art.",
-P!I!'
The variety \!!!ll111 ..... oroue4 Wltll ~.
1t. pollen subjeOte4 to a dcae of 1',000
haploid __I l ' pl._t.
(1_ .!.....
o\b.ar. "... obtataed . . . . 11 plat.
!lOt
1»0,.8.. tile
X-ray
plOduce4
trequea• ., of 5., pel" oeat).
(1_ ,S,..
to11ow1ag tbe reciprocal OtteN with the
alld.th (41) touad that
1" ••
treq_.,.
a
~1'!
t".. ___ 'ot
~
of 14 pe oet)
po1l.ea _bj• •4 to
'trble moaocoooum
It
diet
fNQ.ueJ'lO,. of hllJ,tlo14y UDd.r tbe ooadlt10•• or hi.
ezporS.JJ.JI\.
llapl.o1da heave Mal o"1"t'ed to ooour In n1attoa 111 th
in yer1ou••
pect...
0_ of the
_l'8 ext...... atucU•• of
po1"'~
twi.
~.
. . OOD4ue'te4 by Barlaa4 (!2) \1.1118 Sea XalaD.4 eat! Ul)l.a4 ' ' ' " of
ao••m. ~ x .2.. bel"b!A....
QIJOIlC _'1'...1 thoWlAlld ....s. of tbe Sea
oottQll ad 'th. hJ'ltrl4
t~
DM4.
He toua4 10
leland etratn
V. 13tS ill -hioh OM of the _brJo8 . . uwe11y moh saall8l" ,him tb.
otlllfu".
?rom the•• he . _ able to rd . . 14 haplo-diplold pairs
aeturlt7_
He elso obtaiaa4 a hftplo-41plo1d \mll f'l'I::lJ1
(,i. hiJ'8Utwa) and
ob.~
\~
to
t1plea4
ootta
cue. of \Wia .ib2'708 til strel. of m1:D4
11
g,.
b1r~_...
g.
'Oe.rlJft4go flDOOnl'J'. at leest one ot
diploid twin p81r.
n.
~eh
• • a haplo-
In:'Oneloaally 00110100«1 tb.at Dot only
baa18 «daW that ooatro11_ \ke trequeaoy
dlt'teftmt ftftiu. 'but tha, the deeree ot
8.
&e_t10
ot oeourreaoe of haploid. 1ll
t8"111 '7 ot
tbe lIaplold.
waa
a180 a twaoUoa or p_Uo 801UI'I'u"0••
Cooper (') atucl1e4 the laeSA_o. aM relattft
4.vela_at ot twta
ea'blrJ08. oat cUplotA _4 oa. haploid, 18 ...... ape01•• ot t i l l . at
esr17 staaea of
L!113!! ~.
-~.et..
1.-
u4 1:,- l!U!tlo,!!!.
'ftl.e . .Vlua lI],'Mto1e. 1nolu4ed tU tollowl. . :
"e\'t. 1:,. ear:1t!ft. .L..
418pg!.
!,;.- Re!!B. 1.:.- if!B!2B!.
at obaCl1"ftt4 lOa 1_ _... Of' dnelopba. J:ulpl0-41 S)lot4
twta pairs from 9,500 oTUle. atu41e4.
Que
llaplo-t:riplold twill pair was
8ml!!! I!!tlsga,a
also oltta1ae4 1J1 U. f'l"om the oro••,
:x;
1_
t!l'b~OUL
Aa a :relNlt of hi. stud,., Cooper QOD.oluded that tH -b17C a'l"181as tro.
\he ..,..rg14 4nelepa sloWl1 alld uaual17 4tatategrato••, _
eerly stap
of 4."..10,..'.
ft.
Ia DaWl"'ft•• f~att.e.
of atoh
Ble.alee.
a hapl0-41plot4 pelr_
'I haploid. tJ'OIl Marl,. 200
Bct8'Nr.
301!E!
Ila4 A..,.."., (40) tood "
twin _na, . -
Cil o'btaUed
Obr18tenaaa aD4 leator4
'1!4. a..a:liap of qaRIlOUll 9!!1! var. 'br1A
Sla11v twS.a pai,N he. . be_ toWld I. the tollowtac .,..tee:
!l!!Ml\! x.!.
48), Sec!!!
.p.
)!~ (1.amm,
._1"
(Jro1"4_klold. 3'7), 8Ild
Iaatanoea 18. wl'atoh both
also 'bHa reported.
!ede 4;t.tlt\9!
1f"1 I!Z!. (ChNe I ..) .
SO),
ot'
~.
_Wi. pair
1W1n ....11.,... 'both of' *loll
1JV8
haplolds ll....
_1'9 haploids, were
o'bt81ae4 l»y f6,z1lll (:sa) frena the auwteVCl)'Jl.ol4 DutZl,.et ot _lu IIQIlOpl.ot.4 twtu was cUaoo__4 _4
(~to1'P1
~
l\omel"8~.
...
Fora
!bue It ..,. ___ en that f'oult type- of' 'Wia pa:1" bave be_
dou'ble .»1014, hapl0-41p101d.
_plo-tr1~o14,
A
(15).
~J
aad 4ou'ble 41pl014.
11
!l\u41... have __ OGDdnoted 011 the orta1a ot polJUSbr)'OJl7.
by Cooper (,) has preyloual,. be _ _ tic_d.
pol,embJ'7Oll7 1A
AtE! rMOI!.
(t') beU....d tAet
After _
1Ih1 ob " prot- -
-brJOa _,.
81"1 ••
troa
taw.tip'toD o.t
-,. epold.X1••
a 41p1014
Tbe work
Woo~h
toa.4 tollowJ.ac
G• •
0lI18.1oa of the ftduotloa tiV1aiOD 8D4 by ....11e1" bu44Sac. or develop...
Mat of . . .'UpoCIala. "'1"8148. or tbe eatloapt1'!h
..Sala. (36) aoulu.de4 'that alltlple __)lOtI ot
ebro._ ... __
art.. hrom:
rodueed
~
aao 08il'2"1tDC
(1) t.lle a,. sad • .,...,.,,4 Of . . ~
,lie
. r ot the au appe.ra.a deftlo.,1Ds
~
pa~tl..u7
" " . .• ..-,
_ _. -
or
o.f t.h. othel1 (e) two ~
tile n1slatt•• • tten
a.04I 1a wtdoh
~
ot twtl1lut10a
. . . (or the e . ot . .
aad the Qlm"114 01' the ~lt.eJ') ~ the paetopllJ\10 ~~
~r had
\
't
Mea terUUae4J (3)
nad a 81hrs14
tu
--I'JO . . . that.
. . . (or tbe 8M 8illd ..
~c14)
uateftUIMd ..... or troll an . .
&rOM _
ot
.omaUo aposporn (4) hoII
"1"70 . . . 4ewlopo4 \hrouah _ t l 0
&p01JP017. OM to1"ll'l1lll a 41:p).o1cl ...,., . . _. other a t.traploid
aomeUo doubllD8 of ,be 411141_ .,.,_, (I) two
. . aa4 • """1'814 \aat
8.1'0"
au oel.la or
lJ1
tI'OII aa
'b7 _t.l0 apo8JOl7 OM ot ,*1oJl . . t ....
tll1aed. th8 ot'bor ......lop12lC partheaoceeUoell.7, a4 (6) hoa the
tel"tl11M4 etc i.rlnd troa a
reauoed aaoJOQO_
aposporoue -brJO aao.
nwu. ..
oattnc _
'.l'M..
u4 tl'OIl . . . . ot ..
are of 1a1Ie'NR aa po••lbl,. 1a41-
It'.tOIle of' orla1a or abel- baplot, ....)'08.
~t4.
are :t'JtequeaU,. nerile a.ue
w
\he
obJIo. . . . . era4 the reeultaat tatlue of! .,...1••
tt1 1s WlUal17 quite :td.p
Blenal. . ,
00 per
.!l• .!l..
l)~
ana_ or hoIJoloaou
fte 4epM
bas __ toua4 to ft1"1
wI_
~
nerll-
the apeet•••
(1) toua<! tb&t haploid ~~ .~UIl plea_ wre
oemt polleD ste1'11e. A baplold
-.at
pl_' alt._bled
by oroH1ltc
18
'rrl tieua !g!!p!ct!!
-, 1"!,I11oR!- gzl1.D4rtca was reponed to be
W1!21d~~1
99.8 per eot eelt 81-&1"110 by QftlU aa4 Au. (1").
__
'Y Harlan4 (82),
~n.4
3pOllttllMO'Wl
eomatio
parM aa4 tbe produ.otion
'be. .
ftpone4.
Bee __
1WU.. • .....uIMln
Ko.~tt
(29). fJ.'Ad
S1m11o.y dau have
0_1.\".
aJ'01U08l_
40Gb1"'. of ..otor. ot the tlD1"8l
or
procea1- toll.ow:l.q ••1t1lag 116.
41plot4
of tb18
'~t
LlD4atrom a.a4 }tooa UD..)
IhlltiDa haploid §Ol9H lZPI!UlH pleta.
ltoU1aga'bead (23) • • qui te WGOM.tul la .1riaS a haploid Crul._
plant tor tM . . .
~n.
Spoa'taneoue .:>_tte d1ploldlzaUOll 1& moaoplold alae pluta
ocoufl'W4
~ueatl.'1
-.ouCh to pem1t m... Us) to o'btfdrt. P'1"Ogal•• f'roa
prod..
10 per Oftt 01' uatHaW IIIDllOplo148 tollo1t'S.Da ..It-ranUla''1o. .
31JlOa 1t 1. hlPl1' be1ra'ble \It,at IIODOplo14a
gem_,
0010hl01ne "NOlm1qu.. lw'Wt .... emp1o,.ct. 11'1 .. attellpt to
illoreue the tN<'lu• .., of
h.
..1M pro-
~'10
4ou'bltq Of
~
OllrollOao.
~.
rm... of the 4 ...10,....t. _tic.... ea4 _thode or tM ooloh101ae
teo1mlque 18 DOt _carted •• __ ~ slll. . thia ftS_t baa W __
rata.:r wtdel, • •4
aWN ot the
tor
tMe pvpo...
au_je.t bu
thlt litera. . .
u.
,"8ft
An ace11_' MYl... of tAa 11tar-
pu\Jl.lalle4 'b7 DeBe. (10) _4 a llattq of
bee OOSJl1e4 'b111pU (ll).
Cu. (3) lui_ted that tbe laJeOUCII of' 0.06 pel' oeu oo1oldo1ae
limA 10 per _ , &l,oerl» Sa .. aqueous SOlutioR lato iM aouteJ.lu 804.
ot
j'O\18C
"0,1014 - '. .
_.uta,.
iDOft#.alag the nte or 4ou'11. u4
CI:aJtoa)aome
dou~tDl
ot 41pl01d -.d ehtae1'8l
i:.!!I!SlorttV· t I-
triPlex,
pirOIItslq
re4-taa
toM
1.
TIN..,
fl.
a M'lb.od tor
aepoee of ..It.... 1'J.11',..
• • be_ towd to oeltlr qul_
NO'- of .plold8 o'f . . . speeler..
Qat
appe~
~u_:U,.
Sa , .
1..cpott (29) lined tba per
1'00'. or the haplotd ple' of .100t1_
mtuwe. I..
tab,\!oum. end
CnB. !!il11arl.
14.
q
M1Jl& I per oeat, U per out, M.4 per cot. 56.6 pno cent. m1d 63.6
per oat. reepeetlwl,.. '.fberetoft, It la &p.r••"
aatiM the
~ue_,.
el"l"O!'8 ta .arU-
'l'u~t
of tbe OO«tnWaoe Of baplolda . , . be
eb.rOmOeome fIOUIl'ta of 1"00\ tip 08118
8.1"8
1B~
.ere
WJe4 1'07 ])081'1... t4entlt10e.tl<m.
ot pu:"'tlft .ploU••
Tba nr10ua .,,41•• oa .p101d pl. _ _ -7 be auaa1"'lsed .. follows:
(1) haploid. !left bee. obstn"Yed to ooour In DfJll7 .,..01•• of -plant_
but WlUally ftt a ..., low trectU8aOJ';
tal
haploid pleats
II1p~_
aN
waull,. amellor _4 baft a
mo:M
ctellca:t.
tb. otJll'i)8ra'-le diploid })l"a,
(3) X-reya. kYb1"l41zatloD, u4 abaoraU,. low or high tel!aperat:u:N.
lu.\.,. be_
auc••W
.a ia. . .~ ,,.
"t_ of
the 8p,.8:r8, -
of haploids,
(4) . .__1081 atlD1lst1aa, on.em1eal 'reat._., sad delaJ8f!
po111_\1011
na....
appe8.I'8D08 Of
art be_
~UD4
to l'1Oftls•• the " , . of ...
_pl.oI4. 1a 1IlO8' wteaoea;
(5) haplo14a haft . n fbad t.o oeeur 111 pol,..s!'J'01l1e aee4a1 ....
(&) haploids are uual17 h1*I,.
8terile but some daubltac of ohro-
JIOaoaee 1• .,C Nproductlw _4 't'eaeta'Un tlsa_ ocours ad
tM.• -1 \Ie 1DoN• ..a tt,. oolehlo1ae trea'-."'.
11
.,..t_
oan"11ac uaWlUal 40mtaaat IBrbr geDG
'!'he polle. JI!ll"ftt.
. .looted tor _e 1. ttda .-<iT l'1ere . . 811_ lDbn4 11_. 38-11
(A Pu l
~) • •
browa 11_ (8 B PI B). aad a cagle oro•• 111\'t1.4, lJrowa
.toot.
x 'J,lU1"Ple (A B Pl R).
1h8
_t~mal
TartH orlclL
uaed aa IlOIlOplo1.4 aouroe ._aa1
10111" elite
1~
_1'.
or
liMa that ••• be8B d.ftelope4 b7
OOB'ND.tloaal bree41_ _ tb.ou aad fOUl" 1a'red UBe. 4eftlope4
1IG8O,1014 methOd _re utilized 1. _
1:17 . .
a''Mapt to 00J'0PB.ft the moaoplo14
t:L"eq,aeaelu 1. proa-f,•• of 1abNd 1laH 4....10pe4 by tM two _\bode.
T» tour
.~
lDwd 11....
u..s ._
88-11,
~m.
aa,
and '16.
1'ba Jalmtd 1 . . . . . .ftlOlMtd b7 i!1e IIIOII.OPlo14 _thod _re H'13,
snl,
mu.,
tbe latter
u4 U.s.
Two fI1acl.. aoaa ll1brlda IJlY01'f'1D& three
or
11Ma wre We WI84.
nn.",.
All opu,..po11Ulate4
:Jhoup's
~.
aad tM f'tJ'at 11114 . .ooa4
cycle QIlthnlft developed t'l"an it -, reourftJl' ae1 ..t:l.on
'Wre un4
811
_"'nal p8ftIlta to attd,.
t;he
upoa the ~u.u". of .nlOpl.o14 oeourftll104h
.tteot or
~.
delayed polliae\loa
'.!!U •• mete!"8fl1 .oure. Tar-
let •• were alae 'Wht4 to atudy tbe erreot or tu reovreat eeleot1tm
lIrOad1n.g _ _oct upon lI.01loplo1tl
~ed
mqueJl07.
••ed obtaiaed fltt,. Dr. S. S. Cboae. Iowe 3tete COllege • • •
also toate' dur1113 the COVM or . . ."""_
~het1e
'1'h1& materiel - . J'Alrly
x _; m1Dd dente. 1108\17 of St1tt Stalk s,ut'Utl0 origin. x VI
mixed tbJtee a. tour _y
01'08"8,
allot !3tttt SUJ.k S,.'t_tle or1pn.
x VI _4 (BIl4 x 114) x Q.-Aa.
'ftle ataa4aN utb.od
~
IOUro. x D'lU"ur
or
oODUolled Ol'O_polUaa\1oa • • lIM4 1.
O~."8_
Del.,.. poll1aat:l.ou. aeslgaa'ted
q
16
laM in the S\UBItU"1zod. reaut"',
aft.. the t1p. ot
4_10."04.
The..
befirilag kel"Dela
00' . _
81"8 . _
,hit enrwse4 port!
"1'8 ftre
OIl
\)7 pollJ.aBtblc the aeed
of the a1lia
118ft
~t
oomplete17
ch. .ed a' tbe t1_ of harTeat and aa),
tn a po at ti011 ot)aer tho on tbe proxiDael Mlt ot "-
dieoarded.
TAe mtU"ke<1 .l'M1a WN PJ'III1nflted Sa .. ali.ted prmUaaWr at
seO
C. aad . . . . .4 ea_
...utaa
OIl the
4.,. tor the flPPH1"It.Me ot the
_ral'
pMaotJPe
mota. All aeedlt_.. oxh1lt1\f.llg the l!'iItHer eh8.S"ftOt...
lat18 were dta061'4ed at o.n ez8ldJIMtioa.
Cl..aitl~iulUO." OOJl)leted.
with lfl()St of tho _terial, C_1_ .eYe 4&,. aft.r tbe material Old"tlt.e CON1aat01" •
.At
,lie t1M ot \he tlrat ll.aa1t1oatloa, a Olileok • • t4eo ...so tor
Jc:8'Atia N8Ul t1q f'JtoIa ..Old_tal selt.. 01" cl"Oaa-pollillattou.
pollan _tOOk.
(Al AI
1na
As
mo.
_ _ the
tNq,\tut17 ueed OAl"rla4 the . . . . tel' P\tI"Ple eleuroae
C it 1. Pr) as _11 ae _. )lot .rker ae••• , terael. 1\Ot abo...
tho aletarODe -rkfto pheaotype were diacmr4e4.
Root t11'8 tfYr eytolog1oal stud}' lieN 'klcea tro. . . .h :patetlft ..,..
1'1014 8.-11_ atwr -..
.~
proeeas . . OOIIplete4 _4 Jc:111ecl
J'orme.l Acetic A1ot>hol or 3:1 Aloobo1-A.c:tet10 ao14 eolutioa.
oout.
_1"8 1ISd•
s.a
Obl'OllOeoae
• tiar tbe Jt)ot tips ..,.. aqWlllbedend Iftalae4 wl:th
aeeto-oe.rmtna or ,l"Oplno-earmlM. 0010hl01ae tNe"at • • oe.l"1'1e4 out
beton the ...n.ll!(C8 _ " potted by lajoo'Uoa SAto tlw sou.U.. aode ot
_
fA.q't»Ou. IJOluUOIl of co1oh1c1ae pl_ 10 par . ., clJ'f,teJ'ine.
tioM of 0.00. 0.10, .... 0.16 lMtl" oent eolohi.as
_1"8 Wte4.
'l'lae put.t1..., IDJlOploici ...utD&8 -.re tl"llMf'e1'N4 W . .
after
,be
","8"" tor morpbolol1oal oblMnet1oa.
fl.,..' aeedl1D(lC leat ••
~
Coaoeatl'8-
~
'lb.. 1_ph aDd 1d.dtll ot
etter ' b leat . . tully extailed.
l'
1'tl& upper surteoe ot .18 leat _. t\n"tlar ue4 tor _.1'08ootdO d.eter-
1I1_'to. ot tlle .Ize ead l'lUJIII)er ot stoata.
b7
t~
cOtmta
_ft
ue ot
lID
of
oo4M1lu at.".._r at " __tn'-toa ot 440%.
!Itl4e 'by
to Ole 41%'8etlOll
sto.'Al 81ae _. _aeuN4
ot
4.~g
Ye1...ttoa
stoia_
the JIQ1I'MI.r of noata 11l a J"O'\f parallel
of
the mtaroaoopto field .'t 4ftOX.
'* leat
80r0. .
'be axt_m
d1~
!'.:!!sueag !! !!1!"!l014,
Noaoplol4 plana _1"8 1'01olD4
tn all of the iIIl8Mrl.a1 Hekel.
",,0
".ftle .... tMf:l\l8J&07 of .,.,ploldJ', . .
al't.ow.Il tn ......1. 1, . . 0.81 per
~4
an4 1bU'..,. oro..... all of s'Urt
tal'lftld 11._ ct....lope4
tour ".....4
pl." .aW. !M !!d.ze4 th1'M
~lk
u.. 4.....lope4 by 1088-'_
._nftl
Wilted tdJen ua4
m.oaoplolda pel' thouaaad
a,&the'tl0 crista. 'ho of tJae
1t7 _ • .aoplG14 __Oct,
)1&, 8a'ft tJoeql8lot•• ooui4eftbl7
all
_-'1" at :tela'lI. .I,. low treque:ao1ea
8.8
aelt1ac,
m,
S-U. 1ft2. ea4
""".1' thu tbe _ 0 tr.queaq
tor
!he treq,aeDe7 of 1.43
.1JM4 noOk..
pl_'. ot
H?3 ad SUO, aa4 tU
1Iba ~ 0 t thllt 01'0" tr13 Z
(1Sntc x Purple) 18 .lpitt __, at tlle ODII pel" . ., leYel U Maau1'8C!
"" tile Chi-square teat tor Faa-a. of ftt to thll __ trequeM,. fo'J'
all matetsl wned.
plaD_ of the
'l'be t'fttqu.o,. of 1.13 IID_plold_ per thoua4
pro.-,. of ...
01"0" _'fOl'f'1lag
the 1Id.DC1
t .... _4 tour
.,. Cl!'08.... all of 3\ltt Stalk s,atMUo 0'1'1• • x V 1. alp1tloaat
at tbe t1w ,.. . . t leYel ..
~
...,. the _e tat.
The data ..... f'urthltr elaeAdfte4 tato tour fP."OUpa .eord!. . 1»
01181- of the aMC1 pereata.
deftloped '" loq-t. .
pl014 IIft1»4 u4
fte tour SJ'OUpa _1'11:
-lttaa,
(1) ta'bnd 11...
(2) labJ184 11._ developed by the
01'0." laYol'¥1.a& tM_ lu.••
JIDIl.O-
($) material orista-
a'lac hoa S$itt sw.k s,a"t1e, a.d (4) _teriel o:rJ.C1utlq f'l"aI.
'u
opa-JOll1Dated
Tart"'"
pt'OduMd l'IQAoplolcl•• ,
11
Shoup'. ~..
•
lae' POUt> -..tl0M4
tNqueaoy eonatcleftbly 101llU' t2um the . _
trequ.or tor ell _'-1'144 te_te4. Tla:I.. dSf"tel'ttIlGe 1. alp1ttoaat at
the oae pal" oem' 1 ••01 or ];1l"Obald.llt'J .a .atRD."ed .". the
teat tor flOOd.... of ftt.
Oht-aqtl8ft
19
Table
a preaeat.
va11.."7 _
the tn_BOY dB.". tor the op_ -])01liMte« ltl'u8
the tlrn .4 MOOD4 GJOl. 8,Y'JIthetto yvletl_ 4eYelope4
from It 1Q' reenrl'lltat
a.l"tto.
p'ftUledlU'O..
'fta.. aozaoplo148 are ol&.81-
tied .a Ol'1l!MtiDg III pro. .t •• NRl t1Dc :from "1'1,. or late poUlaatiD'"
'ftt.e 411ft_.... in trequ.etee of moaoplo14,.
8ft
aot etPdt'1oat
at the ti" pC' . . lewl of p,J.'OMldllt,..
Ii.
_urial, ..at
ponioa of tise
pa1"88t. wa. 41aea'l'4e4 _toft
laok of a::,prufl1oa
ot
.1'
the __
~
IllYOb'1. 3I-U .e _. polle.
lJ1Itdy waa
CJOmJJleW beee.u.8e of the
}W0P0l"tlo. of tM
~
tertnl0 of'
at qJ' U. betoft 8I1'Il1aatlOC
~ll
A relatlye171a'f'8e
Olanot.riRie.
dtd aot eXhIbit . .
~
-pluale _ . . . .
MY_ aa,. of
or aner
ae1."ld.ttoa. Both ot the ....ft'Al paNllte 'fbI' ",bleb. 4ate. Oil _. pzopale. . . . . 1"fJIooJlle4, Talale 3, are ,.llew dtmt
tI1ll'61...
Both
dlttered a118tflo.tl,. (P 1••• ta•• 0.01) In tl» • •ber of'
.ro....
})~
DOt
e'dl1'blttaa tM liftU'1)le plUlUle . .raner ., .,. t1_. !tit t:toeq...,- of
!IODOJ101d7 -a DOt dawal_ tor tMs. uoe . . .
-brJo.
'1'W1a . .r,oa - "
nneea twID
110' to'UIIId too be reliable
were aot.ed 4uriDe ,_
lDd1 vld.1lalA ot the twi. pall"
the...
_ftI pJ'OftD to
8O\1I'.a of __plot«a.
1iOUl"88
or W.
a"7. .th
'be 41plott la tolU'Uaa of
The :reM1aSaa twtD --l'JU . . of 1m. 40'*1. monoploid
~.
-
~,.
of'
: -..1"
?ella PArfIlu:
:
t
I1a!ber
of
-
--
._.
:
:
.
,.1"
l'Nqu-07
---~-----
'lotala
~-
--_._.
'fai us
•
:
:
·:
·
,
·
·
·
.. _-,.
-
-
".---~
_._ ..---
a
...
:
,
-~
,
5J pol.,:
__.- ---.- --
at the
_.. -..----
...
·
•• Slpltlo.' Chi-square value lilt the 1" poiat.
, Slgn'1ftout Ohl-8<lulU"e
••
In''
__________.
"
· ,..
lU,881.
:
:
·
·
18
•
"
""
"
"
, .. . .
;
I
:
_.,----_
,
,
0.82
1."
0."
____t._________~ ______~ ___~___Prc~... ~~~s.~._.__ ...1. ___~• •
:
:
:
:
..9,100
0 ••
3
Earll SlDtbetl0
:
»
•
•
•
'U2*! d.""., mostly of stitt :
s
:
:
...11,5«)
10
O.ao
St8.1k SJDtbet14 01'181_
:
V
•
:
:
!id.xe4 three &114 fbur -7
:
ertHta.Il, all of SUr.r
:
••
Y
30,000
tJf1
S1&lk Sptlw\l. orlaiD
:
:
1.13*
t
•
1,'100
1
•
..
:
~
•
mu.x
•
t
:
:
s
; . " . x Purple
•.
0.40
10,000
3houi'" lrU.c O.P.
•
••
. Iro1iIa x PUlIple
10,000
O.SO
Shoup'. xruc s,n. I
:5
J
.
~:It Ptlrpl.e
:
10,000
0.40
Shoup'. 1\J."Ug Sya. II
:
.
••
ft,OOO
1
0.10
Shoup'. K"I S7D. II
•
••
:
I
:
•• ~ x Purple :
:
3,000
1.33
m
s
.•
N-l1
t
BJoowa x PQrple ••
3
1.fOO
.•
.
Bro_
3,000
:
1• •
:
:
"
aro_
3,000
:
J
1.SS
.6
•"
•
:
I
:
.-•
810_
x
Purple
12
2.4.3*·
4,H6
s
1
Hf3
BTo_
x
Purple
.,018
I
3
11111
O.'S
s
"•
...
an4
1,800
0
0.00
: Brotm x ~rple :
".
.•
1.31
:
3,050
..
HI.
t
Browax~
•
•
•
•.
:
"
Bro_
•"•
•..
1
0.33
S,OCO
:
RSe x H!l!
:
•
0
0.00
:
:
3,000
De x HIl4
BftJc
•
•
.. ____. _____ _..s_._
_~~
::
of' the tr.queae,. of ooour. . . ot lIOIloploida III tbe 4ent a'tooks tenecl.
Seed Pareatfl
TIlble 1.
8
21
:::
.
§
•
•
!
11
1 ~I
l.:i
If.
~
I
I..
~
•• •• ••
I
'Il~I'
"·-tt
:,11
1
""~k
0
..
I.~ I
I
k~l;
j
A l.3
•
§
•
e'l
i...
I
•
l:l.
0
~
IJ
§
•
M
~t
"
q
§
•
~•
~
•
8
•
o.t
.t
~
....... ....... .......
• .... ....c.,
.. .. . ....... .. .. ..
••
III!
".: J
I
~
.:t.1 t r... E.
J• •
11£i I
........
11~1
j.'4."
0
.. - ...
§
... _.. ........ .. ..... .. ....
M
AM,!
•
Oil
iI--r·'
. J~. ,.
!I~
g
•
M
.
....
.......
h
'i!
I
....
.. .
.. .. .. .. ..
0
....... .........
....'"
«t
.....
I
I
1
Table....L..
A teet of the expreea10a of tlle 11-11 me.rkor otaraotel"1.t10
whim _1'JI)8 we:re ol._1t1ed thz'OUp the ),18r10arp ad aft. .
. . . . 4ap of ~101l.
: . 1btal
seed PareJlt. :
Ii ... _
:
••
Pro..,- :
.:
t
:
:
Orl6-l
x
;
:
~
.:•
Kn&g8,..I
J
r
~
:
:
,..
t
I
3,018
:
:
:
:
•
=sa
($I
I
:
.After
.
g , :Oe
Per
Per at
:
:
:
81.9
:
:
f!1: 1
I
.,.0
•.
\0
1,1'.
!!Jir !J!2E!!t
·•
•
51.1
I
:
..4
:
;
:
:
:
:
s
:
•
1'0 ap%'M8ioa
Pel'
o.t
6.1
t, •
s
t
••
10••
Seedlbc IIOrtallt, rat.. WN 'iulte
atW!1.d.
hi.
in all of the -te1:1.al
Table" pre.eat. 'ba ISOl'kllt1' rate. toJi ,.tatl•• 1DOJloploU
Stitt 8'klk s,athet18 .....11Ap wi ttl 1184 wi . . .' ocloMot_ Vea_ _t.
~.OIE
1IO'ft4
eoaata _1'8 aot . . . oa th18 .terlal. _\11 \U pl_te _re
troa _. Cfttabo__
to • •
t1814. '.Pberetore.... oturo.MIOM
DUJJ1'Ier of the pleat• •".,... prior to tllla U_ ere
. . .a bet. . . the
~lty
DOt
lato-.
nt.tter-
"tIN 4_ to taataeats aN act 8:lplt1_t
at tbe tift per oct leftl ot p1'OkWlt,. u "8IUJ'04 by the 0bi-acpt8N
,.at ot
~II
of fit to the _ _ fr'oq-aor O"Nr all
L1kewl_. the 41tte'fttaoe lMttwMa the
_19
~altt7
Jlot laJe0W4 . . the tnjftoted 8l"O'lP
by the . . . Wft.
!be mea.
Vee"'_.
ret•• of' the pleta that
II JIOt
at_tI8..'
.8 __and
AllprobAb111 ty vel._ obtala04 wre 0.1 or lergel'.
~ltJ'
rates ot ..,pl.o14 • ..altA,. from t.be el.eYea
..." pcmtna 11.t84 1a 1!eble 3 te tbl tIll"M levels 01" co.....'ratlO1l
01" ooloblot_ u.d ere BOt ataa1t18aot at tht» ttv. per .at 1.....1 ot
) __b111'\7.
UOrtallty rateD ot PUtative IIlOIIOplot4 _edllBfP1 f'1"oa Il1xea
tl:l:ree a.ra4 fOur Wff'I oro",., all of stitt Ste.lk S".thetl0
oriclB x V pareataaa.
Talale....!...
:
I
Tn~
HoM
I
I
t
y
: O.~
OolOb1o:lae
l11jeote4 Into
I
,,"tellar BOde
~
-
.
•
O.~
··••
·
:•
·•
lJital
*
:
• O.lo.c t
: 0 •.1_ :
'1\)tala
·:•
I!~!bva
0
s
:
~ot
.
139
i..
•
••
it
·;•
00
10
·
•
••
•<i!
10
191
••
•
iiii&fi:
o t a....sUq8
m1!. aYl.tlvl!I
11
:
33.1
30.0
10.0
f
6
t
..
3S
I
:
I
.~
·
•
,_~
••
••
51
10.0
'I"
. ..
·
·
II Goloh!o1_ at the ooneeatratiou laclie.ted arI'l 10 per oeld
til
"
,
&l.,..nne 1a _
I" t
~
8Olut10D.
~
101
• I_
,.
11.1
I
:
•
1
1
:
: Uorte1ttJ'
•• ~ o..t:. ~,~
:
• ... #I
'l'a:.le
..J...
ot .,l1oplo14. :trom . , . 4eat stoOke f'Ollo1llD.8
laJo.tl.oll ot eel.hiela. nt tbe la41cl'1ted oo. . . . .'loaa
IDto tbe ~la1" DOd.. ot tbe J'OUIl« seed11..._
~al.l''1
SM4 Pal'8l1 t
•«
:
·
·•..
..
..
s
:
•
••
:
••
O.~
,
... ....
:
s
O.l~
··
• '10. t ){o.
lfo. ••
10.
: Trea\!,C' PJ!4, . ..:~.t~: Dl- oc1
Sl'le\\-p '" Kl'ug 0 .. P.
Sb.olIp'" XJou.s I
3houp'. Kru.g II
m
..
t
:
:
I
"
1
I
:
••
2
t
0
.
·
1
•.
1
0
0
••
•
:
I
··••• ,. ·
.1
1f6
:
H'13
:
:
'8!l1
rm.
I
%
·••
H!la
·
,
:
"
'1'oMl..
Mol'tnll
:
t,.
.•
..•
(Pel"ONlt)
~
iii
,
•
I
·•.
6
;
2
•
1
ao
0
1
••
...
.
•
:
I
-
•"
·
0
.•
0
··•
B
25.0
.. ........
, , ..........
'
1
1
2
0
•
~
I
:
38-11
H2.
II
·••
2
*
...
t
1
:
..•
...
..
·•
•
.•
:
!
•a
1-
..
·•
1
l
t
•.
:
:
0
1
•.
•
••
:
:
·
··•
1
.
t
1
0
....
0
l
a
••
.·
1
%
0
·
··
·
•
1
~
·:•
·•••
1
:
!
1
·:
·•
·
....
~
:
,
'" ,
....
0
...
I
0
].
.•
0
:
.•
•
-
t
3
.,
"
:
.
81.3
~
1
J
:
11
i
0
1
•
t
0
a
:
•:
••
•.
1
:
:
..•
- · ..
:
"".8
'u
JfO.
Died
s't'ree:W·
,
....
• •• t
%
•
••
1
1
:
.
:
. 1 •
0
"
:
•
1
1
a
·.
No.
t
:
·
·•..• • 1. ·
•
M
.:
O.l~
, ,
1
.
I
1'.be -Jort ty 01' tbe lIIIOaoploid pleat-a ob\.1_d throua'hout th. couree
&1....
of thle atu4y _ " coll,Ple.ly 8tel"11e.
trom 1f'1'biD ,he
'but b
SolIe Jl,aate lla4 natbere extJ'ude4
DO • • •
were theN more 'llu!la t1 n . .
poll_ ooata1al_ autbu'a proclu0e4 OIl a plgt aa4 1. 1lO8' l . .te.aoea
where }JOlla • •
It . . troa
:J)~
ODe to ttl. .
_then. 'AeN . .
e _I'ked _de.ay tor the tIOnoplolda. MS_le. . of' their ortda. to be
pl"O~J'O_.
",. 4.,. em
~e
te1"tll1t,. at a &rOUt' 01' lI01t.OJlot4a
or ltn
or181Jl
Col.hiol. . 'trea'-t' .,:peU'lltd to tUft...
are preaellW ta ttalQ.. G.
the pere.-.. ot pl._t. produe1Dg self_ _ _ , but the 41ft. ._
. . . . tha ooleldClae 'treaW4
18 llOt .1p1tl. ., at the
croup AD4 .... plaana
t1ve;~
.a' 18..el
b7 the Ob1-aqun ••t.
f.fo.U18~
1.....,.. fit . ....nN.oa
01' .1011101_.
'ftle __ JIJIa'bft 01'
alUl..
!'eOel"f'1.ng DO
be-
trea""
01' pro_bl1t • • • .-ave4
tftlld . . ~ tor the 4tttelWa'
tlnel0pe4 per t-"11e pl. __ . .
Jd.p..r
t. tbe ...-laJa_ted poup __ Sa tbe PO. . 1Ib.10b _eetYe4 tajeotlou.
Aa
--race ot
1... t . . cme lEe1"llll1 01' elte4
...a },'J8J"
pleat . . ob-
tataed O'fft all . . . . . . . 1t111. fbI' ,be :tenlle plata , . __ JlUI'Il'ber
of . . . .
~
hom 0 to '1 per pla't.
---_.. --------:--------------_
..
·
_.
·•
•
I
I
:
•
. __
··••
.•
~"
""
MIs lIUIIlbel" of Draela
per te.nllAt
pl.-.,
\rea""
Me_ JlllUlber of brule
~r :plant
,
...
.,........
• .,
1/ Oolct4c1M
):1.
:
:
:
:
•
·•.
:
I
t
II
M:
..•
:
·
••
.
,.
•
••
·
'1.0 ..•
4.0 ,
•
.
...
•.. 11.8 :
•r
... :
t
..........
'
I.
:r.
:
1
Od :
:
!
• •
•
15.0 ••
··
0.0:
1.0
0.0 :
•
·
1.0 :
.•
·:•
t
·•
·
f
0.4 •
I
:
40.0:
1
:
I
50.0
.1·:•".. .
•
#
.,
·:
0.0
··
1.0
.1
1.0
••
at \lie ooaeeat",lou tadloa,,", . . 10 per 08U' sl7MrDe 1Il .. aqaeowa aoluUoa laj. . . . 18_ the . . .Ue l»4e.
An
8t~
• • .ad. to "eterld._ "1'1))aG10I1oa1 ebar&otera ot the
tira' .ee4llDl le..,.. that eou14
lie ue«l u
a reptd aethoCl ot d1n1a-
au1alWac ,.,-_ llOaoplold &J1d diploid ....uillP. '!!.be data obtelU4
tor
tu
_~R8
atu4le4 are pr....W
npre8 1,. I, 3 ••, . . 6. !he aawJ'l&lll.
ta t'a\)l . . '1.
e,
aa4 patena1
petre'" aeeI.1tap
_re JIOt lJa,1eot.ed With ooleb1ciaa .ere.. tbe :pN8ItD7
1'M
pa'N.~
~1:Al.
!be leaf
a1ifl tor
iJr'rolftDl the I:rU&
1».~ • •e
PI'O.,., .m
8O\IMU ••
",II. .....
...nail le. . .
apart; tbalt
pa~t.
an . ._tea 18 ffab1a ,.
aaa-1Jljene4 :p8l'at.. , . 1.......
w.t tk ftapeet to tMi.!' laPh
"",'el ..ota.
the 1. . . . of the
eros... lDY01""1I8 the tour all t. 1almtd
wn
the
lar... ad tutller
pro,.,..
11M_,
tIl_
'DIe.-.u of the
Gf the paNIltal plat. . . . .
_N .0_ or
1IIpoM4
'1014 ea4 41»1014, baa
aa4.J'l8I'I'OW!' ]a. . . . . . . . . .
1t~ _1'8 b1'O~
vee_at . .
...aliup of . . 01'0'"
p61"111ltlll
tH HtI4
tDJene4 ,..117,
la.8. ot tilI8 e&eIl1liap of the
tiNt
lajeote4.
to ..... of the :Pl"Ofl88Y l'IUt ... Saol..... to pobt out . .
upoa • • PrO...,.
ot tba
Wft
...ali. . 1&at' ellaftCters. __ rl!ltoN, .... DOt dtreetl,.
4itteraa08. _toll _re toad whtm tbe lajeet10a
shOrter
aa4 9 and 1Jl
'!'he data
OIl
'the
Table 8, and the 1Ja1Jn4
11nH dftelo}1e4 '07 . . 18OIlOPlO14 . ..-4. '1'allla 9. follow the tJlU4a
lllutJ'atect ...,. the Eftla . ..,rt81.
The tftq__., 41nrtllutlou ot 'tile ...allas 1eat ••zouto,.. of
''M
~
wkl.h 41"
DOt
.zhUd.t
1;he . . . . .1' ~wr1atl.a
),'tU'e4 la nlllN. 1, I, 3 • • , I.'Ild 5.
are
00. .
'!be _ _:plold 41nnbatlou are
baH4 upoa data boll -'l1"t7-ttve 8H41la&e aa4 the Atp1014 41atr11NttOD. .poIl 48. t:roa t1ellt,-toUl' aee41bap.
..,. aee4llDp o1."1g1.aaW
mill the uo ••• 11ated 1n Table.
'I, 8. _4 9.
'l'he lear
.tu41e4 were qnt U YfU'18ble 1a both populatlou.
ohaneter8
'l'hne . . l1ttle
4itter... 1n tlii8 rape of tba t_ \)tpe8 of proaeay aa4 tile lI04al
el.....
De
SlOt
_daly _paraW ill
all,.. or the
tin tiatrlbutiou.
panJl\
pro_,.
~
30.0
69.2
M.S
~
8.00
.49
1.98
-t--
4.96
6.50
~ a.u
.49
6.50
6'1.8 ,1'1 ••
30.0
:
.
:
·: ".1... 11.56
--
:
l
t
118.8
31.1
35.0
U.s ~ a.l'
19.a
.u
,
,
:
:
I
t
.:•
I
t
:
I
I
:
J
::
~
•
I
P",
t.
1.!JO
:
;
I
:
3..
i.
3.10
6.66
4..18
.16
.12
~ .so
1
t..
l
--10.0
15.'
U.S
If'
•
1
:
:
:
t
...----.~-".
.1t
.14
..... -
t
4.43
••ea ~
al.o
31.1
-
- -..
••
.1'
.M
.20
t.
t.
-- ..
: 1.80
: 1.10
:
1 &.33
3.44
.2'1
.16
.US
. -~ -
~
.1'1
.13
.PA
at <KG
1.80 I-_ .2f1
I.!? t .24
L4'
I.!?
:.S.l'
4.18
3.50
5.83
La~
2.8'1
. - ... - - - - - -
25.8
.'.8
t. 11.10
36.1 & 1.3'
26.9
11.10 :
1.48 :
:
:
t
::
:
t
::
:
n.w
~
..,.~U ..
a__l.
:ia
t.
... 1
t. I.":
t.
.00
3.fiO
1.80
.91
:It."
3.lI'1
34.6 ~ 1.48
U.s
'I!1.1
C
a.·'Zl.48
4a.0
81..
u
•" 4'1.8
: 34.6
I
:
-2-1.se ·
3.48
; S.oo
:
.'5 ::
.38
.'1tS
.19
4.43
5.68 t- .12
t.
~
.14
x Pu.rple)
3. '10
2.'5
ltrua II z arc_
~
.. -..
9.0 C1.00
15.4
11.2
:
:
:
¥ (Bftnfa
Krus It
.".10
.86:
.50:
11.5
10.5
t
t
:
:
:
,
~ Wid».J
x (Bro_ x Purpl.)
.89
.80
P.
•.
lCru& I x (B1'o_ x t\trple )
1'.0 ~ .2B , 4.00 ~ .14
13.1
.10: 5.18
.121
9.0
10.0
lSoll
12.1
I':rua o.
.t
Pl"ofJUa.7 t'8Hlft4 ooleh1Ol
. ._treataalt.
lbaoplotd~:
MIllteraal p ......t
pat6rD.al pareat
otp,101d Pro_7
Diploid pro• .,.
UDaoplold PIO(C8B7
:
Matel'DBl , ...at
Paternal plreDt
Wi':\\:
-T-~l"t ··---m-'fo--ot~-:-----.~l&- ~~u-s-
......
- - - - ,
1.... :
U
leat
6.2
45.8
•
• •5
"•
:
pro..,. ·
par_t
:
t
·
hterael plU"ftt
Diploid Pl"08811J'
MoDoJl,old
Mftte!'lllll
Diploid ,ro. .7
Monopl.old
:
•
!
:
llwolYiq the opea-polUaate4 Yfll"1e'Y, Il!'U&.. . . the tiNt _4 .00114 .,..1 •
4ert.ed ft'ca It. jJ
perea_,
X
Smalr'1 of the . . . . eat .~ .non of .......1". Of tM tim 8..all»& lear of
en4 JIIOBOPl.o14 8114 41pl.old pJ'OpldH .,t a:h11d.Uac the _rJano evaeMr181d.8 hom ft'O . . . .
Patanal. paftat
_.l'D8l
• gO
'l'a'1•...!...
!
-e •
Pros..,
Jarem
y
33.0
:
:
S
1.18
. . 2-
M.5
"'.3
--t -.12
2.48.J. .28
;
34.6
:
:
:
~
.as
• 20
d8
.x2___
9.5
:
:
t 1.04 ·:
.38
~ .M
_.....
1:;.6
11.&
:
&
I
:
.11
-,.-
.40
.14
.24
.03
.79
--..
~
5.58 "
4••
4.07
2.so
I..
.15
~.)
~M~
4.:m
x
2.41
(BI."o_
.20
x
8. x Bro. .
:
11.2
: 15.2
• 10.'
: 11.0
38-11
I
••
,·
""'
25.8
25.'
36.lI
34.6
31.9
~
25.'
•• 33.1
• 56.1.
;
I
t
••
::
a."
·
21.•
.....
C 3.00
1.59
11.1'
:
• 11.&
-
,
1
.12
.1$
.16
.08
.82
.K
.19
.06
S.M t -.M
2.1f
1.20
3.1.6
~
i
I.
.. t--.10
2.27
3.00
3.21
•• 8.80
: 1.10
1.1' •
••
1.24
.20
.Sf.
i -.58
2.75 ~
: a.1? l
; 4.~
:
1.17
1• • I
i
-t -1.14
e2.4'
t. 1• •
u
:
.h t 4.98 ~ .16 I M.$
.14 : 36.1
1.39 :
.58 •. 4.M
••
t
:
......
1.95 _ 1.13 I -'9-'1_ 5.50 t
14..50: 11.54 .60
1.08
1.58
~ 2.61
....t. -
5'1.2
05.1
~ ~
~ .to
18.5
::
-
e .38:
16.2
Za.OZ ·
1.96
....9
4.36
,to. Width
~ta
: iflomata ii1ii'IIiei
...,!It.. :~lt.t1.l!l<! .~t.~_I
s la-I ~ .20 ; ti.68 ~ .12 : 34:.6
: 62.0 ~ 1.151 •• 12.$
..
I
:
:
t
83.3
~
:
x (Bl'own x Purple)
ma.
ill"
..".
:
•
leat· :ratio or l4J"Jatiiiu '"
Pro88ll7 1'8oelft4 oolah1C1ne trea-.at,.
pateJ.lll8l paNllt
Dtplo14 P1'OpD7
_ _ plaid p~
Uilte'l"'lUll
_plo14 PJ'08IN17
Diploid
PatwDal 1'8;"8'
_unal..,...
t
69.a
45.0
4.51
.49
t. 4.54
t.
--t -
30.0
40.0
69.2
•• 81.2
1
:
1
pro..,- •
~
&I:moplo14
Diploid
Materael ]IBl!'eIlt
Patomal iJB.1"8Ilt
PaterJ'.l81 pareat
Diploid :P1'O&8B7
l»nop].cld },U'O&'8J17
..
._.J...._~_.!~M\1\
; . 'reat
!I
~r1 of the _em. aa4 sYadaJ'd el"Z'01'8 ar ~er. or the t1rat PtKll1Da leet of J8ftJlu,
8JIl lIlOJloplold aad diploid ~ _, dhUdUq ,he _rk.. .-ftlOterlfltlo8 t'ra 01'0".
1BTolvl83 tm.tr ell". l.bred Une-.......1ope4 by ___al"4 _It-polllDa'101l ,ro.turea.
Ma.mal par_t
____ .
Table
~
ot
:
thr. .
ot
...
. 'leaf
,
the._ l1aea.
}j
...
leat
t "t'l 0 0
i
l ...st'll:
pal"8!lt
JlII"08-"
~o1d
1)1'0.,.,
paterDal
Diploid proos-,.
_'eftsl paaa'
par_'
._plold~
D1plo14 pm...,-
Pa'\eraal pare.'
*_aal
MoaOplot4
Diploid pro...,
Ma"'mal pareDt
Pat.naal parent
:
:
••
;
I
:
•
.
•
•·
2
t
:
;
I
I
11.50:
3.00 :
1.011
.-41
.u.&
...
-
....
II'
,
:
1.10 ,
J
:
.'.1
~ .'3
.49
'9.a
1'1.5
19.0
~
34.8
G9.1.
1
~
.38 I
• 20 :
.'15 :
.44 :
3.06
3.46
4.09
5 ••
~
~
.IM
.80
.00
1.00
,
2.88
~ .10
.10
10.5
• 00
... ... ..
1'.0
18.1
M.
5.68
: 4.•
...
••
1
,
......
~
.44
.12
••
:
:
:
I
.04 :
1• •
.38
2
.
•
:
.M :
I
OIl! :
.<&0 ;
.18
.1&
3." 2 .12
••
•• 5.68
: 3.33
m4 x (ko_ x PtJ.tpl.a)
,~'1.0
10.4
12.2
8.1
Ull x (lIN. x Purple )
9.6
14.8
12.1
9 .. 0
1mS x (Browa x P\1!'Pl.-)
~
8.22
1.48
.00
'.35
.00
~ 1.48
.88
1.09
1.38
I."
"
•
.1'1
.18
...
2.2'
~
.a
.11
...
.....
~ .33
1.M
a.53 f.
S.I'
3.66
~
.1&
.M
2."
~ ••
2.17
.M
1.08
2
2.0'1 .;.
2.2'
nel4 at 440 X
.,omata .~
: 3.50
:
1
t
:
:
:
:
I
·•
I
. " . tin
.
~ 1.48
l.st •
... _...
:19.0
34.6
1t.4
:19.0
M.&
3St.2
89.4
3'1.11
34.6
11.2
rI.8
u
-.!!~
_to_1o.
Sulwaer1 of' ~. ~ a.a ~l'd QT01'II or ahsnotere or tl!e tiret aeedltac leaf of P'lI'8W.
a.Bd aoaoplotd aJld diploid pro88ate. aot aD1hlt1q the _rker ob.Arao'terletle_ tl"oIl ero ....
lJlY01'f'1D8 tour labftd 11• • d..,.lOpe4 1lY' ~ BlOIloplot4 metbo4 aDd two atalle Ol'O•• ht'rl4.
______________. .__~}e~Na~;h_____
: ___ ~.~4~1k
____~I~._t~o_.~~4~.~~;~.
Table...!....
"\'~...
0
<to.
P-
H
0
~
~
~
Fisure
t;
(l)
Q)
g
~
~
0
0
5
I
I
I
I
l
10
15
20
\
\
\
I
I
/
I
30
1
'I
\
I
,,
I
,
I
;5
\
50 . 55
~
60
\
65
thp first seedling leaf of monoploid and
45
~,
___ Monoploid progeny
Diploid progeny
cl~tlrRcterist1cs.
0:
m~.
40
\
\
,,
,,
,,
I
I
I
V,,
I
,,
,,
,
Class means in
?5
V
/
/
/
l
I
1
I
I
. I
I
I
I
I
I I
I
Frp.quency distribution of the lp~gth
diploid proeAny not ex..."'1ib1t1ng tbe rr.arker
No.~.
o,
2
4
':)
I"
e
10
I
I
12
1\
I\
I \
I
I
I
I
I
I
14
16
18
20
22
2h
26
?8
30 ,
I,ft
Vl
ex
~
"..0
>-.
'-l
\t!...
('l
'H
I:..
0
Q,
F1gur~ 110.
~
(!)
~0'
~
0
4
5
6
7
g
10
11
class .mean in rum.
9
,"
,
,
•_____ 1
\
\
\
,,
,,
'\
,
12
\
,,
\
\
,",
13
15
16
17
___ Mor.':rp1oid progE'!ny
- - Diploid progeny
2 • Fre'l.uen~y distribution of the width of the first seedling leaf of monoploid and
diploid progeny not exhibiting thE'! marker characteristics.
0
2
4
0
""
8
10
12
Ih
1""I')
1S
2G
--'-
'"
?l~
,.,""
c~
28
30
~
Fi';-'1.!"e
Ij
re.
/
/
/
3. 13 ). ,8 3.63 3.gg :~.l3 4.38 4.63 h.8S 5. 13 5.313 5.63 5.8g h.13
,
~·1pa!ls of cleFs in tervtls
;:>.gg
,,
/ __ J
3 • Frea'.lenc~· d.htri:;'ltirln ~f the retio of leneth to width of the first I'!eedllng leaf of
--;O;opbU -!1:;~:i ;1i~1oid. :rroben~r !lO+; "'x~iblt1r.f, the ::JE..rkfOlr cra!"[tctp,ristics.
.
1.3 8 1.63 1/.B8 2.13 2.}8 ?63
\,
,,
\
\
___ Monoplo id progeny
_____ Diploid progeny
c.n
~
31)
Jlgure No.4.
lrequency distribution of the
siz~
of the stomata
~the first aeedling leaf of monoploid and diploid pr0geny
not exhibiting the marker characteristics.
I,
35
/
I
30
25
-
I
____ Diploid progeny
\
,
\
\
___ Monoploid progf'ny
\
\
\
/
20
\
I
\
I
15
I
10
5
o
I
I
I
I
/
12.5
/
\
I
\
\
I
20.5
.
"u
"5
Class
1:'"
.. ::>.
menn~
in wicrons
5
18
1:1)
-
'~I
0
2
4
;:)
,..
g
10
?oo
\
-.J
\
\
\
\
\
\
\
\
\
\
\
\
\
\
,
\
\
\
\
\
~.67
\
l
\
4.00
4.~3
,
"'5. 1i 0 5.::q 5.67
r .... --~--~,
,r~f"!"!l7
"8rJi~-~n:1
___ :-!ol1o,!)loid
___ :Jir;,:"oid
Number of stomata
2.33 2.67 3.00 3.33
\
,,
I
,,
,,,
, ,,'
I
I
I
I
\
,
,,, ,
I
t,
5
Frequency dietribution of the ~~ber of stomata of thp. first seed1ii~ 1e8: of hlonop10id and diploid progeny not exhibiting the marker characteristics. Class values based
upon the means of three counts acrORS tee diameter of the microscopic field at a magnification of 440 x.
0.67 1.00 1.33
Figure No.
'\,Q..
~ 012
Q)
:;j
Q)
c'
HIt't
0' Po
~
~ ~ 10
>. ,..
...-
20
22
..,1.
,- '+
?6
28
<:"
~"
,..
0.0(:
(jj
..;J
'ftIe tNquea07 of the ~.. of anoplolaa. al'~ quite lew
_ta taoto!'
40ea !lOt appear to he 'the
lta1'IIII , . 'dllha'lOJl of tld.8
_th04 IO'r . . e.nabll.-" ot la1tl'e4
~
It la 1lJ:ltbl"t...... that of tM
u ••
aouN8
rer ... 1. a .....1. . 1UO-
_tarial 1aOlu4e4 111 _Ie
-1147. that tJoa 1IIIaIAIh a plat ltree4er _14 be . . . 1a_nete4 Sa
o1nellda, 181'84 1u.a PI04l1084 aoaoplo14••, .. a18D1tl.-17 1o_r-
t:requeJl107 iIba.l\ . . CIt!»r
of
~14.,.
sa ,_ OJ- polUnatat.
.....U,. . . . . .
4e~
t_
~."
'f'Wletlea oou14 M lltonaal 1IJ\1l.cl
the 't'Glue of the _thoCl tor ..-1"'108
Jl'O..... "Itlid
ta •
~
pol11aaUoa 1m' the apoaoado .le'ot101l
U .... la 'be .....lo..-t Of
laoJeUedi tile
A:IJ:I' '-Jm1que .....", . .
tJOVOae.
tN4t--,. of
tint aa4 . . . . e,ele JtruI
1IODOpl.0147 OftI' that
~
.."n.',,,
of . . utlllaal ftrl.e"
polltaa'hd • • tIM ptaUllaie lJIt':l.cma. . . . . . _re fullJ' deYelo]Ml4.
!lie low heqt1eaf17 of
o~..
of _Jl.oI48 la tAle O'p8Il-fOW.W
TUf.ett•• _y p4ll"lPlpa '- elt«IiIWIll'W 1Q'
_tao..
~III
llaea 1Ib1eb
aft
lmowa ., Uw • htlh tN• ...,.. . . 1'aztaty u4 re. . .r1ac 1ImOpl.ol48
t:roII
t,'be
,1'OpDla..
\'!d,a 1IDtIl4. __..el'. tmpo- the ......t7
of powbs at leaa" two a4dltloaal ".I'dl. .
1al aDd -ti4
OO~I,DC17
4 . . . . . . tbe
to proT14.
a••lroUl",
eouroe _tar-
Of the -'1104.
It .,.... that .,. .ee4llD& Pl'D8IPV aot utdld.t1llC ,be muDr
ol&aNnenatt. . . .lilA be
~
• 'itoa ot all JIIODO)Ilou...
aul - - . - to? poalttw lb..tt1•
four of , . tint
tin,.......
oh.'roJDo. . .
4e1um'a1aaticu IIfl4e OD _ _ploW ....U _ 4u.J1.as ... ocurrae ot tbl8
• ....,. wre 1a enor
8.8
Ja4&M 'b7 the -1'l1b01o.,.
e4 tentllt,. of the
___a plaat.. All or the .rar. __ fa ole.. .'tJ1.ac pl_'o ..
4tpl.o14a wbt. thel' _ " la
oatl_ of •
_eli,,,
raqu...,. .tWhloh
IIOIIDplol.4a.
T1da may 'be . . lAdS-
dou'blSac ot \he
.~
. . . .r
QCeun in \!J<aaU. ,laue or rtODoplold eon eeedlin"..
!be hiab .,!"\aUty rate 'Or tbe IIOllOploiCl aeedll:eg8 and their low
8..
tertUlty are the major pro_lema lJ:'.J'OIlt In tho fIOJ'lopl01d _thod
uaed In this study.
to
"cur
Tho low
tnqUeMy
at eloh . .op10148 _N touad
would Dot I:)l'Oh11)lt oxteaatw 'lIM Of the _thod ill a brMdl1r8
,1"0«"- it 1t _1"8 PQpfble to ob'aia ..lted eee4 on all
~~lotd.
OJ'
.oarly all
deteoted.
It 18 _148ft' that 0.08, O.lO. flD4 0.1& per cen' aololdot . . ta3"t10... are aot a4eq'lllte ~a. dOll\)11...
tlou otber
~
tbe. . , d.ll'te1"'8Bt __01.'1.8 of applloatiOl'l,
obllldOAla may pJ'Ove lIOre a.tlshotory.
"PP1toatioa .....1"17 iajured so_
A! tho'Q&h \his bljun • •
the IIIO'rtallt1
'H8"' •.
J."8.w.
a~
...
In.1••t1oll
or the . . .UiJ'lp
Goaoe......
otMr
Q1'
_thod of
8U-3~
w
1t.
retleotellt \), a _S&altloaat 1neRaae 1a
It wOtUd __ d ••lft\)l. to UN • _tlIod of
applloa tioa wh1ch would ruu1" blle•• 4 ..... to tb. . .elll. . .
mule 1 t 18 quite osay to dllltlngu1ab. JIOROplold hom diplol4
plats _
the bfuJ18 of the
_~108J
_4 tonUlt,
or
t1t.e _tUl"lt
pleats, it we DOt ,08811)18 to distin.,a be","n tbaa In
line . . . . wad.er the ooaU'Uona
Hi
oba~owriatl.o
or
tbt. experSaez1l.
,he . . .-
It .. IDrpholoal-
cum be tolJDd whloh woUld. ,a!'lltt rapid _d po.ltS:"
ld.tlt1C8\ioa Of maaoplold ..e4l1nges, It would be
.:~dl"tlllllt17
'1'he elfJlld.natlO1l of' the t1mG
t~_
O~D8
eat exaettq
or
ftluable.
-lc:t.Da
o)&roa:)aO!e oouat. at the /putauvG Jdlol;>lolc1 eee4l1l\P would pttmit
~.
8On4llWla of ...h 18.1'.1' populatloU.
It 1t were to be of ..,.
mar1r:ed . .ent, meb. e. morpbolostcal chanot.,latle 'IJOuld Dlu!••aarlly
ha'h to be ....tdent .tore tft&tl1lsta to lnorea.. the tenUity 01'
])101d8 1181'8 appl.1e4.
JDCIIDo-
Such II ohft1!'aot erlatlc .,. DOt exiat .~t\1ral11.
hownM', It .,. 'be poaa1'ba to lnduoe Its 4...ekrpl8eJ'lt 'by the d.lfferentlal
rcutioa
ot llOaop101da aDd 41plold. to 80_ type ot troat_nt.
At le••' three . . . .atloa. are l"tIqull"Od tor tbe •• tebllaltmotof
awed 11• • by
1n«:
~.
aoJlOplot4 l'I.Ie'tllod..
Thea. g..-.t!"•• are tbe follow-
(1) the , . .ratioa .M.auy to ~du" the 1Ml.l"ktMt
'1
_ed.
(t) the ...-"t101t 1n wblob. the moaoplOl48 ere 4e'teotM, 11"0- to
ptul!':lty and . . It.... aDd (3) a d1p1014 g-.eratiOlt to RPUre an aBlPln})pl,. or Belted ......
1.'••,s
ot the oombll1111«
-'nitt••
ot the 11...
••'&bIlflJh" 'by the JlSCaoplold _tlao<l OSDIlOt begta wuU e.1I1 lee.st aae
__ ratiO.
or
~_
.u'plo14a brlw _ . Jr04uced.
'!'be
Det MTinS
In tl_ in tbe pzo4uot1os at .ated pu.re lu."ee41. liD.. 'f':rooa uat ••te4
.artal .111 not, tUNtON, be
ft8
caat
q
a,. ...m app&NBt
upoa
au;,ertlo1 a l e'XflU1aa'101l ot tlM metb04.
or
'lhe lIIJIloplold Butlhod. In 1t_ ....eat .t...
aot appear 'to
\)8 811
ernot.,
4.....1o:caeat. 40_
aubatf. tuM tor. 10118-tl- ..ltiq 1n ..,.
el:__ 1... "fU'Oh tor tll...4 l1a•• to replaee tho.. preaftU,. ue4 ta
1\tbri4 comltbuR iona.
The 11• • developel 1IJ' tha soaoplo14
_~
_,..
t.-
_ . .sr, M valtla'l. tn oenala ape.tal «_.t1o Uld bZ"M41ng .tu41...
It the t'requene,. ot . .
orea" or the
~11tty
()~_
ot moaoplolu
leyel iDoftU" or It
e.t8b11a.4 hOm lIOaog1cld8 prow to be
than tho•• eateh11ahe4 b,. 10...,1_
uaume l.renIn,. tmpor'te... a8
til
"h~
~tl.J
Mlt~t
GfUl
be sreatl,.
1Illn...4 1ta..
8Upe1"lor
~.all"
the method will Wl4oubte4l,.
'brM41D1 \eobalQ.Ue.
AJProztaatel,. os. baD4ft4 "'t)" .......... ia4t..rldul - . _ "......
_DUne __
IID8Op1eU
q....,. of
. _ 1n
propal. . of . ,___
1PO~8.
t_
0.88 per
.a.... of eo. _t'e ....saM
~4.
~J14
__ fen_t to OOGV at a __ fa-
•••US-. . . . . .aH.
tnquaq Of . .
81aIdftoa' 41tte:r-
00....,.... of __plol.4a . . . . to....
Del.,., poUiaaUoa aa4 lSJdW ap,vaOld.o aaleoUoa 414 JlOt
•
to-,
1_"_
traq_07 or .,.oplolq 1• ..- opa-POUiaate4 YUl-,., D'\tC,
.a
1t. orin'U••••
!lut )d.g)l 8n'tel1 tJ' ftte aal 10" aalt-tes-Ul"r 1.....1 ot --)dot..
we.
fbad _ . . . . . . taeten U .•"'. . . . . . of the .,..,1014
.tllo4 fbrlnal'....... ta -'ab11atdaa lDltre4 l.1uI ot _ . . Iaj....
'tema or 0.08, 0.10, au4 0.11 per oct oo18h1olae .,lutlOllI uto . .
. .tallar BOd•• Of the 7OU8S JlI'3IIOplo14 •••dU ... 414 SlOt alcatnnau,-
laoreaaa -'11' aelt-t'e:nlU,,..
~1l
of . . ttnt NI4!11Dc leaf. leaf w.t4_, the ratio ot leaf
1...-. to laat "4th,
.~,.
81-, _at noaa trequea07 . a aot
tout to 411_.17 a. tfcseatlate . .opl_a
!be
-1IIOJl.OS4
npe10r to tM
~
aeibod. .. __ Sa thi.
loaa-'s. ael.t:I.JIa
41Jlo14 .-4l1qa.
8"", •• sao' t'o1Ial to ,.
.'.4 DOW 'a . . tor . .
tNIkJItU....
- ' of 1arp . . . . or I ....... 11._ tro. ,aY1ou1l1J' . . . . . . at.rial.
a.
4.
1Joo_.34:113-16&.
Meta SUoll.. Haploid ,1__ tn
IMS.
Obe_.
s.
_t_.
MoDoplold tNct_IlO1•• ta is 1O__ rotal 40Uble
aDd 111 1ta eoapc....' 111 ICLe . . . .
hierl_ aa4 til.... 1_a. a.:aett.. 361318-881. IM9.
fl'Mn!at
0l"0•• ~4
S.
Aero.. 44: .1-11'..
.a
PJto4"oUoa ot hoaIo:qgowa 41ploUa o't "'se trcm
aoaoplo14a.
JOur..Aaer. 900,
a.
0lt.J1........ BUct. It.. , aac1 a-tol'd, Roaal4.
e.
DaYt., BJ'adle),
t.
:tour. Heredlt,.
aor~.
..«l11_ ot ~ft.
M:99-104. Ita.
2!"'oua __am
L.
1918.
Haplol4. la
J'our, Bertlil1t7
at,.,.
o,-tat. _,101d.
c......
.Al:l.ww. JIB,. 66dlt33-H!. IHl..
,
l.lJllcarat. a. 0. '!be .,..10.,. _a .-.'1" of •
haPI"OU aport f'Jltom SZ!!I!!MJ:! .t!!J!!b!!M' OIaetl..
so.
tJrom
'\0
)8001"\'~!I!'
ad
11:01-80. 1930.
10.
De~••
12.
1£1...,. 1.
13.
lilD.8wller, S. L.. end Rllttle. •• t. ID4uee4 po17Plol47 1a tlor1.11;QI"'" Amar.
ftl31o-:sal. 1941.
14.
_1"8Oa, S. n. 'Dae re4uotloa cUd.on tn .. haploid O!l!;t_ ••
La Cellule 3':109-115. 1929.
15.
Pam,
16.
J'OrtUDO. 1. Valez. UD;publlellecl M. s. 'the..
OolUllllda, W.880url. 19• •
1'.
R. OolOld.otaa polJ'Plo14,. aa4 tecdmlque,
I: H9-6Si. 1t4O.
19ot. Rft'.
OhJ!lOm08OM 1a«_ 1a relation t(J t1'ftB811eff1on treQ.....,.
111 mal_ 'b1.aoaes. GeeU. . .:349-364. 1M3.
1'&'.
LN. A. . , . . .. ."1. etuc17 ot a ..t of _ _ lIIOIloplo14 'bU••
OeMtl.. 3'1,580. 1958.
••-.n.
OBi• •,~. 1'.,
lour. lot.
a:a4 A.uct. R. C.
13t~.
t1a1 ftZ'dty ot
A haploid 'tIIbetn ,1_'_
1.86.
/litter.
'our.
A."
a. R., a.Bd. OOo4ld.a. It. M.
Up101d 2.!IP'lP" 1f1th . , .
couldera\lou OIl haplo147 18 plea'. am\ ~l..
83:123-156. 1taO.
18.
Oatea.
11.
o.re.a~,
10.
Quatat8llOll, Ate. Studt_ on 'he _ebud... ot pl'lrtaleaoa. . .l ..
Ben4l... ntl-W. 1'-'
81.
Raberlaattt.
08.".8
Cr!Jdl
a.
kparia&te11 eJlhalw. _loltle Pfl.amIe YOA
Plaata Jlh&H-70a. 1936.
5!JElB L.
o.
ZUr ZJ'ologte l&'IId PllJatologl& tie. _1,.,11. . .
YOZI 0!!2!M!:-'. 51 tab. pl"ftaa. .Ak8d. • •••
a-~
7:-....&'.
M.
1929.
Berltlld. S. O. Jfaploid. 1A pol,.-l"JOD1o . . . . ot sea lela
oo'toa. lour. lJeftd1\7 .,:2...I3l.. ltH.
uat.,.
13. ltclll. . . . . , L1ll1-. It. oTiOlosteal ....7 of haploid ~
~. plata.
CaUfoaln PtIbl. Apt. ~
1
- lSi. 1980.
K.
xap..s.,,'al ~UO.. of haploids
o...tl_ lO.ltD-3f'1. 1938.
IT88O\'. Y. It.
l'U1!&__
IS. Zoh.autta. Doaal4 It. Haploid. b
?roe. 1O:ts-l00. 19M.
~
1.
!!.tt~
'ftllS!J!.....
Aoad. ,.1.
l.oJaI,le,..
24.
.1"_, .1eak1a 1'•• q,4
A. E. a.J111~ alll1 -.rnat olll'omoao. ......... 041l01'O tU:ld 0-..1" ftl"l.tl .. of noe. lour.
Ag:r. Ree. U:.1..... 1141.
2'1.
b-,.... To.1_.
m.
XOstoff. Doatoho.
at.
~
Haploid tol'tlllUtOli by r,...rqe 11\
51~. 11M.
Oytololi.
Dl~
'.hl.
h....
'lY1t1~
ftll'1.ftttou OOJdU10D!8d euplo14
ohro.,ao_ 6U.tere.tioaa. .Tour. 0IM'1.. :S6:4t4'-4G8. 195'.
1ba pft>'lem ot .plo147. 111>110.
IMl.
, lStl-U8.
. ,• • OIl • haplotd potato h,....14.
a...t.
30.
tAtIral, RoU"....
Il..
L1D4at1'O.ll.:I.
8114 1008. ttat_lDe. 07'o-eeaetle lll'ft_tiptloas
ot 11 luIJlo14 1Ioaato aad ita 41plot4 ad tnftpl014 JlI'O&UJ'.
~. 1_1'. Dot. 18:.....uO. 19m.
sa.
MIIOI'aJ", 1rea018 A. Aaotlwr -,1014 Mootl_
Bot_ au. 93,.rt-I3O. 1931.
33.
6'.laa.
34••1-396.
'I.,
AnIe.
ll;!Mftta4I
1936.
lfn'eil\u
t~'bt1cIua
plan'.
Cbartlot.rtattu of two haploid twiDe in
lifted!'" ShlM-l40. Ito.
E!!)z11l
34.
..".,.tn. M. Chromattn __ e4 .eU
3&.
X1elaa. ftlU' 1.. Bnetiag bellaT10r ftD! ~_ _ .....l"I ill
pres-t.. t.oa t.D u4 tnpl..n plaiR. of l!!!.11'8.s1,.'
lot, Oaa. 108:16-10. IMI,
U81. oaJ.ltom1a Publ. AfI,r.
sot.
'IOlute ta related epeoles.
1:10"1-830.
1931.
sa. ........
(J. R. !be plaJalo10Q ot polY,Plo14y In pllJ.llta. I. R......
of tM literature. u.o,Ua 9&153-1'13. 19".
3". lfOJ'deDakto14. lW4a. stu41. ot • haploid 'IlJfJ -pl_t. Heft41taa
••
II: lOf.-alO.
19• •
at...
ltaattolJh. L. 11'. 30M street. of hlch ~fttUN Oil po11l)lot47
aa4 otlliDr nr1l1ltloa 1».
Proo• •". AGad. Sel •
•• asa-aa. 1932•
• ftl1411Ml\er. H. 'E.
--"Pil'l"'pl""-oida ta '''''pl01<1.
15: 1&1-161, 1939.
mat...
-rM
~_
or
~hnoceae'l.
Pro•• N... Aea4. 501.
Sd1aa. !l•• l1ue81••• A, 7 •• aDd AverT. A. O. Bal. . . . . . . __
bela._ haploid. 1ft D!!Uft. J'OU1". ReNdI',. 28:1ts-1O!. 1"',
41.
Smi_.IMI,
l.tlth.r.Haploldy
41.
8\841_. L. 1.
<M,
Vb 0'Ye1"1Nlek. l' t Ooak11D. Marie
U.
W.tte_ta, '1.
lour. AI/Ir. Rea. '15:191-301.
1a e1akoftl.
'DIe C'.'l'peJli..nal. 8>4U1-_oa of lIel!'e41tr 1D
,leata. SOl. Ap. 11.5.-I'Ia. 1t8l.
_1"'.
&.,
IID4 BlakeelAM. A. If
ClOp
OMalcal
.tiDll.ttoa of cm&l.e 4tmtlosa-' aM ita poIlet...l. matioa to
~g. . .l..
lour • ...,.. 18.&4'-654. 1941..
1IfDrpbo1ope UD4 Plt.:7elo1olie
. . . aut . . . ., __ GJuadlaae I.
u. VeNft. 33: 1....,6. ItM•
• ,. ;too4-"lta ROMn H. OytlO1ocleal
1~8. . .1. _d
891~. 1930.
4'. Yetelk1a. A.
K..
«..
1C.1.......l. 4el"
zett. .r. Ia4uok. ANtilla.
_.n..
])01,.,,1",0117 1n
1a the Betv1.....
J9.a!! ......
lot.
ad vae11jeY. B. I. Ant notal 1na.tloJl ot
b.aplol4 4U1'U11 . . . _ .,. polU.'loD with £o.~ JOll_.
!k111. A»Jl,.
Oer.Iet. nut INedlB1. actr. II, • • ,:
Do'_
_ .... 1'••
III.
ou.
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