Kirhlvn M Urn. thrector
Mkhrl k DNIS
I b d C frrm
John J Fmrr
brr) A I HolUnk
Dmccl B Houron
W r m a F k
J . m n H Muun
Rmct G McKLnry
Karl E N w . Jr
Wlima L Wkr
John G.Sueacr
K e n d l W Thcil
T. Craig Wtdcnvul
Lynn B. W i l h
Rogr N. W W
lbba R. I%*. Cbur
Jmnu H . B m n . Ex dhcw
TABLE OF CONTENlS
Ovmrkw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .i .
lamrduaion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1
Brid History d Soybean and Phymphhn
Roc inOhio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I
Orrvtic Control d F'hytqphchon Roc . . . . . . . . . . . . . . . . . . I
Whok Plant Resistance m Phyaopblhorr Rot . . . . . ..'. . . . .2
TolmMce to P h y t o p h h Roc . . . . . . . . . . . . . . . . . . . . . . 2.
Rre Specif~ityof Tblcruwx . . . . . . . . . . . . . . . . . . . . . . . 4.
Phytophthon Genetics and Stability . . . . . . . . . . . . . . . . . . .5
T o l c r ~ ~ ~ etsT .c.s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6.
T o k m in Cultinr Ripley . . . . . . . . . . . . . . . . . . . . . . . .8.
Recurrent Selection for Tolerance . . . . . . . . . . . . . . . . . . .10
.
Objectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
.
.
Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
a. Rrformancc of Lines h r n Cycles 2 and 3 of
Recurrent Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . .12
b. Rrfwmance of Cycles 4 S, tines Selected for
Yield and Tolerance . . . . . . . . . . . . . . . . . . . . . . . . . . . . I3
.
Rtsults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15
a . RrlOrm~ceof Lines from Cycles 2 and 3 of
Recumnt S
ekctm . . . . . . . . . . . . . . . . . . . . . . . . . . . .15.
b.
of Cycles 4 S, Lints Selected for
Yild and Tolerance . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18
DiEcussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20
,
,
Overview
A a a p r p n U o a d Q b m d ~ q (ollgmxawmio.daaccud
raeurh rr MRDC-OSUhu bcea
yddMpwdad~lppm
brtbcncwcarcpbcdamEc
fk&krprmlduqma.W
Sayban hr bsca w
s mrpr
y r l d ~ qmcunlad deMc (m * l u k ~ & m r m w q r n w uJl r m p lina Wgl. Phyrrphdwn mcupslfir) d u w r ud pcnn- c a d u l ~ I c & n w .
lkcaulr,dlbacmdervon
r m h u k c n f k m n o ~ p(um ThubuUcflndsnbtr (I)(
dime &ding pmductum droy
wrc:(I)rkc*rdmtdqd
new concept concerntrq m e
bw in Ohm since M5.R e s i i
mnrpae~frctyla lolaurr d my
i)th~mdiliamtypmd
culdw, devctcpd In dha lllln bCM m PhYIIPh6*AI a,(21 fk
daurc,ii) fk rduomlup berad rt the Ohlo Agricultunl
evdnwe Rlppntlq the m am. mm ldmnx dm h kld,
Rrncuch Inti PKlopmca C m r .
aprdcnwdfmn~Jraudyddvrc grcmbowc, and smwlb chmber
racohiosac UaiHnity (MRDC. mahnktomusuntdcrrarr,ud [email protected]
OSU), )WO prcndrd p o d amtml
( 1 i ) m l l h c n b W r n d y d ~ daracc,ud iii) fk inhniancc d
until infrequent nccc hacrmc
a I my m
k
m a modentrly
0mrypeddcnna;Mda)fk
dcvrrlopneot d Pprnor [email protected] pdpmknc and muledin d
m
i im
n k l l f i t cw Md KkaHrn Of 8
p n p l u m n k (lntduclm), m a ~ w i t h d l ~ p r d
M
y mbMl cultinn. A mm
lObk (nu ~ ~ p c c i f rp
)m
, c
M d ( 1 J m l u r t m s n d ~ u f ~~ u s i q u i m p m c d n c u r mxvbcaonlocdnurtJauud
mt rkaioD mdmldqy.
m m l wr, dtrmd.
Evaluations of Recurrent Selection for Phytophthora
'Iblerance
B. A. McBlrin d A. F. Schminhtan#
Dqmlmats d A g m m y and Rnnt RLhology
arcocdodpnduawnkwhIy(6g(uo (;cac(lr Coatid d PbybphthonRd
h)(lhuckr. I9471 ohti Mlyuyhun
WWdSoyband
paduabnm198B(rmslmyur
Cnruc m n d I$ Rnr( in wyban
Rot la Ohb
brwhrhW&mrrC~rwc
hr Crm r b t f m l cn wed In knir
1b mtllm ha (Runy a J. WZWL
The soybun I G I p u nor IL t
(nit mrrurtly uc'lunnl, uyn
M m 1, wu ~ntmducedm North dthenopedua!muccnt11(
urlullng (I)mrwc
1 6 5 0 r m l l m d l y ( w ~ ~* p~ h - k ' h m p i b l c
Amcnu tn 1%5 (Hymcllvt~ud
pmdwwmdprredI.uhn
Hrrtm, 1982) but remined a
mr ( a mnxn aue durn)
IKIuJhnun ud kdanm. W8).
bmmulcu~udmutymuxw yw\) 'kuyWM*rpll
ly t 9 4 7 n k a m ~ o r c m p t n o h M ~ (2) w h w Ikhmmhaurr ud
Iycmp~MMlOhaUSpmduflm
murchn. W7) ud lurc Nu7 hu WLn. m)' (71 m cakryl
In 1907)uml tt w dcvclcrpnl r s M
brm mon wdcly g m n than nnn
alvalcnlp (Pcobsc ud JuW. Wn)
mmm (dtw nsuyl) (%)Icy m
d
b r $ ~ ~ R u n y U IWIVt
d.
The tnnsfwMrwn d h scyhcan
Gnu.,)!%I
ud (4) mot r u r w r c
k r r m n l l y ~ c r o p m p hccmkrrtng Imv ~nbxluin,hl wl
fm m ~ntmducedcnv 108 nupr
s o y b u n ~ a d ~ ~ ~ l n l tmwntty
h v l ~ k1 tncon~pt~hlc
crop In t
k U S u a rnbutc to the
m r r In
t ~ n o u sMIborn
I
d ~ m vha&
dfom d U m d Sutn DcparmnI
the nxn ~nd~cvtrd
by hydnpnlc
becow s mpr Ohsi pndurtton
vf Agncultun (USDA) a d Sutc
~mtulatiiinpmcalutt (Ktlena al.
poblem In r hmlq~ullyuhor!
Agncultunl tlrprlmcnt SUtwu
1974. Ktlcn. IYM)
p c n ~ d111 time Thc pnthtigcn
A dtgrnhttin kl ddiw l c m I#
( S A E S ) m r d m 7hcnrcvrbrr
appvprurc A compltiblc w can
~hatIhcsoybunwugnwntyolr fhpp/uhc)rn mtgurpimu Dmh
f ~pblvnnra Kur d tnvtn
1nbcc1,cdanrrc, rd pnulbly ktll r
tdtvduJ m Ohw at a d y o 1853
( h g ) wu lhuvn to be the cauarl
rurepthle hnt plrnl lhvl wmc
(Robst ud Judd. W31,ud apcn
race a ~nmpattbkwrlh 4 knl
macs wtth vyban date fmm 111112 qpnnfadM,danqnngoff,cud
ma d rtcm nit vf ~unrep~hlc whvh hu c l h r whole plant w mn(
(Thuchn; W7) wllh the lWMtIvn
&nnd*
~Schmnhcnm, mtruncc y s l f i c for lhnt nee In
(rind c i m w J y una I892 a h
1985) Thc dtacaxc war flnl
the true of whok.plmt mtrtuue.
nloauon)d the Ohlo A p u L n J
Ihc plrnr u unmune to mcmparbk
ErpauncnSmn,mcberunrad obrnadmohmm WI(Suha*clry
~a bsuuc II hrs r hypmcmttiw
lhcOhK,AgncultunlRcscurhud ud khm~nhcnm.1955) 11 16 nw
common In Ohm f i b , caumt~up
mpow whrh kllr the pnhgen
Dndoprmr Cater, dTk Ohm
to I SSO mtllm mu1 IOID
(0
udplrnturutuoupdthc~nbaron
Sole U ~ ~ l r t (MRDC-OSU).
ly
c ~
spprm~~tcly 81ce c n w i a mmtr flak Rtxn
m m C d u m b e ~ U h r a ( W ~ g f ~ Nntroawdc,
16 mtllloa ha uc ~nleEtrd(khnur~unurc
m lrmrlum-lya Mkcr
ud wkm. 190) Oluo prodwim
thcaner. 1985) The Soybtmud khnuahenm, W81e) cur
d
w 21300 ha tn 1916
~ d ~ l a P m U o D ncultn
l ~ tn a lnuoa h wtth very
(WJIIP~~ md ~ r k ,fin) ud
r my p a # symbtous wNh lralc
lunnrd~n(thCpmoOmawnmcnrred a 52a00 tm by 1923
pdmmnccx~ltwmlforuycocvolulton but wtth many
( b , 1 9 U ) ~d~
i ~ rmratpulat~on b w , w & m u w h o l t ~ a m d g l t m ~ & d d m e P l p ~ i b t l ~ l for
~ C u l t u m I p r t r c s d h D c l t mtnsacr (A F k-1,
bcpmQF)3(P,,udml938chr
rn). obleMlmdrm(rau(uweRpc2
p o s c a m c u a m o W b r ~ gcaarcl(-,
lntmduction
daenkd by k,mi) Ruoc
d & ad 12 gna d
ccrlrhaau~wlnmoher
nMc or wry IMk nnn td on m
d~vdurlp l u u 11dro hu r conus
wn r n r d u l t cnpmcc bnvan
m t m (dl dm) ud runceplblc
Ul dud) Thu 18, ame plu#r wr
[email protected], @,-b.
bl*.
&I-k.
@,a @,c.
loci
@J,
b,d. @.- b 9
~rd
RpJd which cmddion difiaeo-
uJnur;lmalothosmsinsay
hPn.uucnpxbdinra~ar*
(Mm. 1981; Pnpcr d d, 198%
w w u d ~ ~ m ~ p L L l d r d ~ ~ w h m rLyan
t K a d. 1986) lRMc 1). @I
(Kihad, m;Kilm. 1986) mnhypaocyl u IMllduupq qj
dbbamrmPamWI).~
&&rlducsudnnbcuryad
unchnanndmudmarc
by nnn tauculalcon p l r d u n h
tnwnurludl4!agminmegmn(bkn, 1986) lWufmm expna
plrun lin HW79149 mkssbd by
tm dr Ini~
fnnn p l m m plrnt In
(MRDC-OSU W r a d, l984b)
hanatypuc luw* u albd m
ud 'ChrpNn' (McBLin a d.
complcP pmfruwe ) The term
m).l b i F h m gem havc brm
"mlc drrducuyl mutrooc"tr dmou
exclurlvely reserved
IS
an
dcrcnhc
duam m
ud unount or
~pnrulrtnmand lacon u7r As In
*nded w1rh FBI#. It m s li slower
nr [email protected] &m't m I y
tndlcrv iowrr colm~urron or
rq,mductkn the r u Q n p n w
rrm "IIow ~tlrl("u Imore
rpp"rpnuc chnrruDlwm t f k
lapn*s I n & m k n m b u r
daad wnh h18hly rpalfic prcor
wage Skw nlllng [email protected] hr been-
cpdnnmlpsrd lcnn
*
lo
hnnrr, %, OM uul Schmlrhnncr, 1985) a d m l y In dlf
lercnrrr (I-!d) In rtK rbne of death
d dtfknnl suxrpl,Mc Ilac.\ 10
hypncUyl Ml,wllh vmilenl MS
(R L. Bannl, Unrv dlUmm, IllMI, 1990 phone ~~rnrsum)
UradRlandullyIW~mnaol
d!4qunil1hc~uncMclled
by hrmcrly ~nmqucllcannpnribk
mm d & which bmune fnguerrl
dmuMindipg.7hC'Mubdcn"
F.
RP,, p
m
v
w Bood h
*mbrlrcmearltinnIw
0hiOpmdudimbrw;m~yars
(Schmmhcmur. 19n) buc has had
IitlkormducinOhiosinaabout
WR.Thes&ad~dcplqrdwrr
&
whjch
,i,
is nar d umu d
uc
only m d
m Ohio W which hsK
lnkamn;r4.On~odnhnnd.
t i dl p d e s good a
d In
A r k M m md nrlghbonl\g swcs
(C. L. C N W Uaiv, d ArLamPs.
hycthrilk. A h , npon u h
I989 Sqbmn Brmkr's Wbhcp).
the gcae e l - k has multi-rre
nsiancer,the~naalypmkn!
nxs m Ohio. It's firs Rkae in a
mnkm&wwm'WIuPmsSZ'
'Jr-mopntmEyhbpd
wim R. L. Caopr, USM-ART,
m m , a ~ wR. Pdr.
.
Wd4PxmY.IwSrc
unhiv(Coapclad, m,k
a d, m. mh, 1988b; Mcghm
a 4 ~ 4 6 ~ 9; 1 q b ; d ~ ~ l h r
a d , l986).Thmmhpri\r
alith*gravniqOhawhich
hsK llwgenc (a rtuirtlaa. Ohio
w y b a productioo is a1 risk for
imtPsalhnFnrydaumltly~~
rpfg (12, M 19.20 a
d 25; 'hMe
1)dhIgwhichaaddatthisp
(no new iW d rre 20 has ban
~~incebociginrl~losl).
Rm compatibk~ceswnc isolated
fmnOhiofieldspriorlolhcnlase
of most of tkm cultim (Hobc.
1981). Onc w ~ b
y ddnss the risk
is thc pyNnidiq d genes such as
in 'Kcller' (I)psls.Rps,). 'Miuni'
[email protected],c*,),'W*
(b,-h.
b ) ) whew CI a, E J @ ~ , ~u-i.C ) ,
'W (Rpsl.b. @q (Nickel.
1990). H w m r , nre mm (10, U.
17, 19,20,24 or 12, @,20 ud 22.
WI)ucstilIIckad.la~
tmdthisiisk,adTmu.~W
m~-OSUodmlopm~cc
spFihcgaeticpmcrioo~~~
b h (SchmiabcaaerrodWr.
m).
It wm a*cd (Sdunmtunan and
Whr, W19) thrr nm dtivus
which arc kilkd by the bypoco(y1
hacuLQnmmcsvenlydsrmead
ME).hhsr
~ b c e n i a m p n r d i m m u y by~inmeWmdtbavLiaL
aJliMrdPadty SAESsllrhdng
yiddlaa,dmcbmanidchgbrm
x m m r G ' c a M y & ( : ' ~ g ] : dcKd!ula5dcmcc,~mcbod
(-8ndCnmms
'Flyer', 'Gnome 85: 'GRE836',
-36,
'Hrprgl:
'W
'w,
dehPitioadMuPcn(lW)Mdis
squinkdcldermaI6Mimdby
sr.'fklb86:'RcsnJr'adSpLV, Sdaniltbamu Ipd wku (l979)
mwen*adPwmmdapb
whichindudts~dow~
WdmhpdaQWX:m,
lmdeyadGmJ. BBZ),ajmemd
-
lWa 1. C
3
e
m IOIfwstnncelo PC
I-
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d h g on a plutl wtth I)Y JIWmbug tmt my be l q c r thao on a
plurwhlchJhwelutledavmwlg
a t -b
SY
Imord~s.
prwdes k doad m m
s
d prcduc*, fewer oospom.)
polentlrl for lncrersed P r n ~
repmdualon on dm mrq mu
pcarnrlllhehctlmcdtheobacmlW mans hat slow m a l q In the host
mrlhe~RpmducuondARon apanullyrkcuverobwrdtk
dcvelopnent of LSI qgmu F+q
the hou phm durmg an entlrt
popllabom wtuch prol~bnteIn the
gmwlng nernon Ewn l l h e w p m
RwSprUld(ydTdancc &Ion of the slow rotting trill
mle of rwl pruners on slow-rott~ng
roleMa ronglarlly pclpoacd appmchen ma rcalmnce In there toknnt hosts Thtse conceptr art
new ud have not been estal
o r n n - r n ~ ~ i r ~ c m u be~ng
d wry llale observed rot, and
thcutx*mntdmdocs~ndKovmuch Rwcprchcrs havt not been aware of
dhq iSchm~nhennaand Wdker.
um
a n UYEdO(BI
WlP) H o u l ~ c r , n u n a v m w g w n l r a k r c d c n k m d ~ n c r b y ~ thev p ~ b l l ~ m
rt MY polnt In umc. tt st111m
y not
cvldencc for or q m s t these conthat potentlrlly rric.apcctht
k ~ l a 3 1 v over
c a s m o n tf W l
cep(a s na On6 such repon ls hat
mtuntnma a n ~n.lurleQtn the
former h q d ~ u s enumnes In
(wpmproducuon IS as great as m
b m d dctindlon Tolant uybean
hst mtlng plant. Slushcr ud
M~aslsalpp~
no longer arc useful tn
unrnv hn* one or m n nf thc
B r h r (Wn), fwnd equal nwporc
~dentlfylngevcnthc mosi rntolemnt
M-m
rmtrauwre(wh
productm cmtm tolemt (dow mmg)
soybean strains lfrcr years of gmw
u 'CNS: Rp2.K I ~ M ,I986 m
d
'Wlyne' (nm m t or whole plan1
mng conunuous q b c a n s (Hanwlg
pors~bly 'Yi~rk' and 'Ware'.
m a n ! ) as on buxzpubk t n m k m
and K ~ k n .USM-ARS. Gtonev~llc.
T h 0 m l ~ nH 11, 19118). rltw rtmtng
MISS. a p o n rt 1989 Soyhean
('Rllr'. W k c r and Schm~nhmner. 'Amscy' but only a few oosporc$
wen (ounl In 'Amsoy 71' whlch has
Br&r'b HbkJop) Whether lh~?
WW, lndtcrlnl by low Rcld wort.
nb9:f+qbndKdhbeks?.
mdgmdgcrenhuaewc~~:w~thmn &sI and I\ whole plant a s l r m t to
mr I (911 1mulatal w1lh n c e I)
npgnsslvt, or ahcr 5011 mlcrdm
tnodam), or lbd~tylo endun the
H h , n a1 (1985) b n d hat an In.
wtuch art phgcruc on h g have
mkllon ('Annor', [email protected] 21
conlptrm~blew e colonurd thc enua
p r n t l w r yteW lov M 'Slom'
~ m ~ , W , o r s o m e
nmcx d ra~.twr
plant. near the m- &r palthough 11 hd thc wm iidd UXW
rr unalvcd IS not
ktm stte hut dd mn enter the stek
for uwn ymp(c1ms ar 'Sloan'.
kmm It IS clear that the prcvwu\
W k c r and Schnlttrhcoacr. 19W)
as d ~ da comptlblc we, whlch
ux cd h e term tolerance (Schmltklllcd Ihe plan& But In lhts study,
t h m r nnd U r . 1979) for nonRr c n d u m r type oftolcnnce s
G l ~ e o l mI ( w y b n phytoalcxm,
spccltic tolerance m c c b s m s is Ina mon ~ m swe x (olerum as
lNIr bl F q ) Imls wn b h m thc
sccurPac Rrm nslstsact s race
d a l i hy Schkr (lQn) tk rhlllty
wnu ofmshnl plam dmay afto perform better wlth equal
spcltic. endurn e not race speckaAUpRpmducoon Thcevdcnr
p n l h p dcwlopmcnt Only an
Ifr,ddavmaulgnuy,amaym
ndunnce nrcchurtsm 1s truly nons u s s s t r ~ R n g P e p m d u c o m ~ ~b c . r a c e s p s l ~ ~ o n t h c
prUculu mechuum unrdnd
aclectlu In lem5 of unlnhthltcd
lffslbdbyslcmrolungmsome
cultlvnrs but 1s armnly affected by
IhcpFaposdacwmakldgnQc
Wopmclu of he p a t q e n and
whole plant nstsmncc If lnfocfad
Ihueb IS n c o n m p e c l f ~ ty
maold~lrrludcsvmokplsa
NXXI m u n w to suppon oospon
definltuin
rrsamre,mlbmsdsagclolauDc
A d p d lhe pwentld for nonpmducu~nto a l m m l u k n t for an
.sm,brlolauDclspnaoaed
m spc~tic~ty
m Ihe slow mnuyl
~saroaonalugc.anzlarully l r m r n ~ r b a r ~ d
trait IB m dlficdt h n for
#~dothmrlscWlhybosl a t h a a x d h a p d a c c s p m f r
mdunaa.'Ihcdmmnratlngtypd malys*m,lhemldreproductloa
mnrmnrrmrabcdedndfRadJ
lecmaCe(l(lkn, 1986) Thubmd
ddbuad~mubrdfnlhc
amqpnaccmoddbdd
f ! q ~ b n m c ~ d
pdsllon on lhe butt d rurvlvll
uadcr d l ~ l r eprelbun (field
nruturc) rcrlrrua luwl whkh
h,
a d m n t Ima, v h r h do mt
bm, whole-plan nrrdurc (u
-bybFV1wuh fhs p.rrvJaa nus m t fuld)
lalnurcurmarrednrreductron
mtheunxlntdmcobremdmn
glvcn pcrlod of ttrnc and 1s
p~f t a J l y r L a r u m I h a n q m
rn&btt!hcgmvrhdalersrm
u u n d F h g HuWw,mcdrmobvMddoamcnacesllnly
Ud~ferturlrmShdPkyhYphv
~ p m m u d h q f n o c R u r TOnq' ml 'Srm' twc
r a D d d m a n m p m ~ m a a n I b c pnblypxr*--slm
pramodel$Ddclr~hnuond
fmldraaudyrld141ud43
dr~maarrltlkdc~lceuua -udlImdr\pwrmryrld
mddvlrpacaulrhvcdka
lowes, m(snlW4~)Hourwr, dlc
carradlk* Hamr,flmumw d vyor ( d y l
r m x u d ~ k a w n d rdnua)
~ ~ d t a r u l a d Amwy 71.
m y a b e m u n p l c ~ m wmpmlaCnrqudSlonnua
[email protected] d unvwrulral &cb.
P unplicd by the m d l ,busd
W u ( l ~ f k d yh & ? tjm I8 ud
mkmwnphnammr 'Ihcunuura
d m o b r c d w ~ l m r n w n c r 18 percent, q u a m l y ) Comprlag ~ m u y m'8acm'ud'Rllr'
n
may hc afkaad by rhc other
mahrugns(rhc~rmrnol~o(~nt k ~ rfield a k m lnm wrc
m dcgm d penctnnr of rhc
q~fdyhmr(41.33ud28
M WIT k l r ytckl
W W ~ m p o ~ c m y d c prsrpravely),
~ ~
(67. 50, and 40 p e ~ c m .
~ p a ~ u x l r m ~ w r u w c wkuca
d
rapn.uwly) ~ n d ~ l l rnupenor
q
muL rhe other mechn~smbw h
dmnrctnnpndoAmuyn,m
~mnnpll~blc
mr uc uud Endunncc and shnv nmtng rypr d
ualllng coknnct rnlngI WIT
l d c ~ c cmry bc lnhcnlcd in~ g n ~ f r u apurn
l y (a &crtm and
d e p a d a x l y h u c b e y ~ y a f h t r c h Ibc run hw kllr w h cmpusd
a h e r ' l d m n r e m ForenbAwyncm3Y.lndI~rrm.
q l c , mom v~prouaw g m ~ ~ w rsrpacuwly) Thur ~m cult~nm,
gnwh wnh Iglwn m n l d mn
nunpl~fdty Amwy 71, hrvr
moderne wknnce u radllngn bul
m ~ c l u r u r em
)y suppnn mon
vlgomus ddmw (slow mntng) m l
Ilaktolmnauolderpl.nca Sane
rm g d . wll In more lola1dry
dllnn alvl hm pm Ybdltng
m c r , ud more uad Funhcr, dlc
ron~ruchuBcaon)mmprmdco
tbnr f ~ l tuknncc
d
ud y tcld ltll
rclntlw capresalon of BIm
O
n gW
lba hu been dwMd hy OK
ud endume a not consinn1
t h r m g h a a d r ~ r a n o n m t mlhon tn odw cubnn
ww anua WgC-reLcd nqaws)
mWy.'A3121'(rmqMpmrd
Thc~d~me;lrdun(H
Mdmn-dnpkdculltvan)
w*, aury4nnd Rararch In'l
h3U7hraknnrracsmlylJd
w%lllhyarIndnn,dTIWen, [email protected]
dunn (fiw )un,bur buuaarl
USM-AR$ Suncvllk. Mwnupp.
wbmmnha$RcDla&rdr
scybPn Brccda's Wa(ohop, 1990)
b m o b s a x d d l ~ k K b d ladlylgse~morcthntuowclts
damce(Javmrmgdadur.
*amgm)
mrr)@Pmg m ~ h c w n c u h v l r n ~ d w w - d ~ r f i a a d
(OC *I)
A3M
, yrlddd rlBmhanly
Am
rqr71mwymdcnamfwld(ab
kulhra&hlghlylsLPd(<95
wbmnccs4ud7mprmkm
pran)raa~adamlum(40
Uakr~ccitqparurearcasd md M prcmt Lu, nrpwvcly)
4 . 2 ( l p a Y J p r r r m d d ~ d mdugufrand&mcamIML
~ ~ ) m d 6 7 p r r m ~ v m
l d obwmd Thc 6cl that
losr~ud-.
--cbulrdunqlw
W C ) PranlbcumemddmI.
devdopmxmy~~pr
lxlnhmlbrmhqfh$manymardan,dQd
plmu tn MUIFE~II~~C PUI~IW
dun*hbrurmwldIcluucRq
~ k d a k n n m p d w ~
cm tnka I( any
d plrnr
dnclopnsmpmVkdWth0nil
rurhallkldupw1ty kw24u48
hi I m p i m u c w r r m wh*bwm
u& a curMtd ud mJntrin
wnunl w11l for 1 n ~ lwk r
L~dbdnxnruklnmrr~
ncnanudmrrlnisJldw
yrld ~Schnulhm,unpubluhrd
l h d y In prrpm(lun) Thc rhmlcm d lup krrms ud ~uhropuda
durhdWur I M d u k l p h d ~ ~ ~
Ulcmcubinn nuch u 'H~mdy'
rpprmtly doa na npmern k
mkuon, but ddlybd yrnpnn
dmlapmrn Inu d tdmnrr
durlng npmducttnn t u c haen
dwMd In [#her w y h l l m ~
( H A ~ Wand
W Dkn, USDA-ARS,
Slarvtllc. M~uuupp~,
rqnn II
saybemB~IUbrWnR1990)
11 II apparent UII
yr-nlnu!d
rupmur my k utpcnmpouwlon
vnnc k~mttl slow mlng ud ndume, ctnnplrnuw racnrch on
ud~lcaHrnIorrJLFnncc Thenew
mdclpnmmvrrmaipru~in
M ~ n lrulr
q urmlwd m Idem
H m r , m k q ud ynr~~mu:
typ~dl~vcdlki,ud~nlr*dchrngamlolarnrc!mmbe
m x g n d u unnplrutlng bcton
when mrrunng Ute I~V~IUII
mm
undvad m full-lolmncc.
&I*
.
-,*m
l
clrpbqn,ndtbcnadnomt. mplracaathacaaaphtbtdw
b m , Md mucllvm d viruhuc
sg.ban-Phvsrhthonplhoryacm
~ ~ a J l e r e p o p u ~ ~urlude
n C C rccuvc lnhcntancc d
vlmlence and the theoret~crl
pacaadtbchnrl l t a l u w r m l h
RanberrdncadCpadtqyuc
OMo unlr hrvc urnr hurtm d C number of hoar
nadhqwhrhwweeocnprumuoncc gmr I f vtruknce e
Mewhla~nxga*rba&r!ltrtr
~lvc(up~byChcgene
dcpbymnrmlbcMd~Habc,1981)
hu-gene concep). lhe rate d tn~~bk~e*ucrMeognnu
Md podW OapOlW Ul8 ~lllllkd o u u m C f r c q u m y d r r r w m c t
wtll be hater ~f Pmg UMO( lnw
dcgm on rnrrtllu pluub (Slulher
d
le a RCyOYbCUI md ud
ud Smclur, 1973, Bagk R l l r u ~ l male m
dtlute thc rccesstve humazyBota
ud Riukr, 1983) 11& 1110 k m n
U u t m b c l l o n d ~ p l u u b y M m n H y W hg mapom k bear
my m (10') m wtm (R Bbt.
p u b k m l ~ncompnble nces
raulu in C gmwth dlhe c a n p t ~ WuX-mu. Uhanr, ohm, Ph D
Thcxtr m pngms) Further, h g IS
Me nce d y M r d , 191131 p n
tht~ghlo be dcpcndelu upon wy
wnubly miuctng opponunctles for
p ? n a * ~ b a w a n , a m l ban for g r m h and npmduct~onIn
puMe Md lnnxnptltlk ms Thc
typt~al cropplng systems, bul
het tlvl the ~ncompstklcncc ducr
although ltnlc IS actudly kmnm of
IU ~ h l l ~10
t ygrow and male m the
nu lnducc nalrwKr m thc htwt to
mol\ oftKher $ppales or tn the so11
C comprltble n c wwld nuppon
C upmerit that mv iompublc
Hmvtr. Pm! may bc hetemkar
ram w l l bulki up ~nhpenkntlyol
p ~ (Long
c
and Kecn. 1977) and
Ihc me d mpNducl~ond thc In
thus recomh~nrt~onwlthtn a
mmptlble mea Thd dllut~ontd
ampmblc rsx g m w q m a wrep
canpllible wes w~lhhrompat~blc lrblc host may mull In new races
om hu no eflat ~lncathe remmt
Cm~~denblc
h&n gnetlc vanabl wtll sclmt only compt~blc
[ton br hort spc~fictty1s pos~ble
ram ~ f l h Ire
y P a m The ab~l~t) ~fhq 13 hctemkarymc lluspo*n
of ~nrompt~bk
rnccs ID qrcdum
llrl hddcn vm*lon has neg&ttvcunu
~
d 1fgenacwuchMgC
y
pllrnt~onsfor stahtllty of rcs~stnnce
k l w n urolnpruble lad compt
and m s p a c ~tolerance
f~
because
~blem s SI froquent Elmuse err
ampatibk heumtarymc nces an
lnlatnvl rrwia In gmwth of the
u v m e In susccp~bkhost plana
c m p l t b l ~ncc only, there appears
Slow-mtng has& may pen11
b be 1 [email protected] oppor(uNty bt ccmp w e r toml reproducuon of thc
pttbk ud ~nmmpllblerncw of
plhogen but dso m b m u o n
f'q to t o m u * unlgs they cnn Q
- w W n r m y ~
m w a d c d r aqbmroa
fuwson +nous
p l c s ud
Mlmgbuc (IV83) ~nclwkdlhe
conoluuon my k in C L m n
ofr mmor mmt Cocvolutm has
Soyt#m-Phy(DphhnpthDsyrtun
in IIta dp u h ~ e m uhrblcu\g
s
m hod urn lo llhgenes in soy.
Flor's pnc-bar-gcne c ~ m pfor
OYbCUImdAngmberkctcdmd
lWSUW urd V I N ~ ThU C*
dinanmurcphosysna Alm.
c c p u a ~ m t h c r m h u c nun hrc been mtervuung lad
m-RuIhw pthayam. Tbc dingnugwx gurs m the
har.hghmmegeae(rpc4Mid
0cmrcancwholcpLndmd
~ d i n d c c d , a * v m x n
have become prevalent once
~
~
M
Wbaher Ihc gau-br-lla~carcp
lpplitaafhgincbeusedrkrv
- mtiag ud whether hg hr the
~ p a n c i r l l o ~ ~ t e f o r
&nu d a g m unimpwurn. The
important question concerning
stability of a urublc Iml of
derurc is whdhn some types of
dow rontng uc ehhcr non-sckctiu.
or aelcaivc for bss rggrssivc Pmg.
~~ceisnoltsdecavcbydduutlm buf endunm alone may nu be
~ufticiuu
10 pelail adcqullte EoIIV~
ud qua1 yieM mdcr conditlom of
both SKR ud ndiscas pressure.
AnumberdmdhoQ,torlrrn
soybeam lar tokmce to phytoph-
thonmthrvcbmRpbrPpdinC
lirnsarrcmpnd-r.
1984; J i i ud Lockwmd. 1980;
lrwinndLsr(gQI.IPg2;eumdlsrd
Anderson. 1982; and OM ud
!am&mm, W).Fvld scrrmirq
brRngderaaharbandu
pndomuwrmbod.Ahhoughw
~diasmedvpmgsd
lnmwiqfull-saooarlZrs~is
&tidy inaparrivc n run. 11 has
smral assochtrd probkms. Thuc
include: (I) nonunifwmify in the
~ K Mof h g density, i m k a&
gressivenss, d roil conditions
~hunrg
~ tbc
~ p & p ; (2) aoaunibrmityinhaquuwydncawithinthc
W,(3)lackdmndaionkroran
as ud aggrcssiveoas of Pkg
popllaimrinlhcWdmoSein
the Ecybcvlproductronafca;(4) inrbiliadishwbcht~
n
adeobnr*)muns. (Sllrrmemn
n&avhcnROLIh)dad
~mnrLanmnw4lMd
mnuqkrramqapnmnn
cuh1w ([email protected] lMtunty*\ I). u m d m d l y ( r . . Q n P < W )
8 ud Wr
yew, ud I61 plcndhl Ernhad
m c h d ~ i m u l w y ( W ) ~ ~ k h dnWDI
tmknnce ud plur death wnh
Rapwcrbk*r Iwmtn. plqluuamobu**avdrrraldr
5 * f ~ l y
pm dal lnalunly &al rnld ldsnox
aufr m n t y
The pcobkmr u w c u n l wtth
ma (McnlAulnd SdmnmawPr,
rJrm.wtr((rypldM d s c m n l q b f ~ ~ l a lbrm,rdI)lllpnLkUdan
uapubl~lhal,McBlun n d, Wld)
rcvuctvn m tmrslllplc ~dnmtny .Rn -r
mmrynWkmuJSdmdrna
Imulum-lrytr
(t9Wc) eumtrulum d t a u f m t
ud grmJwuv lnknncr rwl(g uknnrc wwrn wen mwdal m
mrcunl\p II Nrm vrnu he lml
pmadura.oflhrsrrmoprmlhuhi 8gn'wfl plmPlm a pcr
cumr*ly m
m
l bk. t lmculum.
I!~ I ksml
M by W b r
mapnrrluaf k l l k u ~ n nutunpl
~
nxrrds In w h prrypc m In
lryn rln-bnd *s dkr pu]
m l khinmnhenncr 11%) lhtmd
p r r r wmhnms bctvera k l d
pafildkn uyhun M l n p Thc
W h r ud Schmn(hmm. )9n*
txulum-lryn u*l ( W h r and
mm ud the ~mrulumlycrm1.l
slwbanlmaurmwmndm
Schmmhmner. W
M l d h plan&
d t kngh d t ~ u cr4rmrurwm
r
mhcaly p n u p ( r ~ U A 0 ~ u t k
upvud fnnn hc pun r f wnul~n(l bcr gmtp Ir-QIP)) A umrlu bul
m be rrccml qu~kly(1428 [email protected]%)
mnlal tn m d b m cm gnwh
m r u w c h y p w m ~ m t b
m a c w I n ) l M cm!mumu d
d l w kx contml d we c ~ n . c ~ g n w n p l u a l a m O l a h u d u r l w . tnprm#(u*, url Irtul
ptm Thc mn d
y k l - khrmnhcnncr 119BS1
Mlu~pnxlpollSmntiunrq
upod 6lm.bwd rn (Olah Ill
bmathemallhtltughkk~a d r 4 5 a a l fhrlblraplrwrbr
SLhm~aheMcr. r)85) d l t m hc
g m Ihtgh 4 u c In t ~nrulum Ehac d D ~ ~ ' b k(1)d 1lJUr)E
@r w d ltw wlua m t cuhcf
bawrm gm kn lmknm ud
m a s u m l ol hdcnuxx rtlallwly
Munty p e n ar p k n t n ~d
qu~k1)(ahmil tw w
u
r
e
h
)In u crbn
lrmr nnflnlpt~vr
ctsrclmwm d hc
w r n y gmtr, (21 yammw bun
Pnlkdm~~mnenItrdu, dkruit t ~ tnruluni lyer dru with t dun
kIlumul try cunfunlrq v(lPn cluc u1
~ m n d p l o n u m b e n s n v d d fmm he c h r her Ih l u a hud
m m wgnificandy c r ~ r e l d r~r~bnerwnhvpudueru
E p - l c d , w k w pmmc pm
w n l ~ tbc f ~ l dtnlcmncc wtmr
L* muunty m plur he&) b,
ccduns do not Thcsc pnxdurrr
haw heen d t W r m rty war
(rm.041. P < O M , uul rr049,
whrh cunuinm hmulrc cm he
d m d . (11lrutwrg lurr wn hang
u n p a n d b f r l d u ~ W k l k r P<ULX)I, nrpauvcl~)Thc c m l v
tnnrc wrc m( u pxd A$ cxpauxl
nrm cltnrly nlu(ed ur rvNln
pndSctatuahmrrr.m.OWIand
prnnul or vnicrtml
hned on Walker lud khm~nhenncr ~nlolenn~
schmahm. l98.5)
NIIIVW.
i
d (4) tk M N ~ U J D
(F1114c) H
~t %&In
~ bunl,M
The Ihm pmcdurtr (field.
ladt~mrulumlyerlarcomlred pnrrnr nwklng v w m url olenacc
1n0Culum-lyer, uui b h l borrd)
u n c o m p r c d u a w g h e w u l ~ m dl with coeh other (r-477.
1fmelolaurcdmrrccdlcclDd
'hmdy was p n of m Mawr'r
P<OM)I) The uoculwn Lyer cml
dumg a &r Muntnn Thc fm
I)aeeaplvuumrwmardMd
k t s (Hacker. 198'3) and h u bxn
ud the ~IJIIhanl
I
tea *umh w n
dlvnurd Thc frvld lcll pardure
publlrhcdelsewkt (McBlarn a d.
nd omW wuh r c h u mulunty
uwrlmlhcneodmvrdhepbb
19916) Only a bnd dwnptlon d
but t feld lolmnce mm m
(r=-046, P<M02) The ~no~ulum w h a r k h ~ d m n t h a l h r n w i
tbc nsurch u provdcd hen
ThecondaumdvacrfmmIhc
I r ) a U d ~ h u d ~ r o m ~ #nmWJ]mudSmnutmana
rdmd wcll w~(htk &Id dnwcr
rwld, uwrulum-lyer, ud slamm ~lppar
, to bt ~ r ya
bnrdmdWrwtthachohrd
KancmfIullIhch*lc~l5o( - k h d un daurc rora A p p m d y
t b c r e L t l v c v a l u e o f t b c ~ m tb 45 urum I& (1-074,
P<W 1Pd r4.076 P<OaOl,
damreunaayr,mrcmtbc
W
u
g
l br unpcoved Iokdenacc
raps6rdy) Nehermcarlwlnur
WarcIhcpbnlaKlmdpdr
wac damuncd wag 45, loybeul
r m u a w b s c b u e ~ t o ~ at b C L * a ~ ' 8 f w l d r 0 n r O X - ~ b y J b v ( l t l ) m d ( & m o n
4.FidddamDemnrwlacbad nkdwhmaury*&numcm- natFcIhcplaa,mCmrr:LW
aacorarthunbrdaracc.
rn~munLOdvtgor,Rd~qsl --IrnphacdmC
n
y W q line hrm r cmr, if aland endurance. Howtver, the
arpabrpolypcnicbaitthrtvllr
m d u r of
~ r~line
~ will be mukcd
~mapsiticrbdionpcourc.
ifitnarlautcetrbsuulhacd
QrtmftcJd cmrmurra! fn dun.
n a b e c l y u g h m l o ~ n d u c b d dhrppmro1Ilobelughly
tbn,motuymsdlum,udmOdfof
aokrmt. la k t . R @ y h, more
vlgnr CVCM ~f thc Itne hu link
moculdm Punkr, fvld tctl drtl
ldsmccdunthcrmadrnarlina
o a r e l * b a a w n h y ~ c l d u n d t r m e n d u r n . Thc m u r u m t d
dillanbmnrypkuhdaufm
~ u ~ a m l w c d l i i l e r l m y n a drrclapd by W r ud SchnutthePncr (I984a.e)usiw thm qcla
cornspad wll lo thc full-gcuon
the uwrculum-lyerurt W k e r uwl
dnammsebctionrpcrifrrl3.br
nynnv in thc fuld. Superior perSctarrclhaacr.
ThcpcW
cha tnit. Thus Ripky was thought
h r n In the fuld with multiple
w u l m n obacobcned tncluda whole
lo passers one or r few majar genes
ma in combuutum wnh @-rim
plrrr fattmu,mu rcsua*, sluv
Ud v l p n KaKs fm me inaulumfor lolnum.
mnuy uwl endunre If m t h n
'IbkmrcmRgkymsmdi~(
o n c d l h a e r r p ~ l l ~ ~ n I ~ n e . l h nlyer
e !en h l d lndiutc wpenar
pldrklrstr'sThtris(Ziwmiy.
wtll be muk~ngof the other t n l e
m m n y s ~ f i cloknmx i f whole
1988) reported in McBlun et 11.
p h t or rant g m r are not present.
fbr ntolpk. III whole plrm
1987 and published elsewhere
If rcslsture is prcsenl, and onc or
mtmnce #me tr p n r m in r line
more compatible ram art common
(McBlain a al. IWlc). A brief
ud m compt~bkme extrll In the
mthcf~kl.thcwdthcaxnplibk review is pmvidcd herein. Thc
fvld, the Ilr wtll a p p l r no dlf
kmH l h n a IINwtth a hvh lcwl
nccr in the inoculum-lgr lest
cultivar Ripley was fwnd to yield
of race nnnrpeclftc tolcrrnic
h l d allow enlurtion of slow
well a Northwestern Branch.
whclhcr 11 a d the slow.mt~ngnr
(URDC-OSU, in spite of there
rolling. Progeny lests from a
a d u ~ c lcy p I f lhc 0b)ccIlv~a to
ICWKISS will bc needed to enluatc
hiq been a ,were avenge lml
a linc If mistanv IL>rlIL;nmlaces
of Alp damage b all other suscep
&lop llnea w~th~npnwcdm e
m p x ~ f Idennrc,
c
UIC field oul
IS prcscnt and obscurtng race
tible entries In hnt lest (Ripley IS
a MI dequrtc Htnucvcr. 11 w ~ l l ncmrpectfic tulernnw. Evaluation o l
sufiaptlbk)mat 2). F i eJ*F and
e n l w o Ilnc wry w d lor ovcnll
cndurancc In ~ h cpresence o l
moculum-lyer sod s h t board tab
resp)nsc UI A# q a n t l e v d
rtsiarlann or wry slaw rotttng and
withm4also~kdR1plply~
mchantnm ~f hr d m an taken
mawring ~gcnlalFd respomes will
wry lokml. Ripley plantr s h w d
q u i r t nnv merhodolopies.
b t h nutuntlon bqbs
a wry narmw mge of phenotypes
with thc slant hurd technique of
Ihr Nllwlron of slaw m ~ n and
g
lMemnce In Cultiwr Ripley Olnh and Schmitdnmr (M)
endunl~c requtnr tc\tlng for
using
m n l of mot m The qlant-hoard
In P85, rhc culuvar R~pleywas
tact 4 (0 to 2 mm of KN); whercas
en using r mmpttbk nrc wtll
relnrvd by thc USM-ARS and
'Harper' ( W m t l i ~ and
s Mr. 15'74)
enlurk! Lhe dm-nuttrq l r ~ lrnlnuh
l
(W1Ixau(Cmpa a al, m,. M a m c p n o f 4 5 m r n o f m c o m my tnfluenr !ha1the ep&rnl~\ or
I t s u n i p u c i n i r r c o i n b h h o f ~ h porad to 50 mm for 'Slorn' (very
m n c x ~ y h v cInordarlotcstm
yield, shon suture. and h~gh i n ~ ~ k m tk
).
phenotypic difdunocr. thc plplrn~sgrowth In
toknna lo Alp. 11appsrrd nsisk r e m supported thc rDrmp to
~ ~ p ~ s e t o ~ m u ~ u d u m u [email protected]
l owdcr
f
heny di*
pressure In
dunify mjor gws for denocr in
m IS necdal Unfnnurutcly thl\ IS
a 1984y'itldastatthcNonhwa*rn
thc F2 generation of the c m of
ahydmp(wl~*~1,wChe~mwihdB ~ n c Seslan
h
ofthc Q9RDC-OSU.
Harper x Ripley. More m n t tests
shnwsumllfFF*dmuchbymx
nur C u m . Ohio. But Ripley ha
shoorirhaamnaiamanrponsc
M K d u c m g m r ~ , u n l a s t h c prwcn susceptible lo hMacayl
similu to Rps, for na 4 CRbk 3.
m IS a ~ l rIn conmst, thc
sdrpsd hMcBlain a d, l991c)
inaulumnwith wed mxs (1 Ud
mmulum-lycrm whtch Nzlwm
4. W~lmx, 1986; .nd 7 and 10 in
knisrort~linthemot~
rhmc napMIP w, will lllw thc
Ohio Yield Trials. unpublished).
Rpt2 is in Huoroy k k g r w r d
d d l t m of mot-m mmgs Un*
Neidrr Rrpky, nor its parcnb. woc
( M 9 4 ) .'W:r gnabpnn d
crmplabk rm, ttur ulll pama
sek*cd d i i y for I d e r n . It
Ripky, was rhovn io bm naIm?srobcmlurd&slowroccurg
lepurr unlikely thr the tugtlca
spaifr eiemce w h imcul.pd
7hcf&lcrtucbrrthrnthc
-*[email protected]~
utknxn(r~hmuoaacl.I9881
R O l c y q p a n @ ~ ~ ~ @ =
br rmt tautracc
No~ckPtrcauklbc
dwqwbd~adwdmbumnd
FI,hrmtymcrmaerthPIhc&m
bmdorMdIdau*r~csudl
aa*gucrmuvr,Wqudtbmrcn
bcnhaa d tokmrc In this cnxr
~plurrcmhCrJuabaud *a urn m mlubk pcallcar d U u r prqpwr' pcrlormrarr
tnher m Ihc dm! burd a in lhr
fdd Thnr coneblm mdfrmn
Md h m d rcmc hcnMnuh (hl)
wrr (r* 57, N= 34. P<OI, ud
r- IQ N=DI. P<DR rrspar~dy)
MuydmcFzphaqpMdnbc
apcs.swnngOmmmtmlhctrq
ya hmq h~ghpngcny mutt 11 16
drffvult to undershnd hnw ewqm
aniM occur plvm Ihc ~nlunakurn.
w of wund ud ~nonrlum kx
thcunFlplua~mm.
s k u bunl a c r m wa more hlghly
hcnuble (r=MN= 54. P< 01)
Rlply apnn lo haw a bnn d
ma ralsrnm mlu 0 @Wac!,ud
sohthramdlau~thradrothoughl
hl bc qrulrtouvcly uJwmd I
lk
49 rnfauolkdgravmgd
uoDs rrd m
r
u
mpmdum d
lhcrLnbonrdshnrqueudrhc
8*ryb-JlplPmrhewg
~10kpnlecrrndasllhc
@man scgrcgrtcoa d a t q k
g~brrmtlaoPacclJl~ur~m,
~,nwth?rIhc~b*rdmr
dwhcldap~nwtlcd~
Dmddamnmaraaonrdm
~~hPcs.Dmahnmrnt030
planandasld~ffaarcsdJa
blmmdmmr Im501cakm
tkrlrmbmdw(McBLaad,
198n.rvbrb~Wpcrmpu~mJ
s~bneet4.7andloud~krmcrrn~r~4olRlpky
and ch&d u mruutrd by hypooayl Imulmbn, Imulum-!qr
and dm-bwdm u f # % !
Iky ioknnx. Rwy h n d that
either a quMtiWive gncnhwac or
r viuul fzld evlluuion rmld bc dhctive in m n i n g for lincs which
yielded mll in the prraence d the
diaure. They also found that
tokrurc ww highly hrillble, at a
k t c~mpsl~ble
to plan( hetght and
~lunt(
y MI
Allhargh
. they wncludaf t h l this continuous, qunntilllive variaticrn uas polyscnic, it is
possible hat a few, relatively major
genes for tolerance exist. This is a
w i h l e o;plnnumn for obgining an
entry-man herimhil~tyof 0.n In
any cau, they made gtxd progress
in thm cycles d recurrent seluqion
for tolerance in a population
d m l @ using H) high-yielding.
tolerant parents bxsed on replicated
yield tern of cycles 0, 1. 2 and 3 at
the Vickery, Ohlo. Phytophrhora
nurscry (Fullon silty clay, line.
mcric, scric OchnyualO which h d
predominately w e s 1. 3, 4, and 7
at lhaf tlmc. The msult was a grmplasm nlcasc (1984s~)with an
impmved level of tolenna. In
suhsequenr rescrrch, dmlopment
anl selation of 1 1 ~fmm
s each of
the threr cycles of w u m n t selection was initiated in an attemp to
pmduce high yielding Pmg tolerpnt
l i m for release as cultivars. The dfon w r ~remarkably unsuccessful
and ncedcd study. T I C d u a h d
lines fnnn cycks ? and 3 and pqnblions Jevelopcd tium cycle 4 after
malthcrtlon ofthe m m t .sclec~prqlnmforhgDduMamn
W r ud S c h m i t t h w r (m)
Nllwted ~dcrindpopu&"$
h o m q c l a Q 1.2.and30frrmr.
nn selccmn for tdenocc and
p r o a d u ~rhey usad cooli O& r n r
pcr cycle ud usad evalurtions; d
tolerance based on vgaative vigor
at the seedling ltagc of field gmwn
St's rc the basis lor sekcrion of S;S
as prrenls of the next cycle. The ten
original cycle 0 parents used wcrr
lugti-yieldtng, tdcrant bRCdq linn
and cultivars a d it wds presumed
that tolerance oauld be impmved
without aftkcrig genetic varial~on
for yield and other important
agronomic tram. Dam fmm plantrow and yield trials of the
Stderived l i n e of cycles 2 and 3
and of S, &rind lines of cycles I.
2 and 3 in 1984 hdicakd that y~eld
and other agronomic traits (klght.
lodging, and mmrity) had been
negatively a f f a t d by selection for
tolerana only, ad that few, if any.
high-yielding. high-tokme lines
wwld be idemfid. This uneapMedly negntiw cumulative result
d c u l t i w ~lccfionfimn the PMGT
recurrent selection populalions
required funhcr study. Raonsderation of the aurl mi(under rimlimin the PMGT Rcunm ocleaion
m l t e d in the folkwig CMEIUsions: (I) seedling vigor in the
p r r ~ o c eof Ang was undcr d i m
uscdbddamineifthcmodifrsPan
skctionrndsssdliqg~akrurc
MS SCW
indidy; (2) d i m
w likely to incruv success in
sclectlolldsadlhgvigormllso
high-yicldntg m k m t lines
hdina&kaionvigaou,HgCPM
dplllyp=dbburc.Thcstudy
of~yrk4limwthcfirstat1empt gmwdl which anuld sckct lor
~ s e p n c l ~ i n c l e s c m d cgmmacJlyinknr,bggiogpm
~
d n g ud endunaa in
liocs,rod(3)rccdli~s~Ag
ah.rrpa wrs oac wtxa, ma*)
ooybun.
d MI-ldaM
bulmylPldmtcnbnnrrwtl
d f l r r r r p d k hwclru
han tbac ~ l a c l u s mthu Itlc
m l c r ~ l o nprocedure ured fur
daurewslnlUsl,paucop
dua q m n o m d l y sknor Itm.
udddnJrlaaonkfutlexpm
uoadmknnctcmrklu*on
Thc ab~lttyor l ~ h ~ l lofr yRCUIRRI
rek*~onfor A#u>lenvntlo pn!
duct h m supmn lo cunrmumvl
nmres wild rnt~callymlw rhc
uvfulncu of he pnrcdurc hxruw
Ilm produced wlll he useful 13
~rmplum
or cult~mnonly ~fthy
w uiprnar m d e m ud 11l a d
wmpanble In k c t n l u UI thc~r
(#her heromcmponry brad~npllnn
Funher, Plpcr and I;chr tlW)
lhal cmnltonal crnue\
mu t p m r to h r d~llcmrmuc
r n sclert~on
~
pmEatum for yield
That ~tudyalrbllshed convcnt~mal
cmrta a a ,wdrrd for munun
ment of runxsc w~tholhcr b d l g
mnhods The crlt~cald~ffcnnce~
In
thu w d y compared lo hat of P~per
ud khr (1987) (besdes the d ~ f
fe~mIn mu mlmnce, ydd.
k @ t and lodgmg v\ yuld dm)
rmr that mC ltnes fmm mnuonrl
c-mnadlrstly
&acdkrr
~okrnrreto h g Imme ~ndrrccr
JekclKIllby~brywlduould
he apecccd)and Ihu rhc m n ~~In*mrllyurludcddl
mrkMe lrna w prrntc, IKU just
~mmPMGrT&hntrsssybr
Agdemrcdluwsfmmaxmnuaicmmwlulnduromctnslr
wedcocnl~doanplrcIla,
mrnrmtma-dfa~o-
trm/era4dthubdmrbm
uredtrr~mrdecc(~11pnor
lua from mmauocul nmra
*poaqparard%Imfn*nqdcl,Zlod3~
Ihcrs~lmdatbn-
wacthobculiDDIdQe(sd~
Ir*dhWhdSebmrnbam
o k (prbnrrlly M . W ) Po
ccdurn' c m u a (we4) tbwr
(IPMa)wouMmlheuaoufulm
pnannsurpomlamplwn~~ hMhm*lbpcbdbbbenm
cubur ~ y ~ l k m l m r u h * J I . ravlNMvns uwl UktIon crlmrh
m low s yeM. all, md pmr @
r C S,dmwd l t m Ram PM07
A m mm damn p
bdgq, hm INk nluc~nu
p m wksr chy m p 1 m l y ~xkuowurrrdnlbmdrnlhcbu
w b o a u w d h q ~ m d numhcr d u d k llw.hnm L~ACJ
&krmun uua m lhctr plm
I . ? , ud 3 Cllnelad mptnuecl
SelsLlm w~fhmud betwen Ilm
lnnn ro)oaxmdmnr mulug m
drrnrd flW rkard hkIalll Sl
kw yield, I& uatlm, lodply
luws Imm wh cycle d m u w
or m u n l y lhnukl k rrlpDd IIu
I d tm y u r t1( y r k l I c U I ~ud
+lat*m
d r m ~~III
yrlJlq,
d n r t , k x l p q ~ u c w I m r p w d a l nrluuum (IN I((nmmuc I ~ I Ud)
D~
rim wulm ha thnr uua IC
.r munty, k l g k ud kxlgln# worc
m d after ukum fir rtlerurv
r k h l a a h c y k Cauumhtvm
The fimt ~~JUIIK wr hi deter~ m u l b d n ~ n d ~
mine whhcr h e o q ~ murrcnl
d
~yar~yk~m.(Vkh(~n~qk
viearm rhcme fur w l c m c w11h ~ d ~ d d ~ * u l l e Y ) I r y d c l W k
mbKqwnl u k ~ i r m
for high yuld.
5 ) F x h rycL pdum laa br
shon aMlrc, Md I n l g q rrxl*C
c u h ~ w d d q m m aA p m x l p ~
m csch ryk uuuld r e d in nupcma t 1 ( y r W ~ d h h c r l ~ , l m
fnmahc)ckLxrb;tmdp
lma far rcluw a gcrmplu.'~rnor
cult~vnrr The rppmprutc ~hnkh d S,&r~vd l~ncrWM uwd bd ~n
4. S q w ol In(lng ol PMOT md ncm.PMOT Urn
Vsar
E.dy pnwaon lertlng
dngb pod k r a m
N ~ ~ ~ W O TPMOT [email protected],
Non.PMCn
PMOT cycle,
I#rA8*qunrrdUhgdPMaTcydr4
w
d
epk
CI*ndu
ywr
1
lW,l9M
.OW~dMhr,IldluDcrhrl
15
iO(rW tQe67
20
1967
[email protected] wlntnnw~ry
S,gm~nOMaw*arO~rulm
udlhORHthUlWlnlUlll82IWmchS,
30
1
W
Mattdals and Methods
~RrlhrmYrcdkh
~
2
m
l
!kkcnm
30S,,prcS, pram In2row ( O B m
w l w ) pklr 2 4 m brig, a'Iheme(horburdbdKhpmc
d W ~ t 1 ~ i t h 2 8 ~ ~ ~ n r d 5plplbarmbthuq&drrmr.
~ ~ . 1
check each 10 plob
mn rtlavon bf p h y q h h l
S,', with m
o
m lyl. lala of !adgln~pmnr
okmakhxaplb(rdod(H19n
llm compand to 'Flyer' and 'Sparkt
ndschdmm8841m)'Ihc
dlnc8M
p r a a u s d m h r m ~ ~
S,z'8 01 Weclad Sl'o whkh m too tall,
war w; plblr smug A5l)MIP
llltr lodging prone, or wilh pow emecgenw
(hrmAW.mlfarentnaiskaun
.
-
dlrrrdd
.Remaming S, ,'s (19%) hrwskd (w ywld
.S,,r' earlier thm 'Cenlury 84' WW
.Remainder wloW on h i s d ybld ae 0
percon( ot c W se( ot 5 c h w and
malurky, to tMai 95 liner,
35
1988.1W
-
.Thr 95 W
e d S,, llnes and 5 chccltr
loalad using thn Inoculum-lapr techniqus
buf -ring lor plant death, md ml,ud
"w
- L u 15 liner, mod to beg~ncycle 5
.SIM
pmcrdun, 4 WKI to WW early llna ~rwnequal n u m m d rd;Plc)
r
md wldr cwm and uleciiont lrom t h m non-PMaT llrwr an uud
h-wm)
psllLhond40W~~P=
hun~tunryGoupsObIV)(Fehr
d Omz, 197Sk A'F62CUM (fmn
AP6).
(from AP6).
CumbnLnd ('CgRoy' x 'WdI"d)
(lhhmh and Rhr. l9Qb). 'RUI'
(L66LU7( W W s uacr lux) n
caud)(lhtm&dFthr, rn).
w(CIIMxBrmr)Wb
n PI. 1980). and n d r pnmu
culpvus Agnpm 'AP26'(unknown)
(fmm Agnpm, MISSUHI. KS),Northmp King ' S W (A55-5629-4
('Roawlrt' x ' H a w e ) x PI
257435) (fmm Nolrhrup b n g Co
M ~ I sMN).
. Pfittr GeaetKs
.
Y3Um'(Amcoyn~ruda')(from
auly ~ n e n t i o [email protected] (Cwpcr.
1990)OI impm\r the pmb~hilttyof
obtaining rupnor linw for r c l m .
Rw sacoad cbwluofthis sud)
war m h i n e ~f the modifKd
r e c u m s e l m p d u n was
urmdulinvbctiagslpriarLurs.
An upcriment directly mparing
tlw old and m pmtdwrs would
hb#lLhemoslrigo~wlyb
~thirobjstivc.&nrhcprimuy
objcotivcfwthoJc f u n d i ~ ~ m r j o r
portion of this research was
impnrvcd culnvus. The continued
use of an obviously inferior pro
cedurc for producing high-yi e l d q
tolcm culuws which cmsumm
25pmdaUrreourascouldm
bc juslifd. An dysk of s c l d
pux lina as uscd in tht first ob)aaiveuwMnquircrlludsix
mon yurs. G i u n such m inveammd of time d nswras. awnpuiransofS,,lkwitbtlwhe
.
Pfier GC~CIICS
Inc St h a .
MO). ud Rdc 'B2M' (Corsoy x
(fmnRde Co Inc Glen
HKn.WI)Thesewcreacleacdrs
[email protected] parem wnh y d
compnMe or eqcul a thc y~eldd
the he culnws NulrMe tn I977
'w)
.
d v n t h & ~ a c ~ a d 2 b b
4Dma l m S s a l c ( 4 . 2 b 7 0 0 0
tlwaUnatl).uatdIrn10nalc
(McBLmdd. md))h i m hmmc~r$s,--
3
~
rsuaDaoncal.2,KAU.U
Ubl Bllrdurrved24(Mk I)
AIbatmcprmr.So46hrpad
field tolerance rad wrlrble
bPwlrspmc(ryprrldm
ddrk6Gl'MdSWmsflnm
mmbmqmrpMdbslrhutlhr
mdWdwdrvdopdrluam
W & h dT h l h w m
P M 6 Ilrn came fnm hlh p
rararocr)hUdrnpmayaa(J nbnrkhnprbmvnrdtlr
bat PMUT and m-PMUT l l m
-.Nonhnp~~phonc~
K n a m W . R 9 0 ) ~ n h m m r c m p M d m l 9 B b Wud
thc poonu m k m rar d the
1989 Tolennce tu R
t war
plmts Wlker ud Schmlohrnnr.
muwd u q r nmn rg ntlq on
IQ84eJ
plan& g m In a r q * ~ t cthme
Thnc cycb d nmmn~sekctwa Mcplrrtwn ~ m c u l u m - lI~ Mr v~lh
~
d t e d m aelecuon of 10 S, Irm
rn41nthcgrmrhtnueml989ud
w h r h ~ 2 8 c o m p r e d t n 3 1 19PO (a rhe 1988 hnd 1989 ALT
m b l e w rure lor the prema
tala, respect-ivcly RIM wm r o d
M k c r md Schmnhcnm, lPROc\
fmm I-no nun N( kl #)*dl nrm
Thc &at IIIX word 2 5 w h ~ h15n
road Checb uaal I d thclr ~t
mqlnal ,mpmmenr r1 bcrt, ovcr
Inon (o I-MT 4) WIT OXZUR lea
the btllpumt. A76304005. whlch
tmncly ~ntolcnnl). Slt~rn (In
scored 2 The small m m p m m
alnw).PclL ( 5 k w . m lolnurl
r mlenncc r a n in b n c I y cycln
but not root rcslnunl).
of recurrent selection umc at the
WYl-I-LC& (nxn m~mn~),
uxl
upcnse of nllow~ngo h important
'Rlla86' (whok-plm~rcxlrlmt)
agronomic trulr IO dr~R A proper
Tbrsc c k l u h m churocr~ntrc
dunomdthenrmsdlsuncn
~ r c ~ . m n r t c ~ l n o f 7 50.
l o70
aclstrw for tolcrnnce must c o w
1090,351050. 17o3U,md IU
h the performance d Irno
la 1 7, wib thew n n k q mrmd
dcnvcd from sekcrcd SI i ~ ncom
s
( U l c 6) In Iht 1988 md I989 M
pared to llnes obtalntd from
nwd L~ne Tess 1AL.T). the
tndrt~nrlb~puennlcrosses S,
vmmm lor mknncc mtres mrc
ud S,-denved l~nesfrom PMGT
Cycle 3 and F1- and F,dcnved
I ~ n sfrom non PMCT (mostly
btpurmrl cmsscs) w m MllW
and sclstal us~ng a a~nglcpod
dafeot pooeQrc (SPD) (modrfrr~dslaglcsecddcscenpocaium
d h , l%6),drmdfdculy
gearrrclonpoadurc (Em)
d s r
to t
h d Cooper (1990)i l b k 4)
All c y ~ kI rad Ihc cycle 2 l ~ n r
whKh wrc evlltukd by SPD wrc
w k k d by Walker Thus, d y the
+kUimhameyrk3SPDPad
cyckrZIPd3EOTtvcrcmnrdnad
wnadamdouoduyfbebrteder
wbodd f b e u ~ n d p u le m
~-(PMOTmdmtm
*plw-dh
dacwJhqkpdJDMlfmln
c*Id2Or.dl.~pmc,
k~y~.urlmrrdcn*lymkm
la*nd~mlsn-spprnn
di~k1pueLaolhmSPD(19113
k1198(*o m Jrun))I
d w lhr
Qornr,npwcdrnn*mr*
h*blml*lF=d
yield Icst~ng and rlocurm for
rprmunrmpnorbmfir
hbvucrable5) rmanaw
hcnraumlmrwurr~
# ~ r m r a m l d ~
v ~ ~ d ~ r M d q p r
vrxr trl lull pkn a (kKnns tune
~mrrIhrclwuenwryncu
c&ck of eWv entry rcduciq h
(ktri: 'Iknychidu) d
a l i n e U * i c , m a i e O c ~ n . 7 h u ~y~cldlvbnimwptrmdW w i n g to hrlf compurd lo W I J k
ud schmi-,
19&0c).
on nourofdcgmddsmrgc.Clhc
dwgtuuiclleffkdtonvid*lec
d c g n e o ( ~ w s a f l e c c c d m 0 r e rhe h w t PCscore ( m r
rion br aceuive vcgcwin grwlh
VI,Ohwr~(Puka,~ihydymil.mi&
w.
100 pcrcenl of Pella) wa act e q ~ l
byporim~rywthealgcdrhef~ld
lo I and Ihc louenl score (near 0,
by WYPC.)
unollg
a h most p h W M died) was rcc
the hrmsted S, I lines rm baxd
equal lo 10. the remining rom
on yuld ud maturity and 95 ltncs
wn inlerpolakd. l l u s m f o m (6 pertent) wm identifwd for
tion will rault in a
c
e rcwrc
d w l i o l i a~ puvntial prcnu ofcyclc 5 d~6 mm of pure I Ifor ~ MI a I to 10 scale i f a hihly inmluation w~thfive c k b in a t w ~ tdentnt c k k IIIC~ as 0x20-8 (0-10
percent vigor, rorc=l0) dhighly
replication. Lnicc yield trial.
ThF 100 cntncs (95 SI b'! and 5
tolerant andlor ICSI~WI check)
checks) in thc y~eldtrial were also
(%-Dl+
pcrcant vtgor, m = l ) art
t e d in I xparnte t h m repllcatmn
imluded In hiest. The plmt death
score will bc Ibr any pure line with
tandorn~udcomplete block cxperimcnt for tolerance to rnce 4 in thc
rnres~stancofor ruce 4 or acccpprccnhnux In 1989. An anemp was
able s l w - m t h g tolcmce. These
msde to measure thc IIIK
types
Y
of
uc hacrogeneQrvlines, so low plant
k~lerance,nra( mislam. dtnv mdeath scorn 2 to 4 combined with
a t and
~ endurance.
moderate rcu-rot (sol w-mun&!l
T h m tescons wrc determined for
scores of 4 to 4 and madcm vigor
1mr*id3to5mayudluucdrlm
u)lerance u) rnce 4 A$ uslng
number d c m r g a l plmh that died.
uc segttptmngfor d r * i i m e or
m n t of mt m observed (mot
hYKmDdelPlCbkm.Amulnabe
resi~lanccand slow roning), ad
rn-mscore combined with a lowplant vigor (mnl resistum, slow
vigor score should indicate a
nming nnd endunnce). For sake of
moderate lcvcl of slow-rott~np
tnak~ngu s y cornpurlsons betwan
t d e m c ambind w~tha hrgh level
the t h m scorcs, all xorw were scr
dendurnm IOIC~UEC.
A low WHto rhe same I to 10 scale range. For
m scnre and a high-vtgor score
desd p l ~m I s : I=mdcPd plant,.
would indlcnte a )ugh level of slaw!=I to 10 prccnt dead p l m ~ up
~ ,a
mnirg u k m but lit& eodume
10 whrh SI 90t prccnt dead p l m .
tolemforthesdlunxlntdmt
Thc amount of m was scored fmm
lhsl was observed. Rrm resiuam
I-n~m
to W)=all mroltcd.The
u w l d k chuvrerized by very low
original vigor score ( W k c r and
mot-m scwcs (1.5 to 31)) unless
a k a t d c m t h c b u h d d y m ~ ~ n . Schmlthcnncr, 1984~)is in percent
[email protected] U n h a u c l y if t
w
t
rajsmcc is scgnlptipg, it un't be
ty. nhon a medium kigb, gad
andinwncdcompdtothcdcad
l o d g i iso once. gmt a n e r g ~ p l M l a n d n w m s c o n s . T w o
diw\guirhed f m sbw rolling or
and y i i (if obviously poor the line
udunaa tolwure mechanisms.
mat if&^ wtn m w . Fi,
H m , i f a line segregating for
wa dkudal). Visual rlec(ion
the pnduct of numbcr of live plan&
*&kE*dwilatheym
andrmnpluvhclghtforcrhcnay mrcPisranr.~~slowr-mttulg
wain w don,ud the m i n i n g wrsdividsdlytherpmcpmduabr ~ d m d u r u r e ~ i l w i l l h s K
alow(2m4)smrcrrdbescM
lina(19 pcrcew) wn huwdfor
the check cultivar Pella limes 100
yield. A miti& lpplrrimdid not
pcrceclc.~mahodsubaitu*sa [email protected]
tdaraec(4l)aQ).'Ihiriss&
ungk chat la an IlabAad
prrwot=-fmm*r
nlhathadclma.TheKJbn3S,
lina fran cyck 3 wre i w r m u a l
inaprurldtalklmrhewmmetnd
19B5 and 1986. The So sccd
p n d u d ws Knt rl rhe Uniwnrty
d RKm RCI)I mnd ltnn Swc
Univenity wlnter nunny In Puerct~
Rico. The S, wr gmwn at
WRW-OSlI. HCmw in W87
Forty alngk planb werc xelawl
W
o
n d i c r nmmry and rhnner
height t h n Willlunr 82 In cuch of
41 S19r.In R88, m augmented
dcaign (28 repl~ca~lonsd fiw
cherh, w~tha check w r y lcnth
plol). with one. ~womw(0.76 ni
wdL) pla p r entry, 30 S, llm
p r S, ud fiw c k b , wu uaed rm
the Wst Bvdycr firm (Canfield\iIt
kum urd, fine.lnamy, mlnal, meric
Aqu~cP ~ t u d r l futl the OARDC
OSU In Wtw~rter,which had heen
plcntcd to aybenn for five con.
lautluyunuJmwhrchhqM
han & a d .The m w n t of d
milabk frvm siqla plmtr wwld
nnt p r m ~ field
t ulennce wows m
be obolnal in the sune m u m r thc
d i m n nunery In Vickcry. Ohlo.
The maulum.lnyer whnyuc wa\
used to abtlin Pkyp tokmu sons
on the liner atlectcd in dv held b r
yicld ddcsirnble q m k plmt
typ. SI hmilies wn visually
~~
kwny,
k b 0 0 U lhU Mlpc
nbru*arpmtdbuncuc)ck
or ui r scum drlnglc plml rkrt m s Cycle 4 S,, llnel wen
rbFoed on the b r s ~ sof qrrnmmr
prbmunceudakrrnr-6(
rrackctlonof pun llncs for germ
p b m and r u l t l w m l w l r o n ~Thc
10 ridS, Ilmmn tcrlod in
a t w rcpllcauon. r w M (4 a d 7)
rsndomlzed complew block &sun
,
-~pcnmcntd~planb
~dsouwcuklcdmchypmlyl
w~thm
s 4 and 7 to confirm that
[email protected],cnor @s,-k hd been
rcdrrmlly unmduwl d rkcal
In s d d ~ r l UI
~ l t k I332 Si lines
alcacd from PMGT 1360 F!
I~nes,fmm adapted x adapted a d
1030 F, from w~decmsus were
enlunlod in the umc 1988 wrt
Sekalon crltcria wcn !hc samc ad
95 early F2, ltnes fmm cnch of
PMCT (wkcunn tnuns~ry7 pmml)
and non-PMGT (uclccr~onintcnsily
4 pcrccnt)crmsca wem lcslaf ~nan
a d p n t two replica~mf ~ l dr r d In
1989 with t w common check
,
,
,
PMOTImrtEadmttsnqnmc
eQIbcmPMGTlmemmn5,~
daInuyluvnckrk~xdk
~tmmtmdmxnpplbum
NmPMCT luw mam wm: mnl
ha4dmlLrlmuariRdndvn
mcmaouihhPMGrMdm
PMm, bscunc cn mr ~gnxu h w
umPM(TTllmuutwuldh
buRd Im m d P M q mnd lrrl
PMCT Srhvtm kx 41mwrl
lcmnhk k* 3r uon'101\hrl
txlmr H M W w n u r k d g ~
pmcIhnmPM(TTImrmmCm
MdWkmrryrld
N
1 PMGT cylk 2 5, lm ( ~ f
10 Wtuch WIT u d s pum\ I J
c)ek7)prmhl41parmtfm:w
pm: l~nrhan k KT' pnrakth
&brrd(ixreplwatrnmWR7hr
~ApcnrndllllmfWltsd
m ~ ~ ~ ~ ~ b a a l r K 1 1 2 ~ ( f
h Ins (441) w m dcrc)yxd fnmi
mmmnnsanwhhatrpum
bySPD7hu~\trrdhc"uuh'(ll
a d r d al & a m pn&
sdaawnmNqjlRLhM+lyh
NO pOceduIEs
ud SPD) w
-yunh~~ * u x l l m o n d a J ; rnu)
fl ~)CIUII
d ~ y 2kllnea ( H M W
ndHMSRlll)uae+lralhrq
m t k M T m MmanpvnrlIi140
(U p m d ) d lmfnm nm-PM(il
oaM(TaMen T h a r I m w a c
Ibosdvrnccdohngwnpllcslnm
rnbadandrW~diPI
mdh1981yrWw BahPMGr
hmruae-Mgnsparmh
avenge line from wn.PMGT
~norvlsdmhaWRll
frvn yrld, [email protected] HM8887 M n
gad-Inl987,thmPMOTeydc3
popllnmmcdklcdhanc
EXJrpmdwrbr+bmandpuc
Imr I n 1 9 B 8 h w a c 6 2 & 3
hmds*d.a26prrmddrnlI
# * 7 . l b r k m u m d ~ m r * c b d f m m ~ m b O I p Y m r fV W N m ( P M O T ) m u r n n (
[email protected],lowss(Mdm~nd~lIXM-PMOTmnr
In th Ohb h m w d Lim T W (ALT)'
YCIlurdm
Stnin
Mauq
dm
RM
(sep)
M.
w e
(m)
Elgln 87
HMWZB
HM M)29
HM W6
3.0
1.7
2.2
18
2.2
2.0
3.2
2.7
29
14
2.7
0.79
1.9
3.1
18
20
27
17
27
23
2.3
2.5
3.4
2.2
33
2.9
086
0.81
084
083
1.10
1.08
24
083
1.5
n
3.3
a6
as;!
24
28
34
3.4
3.7
4.3
0.91
0.95
0.92
271
57
0.90
1.6
23
2.0
2.2
2.0
25
26
22
28
10.9
A3127 r Charger
Elg~n(5) x L24
A3127 (4) x L24
OT1250 x Wllliclms 82
HW79149 x HW79022
PMGT cycle 2 ( E m
A31a (4) x L24
~ 1 (4) 8x 1.24
P M m cycle 2 (Em)
Wlll x CHllury &4
A79.138012 x A3127
Non-PMGT Llna M a n (n-11)
94
0.74
0.77
0.72
0.78
O
M
9
12
7
10
13
A3l27x Chugcr
Ewn (6) x L24
H
W K Ontury 84
PMCn cycb 3 (SPD)
Zane (4) x HWTB149
Mean
rn
Rank
4
(m)
(ma)
6 kc.
1068 ALT A
Comd
2'
YW
WLodg
2
10
49
26
1
1088 ALT A
Conrad
[email protected]
Wlk
H
W
H W 6
H W 9
1 W ALT 8.
Rmnlk
Flyer
HMW
HI48880
HMBese
4 1oc.
Non-PMGT Line Mern (11.20)
1889 ALT A.
Conrad
Elgin 87
Roanlk
HM8W7
A318 x Charw
[email protected] (5) a l.24
A3127 (4) x 124
AT9138M2 x MI27
H
IlunmrrZAno
m
H M M
HMW8
PMGT cy& 3 (Em)
PMOT cycle3 (Em)
Non-PMW Urn Mean (n-13)
2.55
274
259
255
43
3110
2
2.78
44
n
42
1.3
13
2.8
35
78
211
1989 ALT B
Rmik
wf
HMsen
H W 4
H W 5
HMrn
HMeerr
HM8878
HMrn
HMW
HMWl
HM8982
HMeee3
HMsee4
HMEW5
HM89b7
H M W
HM8992
HM~M
A3127 (4) x 124
mln (4) r 124
PMGT cycle 3 (EGT)
PMGT [email protected] 3(EGT)
PMGT cycle 3 ( € 0
PMGT cycle 3 (Em)
PMGT
3(Em
PMGT cycle 3 (EGT)
PMGT cycle 3 (Em)
PMGT cycle 3 (Em)
HC79-478x BC3F2.812
PMGT cycle 3 (Em)
PMGT cycle 3
PMGT cycle 3 ( E m
PMGT cycle 3 ( E m
PMGT cycle 3 (EGT)
PMGT cycle 3 (EGT)
PMGT cycle 3 (EGT)
~mi23018x min
(Em
Non.PMGT Line Mean (n-11)
,
bah iu S, populations and as S,
lim for two years each. Thus, they
should have good tolerance scorn
for m n s other than eacas~vc
Wgctntivc gmanh. These lineb
should show no mom comlwon
bcMn yield md Idemwe Ihu,
mp-PMOT lias i f the munab an
.-U
A qnihcua w v e
*irerpatcdifBdem
Imp-
yleld. However, the ctrrretn~wnhaween yicW rlhm to r ~ a
~ n v ~ m m nnd
n ~ ) rnlcrnnsc
(me 4. ~nr%vIum-lyycrtes4 wlth
cbnc replicaten) for Ibc I9 PMW
limmspnitive(r4.45P<OM)
iadicating tow iolcrance waa
uroeirnd with high yield. The 24
non-PMOT lina from u c h dl988
dI(W9,Mdmcmmbbrdl~
PMGT Ilm fm bth yarn M
m - s l g n ~ k mor
t w i v ecomb
Im t d c l e n b (r-OM, P.cO.29;
14.37, P<OT#l; and rm4.2tl.
YcOD3, n*patiwlyj indluriag
no rcluvmrhip buuren tolerance
ud yield a thrr high mlenacc
anr rcurimud with h i field.
Twopouiblc~6(thac
unrrorliqp dm m: (I) rclirunee
~ s ~ c d u i o n w i t h m o c
0brc~cionnwould argue hat the
wmc (r-0.50. P< 0111) indicirrinp
y~eldand u J c m d A3127 ud of
H a a rel~live~y
udepadnr ,,-,R
the ahcr parents In the blparenral
PMGT rflatcd y r l d rrgrclvely
dldcomcc. Thc mr wore cornc m nm
~ rehtively udcpndcnl In
The 48 m P M G T llm Lmn hc
laled with msNq bur Ihe v w r
nuhl~vdalla)I~IM
wuh m ~ b w n c E O ( U P ~w~lhc comla~wmfwd with
m d d mc (rPO.20, P< 0.05; and
ud lhac w~thoutThc c w n l a t ~ o n ~ ~hcPMGf cycle 2 and 3 Ims. It is
r 4 . 2 0 , P< 0.09, nspecriwly).
pnasthlc that l~ner with high
d ~ 1 w 1 t h y r k l w c r r-OM.
r
Euly maturing l~nes an! more
u~lcrurcand hlgh ylcld ue less
PC a79, and r < 4US. P<043, hn
hquent than uther cornhinat~ons wcDthcruJpitntob.am.itherc
31 ruscqn~bkud 17 nalwunt Ilwr.
is n cornlation of root gnwth wlth
ud rkcrwn b ~tolcnnx
r
will mull
~ p ~ ~ 1 1 IhF
w l ct~mlualon
y
l a that
sad gunlny d m u n t y w k h may
then Ir rwc ntnmg w~dencct b ~ In even lewrr h~ghyicld~ngIlnes.
explain these d o . 7he dead plant
With~nPMCW cylea 2 and 3 them
n r m r mndlor the brtgnxlnd
scorc cornla~edwell with both n*~
wr a fnnd fi~rm)re lolcrvnf liner
p e r Imm fhc na~runtpYnnls
m and vigor scorn (r=0.71 and
O
I he laur (r=-0.35, P< 0.14) and
nlulod on ruenyc, in lugher ykld\
r-0.81, P< 0.01). Var~ancesof h e
r s ~ p ~ f c apr~ul~vr
nl
ccmclslm with
H ~ w v e r the
, pnltlve comlat~on,
mat and vigor scores w r e comlndginp (r=061. P<0.006). Thc
lfltlamp low u l k m U LXYOCIYINI
conjccturr thal xlwlicln for plant
paniblc (0151 anl065, rrspa.~~vely
I.
w~thhigh y~eld.in Ihe I9 PMGI
l1nc8contnstr w~th uuwcpt~hle v~pornl flowtr~ngas an 1ndlrec.1 hut the range of mc~iscorer was I
to 5 versus I b 2.2 for the vigor
n o w P M d der~vcdllneu In w h ~ ~ h cbtunlntc of tolerance selected
vcgclirtivcly vipnwa. lodging pn?
srwres. It appcsrs thxt some linch
(hccc uar,m ~c~rnlrtun
The parenh
show more [email protected] for a glvcn unxwnt
d PMGT m denml fn~mthc UIIIC
ne, ltw yielding liws IS s u p p o d .
bul nl pnwcn hy the nsullr.
of mol roc ~cndmrncmlcrne) anl
rrwn ~ m r u t nImw\
l uul ruc clouly
The pxlr performance of l~ncs hlthc v w r 10scly t c e r with thr.
alllnl ln ~ I I~ner
U u h d ar prcno
mot M assay il useful In testing en.
In the nnn-PMGI' h~parcntally xlwtcd fnin~PMGT compand to
durance t o l e w e . Thl correlation
hlpannlal Ilnes In the sanle $sb.
devcloprd l ~ w w~th
r
the prr\lhly
of Ihr. recurw h l e cxepnlon of A1127 w h ~ ~ h aupfom n~twl~firntion
b e t w n c~thcrtolerancescores and
rent uleclr~np q r a m to nleasun
yield was not r $ l i f k a ~ at lhc 5 perhm romc stluthcm Iln*n>awcrrlrr
cent level but p s i b l y indtcatcd a
uwl tr Its rclnnd I t m m r unl~kcly tcilenncc n a n d ~ m d yand a) Inh t PMCl mr,1nct1YR1w~thIIIK'S side yeld auing in cach cycle. The
mldual a\urlation ~ L I ykld
W w~th
closnl mturc of the PMGl ppulahigh wlerwe (r=O.L59. P< 0.11) In
that had mlrc unfavwaMc Ilnkngcr
tlon may alw~bc d pn*lcm In mainthe 95 u l w t d cycle 4 Itnes.
him m n p e hut thc dlnr ol A3127
talnlnp >ufftc~cntgenetic gain
Ten c)tk 4 Si lines wae akced
weds to k cxamlned The 18 ncmrrlaltve to c.onwnt~ondlcmsses i f
fmn%evalua*dbssadmtderana:
PMGT lines uh~chd ~ dnol httvc
hlLr lrr a parcnt ~ I W I Jr r ~
cornla
ww gcrmplasm with unique comsndagnmxluc~in1989
IICM
hctwen lolcranct and y~ c l dI t
(btAc8).%mmf~nnkhlnations of yield genes is dcwlIS p w ~ b l cLa1 the culuvlrr A7127
aulum Igrc s ~ wnh
i nre 4 Rw
op*l.Th~apossibil~tywould argue
hu pmu& n pnpan~rularl)valuablc
ox&wl8Pnd oxm30 (full dnms,
for an open murrenl selection
c~~mhtwt~on
of tolcrwc and )tcW
p~pulationm addition to. or instead
arccqurlalhosedresirtarr~.
but 11or l~nexdcnvd f n ~ m11 ended
d,the PMCT pplation.
OX8.5259-2 snd 0x8525923 (full
up u ptnnts of only 25 pcmnt of
Ybs but dso half sibs d OXB609-IS
the selected non PMGT Itner
and-30)hwcsxlrrs~lo
dthough used In 37 perant of the
LOO347-I-6-6 W hr wtde plsn
crau*s It S
I nu*unrlhy that ZS per
rcsisPrc~ncc7ndisIul~
mdthusc llncs wh~chhad A3127
a nce 4 (Schmktbcom,maddm,
or r hlghly rchted (coefT~~ent
of
The cycle 4 lines w r e nlcd for
subpndpspcrinpepprrtm).?
puclaqee a97)line u I pmnt.
b l c m uslng a dead plrnt scon.
Laamsxnccdpmkmnar
M d m m D c ~ f w a g m h 6 0 r m c r o c ~ ~ ~ r e U d a v ~ ( ~ ~ s c o (or ( m
~f u a ~ s a p r c d ~ ~
yst brlldlhem y~ddalwell. Tbcw
in 'hbk 8). Tbe vigor score did art
0ad5dI)prrrllk&TI.
u ponuvcly c w n l r v d wtth yteld
ud (2) ~mpnvlnpP q (dcmIn
-
W k ~ C q ~ l c , W , ~ ~ C ~ k r d m k m d ~ ~ l c * d ~ , . d r t l v r b ~ , , p
~10(PhyOOpMhOmtokruKaQMQT~4)rnl~'
PmORangJ
F)(Wdd8M
EntF/
Nmm
VlClgpl
HMBM'
Mh
HMWW
WJf
h~wdrscrlpion
W1-c,~lom7
Rpr1-k.Rbm4 7
Rpsl-b,Rps,.R1D-4.7
Apsl.MlRlornce4
Rpsl-k,H61 R b ~l4. 7
OX 880618,
OX 8 8 0 6 ~
OX 8524U21
OX 8524025'
OX 852592'
OX 8525918,
OX 8525918'
RM
20
28
30
32
38
27
23
22
25
23
29
29
23
28
28
(axml#lo)
WLodornp~~lwW)wcwl "ifp
(m) (m)
plaM ~ x w rwxvr
om
075
On
080
081
074
067
078
0.83
ON
OR0
067
(Id)
mrrn
rank
25
2.6
22
26
18
24
20
24
25
25
25
18
22
19
23
2so
ZIP
303
203
307
295
280
12
14
2
8
1
sz a
10
10
10
10
10
10
20
32
10
12
10
to
10
15
10
10
13
13
17
13
43
43
27
49
14
13
15
14
14
C3S1-58 X C3Sl.m
5
~ 3 ~ 1 - x5 ~3~1.10
8
13
14
C3S1-58 x C3Sl-M
209
7
14
C3S1-56 x C3Sl-M
282
11
17
C3S1-05 x C3S1.56
14
278 15
C3S1-65 x C3S1-58
XO3
4
30 15
C3Sl-05 x C3S1.50
288
9
30 14
ox ~ 2 s 9 - n ~~ 3 ~ 1 - 6x 5~ 3 ~ 1 . 5 8
om
ZBB
8
27 14
OX 8525925'
C3S165 x C3Sl.58
089
302
3
30 15
OX 8525928'
C3S185 x C3S1.56
074
283
10
37 23
OX 2C4'
Rps,,RpsH?, S to r a w 4.7
97
9 7 100
Palla
Rps,. moderne aleranco S lo races 4.7
10 40 20
L80347-1.151G' Rps,'?,very tolerant, rw 4, 1 wtlh raw 4
13 23 15
M a n of l~nes
23
2~
17 29 tfl
258 on
0.6 10 09
F-lN
P<OOS
02
013
10
NS
03 05 05
P<030
00
OD7
08
024
' R M - W I M mdlurity *em values are afxMed Lufwon Vlc -20 and FI r - 3 6 (Flyns vdur *u ul by Wulon wch
H m . o ~ r l ~~mn" lM e w hsd ~ a l w
m a n 01 un3lMr rnuiu$w c
m
ru
n RMI Lodgly) w e from 1-wm
m 5-pmslram FW Is m F-wpraaud IrW ugnlllcw d i l h w ~N S -F Ucl nol rrgnllkuIl
2
.
1
1
%
W
p
l
u
a
d.BI%
lO-MO%RmR~Saim
l-rnrol(~~MYn),
4MgMScas I - m W
I.hlphe* (pl.nb x heylhl) uon 3.50% mu w r 6.m d mu Wn, 10.0 w m
95?:54&~ E
( g m ScarrhDn1nocukrmlylrWu*npr~14.n47R-RIUVX.
I-n*M/#a hall %t
tW Mwmbw Ihs hypmNyi l M u W &
lhs tmcaed lam
wcmw mpsnm~llelIIM c#, Wtm fmm cyde 3 d a PhylopMha* murrm u*dm twdmm IPLMT)
AithOqhynrdIhe~m
'We9pnmmprabko1csi#mx.
bymmcdueorcnjcncrl ly*
tmducal, multi-me, hypocotyl
misaMt (nrhac Rps,c w @s,.k)
batdonhypm*ylhlswffhkcu4
r m d 7 W 9 ) . Hcmncr, hsppears
~mrcsbtmcc,rbw*wd
enbrclPEcrdasmhwdlbaenim~intixsdmimdlugbly
bbnlaa.AUburdhacILa
(-18d-34adOMIJw-2
(uW.8 1 la 10)
ms7
raw4
mm
Ern
aWsOQ3a'
vw
lcrxs
lcrxs
Wlng
41)
10
3 3 3 0
43
25
27
30
3.0
25
17
27
20
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w
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'Dmo#l ap*&
l i n C,S -XX lines are S u*nrma Imn
3 ol rrcumnc wle*ton (ot-"Raw 7 nrcllon, We tconhnl M h l M a p h n t dam rvheh us urb)ld d rnDMer prbl(In -m)
l h a a drtr lndlcnr il I8 nua(rm lo mce 7 by t h m upunr lrsb u u q lyrr dun borrd and hypadyl pmcbdum
lPm~-Waphmonm w w m a I ap
F.lsd m m F-twgmmrd but a$lnihum dilfrnnca
r R - m m m all p h b WMYI. S - w e c ~ U a . I1I plants d~e I-~n(acmed&a !aaria 1 to 4 of 5 plm w m
wtm kculllrd in t h hypocdy( villh I - d W Rce Number d llvlq plurcs wilh k i s b s and numba d deM
plmO,u.IldSInaWodp*ntllhorm
Discussion
Thc -I
elcctim puccdun
usod hy Wkcr md Schminhenncr
impmved Phyl tolcrancc. The addilion of y~eklwing as a sap in
is now supponed by bnh thc
dcmarnraMy pmm rdrpu pCTIDTmam d cycle 4 S,:, l i m armprrdmsunilulysekc*dliihrm
mumc slcaioa w w on
~ ~ ) m r I w d t O ~ ~obsenmtionsofthehchetcmgk u
qrowmicllly wccpuble lines.
uvous popllr(ions from PMGT m - ~ c m m r a d t k e x i s r m e
T h a c ~ ~ w c r r s k C ( 0 d u d which irdiuted that the mon
d~~nnsoc~dIwyvld
-hrmcOhioMunrdLur
~ ~ m l m s t iwithn b i soltnm. No smog
TWuurlbundmshaarasignifiyield. Thc dditim of yield
evidepoc suppwvd tbc pnnibilicy
aal ccnduiaobr k yield with
lathencwrclltgkctioaw
Um~~hdbCmCumal
w
- -- -
NMm
C m
m
20
19
22
27
am
an
[email protected]
2s
16
om
0110
073
0A
uwcb
Vidwy
NKS a 1 2
~ ~ ~ 1 3 5 '
Connd
Burlmoll
Y~t"[email protected]
RM
30
d O ( " Q m R n t
274
m
2~
2
12
24
22
529
1
3
19
21
23
21
2 74
2M
2w
911
10
284
264
2M
281
7
so
w
la
CompanMs lim frwn adapsd cmsws
OX 867020'
OX 881W-16'
OX 86221.10'
OX 882264'
CENTURY 84 X 52125
P E L U 88 X HOrr
AIW x SIBLEY
SIBLEY x PELLA BB
22
25
24
am
27
on
50
30
OM
2.0
on
22
OW
0 72
o n
O
M
21
24
200
28
072
22
282
03
01
0 14
08
04
1.09
067
4
8
2
CompraMe lines from mde crosws
OX 88117.13'
OX 881204'
OX 88120g'
OX 8812011'
OX 8812014'
OX 8813818'
ZANE X PI291302
ZANE X P14077l0
ZANE X PI407710
UNE x P I ~ ~ O
UNE x P I W ~ O
ELGlN 87 X PI384468
Mean ol I~nes
F-lsd
P<OM
P<O%
'Derides experimental I~ne
'Linen &Wad horn a annparaMe nurnbfot
F-lrd
28
29
29
19
a07
26
24
ZM
7
14
0
4
16
M M o u m r p l M row
r a t ~ t h r u m e c ~ m d ~ w i n p M o ~ m r ~ ~ I W P L # I I ~ L 4 S , ~ l n l
r n t n i 0 ~ ~ * i m 2 m m a n c h c l o , v r k a r y ~ d & n l k oAnd r P . d n a m M c u k M n a d r l c w b n d I r q U m u p m #
RM-Waha malurlty cnwpialrd hun mom mduQ wing Vrkr/=20 and B ~ ~ - 3 1F l )l d Y M F poYQId M
UJniRaM dillerenca
for yield ud tolerance in PMGT
cycle 3 l k which would cominuc
in later cycles, although s vmd in
rhe cycle 4 lines idi a weak
~rsociitioo may cad. Ssupmor
reeurrenl poplluion. Such an open
p q n h o n YT hu bem i n a d .
Tbc PMGT popllalion can be
dbcoruinucd when the rclal~ve
c
u
m cycle bebn vkction for
toknnr. Sanc d W linr my
hmcanbtniavdduredcnoEe
t y p l ~mrArtrirunoc, clav maiag,
~dcloscdandclpcn d ~ J . 7 1 r k m y i i h
L60347-I-U-Ki (dcudopa( fmn I
mm e e l # h poplluvnu on
cultivvrforyiaridb*Rmhm
wrnpnMcorwhca~ism c r r u m d c i n ~ q p r a b h m
do*lopcd ofkr pmgnmr
WdIkmnbhaion'brrheAnt
caapnMe lsja ra Ihe c l d
a i m rhe PMm populaion wps
pDplLMar,iomt-PMmcnnra. aJliwdadDpdbnn&i'W
W.TheK dtivur n p m
whid,wrnbrdMy~b
k PMOT cyck 4 iinm pmvided
KioppommirylOimpweyiLdand l
fumzrcvideDccChrcvlcdioafn lMdlhia d d. 1990c).
tolenaa;lidmarhikrtduciia
~ m d u o u l d j u a i f y ~ o p a yield i nsnvry wilhintwlr n- ~ ~ g d c w p p p b n n i a e
-
damecbuclamormmdthc
dvsetpdldaaccrndhdyield
armpnblcalbhmndrpcdx
prbnaw-.hw
kkrctLvttheykwrv~adncw drpaf cmsrcr. [email protected] l m l s of encould only be idmtifkd in
~ p a e . m ~ d y w 9 m o .
lim with no rod resisture ud
blOafiPrrlyUbarmpbm
d a alw mcing.
-dYu*dby-The pn& intefnctions of the
p r o p s In e~thcr comblnlng
mkwudrJmnraunprw
sg-mthon
p-m
nguln more study to predict the
dfau d hyprayl resistam, mn
sluw-mntq or ratc rel d t l w c c u d ~ h e ~ f m~slpncc,
'
admrbsn,lndcpdm~tf ducing resistance, and durance
~ y p da a l e m e on the stability of
thaamcchruvnwkdevdqwxl
we8 (If h g . Nlrmw KRV (wnscleaive) tolcnrm (~ncluder,endurm c , md pnhbly. some bms of
v l w nmlng) rcmaiw ac the ideal
=So)rbwc-w-*>
mans to contml h g without seryaan u rclmfvely prig ud ha\
kcting mv physidogic m but h e
had lwtk tam &r mewdutlon
ability tcr select lndependcntly for
W
n haw urcncned hy dew1
crpq iuhmn with ns~atanrd i o r
eilher cndunncc or slow m i n g is
mlmm R * r r ~ ~ r C h ~ Imrnlcal
~ ' pby cumnt knowledge of the
p a an p m m In a11
ppLtll)n\
mhrvusmr t f thne rartnw of the
ud can tarcaw w dmqmnfi In
Syhean-Phytophthora p~thosy,sum.
gunc&s wfthin clghl ycan o l re $15
Given thcsc lirni~s,continual selecM gar depkynrnl Brmllng for
tion lo pmducc highly tolerant lines
DkrUKV IL drpndmt upon guxl
rcgardlcb~ of the mechanisms
asay Wluuqucc R l d , ~nnulun~ inn~lved,i s justifm.ble as an alteriyarladdrrrbrudrsyrdlpnnrd native lo whole plant resistance.
usdul a mrurnw ukmx td pa
11m The durn-lq.er bst can be
References
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wv\sarruu\gwlueh~scpus*
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r M&.
adwrnx-*R
Rawullrkr
U.S. lkp d Cmncnx. W. 1.
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-1.wM.
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~ / ~ ' I I M~ .
c
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y
595.
&manl.R.L. Bn.Two major gem
RwtimcdIlowemgand~q
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&mud. R.L snd C.R. C m .
ma -R
or 'wdliuns u
sqrtcan.Cmp Sci. 28:KI27-16328
Bha. R.G.. A.E Smmitmenner, ad
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R
M
~
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pRsa
~
d
Cmpa. R.L.. RJ. Mdu. A.F.
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wpD&&RPl.l(rgnamond
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mib. M.S
Th.a Dep. dPInr Wdqg,
'hhc
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H y m u a . T md 1.R tbim LPB~
*
7h?lllmknmddrnybmk)
--b/Smrl&rwn
m ePrmi;&rryR-flcm
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il
dc*rmmnuq relwve (CKhd r~ld
m h x x m qkrm t r Mph.
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Sctimmbum. R.L.Cmps, lad
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.U).42442S.
Mcl3lun.&A.,RJ F i & S K O
.
Muun. A C.llplkhru. A.P
SE~III-,
R .L. Cugcc, uwl
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Rqnmmd
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aI.wmyh
Cl n p h
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!khm&mm. R I..Ccxtpcr, ad
hd ld ~
tnm ul~~ophrlu,nr
a uw
* ( q R n r h J R
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plPsm ppddm. Crop So. 24.739
%W.W wyhm. Cnlp k
i
nvwprnnu w. alp.P h I
k
64175.778
Y1425426
Mu, W.R.. A.K. W r , A.F.
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roc of soybean csurcd hy
% L h l m h m . R L. cmpr. d
'Rlh 86' q t w n . Cmp k c .
Hpfupluhr~rnsoflc n. w. Phy
R.J Mutin WL1. Rqpru*md
nLtn.
lopalhology 4X:201-2W1.
'GRW.W uyhrm. Cmp $2.
Fchr. W.R., A.K. W r , A.E
Kilen. T.C 19% klntnm$h~phc.
30 42h42l.
Schmiuhmnm, S.R. C i .4
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.
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28:1025.
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26:lIl-712.
Mu. W.R.. A.K. W r . A.E
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W crl
McRLun. BA., R J . hmW, S.K. 9.
c a d major p for r e h ~ l u ~tnm
'Harpn 87' w y h . Cmp Scl.
Miuun. A. <'.lpDuboir, A.P.
phVDphmaran ~n*ybcan. Cnv
28:W
~-LS
R.L. C q J m , ad
%I. l4:260262.
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pIcu
Thc Ohio Stsb Unimsiry. 92p.
McBlun, RA R.J. PnrnS S.K St.
nvgruprrmo f. sp. #I*.
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,
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. .
.!k$m&mm, R.L. calpm, ad
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bcalmdonddrphynabrin
R J. Muwr. 19Yk.
d
Evidmcc b r hanolrrryusta In
Olymlinlinpkccionb~
'Chpnn' *cybcm. Crop W. 31:
nu*m
I.bp.
h y p l p c i n ~ ~ m ~ i n l c d a l &u&hom
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6vminn
vimFhp?pM*m~f.
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'
sp. glycincu. Plant b b l .
A.P. Man-.
IQBI, Irr
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.
.
.
e.
ma6
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'Hilrpr'.C q Sci. 31: in par.
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*.
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l&iltiW~
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.
.
dYndavmondWh
i T a h r x t o ~ ~ .
9:235.252.
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~ n . w j w i
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H.D. Luden d D. Wilkccenson
( c d ~ . ) Pmc.
.
9rh kybcan Seed
bxwrch Conf.. Chicago. Ill..
0-14 Dm.1979. Am. Seed Tradc
k u r . . Mshington, D.C.
schwmk. w.. and T Si. w4.
R r r 4 d ~ ~
mr var. so& bun s y b m s pm
w.
p a d . Plan Dis.
58:
352-354.
Wtmdy, AJ., onl A.F, M a haurr. MS.Soybcnm lflmed by
culy mra.Ohiohnnud
HW Rs.40:M-86
lllwhcr. L.L. M7. SoybePrw ocN,
amjornopin0ho. hHtlcom
m Ohio. Americrm Saybun
1W7 ANYlpl E m tion emua, h.Sojbpn
h.9. LauU MQ p 7 8
Rrmison. P.R.. CA. Thrmui WJ.
KmwMty, m3 M.S Mcimsh.
mEvidared-spe
isntyiodanadaolbm,
~pemf.rp.giycinminscybPn
sudlllp. l'hyqah. 72.m-733.
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