Polyploidy and Aneuploidy in Guayale - ageconsearch.umn.eduageconsearch.umn.edu/record/169955/files/tb918.pdf · McCallum collected ~eds of gnaytJTe (Parthenium aJ'gentatum A. Gray)

2 8 2 5I~ 11111Iiamp = 11111

m 11111 3 2

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IIF6 ~

11- L L~

I~ 0

-shy-shy-shy

11111125 1111114 11111 16

MICROCOPY RESOLUTION TEST CHART NATIONAL BUREAU OF STANDARDSmiddotJ963-A

2 8 2 5 l

~ 11111 11111 10 3 2 22I~ 11111

I ~~~ I i ~~ ~ 14 ULu11-

11111 18

14 1611111middot25 11111 11111

MICROCOPY RESOLUTION TEST CHART NATIONAL BUREAU OF STANDARDSmiddot1963-A

Polyploidy and Aneuploidy in Guayule 1

By A DOROTHY BERGNER

Formerly associate cytogeneticist Special Guayule Research Project Bureau 0Plant Industry Soils and AgriculltLral Engineering Agicultura~ Research Administration

CONTENTS Page Page

Chromosomal range in gllaYllle __ 1 Loss of chromosomes by abershyrants_____________________ _Polyploid series in natural stands 22 and commercial strains______ _ 2 Chromosomal varianLs within

Euploid populations and the pol)ploid series _________ _ 24Btrains________________ _ Oligm __________________ _2 24 Aneuploid stmins_________ _ 3 Hapluids from tetraploids _ 25 Intermediate type______ _ 3 Anellploids from diploids__ _ 25

bull lIaterials and methods________ _ 4 Offtype normals from trip-Chromosome mOlphology______ _ 5 loids and tetraploids___ _ 26 Chromosome behavior in pollen Aberranls from triploids and ~lOther cells ________________ tetraploirls_____________ _7 28 Diploids_________________ _ 7 Offtype normals from abershy~ Tliploids________________ _ rants _________________ _8 30

~ Tetraploids______________ _ 12 Sectorial chimeras________ _ 30i Plants with higher chromo- Factors limiting aneuploidy and so~ nU1~rs---------- 3115 polyploidy in guayule_______ _

18 Summary___________________ _~ HapMds---O ------------34

~llen__~__________________ _ 20 Literature cited ______________ _ 36 7-J ~ ~ c1 vi ~ ~ lttROMOSOMAL RANGE IN GUAYULE

~JIn thl-fall 6f11942 LeRoYJ Powers and W B McCallum collected ~eds of gnaytJTe (Parthenium aJgentatum A Gray) and mariola (p iJcanum H B K) in Mexico and Powers and Walter T Federer Cigtllected seeds of these same species in Texas Early in 1943 many of

these seeds together with those of certain of the commercial strains developed previously through selection by McCallum and probably of Mexican origin were germinated and planted in fields at Salinas Calif Chromosome studies especially the determination of chromoshysome l1umbers were begun as soon as possible in order to facilitate genetic and breeding experiments

After a few months of exploratorystudies it was apparent that a polyploid series exists among natural populations and commercial

bull strains of gnayule Stebbins and Kodani (13)2 working indepenshydently came to the same conclusion at about the same time

1 Submitted for publication February 27 1946 Italic numbers in parentheses refer to Literature Cited p 36

61J0587-46-middot1

2 TECHNICAL BULLETIN 918 U S DEP1 OF AGRICULTUHE

Recognition of this was facilitated by the fact that the whole series was lcprpsented in the seeds collected in Mexico Stebbins und Koelani (13) and Powers and Rollins (10) IOllll(l a similar series ill mariola From papers written by Russians prior to 11)36 (93 5 6) it is now evident that they had diploid and t(truploid strains of muriola and uIso tetraploid strains of guaynle but that they faiLPtl to recognize a polyploid series in their lluttelial

Up to the preslnt no guayule plullts IU1Ve been found with hss than 36 clllOmOSOnHs Ill( higlHst lllllnb(r reported thus fill is 144 among offspring of 108plusmn-cillOlllOSOIlW 3 plants (8) Probably (YPly numlwl betweell 36 and 14-1 eouid be found if el10ugh indiyiduni plauts w(te exauuneu Pr(3umaLly some gll1ic combiuatiol1f are yiable for any particular ChlOlllOSOllW num)tr howtY(r t distlllctioll should 1H made betweell th( predominant cluomosome nUmb(IS of populations (stllnds) of gUfiyule aud thosp of Vtllinllts which OCClll ill sueil populnliolls or which wpre a result of crossps mndl in gl((l1tOlHHS and flplJ plantiJlgs at Salinas At Salinas arC the makinrrs of It gl(tlt variety of eitromoshysome nlllnbl1s beCU1ISl dh-(ls( )[1(P8~of gUfIup whith In natnl( are separated by glographital nnd ttologienL burri(ls hay( h((tl nss(ll1hlld thert 1-101( aneuploids would he foullcl l111((l l1aturnl (oll(lilions wCre ii not for the fnet tlllt fn(ultaliyc npomixis is predominant among the higher chrOlHOSOllll mll11bels of ill( seri(s (~ 10)

POLYPIOlD SERIES 1 lNITRAL STA~DS AXD COMIERCIAL STHALS

ECVLOlD POPGLATlOiS AiD STlUlgtS

Although Stphbins and Iocnnt (13) suggtstcd tlll hfLsie numh(r of r equnlR U for tll( polyploid s(ri(s no plnnts whicb hnyp 2j~ Cqullis 18 haw bcen found 01 llpoIted ENnus of thC fOIJ11ntion of biyrdlnts

in 3G+ -ClllomOSOI11l 4 plnnts nnd of tri-nCnts in 5-plusmn -(tlomosome plnnts tqunling 18 will )p wwcl as till busi( IHlm)(I in lhis buUllin Aecol(lingly ~n elln 1)( (ollsiltill(d til( diploid (ondition 101 lHltUIfllL~ O((UIJillg plps(nt-(Lny populntiollR of guaYlll( As lppoIt((l by Bo(hanls(-n (2) hi 1S tlll( for mltlio1n n18o Thl only ]0(] llumhtr found in j)(lIItnlilllll tlnn fnl is rUI p hlsiClOlJt011I8 L in wili(h 21 equals H for 2 COUP(dOll thalL POWlIS mndp in Tlxns

At Salinas lllllc flrC ) diploill rollp(liollS (21=30+) from s(((ls hmT(stp(i in fl lIlol1l1lninOlls ll(l ~()u(hlst of ~rnpimi Durango llpxico ThtmiddotI( Iln ii triploid (olllCtions (t=ii4plusmn) -- 1 from fin indiYidunl Jllmt growing rithin tlt( population from whieh tIt( diploid (olledioll -12ii) YUS Ohlilillld nnd till ollHl -1 from difrPltnl pla(( Jl(Ill Snn Bllrtolo DlllllllgO Su(h (tlmmllCini stmins ns )[(Cllllurns lOll 111 210 2jii und 2iR nlso 111( triploid Till mnjolity of tltc (oUccliollS nt SnJinll~ hllYP 72plusmn eJuomOSOJ1llS llluL tlHrdol( nrl tPlrnshypJoicl (41=71plusmn TIllt S((ltS (UIIl( ftom bOlh nntmnl stnnds illld lfcCnllmns (ollll11(leinl sl min Only 0 of til( (o1iP(l iO]lt ll1ndp hv Pow(IS lIld )1(CnllulH in [(xi(o tlllUtrllploicL ThCy (nme from nil

3 plusmn uCtpr a J1umlwr inclirnip1 a polyploid rla~ll the indhidual l11pmh(r o(whi(h van in pmrt tllllllhpr of chromo(HlH_ + nft(r 30 imJhnlp t Ill diploid rln 111lt indhidunl 111lt mucrl of hid mly be aueuploid with 0111 or more ndditional (hIOIlIOS(JIlI(

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bull

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3 POLYPLOIDY AND ANEUPLOIDY IN GUAYULcl

shy following 10ciLtions Onr fro111 nn indiyiclual plant glOWillg middotwithin the populntion from which the (ollrction 4255 mlS obtllinr(lj nnothel from secds (oU(eLeel nCilt Snn JUiLll N u(-O Lron j iL third from Catorce San Luis Potesi and tlir lC111nilling (j fOm InjomiL in northtnst Zacatccns Th( tmilS-Pc(Os arCH of J(lxns is ljgtlPselllp(l by 362 collcctiolls Of th(sc (xtcnsin~ eollcctions eytologi(nl work hHs becn done 011 only 55 nnd pollpu studits on 4 mOle JJI hatl72plusmn chromoshysomes Elcven of thl 20 lo(alions in T(xns flOm -1Ii(11 nu (((ls wpre coUr(tltCl hny( 1)(en smnpkd Thus JaJ 25 of [((aUtuns OIllIlHleiI11 stlHins huv( bppu found to han nplusmn ChI0I110S011lPS TllIv 11(~ Nos 3SH 49 180402404 -lOO 40liF HI 413 41U 41R 41l 42ti 42R 480 4an 440 H 1 IAA 4i1 4j(j 4iifJ 591 and 7~j2 fillt l ~[pxinLU sllain (42middot178 in POWPIS Collp(liol1S1 Thus fur no Jwntnllioid or hexnploid llitlult11 stunds 01 (o1l1nwlcinl stmins lulY htJl found High( eltlomosonllt lluml)(lls hny( I)((n limi( to (l(nin indi-i(unls in triplOid nll( tdtoploid populations 11m (mins undlo lhpil o1Tspling Allor the I(xicn11 coll(tioHs hnw )((11 snmpJpd ()-tologi(lllly nnd S0111( hnYo brctt studip(i inttl1Si-lly IHn liSP lhl It01e polyp]oid SPIIIS

is l(prpsPl1Ud in tltPl1l and tllIPJOtP tlllY ufron tllt oppoltunity to wOlk out tht Jrlntion umong liL( l1wmlHls or th( (tw bull

A EUPLOID STHA lXS

bull 8i1(P l(plodudion in It li4plusmn -(It ro nlOSOl1H) nnd 4J (72plusmn-cluo1noshy

SOJlWl popu]ntioll is plpdolllilllllltl) fuCulttltinly Ilpolllietil dO) it hus lui ill SOIlll instmIC(S to till (lstnhlisluHtllt (If Iult nIl lsslItinlly cl01HS 01 tlll(luploid 1 and 1( (ombillat iOllS FOl illStllll(C conuneishyciul strain 210 8((mS to JUlY is chromosom(s the nllrnb(ll thut Stebbins und Jodimi (13) rtpoltpd Jor stHill 111 -Illnins 45n and 735-2 8e0111 to hUn I~ Ill1OmOSOllHs i1 ndditioJl lo Ill(1 pnit of miniashytures which is prtslnt in muny 4r populations

A morphological tYPl Cnll(l inllll1(dint( h PmI S alld deshys(li1)((1 by ]()]~ins tIL) ~tld middotItI(polllling to ill( IgtlOadhU[ld tqH nnmld Prtltlltr1(nt llJlIlif Buttlllt hail 72plusmn chl(lmosoll1lsbull POIshytion or all or u 11111111)(1 of coIlctiolls [mIll d(si(o IIl1d I(ms as w(111 as s(ernl ol [c(nllnms c(lmmtI(iul strains til intllllllltiia(PR The prohuble origill il1oldllg in UogIltssin hylnidizulion dlh mndoIn is dis(llSS(d hy Hollil1gt (11 1tl nlld -((ll111I1S llId Kodllli (13i As ill tlll (IS( or nl1Pllploi(l ([ipIoi1 stands Hud stlninc lClJtOshyduCtioll 11 apomixis hilS ]PeI to tl)( (lstalgtlisl1ll1l1l1 of whnl 111 (SSII1shytinily C]OIHS whi(1t ditlp[ SOll1(Whal t11l01lg 1ItIIlls(k ill 11l0lpltoshylo~i(nl appLltlI11P b((HlISP tlu hayp u is(11 mOIp thall onp ulld in difl0)(nC pIu(s hilt Hll sl1ffic-i(lltly simiIul in nppplUlIHP to WItITHlll clnssifientiol1 us intpt11l(ldiut(s Thl- lIT in ehrolllOSnnll lItllllshy

bull b(l SUH( pInnl with 7172 7l tllld 74 hrOl1JogtOlll(S 111-11 b(lll found (sP( tthl( -~ I J(gnrdlls of IhlOIllOSOI1l llllllll)(t 01 [rt()shy

grnphi~nl origill mu ny ot (h(il poll11 mothpl ((lis lih()wlu~ging dllo7noshyson1lS ut IlIlUphns( I Illl(l ] I alld (jlI( 11( 111111) l11Hlo(ytps mnOIlg

5 POWERl L lfID (OrJJcTTlt1X OF nrnu FRO[ 1 EXHO ID THE TR-I~SshyPECOS AREA (BIn m~XD COlIiTIlY) OF TX jJ 12 [l-nJ)ubth-hlaquod]

4 TECHNICAL BULLETIN 918 ti S DEPT OF AGRICULTURE

malure pol1(n gmins The only observNI (WItI(Il(( nt meitLphnsc~ bdwccll ttintclmedintes nnd the good guayule type was th~ t(JldshytlWy oJ tht n lIterllwdmtes to form 11101C polyvulent assoeHLtlOlls This tendtncy WitS int(lprtt((l us indientin~ stnwlurnl hybridity I(ss homology among the cltromosonlls is (onsistpnt with thcir hybrid an(cstl) ConseqtHntly the presencc 01 lugging (hromosol1lcs nt annphfts( I unci II indi(nLlS difikulty in SlPfUMioll nmong the mlmshyhtls of thlsP polyyalent nssoeitLtiolls Tltl finding of 2 illtlr111eshydiak 4plusmn-Chromoson1l plnnts among the offspring from 110nshy

(olltrolkd polli)ntion of H afi-chlomOSOIlll plitnt grown ut Sulinas mgglsts thut 54plusmn-(hromosome ttint(nn(clitLt(S iUC possible nnd mn) 0(( lit in nnturl

MATERIALS AND METHODS

TIll mntlrin uspd for studying guaYllp (hlomOsomN (ollsisted 111niIl~~ or po1ln moUl(l (elk It wus allgmenkd by tnpCtnl ((11s o((nsionl11~T found in didsion and by root-tip (ells Young flower hel1ds w(re fixPel in lL ( 1 tthsout(-iteohol-nePli(-itcid mixture ilen 11 (PSUllY lhl nutLllinlwus stOI(ltI in 70-1HI(tn t al(otol ]iorlts from mnny ItttHIs of tit SnIl1t plant w(r( dissettd out pool(d and sPleeltd IH(()l(ling to izl _boul It (loz(n dlsk flolds WCIC used for making (Hell nCeto-ollin SlWIU

Sil1(~P 4 pollpn gmins lmi(rospolls) nOlIunlly ori~6nat( in pu(h POll(ll mother cpll (ons(([lllIHlS of the nllml)(l of hlomoomes and of llwiotic IHituvior in pOlllll mothC[ (plls nl( founel in lh( POI(ll grains TIHrliol( addiliollal information (olw(rning the (f1eets of l1wiotie bchnviol wns sought nlHi foulld in lht (xnminution of l)o1en for siz( P(l((ntngl of grains aborlld (empt- or obyiousLy c(g(ll(lshynting) pr(s(lltl of millocytes (millutl lmpty gmins) f1nd nbuolshymnlitits of nny sort Pollpll-grnin pltIHtllltiollS W(I( mnel( hy plueing swd polkn ill n dl0p of (oUon bhll in l~t(to-phenol (l-P(tC(Ilt eotton blue ill (qual pnlts of lnetic aeid phlIlO ~ly(erin nnd waIN) fhe diamct(l of glflins wns llWHSlllccl from ilsi(k wull of intine to inside wall of intilH rSllnlly 100 grains wClt 1ll(tlStlJl1 In ordCr to gCt the pcr(cntuge of grnius ahortN 500 W(IC USl1td~T (oullted

The primary objpet in nll tite c11l01110som( studi(s was the detelmishyllntioll of thcir numbCl ObsClYntiolls cOllccllling t hc meiotic procshy(ss in P011CIl lllothltI ((lis WI( illeititnta Because of paucity of clcar l11Ctnphnse plntcs dtlC to dumping 01 slickin(ss of ehromosom(s lISC WI1S mnde of nny ll1(iotic stng( nt which chromosome counts could b( macle or chromosome associntions studied At first anashyphase al1owIUl(( ]uLd lo b( 1Ilnde for th( tpndency in ((rLain plnnts for uniYal(nts to divide pr(co(iously 01 for the preSCHce of fragIl1Cnts in connection ~ilh single 01 double chromatid bridgCs S((o11(1 teloshyphase counts on til( same slide vnlied bcltnuse of th( preSNlec of such flilb1ncnts O(Cnsionnlly n s((ond anaphase stagp could be us((l For diploids (30+-chromosome plants) nnd to It l(ssC1 extent for tlipl~ids (ii4plusmn-ehlomosoJU( plnllts) dinkillCsis and fLlst metaphaso stages W(lC (x(ellpnt Bt(lwsp of Il1utiVnCl)t formation theso stngls rarely WPIC satisfactory for dlt(lmining (Ilomosomc numbCl -of highel pol~yal(nts hOWlV(l th(y WNe illvnlunblc for reYealing Jl1ciotie beluwiol For (ounls nmong high(l poypoids the interphase

bull

bull

bull

5 POLYPLOIDY AND ANEUPLOIDY IN GUAYULE

or second prophase stage waS satisfadolY sillce the chromosomes were less diffuse aud better staillld thUll tiuS( stagcs in36+- or 54plusmnshychromosome plants Fairly l(liuble dltUlminntiolls could be Inade by averaging connts 111n([lt of scyetal cp11s in thcse stagps on the same slide This wus shown ~whcnpwl f1 c1l(k as possiblp by mpans of n count of somatic ehlomosol1lPs in ltlividing tapctal (fUS The interphase and prophase II stngps ~WCI( m uell mo1C sntisfactory than the second l1l(taphasl h0causp of the tltlHkncy for dllomosol11(S to stick tog0tllPl at meta phasc II

ValilLtioll in m(iotic chromosomc 111)1111gt(1 in th( samc pllU1t Sl0mS to bp limit(d to two SOtU((S (1) Addi tionn1 fragmcnts rcsulting from crossing OWl in in(15ion5 all(l (2) lht (limiuatioll of dU0ll10501ll(S by nb(rrallt plnllls (p 22) Tlwll may OL ma~middot not 1)( a difTltrlW( in CitlOIllOSOl1W llUllllHr hrtW(lll mOtplwlogiltttlly disliuet purt of u plant which is a s(cLorinl ehim(r1

CllRO[()SO[E ~rORpnOLOGY

bull

BcCause of tl1lt r(latiCly smnll size nnd (ompactncss of gunyu]e chlomosoll1rs dUling 111ltiosi little could hc ICalll((1 nuout the difshyfcrcnces in lenpth cand th0 loCation of the insertion ngion No attempt has bllIl madc to work ou t iclio~ral1ls fol thc ynrious collecshytions nne1 stmins HOWCVCI somatic (hromosomes in tap(ltnl-oell 11uclCi (fig 1 A and B) show diflNCllCCS in o-Cr-al1 and 1c)nti-e lcngth of each ann s(me (111 J-shapNl hut tlll mujolit~middot are V-shapcd ~with a mcclinll or suhnwdinll insertioll region In figure 1 1 thple S10m to he multiphs of a fpi hnsil tn)cssueh as n lC)llg chromosome with a suhnwdialt insertioll r(gion a J-shnpNl onc n nwcii1l1l1 (Cjual-arm(d a J)l(diuIll llll(qnal-armlcl a small ltqual-UilllNl and fl sll1ulluJlCltl1lalshyarnwd ns lt11 as fL SlllfLlI t~pe hith tap(r nt OllP (nIl Th( lastshyl1nme(l typl prohahly is th( sfLtcllit(d chromosome whieh (alt he s(rn atlndHd to th( lluelfOlu ill oc(asional fll-orlhlt ]Hppnrntions In root-lip (cllfi Jixtd fLlld stnil)prl flpeolding to )I(~-(IS t(ehniqIH (7) a pail of small sat(llitCd ehro11losomes (an 1)( S((11 in 1G---ClllonlORO11C plants Th(y haxe a snbmedinll (ollstrirtion thC small sfLtlllite )(ing attil(Jed to th( sholt(r finn In lllrlllts 1I1wiap I 01 molC 01110-mOSOll1(S thpl( wil1ll11y HlP s((ral nuckoli of difrPlpnt Ol (qulll size

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Pra(ti(nlly all of the ~W--(hromos()l1le plants thus far cxamincd and (1V mnn of thow with hidl(L chromool1lr numhcrs hnn in adclishytioll 011C to smCrltl Y(IT small c1mullosomps Th( n111111)(1 5CC111S to h0 COl1tnnt for in(lhidtlttl plant hut mn~middot VUlT muong mernhCls of the same (ullm( They ai hptllOPYCllotir RilleC tllPY stain mort dClpb~ than otl1(r ehromosollHs at diukiJtPsls and Ils so at nlltal1hasc I or lutllstfLglfi In root-lip tells ilx(d and staiupd acCording to )[lyels terhniqu( (7) such difftlellliil staining is Jwking If 2 or 1110re (Cl1s are prcspnt pniring mnT 1)( SlNl at dillkilHsis Ot mC1iltphasC 1 fho inscltion rcgion if not tllminiti I ll(Uriy so This is (ollfill1wd hy OhSPIyuti0115 on root-tip (lis SOllwtiI1HS Wll(ll ullpailld tll(~- lLP ul1coil(d nIHl lltllnltllik( at lllltnphnse [ nIHl lalp stUglS Ill) mny divide pncoeiousl- at nnilph(tsl 1 TblY 11m) lH of difr(l(lIt iz( eYCIl in the same crU (ilg 1 C) Similnlity in sizC anti hnp( to urm of V- or J-shaped elnoIl1osol1l(S sllggCsts tlllt UHY OWl their orign to brNlkage of such chromo~()mes 11I(y equnl t1H small(t arm of (l

bullbull bullbull

6 TECENICAL B ULTJETIN 91 SJ 1 S DEPT OF AGRICfLTURE

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(bull ( ~

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pound

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e

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7 POLYPLOIDY -XD ANECPLOIDY IX GtAYlLE

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medium-sized V-shaped chromosome ill length and width (fig 1 A a) Aetual attnchment to regulnr clnomosomCs is rarely seen but is shown in fi~ure I D where l smull chromosom(~ is Httnched to the ends of 2 chromosomes belonging to a (onfigulfltion of 4 Although Stebbiuslnd Kodlni (13) referrcd to such chromosomes as s111n11 they will be cllled miniature chromosomcs G in this bulletin to distinguish them from smull biurmed chlomosom(s and hom ()((asiollal frugments resulting from crossing oveL in illYClsiollS Theil fUlleliol1 i unknown Qut they can be transmitted through the poIitn grain as -ell as through tllO egg cell 1

Cl1RO)roSO~IE BEHAYIOR IX POLLEX ~LOTnER CELLS

DJPLOII)S

InformltiOll about meiosis in pollrn motbpr cells of diploid plnnts is bnsecl on obs(lvations mnltip on plttnts b(longing to tht (in~ 30+shychromosome collelttiolls from )Ipdco f1lld Oil some of the diploid offshyspring from thl nOl1controlLpd polliIltttion of p(lnl plan ts of 1 of these colltctions (425j) Th( diploid Hutme or ~n -- -ell romosome phtn ts is indicated hy the prescnc( of 18 bh-nlftlts nt dinkilllsi and metaphase I fiS pleYiol1~ly rcportld (1) rSllallY thl distribution at flllttplulse I WitS 18 18 although rarely 17 10 eould be found ~ttbhin nnd Kodani (18) also COl1lI1ltut(d on thl remi1rkuble uwiotic rcgnli1rity of diploid 3G---cbrol1losome plants

a liuitltIlre chromoollll arc nor iuclwlcl in the rliromoome lI11rnigt(r Imt arc Tlpf(ltrntCd hy a lttl(lltr-rript folll)vin~ l - i~ll

Hollin IL C rllpllbli~h(d data

EXPLATORY 1EGE~D FOR nilRE I

FIIHm 1-1 The 73middot~ ChrnlllOOml fwm atap(middottal ltpH of a plant of train 593 ote 2 milliat IIfr (hrlaquoJmo~lllll(~ tL and ChrolllhOIll( 0 that ha 1 afm of the ize atH happ of thp miniatl1f( chrorl(l~()mp H Thp 37-1 chrom()~otlp from n tapetal C(I1 of a haplOid from colhctioll 42) otlt JIIiniatllrc chrolllosome al C Two llIiuiaturp chTOm()~Olllpi IU and 3 bivalpJlt~ at mrtaphn-c I from 11 pollen Ilwllwr (Ill of a 72middot1-chroll1o~oIllP plant J) ~ixtl1 hivalpnts 1 uAivalpnt nnrl a eirrl(l of 4 rlrn1l10OIllP with minintllrp chromosomp (Ill atshytached nt ll1(aphnlf I from a pollpn mothr ((11 elf a haploid from collection 42357 B LatNul iew o( rhromoolllp at It1Naphnltl1 Wh(lIl hhalent (ormashytion was alrn()~t cOll1plptr from a polhmiddotn motlwr (tmiddot1l of a haploid from collpction 4267 )0(1 tll 1 tlllh-aput F ix biafgtuh IUHl 24 Ilnhalpnts at metashyphnlte I from a pnll(tl moth cfll uf a baploitl frum coUCctiOll 42108 G Iultipolllr piudl with 4 roJlP~ of rhrnIl1olIme at allaphns( I from a pollu mothpr ((Ill of n 106-lthromolme ab(rnl)1 plant J[ and 1 Chromoom(l in 4 nuclei (J[ and in 2 I nt proplias( II from POlIpll morlHr cell of a 107shychrom()~()me plaut f (hrnrno~()nl( at prophaltfgt II in twin pollpn mother c(l1s partly squepzNI Plt If a ruptured COUl11l0Il dl1e o( a 107-chrolI1oome abtrrant plant otc the 2 nl1rl(i Ul padl c11 and tll(gt 40 (hrQlllo()llllt in one cell and tht 61 in ill( othtmiddotr K ChrOn)~olll(gt of 2 Hudei at prophl1e II from a 78-chro1I10501l1t pollpn lllothpr ((11 of the l07-chrolllo~On1( plant that produced thos shown in J and I r Cllrom()~OJllC of 2 llucpi at prophni 11 from a (imiddotchrolIlosome poll(n lIlo(llPr ((1 of thl 107-chrolllooul( plaut m(Jllioned in fI 1 and K II Chromoollw5 of -1 nnclti at teiophtts( II frlt1l11 a 60-(hrotlll)orue pollen mother cplI of the 106-chrolllooIlW plant Ilwlliiolltd ill G S Tapetal c(~ll (ontaining 46 cbrOIIl050IllfS from n dik forN of an nhtrrnnt plant witb about 108 (hromosom(5 All drawing (Ccpt I ant f arp of tilt ame magnification The shading of some chrolI1osomes has IlO significall(c excpt to illcrcnse clnrity

8 TECHNICAL BULLETIN 018 U S DEPT OF AGRICULTURE

Bivalent formation at metaphase I is not invariable however One plant showed as many as 3 quadrivalents in some poUen mother cells 3 others showed one quadrivalent each in some This multivalent formation indicates either reciprocal translocation or polyploidy and may lead to unequal distribution at anaphase I The presence of inversions was indicated in plants belonging to 4 of tha 36+-chromosome collections by the occasional presence of bridges nt anaphase I or II

Two meiotic irregularities which aflected the quality of pollen werc founel in collection 426=5 In a snmp1e of 2)0 plants 4 had 90 percent or molc of the pol1(n aborted 3 of the 4 were examined cytologically and found to be asynnptic nt metaphase I tlHle were a few but varying number of hival(I1ts plus unival(nts 111() had 5 6 and 7 miniature chlomosom(s rltsplt(th(l~T Th( oth(1 irlltgularity was partial or (omplete failu1( oj cytokinesis fhis wns f01llldin 3 plants which (nm( from noncontlolled pollinations Only 1 was studied at metaphase r It show(d 10 hivnlel1ls plus a cllnin of 4 (or 18 bishyvnl(nts) plus 6 minintllres No oceUlate determinntion of aborted poUeD could he mnd( 80l11e of the grnins were compound consisting of a tetrad of incompletClV sCpalfltNl spores frCquentlv of unequal sizC OthC1 large gmins (1( ohyiously the result of dind instead of tetrnd formntion Giunt grnins the (Cjwiyalent of pollen ll1oth(r cells also were JUUl1ClQtlS

THIPLOIDS

Tnforl11ntion nhOIl t Illliosi ill polltn molhpl (ens of 3l nnd on(ushyploid tl plnntl is hn(d on sumplpH of thp fiyp gt4 plusmn-ltltromOHOlll( eo1shylldiollS from ~(xito und of tIll jmiddotlplusmn-cllomosol1w (0111 111 (1(i 111 stlair~ und nh() OIl Homp of thp i4plusmn-chroll10S011H ofJspIing from lH)lleOJ1shytroll(d pollinnlious of W+-chlomoscmw plonls or (ol1((liOI1 4255 and nO11(OJl I rolled jlollinH IiOllS of i4 plusmn-(]u0]110S011H pIn n Is of (olJ(etion 4203 growiJ1~in Uw Iipl(1 nl ~nlinnH Thl t)middot nn(lIploi(]s whi(h hod ](5S t111111 )middot1 (hlomosonH nnd SOJlW hie]) lwd l110rp thnn 64 (nme from 111(s( nOJl(onllOl1td pollinH t iom nt -nlinns In(llldpd n180 (IC

2 ofltYlw-lIolmn I plull Is [10m n 72 plusmn -ehrOl110S01l1C Ipxns (ol1(elioJ1 (hro1110S01l1lt h(ltnyiol dWllellrislie of triploids wng ohs(1nrJ At

dinkinCsis nlld llwtnphns( T tliyul(nts PIP Ht(I1 hut 111( numl)(1 vUli((1 with It (oJl(spondin~ yurying Jlumlwl of hiynknls nnd u1ishyvnlplltl lwlllY(I t]l( nJllnplJn~( r SU1~l (ould hp obg(lyltd nt Ipast a Iw tl-irnl(ntgt W(I( S((I JJte (x((ptiOllS (1( 111( 2ofrtypltshy110l1nn1 I(XIlS plnnts nne 1 olhll nH of hi(h Hhowld pllrlinl nsynnpshysis At nTlnphnsl T 01( 1llnjolit~middot or pol(11 moUltI ((lIs s])()ld Ingshyging (hI01110H0l1HS ~onH or whi(I divid(c PImiddot((o(()usl~middot 1ho~( polkn 1l10th(I ((lIs of jmiddotI-dumiddotOIllOSOI11( nlHIlI whiltll show(d Inglfing duomoshySOl1)( H(Imiddotng(] ~l lnggels whpnHs t1lO~1lt or 1l1ll1l pJolds nvPog(d 40 hut Ihis higlwr nYIllgt wns (tlllsltd l1lostl~middot h~ onJ~middot n fcw of lh( (plls ill whi(h thp 11111111)(1 of Jngftls WI lUImiddotgp TIlt I11lmigtN of lafg(Js whi(h is glYPH n~ n singh ll111n1gtPI d()(s nol inciieni( how s(trttpl(d they (1( in Ullt (y(()pln~m (1111gt1(1 n ChJPlllatid hliclg(s and smnll fni~nwllls (1( s((11 illflNI1llnlly Lngging (hl01110S0l1HS W(I( gt((11 I1lso nt nllHphilsP I r as I ~h()Jl hy UJ( disllill1ltion 23 2~ ao ~ 27 ~ [pw ((lis -ilo(d Ill( rlcds of Ingging at llOth n1lt11)luHws as in Ihe dislribution 2fi2(i2Wl 2i Log~(ls form

bull

bull

bull


microcytes which later can be seen as minute empty cells among mature pollen grains

T ADLE I-Distribution of chromosomes at anaphase I in pollen mother cells of 8amp and aneuploid 8x (49- to 56-chromos01lle) plants of guayule

[Miniature chromosomes not Included laggers indicated by ccntrnlllguro in serios or 3]

IOhromosomcs In 2 groups Ohromosomes in 2 groups only plus loggers

Group or plants Plants Pollen mothcr Pollen mothcr I fO~f W~ Chromosome Y~1Cil~S CdhisrtOrllllbOuStolomnO

ch~on~~sgrno distribution chromosome distribution dlstrlbuti0n

I Number 1o-u7IIber Number40middotchrolllosomo__ ____ bullbull_bullbullbull__ ___ 1 1 2425 20425

10527 ii1middotchromosomobullbullbullbullbullbullbullbullbullbullbullbullbull______ _ -- -- 201021

23325

1 2320 1 2HI021 1 2428 1 22-126 1 2527 2 21128lti2middotehrolllosome _bullbullbull ___ bullbullbullbullbullbull_bullbullbullbull__ 5 2 2020 23227

24424 25126

2 20~ 1 161021 t ~ _ 1 21131 t __ __ _ 1 21120

fi3-cbrtlmosoIllO bullbullbull_bullbull ____ bull_____bull__ _ 5 - ---- 1 21923 ~ - - - - __ -- - 1 22724 - ~ --- - ----- 2 ~3024

_-- _- ---- - 1 25325~-bull I -- --~ --- - 1 21226

2232 1 23526

2131 1 2middot1120

5 252U 2 2middot1228 3 2727 2 2432754middotchromosomc__ bull_________ ___ bullbull j 8 bull ____ a 25128

I _ ___ __ 1 25227 1 )5326 1 20127 1 20226I 1 25526 1 2531I 2 2fu30 2 20327li6-chromosomobullbull_bullbullbull__ bullbullbull _ __ bullbullbullbullbullj 5

I 1 2720 1 27227 I 2 2828

NIiniaturG chromosomes are less conspicuous as the number of ltlhromosomes increases in the polyploid series because their numbers 00 not increase No 3x natural populations 01 commercial stlfLins showed more than 3 and some seem to be entirely free from them Some of the 3x and aneuploid 3x oflspring from non controlled pollinashytion of plants of collection 4255 however showed as many as 8 Undoubtedly most of them were contributed by the egg nucleus since the seed parents of these plants had (j to 10 miniature chromosomes

bull In the majority of cases determination of chromosome number

oepended upon ch1omosome counts fit anaphase I prophase II and metaphase II stages Since the number of chromosomes in the two groups at prophase II and metaphase II plus any lagging chromosomes present reflects the distribution that occurred at the prccedL1g anashyphase I stage the prophase II and metaphase II records were pooled

OD05S7-middot46--2

10 IECHNICAL BULLETIN 918 U S DEPT OF AGRIClLTUTIE

with the anaphase I records and the totnl was examined to lenlll the nntUlc of the annphas() I disllibution

TIH distribution of ehromosOl1Hs fit fInaphase I in pollen motll(1 cells of 31 ilnd ilneuploid 31 plants wns obsernd in plantsthe chromoshysome ntl1l1bltIs of which ranged from 49 to on (lablt 1) For pollen mo(hltr cells showing no lagging ehlOmOS01lles in54-chrOlllOSOJlle plnnts the thtoletirnl distributioll cltleulalN] (cOIding to th( IHnciom disshytribution of 18 tlivl1lpnts indlldes 10 elnssls from 1830 to 27n Only )0 poll(11 molhel cells W(l( olltiPln(1 or tb(s( 1 show(d 22 32 1 2831 5 2529 and ~ 2727 TIIP p(lCPulnges of obsPIYed flCltjlllllci(s WPIP (ompnr(d with (hosp of cnl(ulntNl flpqupneips Obviously UH obsPlnd ll11mhll of ((lis wns too smull 01 th( eompfilishyson to 11n( tIlY llHllning hOWPlI llplt VUS IlO rlisngl(PJ11lnt bp[ (tn cnleullllpd find olistly((l fl(lt]UlIlri(S In 11Huploids with h$s lhnn 54 chromoso)l1Ps lh(Ip (1( olll 1 lnstpnd of ~ of 1 01 11101( of tIll IS kinds of Chl01l10S0mPR (r= l~) Th(s( 2 Clll0Il10SomR prohn lily fOlm n bh-n(nt nnd sppnrlltl lo oppositl pollS of tIl( s[lindh nt fllUlphnsc I and lh(llby (ontrilHllt town Id all (((lInl distrihution of nIL fliP (hloJ1lo-SOUltS to the 2 dllUghllI llUelli 1(l1t (([unlity in nllmlH1 of phlOJ1loshySOJ1l(S in tlH 2 dnughtll 1111ili is llHlpfOlt ltXl)((t((1 bullllhollgh the Dlnubl1 of poUtll mothl (plls is smnll this is ilHlienltd This is [nlc also fol tIl( j 0-( J)]110 so11 H pllIIts which hlYpound 2 ltxltn lJtomosoJlu1S 3 being llH grpntlR[ dl-inlion from (I(lllltlit~- (2j31 jns(pnd of 2S 28) This too is (OllRis(tllt with lxjlPctntion TI1(s( 2 nd(litiOlltl chlomo-S011le (ithll COIlYl( 2 of tH tljYnlplls into qnndlh-t1ents (4 of fL kind) or (ot1s1ilutt nn ndditiollnl pllil (2 of n killd l III pitht (I15C tlWY dl(](IlSl UIl mnount of odd bnlull(( illld i1HIlflSl t hnt of lnm bnliuHmiddot( thus ilJilulll(ing till (pudlllty towtlld (quul c1istlibutioll

Among tll [lollll1 11lotlll1 (dIs from j4-(ttomosOlllt phLllts that show(tl Ingging ctrnlllOSOl1lS th( distributioll ttl nnnptns( I 1(11 wilhin til( llIlIgC town hy (plls without JU1g(IS Th t1f (11 f(w(t ChlOI1l0OI1ll UIOlI ps wit h moll t lin n 27 (hro1l1oS0J11lS 1gt11 t nol ntOlt gloups willi fls [hall 2middot1 (hlOllHHOlll(S UltllOUgh tIlt Inttll would bl~ eXpt([(11 This 11lt1lt1 lnll also for tht ij(J-(hlomoSOlllt plnllts_ 1lt01 tlH 40- to j~-dlJ(lln(Jollll plunls tll( PIPgt(ll(P of 1lI01( IngflIs IN pOn(1l lllo1h(l (pH llot (lIlly (lpl(ns(d I h( lHI11l1lPl of ehlO)lloOlll( groups with 11101( thltll half or thl somnJi( (hl01110S0111( IlU1l1bPl but fIl~O itH[pas(d tl)( lU1l1l1llI whit-I had l(s lhun hnlf

If thr rhlomoso]))( gJoups from nil or 111(1 polil1l motlH ((lis nrc considlrrd inltljvidually only 11 cOl]sistpd of L(ss than 22 chrOmOSOll1lS For tilt 54-ehromosnnw plnn ls -II) 1)(1(111 t contltil1ld 24 to 29 (11101110shySOI11(S iJl(lush-C F(lI thl middotl- to j~-(h[()mosonw plnnls 71 prJlrnt conlllinld23 10 2R llllOI1l(lSomtl Fol til 5U(hlm)10S01lH plnnts FlO prl((nt (onlnind 2j lo 28 chJl)l1losoll1PS lhpl(fol most of tho poUtn glltins ])lObllhly (ontnin(tl molt thnn 22 lud Pss tltnn ao chrolllosol1lp (PH nflpI nllowing foJ nddiliollnl lnggil1g of (11101110-SOIlWS iLl IlllnphnSl 11 Xot t1ll plOdu(((l yiuhlp l)ollpll nl1d proigtnbly TCI jPW ((Hlhl flltili7(l fUl lgY Illlcltus 1)OW(IA wd Hollins S rflitd 54 plusmn-ehroll1oso])W ph11lls POOl ~l~ ]gtolhl pH 1(11 ts

OIlW dUlIt indi(nting lh( IIIl)lllOlOnH Humber of futletionnl pOlllll glnill~ plOdu((d by 3c plnnls were ootnill((l hy cl(Lelmining th~

8 POWNS L llld nollin~ n C rllpuhli~hcd ltlnla

bull

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11

bull

bull

bull

POLYPLOIDY AND ANEUPLOIDY IN GUAYULE

cluomosomc numbers of aberrants belonging to the triploid collecshytions 4257 and 4264 and aberrant offspring fro111 nOllcolltrolled pollinations of field-gtown plallts belonging to the triploid collection 4203Sinco abclrants arise from thC fertilization of Eggs which have the unreduccd Ch101110S0111C llumber the plObnble number of chromoshysome contributed by thC pollen could be d(t(tmined hy sUhtrncting the llumb(ll contributcd hmiddot the Cggs from thr totnl (Uthlc n For collections 4257 and 4204 the plohnbh cluomosollw numlwls of eflCctiyc pollen lnuged from 2(3 to 33 Pollcll from both 54plusmn- and 72 plusmn -chromosome pItnLil prohn hl- itrlilizrd plnnts bronging to 42(33 as indifntNI b- thc rnngC wh ieh cxtlll(lltl to au Also 1 nbelTfLnt ofrspring flOm I i)4-chlomosoI11P s)HCies hybrid hud Sigt to 87 chlomoshy50mr5 i1Hliculing fertilizutiol1 by n 31- to a3-chromosonlP pollE1l glltin

TAllLF 2- R~Lilllalfrl 1l11111lifr af rhl()IJIOql)ml~ ronrUmrtf [1 pallnt grains irwoted in productioll of (Ibrrrllni plants in three fltlLtlll~ of glUlllle

~_______o_o__o_~ _______0

Sum~fT Sumher ltl 1-32 l ~ 27-31 IU -36

In ot(IPl to Palll whltllll UllY pOlllll glllin that ill plOlhwld by triploid Illlli lin 1 11 IhUll ~Ii (hll)1ll0~()1ll1 (1m Ifr1t i1ltilizflolion fL ((lilt lolhd (10 was llmdt h~ llil1g phllts hlonging to (ollpItion

J

middott)i) (~i- (uomOSOIl1PSl ns 11ll 1(11 PUIPllt illltiplunts lHlonging to (ollt iOll 4~()2 j IU() moo lUIS I 1 deg tIl( pollell PIUPlll The (hlOmOsolllt lUllltlHIS of tl P of t Itt ofisplillg lIIP dltllmineltl TI(il lHlTUI Ht- 14~ 4) -11 -1 and fJ I indi(t] tlII fllliliztt 1011 II pollen glUilli whirmiddoth hlld 2) loH dUOlllOSOmlS TlwlpfOlP on IhI basis of (ot t toUld ttlld 11011(onl01l1d pollinnliolls it nPPlllIS t hnl t 11 plnitls Illolmpd flllllliollnl POIlPll ~11l11H whillt (OIltUillld ~) 10 3B (lIromoshy801ll wiwlltus tlIP lll(ditioll mnd frolll u stllllmiddot of tllltlphtlltl disshytlilmtiolls wu~ ~~ (II ~J tPlltll(middotllt Iy gmius whilll (()tltuiUpd 2 28 or ~4 chLOm()ornl~ Wll( ttouflllllliunal

ToB(l 3-LJislrUtlillft 1) rhrtlmll~I1I~ fI 1IItIflIllbull I ill ]loUo ml1tJlr cElli I)f ~ mlt1 oi-rhrmnll1f plllllt~ IIf (Iflolt

~rjrulltttr~ (l~ri nH~rl$ UII~ Itf uit d lat-middotr lf1b~Vfd by (tntrl1 fl-Urtl in s(~rh f ~

rllrmiddottn(Irtmiddot H12 rlulllt lin

fhrltrtIgtoltm In Jr~m(H plui ~lHS

I nH

I hrumo$rne 1htnbuuQII

43-chromoslIlIW bull bull Ivo 1~3 ~221

2O32l

12 TECHNICAL BULLETIN 918) U S DEPT OF AGRICULlURE

Meiosis was ohsorve(l in one 43-chromosome plant and two 45shychromosome ones At metapluLse I 1 of the laUel showed 19 bivolents plus 7 univalents in a pollen mother cell whereas the other showed 16 bivalents plus 7 univalents plus 2 trivolcl1ts The approximation to equal distribution in 9 pollen mother cells at anaphase I is rather good (table 3)

TETRAPLOIDS

Information about meiosis in pollcil mothel cells of 4x and alleushyploid 4x plants is based on samples of the 9 collections from Me-icoJ15 percent of the extensive Tcxas collections 25 of M cOnllums comshymercial strnills und nOllapomictic progeny of controlled pollinations in experiments rlus sampling of tetmploicls includ(ld 2 morphoshylogical type the so-called good gunyule type W11ich gives no evidence of previous ndmixtuJc with 1l1fllioJa and U1e intelmedinte type which docs show some of the characteristics of mariol and for the origin of which introgrtssive hybridization with l11aliola has been proposed by Rollins (11) Dnd Stebbins and Koul1ni (is)

From such wide sampling a gentlOJizcd over-oIl picture of meiosis in tetroploids was obtained In apomicticlllly produced 4x plunts of both jntelmedintes nnd good Ounyule types quadrivaknts were seen but the number of quuclrivafents and trivalCllts varied among collection some showed very few of 1hem and others more Possibly as many os 6 chromosomes were involved in occasional configUlQtions In generol at llletuphase I plonts which had 72 73 Ot 74 chromosomes showed mostly bivulcnts plus 1 Ol a few quacllivalcnts possibly tlivalents and n few univulents The intermcdiate type seemed to have more quadrivalents and polyvalents than the good guayule type and correspondingly fewer bivalents In view of the origin of intermediates inyolving introgressive hyhridization with ma1iolu some of these polyynJe~lts probably are cutenations of cluomosomes f01111ed os a consequence of structural hyhridity MallY pollen mother cells showed lagging cluomosomes at unnphasc I All of the intermediates ayemgecl higher thon the good guayule type in the number of cells having lagging chromosomes and the 72- and 73shychromosome intermediates nvemgocl morc laggers pel pollen mother cell thnJ the good guayule type of the same cluomosome number The good gultyule type (72 73 and 74 chromosomes) had fewer cells showing loggers than the 54-chromosome plants but the avemge number of laggers pel cell was about the same for both However in proportion to the totul chromosome complement the number of lvggels per cell mnong the 72plusmn-chlomosome plants wus less than among the 54plusmn - chromosome (tobles 1 and 4) Chlomotitl bridges with or without imgments weTe not common but they were seen occasionully in many plants of diverse origin Some lugging at anaphase II wete observed 01so

Meiosis in some of tho plants classified morphologically as offtype normals was inc1isLinguishable froIll norIllQls whereas others showed meiotic illegulolities Two of the illcglllorities observed in a few diploids were found also among aneuploid 4x plants They were partialosynnpsis seen in somo alleuploids the cluomosorne numbers of which ranged from 65 to 71 and less frequently failure of cytokishynesis seen in a few aucuploids the chromosome numbers of which

bull

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bull

13 POLYPLOIDY AND ANEUPLOIDY IN GUAYULJil

TABLE 4--Distributiol1 of chromosomes at anaphase I in polen mothel ceU~ of 4x artd aneltplotd 41) (87- to 74-chromosome) plants of gltayule

[Miniaturo chromosomes not includedj laggers indicatod by contrnl figuro in series ot 31

Ohromosomes in 2 groups Ohromosomes in 2 groups only plus Inggers

Group ot plants Plnnts Pollen mother PoJlon motherI cells with Ohromosome cells with Ohromosomo

cl~~g~gg~lo dIstributIon cl~~~~gig~Je distribution distributIon distrIbUtion

Number NU1IIber NIL11lbcr 67-chromosomo_ __________________ 3331 1 313331 1 33131C--------- 69-chromosomo____________________ 3336 28 1130

- 317313 L---------~-70-chroUlosoUlO ____________________ 3 3337 1 31435

2 3130 1 33235 2 3535 1 34 135shy

71-ehroDlosomo____________________ 353~ 1 29735shy 1 31334 L---------~-

Do _________________________ bullbull_ 3530 2 3313711 L---------~- - __ --- 1 34235shy

7 3438 31338shy12 3637 33331gt

72-chromosomo ______________ bull___ bull 10 36J(l 34137

bull 20 -- 342311

- - --_ 34835-~

- --- - 35 136

3240 1 327333339 1 33138

Do_ bull ________ bull _____ __________ 3138 2 33336 110 3537 33534

3630 34236 - 1 34335tJ 2 31 136

73-chromosomo _________________ bull__ 2 3439 1 35336 3 2 a538 2 36130

middot1 30a7 --- --- _-

3538 1 32530 3037 1 33238

___________ (- --_ _-- 1 334311Do______________ bull 18 - -- __ 2 34138

- --- - 1 35131 ---_ - - _-_ 1 35236 ---_ _------ -------i---_ 1 35335 _ _--------- _- - ---- 1 36136

1 34middot10 1 3314074-chromosomo____ _______________ 1 35 a 1 35435

31138 30 137 8 (I

1 3737 - - -- -

2 31238 Do ___________________________ _ 1 2-1337

1 35138 2 35231 middotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddot-1 1 353311 1 30131

I Intcrmedinto typo

bull rangeJ from 68 to i2 clllomosomes Two offtype normals belonging to the good guayulo type showed many Jaggers at anaphase I and II but llt many miolocytos among maturo pollon glains IowevCl theshyone WIth loss than 72 c4romosomos showCcl 50 pClcent of pollCn aborted and the othel with 72 chromosomes about 98 percent rlfale stelilshy

14 TECHNICAL BULLETIN 91S U S DEPl OF AGRICULTURE

ity us evidenced by scunt sporogenolls tissue WitS found in sevCIfLI plants (69- to 72-chlomosome) Restitution nuclei (scmihptelotypic or pseudohomcotypic) wcro SPCll but thpy WClC confincd largely to plallts ith lcss thfLn 70 e1uomosomcs nnd usually nssociitLcd with IJiLltial fLsYllapsis Often 2 or morc irrcgu1tlhtits wcrc found i[l the same pIunt tspc(ially mnong the 65- to 69-c1uomosomc plnnts MiniaturC chrOlllOSOIl1lS WlJe SC(l1 in somC 4r and fLllCUploicl 4x plnnts AL most they wcrc lirnihd to 2 ThC )lCxienu materjal cxshycept cOllection 4258 scCllwcl to bc without nny tlw sceds fo] 4258 came from IUl individual plauli growing within tho diploid POpuIfLtioll Jlom hich collcction 4255 was obtained (Ill collection 4255 the numbcr of miniutuJts lnngcd from 2 to 10) [ost of the plants from the commereinl stlnins had as n~any fLS 2 hOWeVl1 some plnnts with only 1 W(1(gt found in st1nins 406 nud 44middott

1he clistrjJmtion of c1uomoson1Cs nt nnaphnsC I in pOl1tIl mothcr cells of 4t tlud fllwuploiltl41 plfillts wns obs(JVld ill pln111s thc el1Jomoshysomp numl)Crs of whieh lflllgpd f)om ()7 to 74 (tnblt 4) For (plIs showing no lngging ehrOmOSOIllPS in 72-lhrOlllOSOllH pltltlts helollging to the good gunyuk ty]W 111( ohs(rvpcl mnge or distrihution at nrUlshyphnsc I middotns from 34 38 to 30 30 with tIll following fleql1C]wips 7 showCC1343S 12 3537 und H) 3(j3(J FOt thosc belollging to the intelll1ldintp ty]W it was flom 32 40 to ao 30 with the following fleqtHneies 1 showed 3240 13339 2 3438 10 3537 nnd 5

x230 30 A tlst ShOlWd thnt U1Pse 2 clistlibll tions were not signifishycuntly cliffCtlIlt Tl1l plr(pntagps of ObSllcd fllCfucneies in thCse clnssls Irc)l comrtultd with thl PlJ(clltages of lxpcctld 011lS Cllleushylntcd nccoL(ling to the mndom distlibutioll of IS quadlivnIcnts For the good gUfLYlllc type 111 of t11l obsClvod fl(qlHIl(ies Wlrl in 3 dnsses in which according to cnlculntions only 595 p(lccnt should he for the intclmcclintc typc 894 pClcent WPIP in tlws(gt same 3 clnsses This lack of close fLglClment bctwccn obsCJvld distribution fJequeneies at metnphnse I and thosc cnlclllntcd nceOldillg to the litlldoJ)l distrishybution of 18 qundrivnlents is thl lPsult of [tn obvious tcndtncy townrd equn1 distribution of duomosomcs to t1tt 2 poles of the spindle nt anfLphusc 1 If 72-ehromosome pInnls fOll1led only 3G bivalcnts at melfLphnsc I n 30 36 distribution would be CX]wctccl nt nUfLphasc 1 If they fOlmed only 18 quadlivnlents fLt mcULphnsc I closc agrccment woulll bc expectcd with distributions cnJcuhLt((l necording to the random dislribution of 18 qundlivalents Aetunlly fLll shoed some bivfLlcnts nt metfLphnsp I the good gtliLYulc type ill gCJ1Crnl showing more thnn the interJ1lNlifLtrs Therefore the ObSC1ycd distributions are consistent with thp obslLwd tendency towfLrtl bivnlcnt fOlination at the 4r level in the polyploicl srics

Fot cells showing no Ingging cluomoSOllllS in fLllPUploids with more 01 less Hum 72 cluomosonlts the tCnclcncy towfLrd nn cv(n distribushytion of chromosomcs to cnch pole is cvident nnrt nowhere dops the distribution fit OIlC basltd on qUfLtlrivnlent formation

For po11(11 mother cells showing lagging chromosomes in 72-chromoshysome pliLlltS the muga of the fLl1fLphnsl I clistributioncloscly approxishymated that of POUNl mother cells without lagglls although there wcrc fewer chromosome groupS with morc tlmn 36 and more with less thnn 34 chromosomc~ rhc samc WfLS tme for thG aneuploid

bull

bull

bull

bull

bull

bull

POLYPLOIDY AND ANEUPLOIDY IN GUAYULE 15

4a plants namely fewer groups with more than half of the somatic chromosome number and more with lops than half

If the chromosome groups from all of the pollen mother c(11s which showeel lagging chromosomes arc eonsiderecl indiyicLually only 11 consisted of less tilltll 33 chromosomes For the 72-chromosome plants 886 percent contained from 33 to 36 chromosomes inclusive Even the 67- to 71-chromosome plants had 54 percent of groups with 33 to 36 chromosomes The 73-cluomosomo plants hnd 625 percent of groups with 35 to 36 chromosomes and the 74-chromosome plants had 636 percent of groups with 35 to 37 chromosomes Thereshyforo even after additionnl lagging of chromosomes at anaphase II most of tho pollen grnins probnbly eontaincd not less than 33 or more than 86 ehromosoll1(s

Some data indicnting the chromosome l1Umbrl in funcfionnl pollon grrt1ns was obtained by determining the chromosome llllmber of 50 aberrant plants belonging to 41 collecLiolls and lil10n subtracting the probable numbetmiddot of chromosomes contributed by the egg The rnngo extond(d from 28 to 40 usunlly from 31 to 38 Also 7 abershyrants obtainlcl from controlled pollinations wherc tho chromosome numbcr of the pments also had been detcrmined showed thnt the functional pollen grains contained 31 to 39 chromosomes 1heso data on aberrants ther(fore inclicatrd a widor range of eluomosome numl)(rs (ontninedin funetional vollell grnins tlum WilS estim1ted from the nnnphnse dnta in iho pllclding palmiddotagmph

l)LANTs VrIH HiGHER CIlHOl[OSO~m NUllBEllS

Information about meiosis in pollen mother cells of plants having 81 01 more chromosomes is based almost entiroly on abellant plants which occurred in natural popu1i1tions and commercial strains and among progeny of con trolled and 11oncon trolled pollillations in experiments

Aberrant plants occurring in 54plusmn-chromosome populations and strains and among progeny of 3x seed parents rangeel in chromosome number from 80 to 93 Observations of the metaphase I stage were confined mostly to plants belonging to collection 4257 Some tri shyvillents find polyvalents and a fow uniyalents were seen but bivnlents preclominatrd therefore the tendency toward bivalent fOlmation which was evidenLin 41 plants contillllCc1 on up tluough the polyshyploid series fhe flequrllcics of chromatid bridges and c111omosome lagging n t annvhase I and II were about the same as for normal vlallts in the snme cultures As many as 3 minintulC chromosomcs were seen in some plants somo had fewer nncl othels had none rfwo plants with about 85 chromosomes had scnnt sporogenous tissuo and therefore were practicallynale sterile

The tenlti(ney toward rqual distribution of chromosomes to the 2 poles of the spindle at nllllphnse 1 is apparent ill pollen mother eclls of 81- to 93-chromosome plants (table 5) It was cvid(nt eyon in It

122-chromosome oflspling from a stlf-pollinated 83-dllOmOSOmc plant where 2 cells showed 59 (j~) n]1l GO 62 distributions at IUlaphnsc I Among pollen mother coIls without lagging chromosomes the extreme deviation from equal distribution wns 36 47 (instead of 41 42) and among those with laggers it wns 33 14 39 in Ull 86-cltromosome

------------------------ -------

16 TECHNICAL BULLETIN 918 U S DEPT OF AGHICULTURpound

plant As in the cas( of 31 and 4x plants cells with lagging chromoshysomes had fewer groups of chromosomes consisting of more than a half of the somatic numb (1 as compaTed with those which do not haye laggers

If the chromosome groups from aU of the pollen mother cclls are considered individunJly only 1 consistNl of l(ss than 35 clllomosoI11(s For the Sl-chromosome plants 75 p(rctl1t of the groups consist(cl of 39 to 43 chromosomes il1clush~c Because of more lagging chromoshysomes in the So-chromosome plants th(y had prncti(~ally the same number of groups (71 p(lcent) consisting of 39 to 43 dumiddotomosoI11(s For all of the c(Us 69 percent of the groups eonsist(cl of 39 to 43 chroI11osol11(S find 85 P(lc(l1t 39 to 46 chromosomes Probfib~y most of the pollen gmills contained fit least 3S cluomosomcs bLtt not more than 43

TABLE 5-Distribution of cllr01ll0soll(S at Clnaphase I in pollen molher cells of 81- to 98-chromosomc plants from 8x (51plusmn-chromosollle) seed plants of Ulwyule

[finiaturo chromosomes not incluCled loggers indlcjled by central fl(Urc in scries or 3J

I Chromosomes in 2 groups i Chromosomes in 2 grOups onl I plus laggers

---------------Group oC plants Plants PollcnnlOther iPollen mother

Clll~ with C 1 cells with i Ci indicated I ~rol0s~mc inClicnled hromosome chromosom~ dIstribution chromosome distributionI distribution I i distribution I

Xltmber I Xil7llbcr I I ]o-umber II

il 1 3045 I 1 3S142 v 1 I 3-14 I 1 1 39240

aImiddotchromosomebullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull1 iI gm =I 82-chromosomebullbullbullbullbullbull--bullbullbullbullbullbullbull- bullbulli I I 1 IJ 4141 I 1 I 39241

I S3chromo~omebullbullbullbullbullbullbull bullbullbullbullbullbull__ i

2 i I ~~ 11middotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddot~middotmiddotmiddotmiddotmiddotmiddotmiddot~~ 1 4244 1 I 35942

6 I 1 4343 j 1 I 3U~4386-chromosomcbullbullbullbullbullbull bullbull_bullbull_bullbullbullbull[

+1 1i 3~~~~~~I l middotmiddot~lmiddotmiddotmiddot-middotmiddotmiddot~~~~- 1 I 4_143 88-ch romosome bullbullbullbullbull ____ bullbull__l 1 1 I 4444 bull- bullbullbullbullbullbullbull ---- bullbullbullbullbull I

1 90middotchromosomebullbullbullbullbullbullbullbullbullbullbull--bullbull--1 tI ~~ Jmiddotmiddot-middotmiddot--middotmiddotmiddotrmiddot-middotmiddotmiddotmiddot--middotshyt

g3-chromosome - ----bullbulli 1 1 1( 4084amp I

Aberrant plants occLlrring in 72plusmn-chromosome populations and strains find llmong PlOg(IlY of middotlx sttd parents rang(d in (hlon~os0ll1c numb(l from 90 to lO OhS(lTUio))s of the mltnphas( I sblge showed a prppoll(ltrnnce of hiYlllpllts hoey([ SOIllt plants showed many triyalpnts and quaclrivahJlts alHl othcrs many uniyalents This Yfiried behnyior nlny dp]Hncl upon tlH fnrt thnt some of the abellaI1 ts in populations of the good guayule typt had mariola as tlllu pollon pn1lnt llnd SOUlC Wll~ from CintClllwdiatc populations Chromatid hridgts nt iU1uphnse I w(n S((11 only infIWlUel1t1y xot more than 3 mlllifLturc chromosomes W(lP COlUllld in 1 plant Cluoshy

bull

bull

bull

bull

bull

bull

POLYPLOIDY AND ANEUPLOIDY IN GU~YULE 17

mosome lagging at anaphase I was frtquCut but limitNI to only a fCw chromosomes so that in proportion to tht total it did not materially reduce the number of chromosomes in the 2 groups in (ach POll(ll mother cdl Lagging at anaphase II was s((n only infrlquCntly Malc sterility in the form of scant sporogenolls tisslle was obscrved in a lOO-chromosome plant

The tellcilncy toward equal distribulion of chromosomes in poll(n mother cells of 90- to 109-chromosome plants to tlw 2 poks of the spindle at anaphase I is cvident (table 6) Among ((lis without lagging chlOmosomes the extreme d(viation from (qual distribution was 49 5S (instead of 53 54) and among those with laggers it was 47 4 57 in a lOS-chromosome plan t Cls hich had lagging chromosomes IUld somewhat fewer groups consisting of morc than a half of the somatic number than those which did not haYl~ laggelmiddots

TABLE 6-Di~tribution of chromosomes at ana phase I in pollen mother cells of 99- to 109-chromosome p(mls from 4x (72plusmn-chrol108olltJ) seed lJ1a1lls of gllaYlIle

[lnnilltur~ ehromOSOIl~S not illelt1d~d lO~j(~rS ir1If(ot((1 by e(ntrnl tl~llr~ in s~rles of 3)

I Chromosome~ in 2 groups t Chromosome III 2 groups ol1l~ plus laggers I 1---------------

G roup of plants Plants hOletll~~ther polIN~ olheri

I ~ells Ilh ChrIJIIIOSoIllC cell itll 1 Chromoomc mdlcated j t b t i IndJclI1(u dmiddot lb J

chromosome (IS rI U 1011 bull chromosome l Islr ut onI distribution I llistrJbulloll

---------------- -- --- ---_- ----shy-~---

ggmiddotchromosomc _bullbullbullbull _ _bullbullbull _1 Sumber Imber

Sumber 1

1 I 411350 middott250

lOOmiddotchrolllosome bull _ _bull bullbullbullbull

lO4middotchromOSQIIIC _ _ w ~ ~ bull 1

21 I

1 I

)

t

f 1 I1

I1 I 1 i

4S2f~) 4024Y

481pound5 40IfgtI 51152

lOS-chromosome_ r I2 2

[(Jili

~ - - ~ ~ ~ i

4~IW middotll254 51153

i

I06-chrolllosomcbullbull 2 fl~51 I 1 J

492pound5 51352

lOimiddotchrOIllQSQII1()bull ~ -shy (f

1 IJ

1 a

~ ~ ~

495~ 51 flG 5255 53[1

--

1 1 1 1 2 1 2

4S25i 50IrtI 5O2f5 513[a 52104 522~3 sal5~j

IOSmiddotclIromosomc_

IOUmiddotchromosome

3

I 1 Ii

2 1

[)()5-

filii rJb5

lfi25 H~55

1 J 3

2 L 1

4i-tZJ7 51552 52155

5315 532middot 5-11pound4

If the chromosome groups from all of the pollen mother cells flre considcled indi-idllal1)~ollly 1 consisted of less tban4S (hrol1losoTlles For tbe 107- to 109-chromosome plants 73 prlc(nt oJ the groups consisted of 52 to 5f) ebromosomeB inclusive For all of thc (rUs 62 percent of the groups Consislcd of 52 to 55 chromosomes and 83 pcrshy

GfJ05S middotmiddot010--3

IS TECHNICAL BULLETIN 91 S TT gti UEPT OF AGRICULTURE

cent 49 to 55 Probobly most of the pollen grains contained at least 48 chromosomes but not more thun 55

Aoong the plogcny of aberrant plants belonging to collection 42 1(l3 there were some plonts that showed only 0 few of the morphological characteristiCs of abCITants a1though the ehromosome numbers ranged from 74 to 03 rrhe~r werC eullCd ofHype normals from abelrlwts Pollen mother o(lls of (3 plnnts whose eh0l110S0me Ilumhpts rnnged from 85 to 92 W(gtJO studicd Lagging at anaphu$c I WilS limited to n relatively few clI0l110S0l11es Although the dntn for I1nnphllsC I distribution arC scant (table 7) they show thC sam( tendcncy toward equal distribution to tIlC 2 poles of the spindle that was C-jdent in the abcrmnt 11unts which had npproximatrly the saIlle number of chromosomes (tabl 5) Thf (xLr(J1)(s ill deviation from fquality canle withirl the limits of deviation showlJ by thf abelTants From the standpoil1t of meiosis in pollf1l molhel ((ils thCICfore lhiF group did not difler from othe~ plants having high chromosome numbers

TAllIE 7-VistrilJllion oj cilrOlllosomrs at poundl1ll1plwse I In 7JOllen mother reUs oJ ojJIYlJ( normals jrom uJerrmlt 8li- 10 92-r(rollosome plants oj fllIayule

[llnlnure chromosomes not included tnggers Inellentcel by central llgurc In sPriC$ 01 31

I (hrom()somr~ in l groups ChromnSIlJUIS in 2 (roups ouh plu llg~ers

Plants ~~li ----- r(ll-trtmiddot~---middot--Orou(l of (J13I1t~

((lIs with I ((lIs withltldicltrl IC~rom)()o(lJl~ iltdkJtllt1 Chromosome

rhwtnuflnH ltlbml uti m rltrOIlluSflTtI( dislfflwtion I Jtrlhutlon IlbtrihutiuJI

----1--1-- _middotmiddotmiddotY1I1IIber i

85-ehrol1loS(JlIl( - - _ - - - I l 1l 21 lU11

42-1 r

OI-ihro1osome bull

8r-ebrolltosom bull

I I

1

I

middot1~49 -1-117 i

3U5-tfi lt1-11+1

middot121middot1$ middotJ5145

02middotdlromOSOJIW bull 1 1

4(1250 -1-1117

HAPLOIDS

As plcviOl1Rly ](porpd (1) [) f(w plants with 36+ (hlomOsomC5 WNe found in 7~plusmn-chromoS(lnH populnlioI1s Infonnntioll nbout llwiosis in poll(J) lJ1oll(r (p]J of sllh Imploid plnnls is hiwd on oh3pl-nlions mfl(( on n of llHm widell (1( sl ud iea eylologi(illly 1 had ~G+l chromosomes i luH1 7+I nnd 1 Iud 17+2 III t1 lap(tnl ((ll of thc hnploicl from conNlion 4~~ii 17+1 eitromosonHS W01( C0111t(l(1 (fig 1 B) All tht hUl)loids ht)(l at lenst1 llliuintun th 1011l0S() l1lC 8rlO log(11011S li-st1P was srttnt (lnti tll( tIls of l)olhll motlHl c(lIs W(lP (xtrtmCly thill Ill( l1wtnphns( I stLl ns obs(lYtd in S plnnls 1111 of whi(h howPll some hjYIPnts 2 pinnls sho(d only ( ft itnd 4 pm(ticnlly all ([tg 1 f) h ftnd rl (ilplp of lt1 (no11105011(S wns S((I[l in SOIllt pollell lllotlH1 (middot(lIs of tht huploid from T(~middotHlllS Tpx (fig I D) and 11 dmin or 4 in (PUs of the Jutpioill from train laO TliCsc collfigllllttions of 4 chlotnosollllS mny indi(nt(~ leeiploenl

bull

bull

bull

19

bull

bull

bull


translocation In 1 cell the miniature chromosome was attached to the circle of 4 (fig 1 D) Distribution of chromosomes at allophase I in 7 haploid plants (table 8) was less regular than the distribution in diploid 36+-chromosome plants One pollen mother cell at anaphase II WitS observed the distribution 18 18 17 19 indicated an ill(gshyulurity at this stage also Practically nIl of these haploids produced very little pollen or the pollen grains were mostly empty

TA BlE 8-])i~triblllio7t of chromosomes In 1Jollen mother cells at anaphase J of hapshyloid from 4z (86- and 87-chrOlllosome) seed planls oj guayule bull

[Miniature chrOlf osomcs not iDClud~d Inggers iuUicnied by central figure in ~erics of 3]

ChlOlllOSOllllS ill 2 groups Chrolllogomp~ in 2 group ortly 1)lu5 laggers---------- --_-----_--

I

010111 of plnltls jlnuls Poll~tlmOihr Pollen mother i cells With Chromosome ~(Il with Chromosome t c~~~g~g~c dllrihutotl cl~~~~~~~C tlistribution

distribution ditrfbutloll I- ~-~ ~---- - shy

XIIIIberxumber )620

30-chrorno$onlC bull _ 2 1710 bull _ 118

37-chromosomo __________ bull-_ ___ i

1 1 17218l~~l 1 ISl18I degi

--------------------_ --- Bivalent formation in these haploids demonstrates that there is

consiclemble homology amo~g the 4 sets of cllJomosomcs ill the 72plusmnshycluomosomc plants from wIndl they arose Howept- such homology ought not bc used as evidence fot autopolyploidy as opposed to alloshypolyploidy since it is known that some puidng OCellIS even between gunyule and mariola cluomosomes in hybrids between these 2 species For instancc two 54-chlOmosome naturnJ hybrids were the result of fertilization of 18-chlomosome gultyule (ggs by 36-chromosome mariola pollen gmins In 1 hybrid a maximum of only 10 uni~a1ents was se(n whereas in the other where the ma-imllm of unhalents was only 6 from 1 to scyeral triyalents were observed

These studies of chromosome behavior hayo centered around associations at metnphase I distribution at anaphase I and for plants haying 54plusmn and 72plusmn chromosomes the (fflcts of this distribushytion on the production of functional pollen grains Throughout this series from 36 (both diploids nnd haploids from tetraploid) to 120plusmn it is evident that meiosis in pollcll moth(t cells in most gunyule plants at all levels in the polyploid series is complete reduction ot chromoshysome numbCl is effect(lti and a qUiLrLetof miclogtporelt is formed within the cell by division of 1h( cYlopliLsm after telophnse II_ Itl only a few variltnts hltve exeeptions to tltis llOlmlt1 bllmio[ been found (such us lack of sporogenolls tissue ilboltion of aU pollen Ot other meiotic uTcgulality)

In geneml thelc was a tendency towurd equal ditlilmtion of chromosomes at ilnaphlSe I throughout the polyploid selips llgllrdshyless of whetheL the plnnts were of Itmphimittic or apomictic otigin Itlld legnLdless of the llllmbN of dllomosOIM Among 8li+-chlomoshysome diploids ill1ltphase I ditlibulion 18 ptedominttntly 18 18

20 TECHNICAL BULLETIN 91S 1 S DEPT OF AGRICULTURE

Among 36- lllld 37-ChlOIllOSOI1H haploids from 72plusmn -chromosome POPUllltiolls the distribution wus kss regulul Among triploids ana-shyphase I distribution US not in disng[(gt(IlHllt with eukulntions b05((1 on tb( rnndom distribution of 18 triYnl(nts uJthongh the number of obs(lYotions wus too slllull for stothticul unulysis Among tCtlUshyploids at ullupbusC I th(tC was au obvious tllldl[wY toward equnl distllbu tion Among p1tlnts with morl thun 81 (h1omosoll1(s there was n prolloune(d t(n([(ncY towurd (quill distribution Tltis t(ndshyency was Cvltknt ulso in thosC (ClIs from ab(llants whieh Contnin((l few(1 chromosomCs than tllC lUuub(I L)1)icnl of tIl( nhCITtlnts ill which they were produced us well us in tlw umphimicti(ully produc(( offshyspring of abCtTunts ThC inequnlity of distribution in the highet polyploids wns no grCnt(1 than thnt in triploids und in proportion to the total numbC[ of chromosolllCs inyolyCd wus much Itss This tendC[]cy townrd Cqunl distribution of ehromosonHs at anuphusC I wus illterpretCd as bCing thC 1Csult of tIl( tClldency towunl biyulCnt formation at metnphas( LCsuully there wCre 18 bivnlcuts in POI1Ctl mothC[ cells of 36+-chlOlllOSOllW diploids Among 3(j- 1111(137-chlomoshysomC haploids howevCr thC numhc[ of biYalCnts was vnlinbk rrhe numbcr of LiyltlCnts shown by lCtmpioids YiLliCcl mnong thC cQlIections the lltnubC[ bCing gr(tLtC[ for tllt good gUIIyuk type Among plants with higbC[ chromosomc llumbCls WtClC polyvalents were expCct((I ~iyalCl1t fOlmation was pronounccd

POLLEN EXfLminntion of shed pollCn in mUI1Y instances p[CcCded u study of

llwiosis in pollctt 1ll0thCl cclls bCcuuse of thC ense and speNt with which pollcll call be obsclYCC1 For instanc( mictOcytcs W(IC obshysCrwlti in thC polkn of intctl11(dil1tp plfLtllS bdoll sLudies of nwiosis III POIlC1l mothcr cClls of tll(sC punts wCre begun I1nd a callsn1lClntion bctwe(1l lilgging clt[Omosolll(s nt tIl( Iwaphnscs and rniCloCylcs in pollln wus sugg(stcd The ronel lIsion thnt lho diamCt(L of policu grnins is propOltiollil1 to thl munb(L of ChIOlllOSOJ1lPS within the grnin WitS l(n(hed n[le[ tl1( palnlld detCrminaLion of chromosome l111mlwl nnel annplHtsC disUiigtulions in pollCn mothCl cdIs and size of POll(1l gmins of many indiyidunl plants throughout the polyploid series (8 tablel

The fiyc diploid coUpctions diITClCd in a(Luge ilmOlLl1 t of abortCd pollen un To whnt pxtPl1t nllCuploidr iIlY(Jsions nnd nondisshyjunction W(IC 1Cf1PCilt(l in th( nnlounJ of nboJtNl POllPll tnnnot be stated As not(11 on png( S lhlPC plll ts tiClt ])[oduC(( mostly 1lgtort((1 ~rnim sliowpd asynapsis at IlwtnphnsC 1 Certnilll tIllJC Wus lIO correlation bCl((11 mnount of abortion nnd lllmd)(L of mininlmC ehlol1)osoJ11PS (tiLgtl ~n ~1iCIO(TtCS (IC Cntillly nbs(ut or mlt in all d iploit pIn nls cxnmil1Nl

POlll1l of lt plantgt may 1)( dislinguisllld [OllI thnt of ~r pinnts by (tp pIPS([)(P ofm i(Io(ytlts II llltl by tillt Ylllin hl( siz( of lllp good glnins nttJwl Ihnn by thp P([(lllllgP of poll(n gluins abo[I11 Ill IW(lnge PPI((ntltlW of Illl InttpI is sOllHwhal highlr IlInl in diploids llJ tubln 11 In ]gtoU(1l f[olll [llnnls 1Hlollging to til( liy tollp(tion frOlll ~I(xi(o nlld foul (onlllwl(inl (mills th WlitI found YIlIing 1)(1shy

bull

bull

bull ccnlngs of g[nins nilol((1 rllngillg fOm [i to (iii (H(((nt nltllploid 31 plants nvPJngrd higlHl lt( rung PXlcIHkd front [i) to OR ]gtll(tut

bull

bull

bull


the latter being a eonsequcnce of partial asynapsis at metaphase I The variable size of good gmins is shown statistically by Powers (8 table 3) Some gmins i1le small like those prociuc(d by 2x plants others are larger lik~ thosc produced by 4x plants and sttll others are intermediate in SIze TABLE 9-0cctlrrciumiddotc of aborted POllCl1 grains 1I11d miniature chromosomes in 88

1Jlantl of five 86+-chrolllosome (diploicl) collcclio1~

I I

Ii

poli~n mins liniuturll i Colieclon No IPolien gmills I [jolnturoCollection No abort cd cbromQsomc~ I nbortcd chromosomes---------- _----- ----

I Percent NU1IIber Perctnl Zumbcr 53 3 ( 3

4UObullbull bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull j 50 2 lgt5 I gj 8

4 S 11 2 10112

0 IH 6i~ 0 15 I middot-middotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddotmiddot11 2) i 2 lIS I 4 15 I 84255bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull I

1 I101 1 Ui i 10 31 i 1 116 lrobullbull 11 43 l 1 f 21 6 4() bull 1 J 24 2 iO I 311 Z

I 54 I3) 0 1gt1 4 Il 2 15S 6

]0 t 2 95 i 12 I 0~1 314 30 () 42 5J 2

1 Based on only 100 groins ellch determinations by E J Gardncr

Pollen of 4x plants may be distinguisluci from that of 2x plants by the Inrger size of the good gruills (8) find from that of 31 plants by tbeir morculliform sizC The intermediate type shows more microcytes tlHUl the 3x but the good gUfiyule typ( shows fewel For the collections included in the study made by Powers and Gardner (0) tlw range of p(lc(ntage of grains aborted by the good guayu]e type coincided with that of tIw 3G+-chrolllosome collections The raIOC for the Texas Hintermediate type WI15 somewhat in ex((ss of thut of the 54+-chromosome collections The tt in termeciiutes averaged 2 to 5 tim(s tbe number of microcytes in the good guayule type The l1ug( of the 4 most commonly grown commercial strains was betw(ell tbnt of til( good gunyul( type aad the int(rmediatell

Texas type Only 1 ~r(lxjcull coll((tion was included in Powers and Gardners stuchmiddot

Pollen sampf~s of all nine of the ~Iexi(an collections wcro examshyin(d by the writer Only one of the good guayulc type Ilveraged highCL ill pelc(ntagc of pollen nbOlmiddotted thlln the collection included in Powers and Gardners studY (01 but foul IlYe1llged 10WN The amount of aborted POUtll of H ll1 ternlCdiates wus consistently higher than that of the good guayule type in those coll(cOo11s where both types occurred The lHcrngrs of llumber of microcytes for the ~Icxicnll colle(tions ellUl( within the lilllge of the fiyerages for the Texas collections In ey(ry instnnce the II interm((liatlslJ could bc distinguished by tlI(ir mfiny miclo(Yles

Sonlc of the -less known conunel(ial strains w(1e sfimpcd by the wTiter FiYe strnins of the good guayue type produced wry little

22 lJJ(HXICAL BFLLETIX llSJ -C ~ DEPT OF GU(TInllJo

UbOll(d poll l 11 whlIpns fiyp ntlw pIOdU((1 pl(llty of pollln plnelishyttlly all ()f which wu nbortl(1 Thi- -Olt oj mnl( 1liliIY i diJlIllll flOiil tH Jnek or sporo~l1l011~ ti~Sll( showll by s(lI~d nlHuploid ollty]ll J)ojlllnl from h(l TlXlt- (ollld iOllsFll rom of tho inlllllldin(p ontllwliul stluilh about Lj PI(Plll or (Ill gluins us llhoJl(L g-nill IIH IlYPIUgP IIIIII I )(1 of miCIo(yt(s Ijllgpd frolll two (0 fln limt IluLl 101 tlll gone glln~1I11 I)P(

TIH 1I1l10WI of n]olld llolllll Plotillcld h 1111111(- (lassiliP(l ns olfshytypt nOllllnls (jl 10 j ChlOIllOOJl)l 1 llIllgld froll lj tu lIl I)(l(lnt Tlt(l polll1l plodll((d II -)j ]gt11((111 of thls plllllh (ollsjt(ld or IIS tlmlt ~j 1HIIIIl of nhollll gluills hllpn-l Ihl ]lollpll [lIOdll(ld by ill P(J(nl of I hp1 phillis ol-lild Ill iO lllIClIt (II OWII of n)oJlaquo(] gmill-l III ~lllI-Illl tllO( pllI1l-l 1111 pollll of whi(11 ((lllsi-ltp(l of 75 1)(J((1I1 (II JllOlP nho)I(tl glnin hud TO 01 1( (h lOIllO() 1111 Till most trile I1g l((Optioll wa-l n ~-(hl()nHhOJnl pluntllt( ]lollllI of hiIll ((jbil or Ilh(ll 1-- 1)(I(II1( of nllllllld glHill

P01l11 of pl1Il1S ih ltig-II (hlOIllOSOJ1H 1111I1lh() lllny )( J(middot(middotOl-rlliz(middotd

1gt t Iw Inl1Imiddot i 1[ lIll good gltill -111H gldlJ 1J1 [(lulJd hUI 110-t of ttlt11l Ill IlIl lli HI ill shllpl IHlwlJ hy POPI-l IS table 3)

1lllll I n gllnlli I i ill thnn rOl plnnls ith T~ 0 I(-ls (IJ()-

1llJUII Thi j 1)11 1I to llll llring lJllJllIHI (If (hlOIllOSIlIll(S

within lit gllil1 1I11(IIH11II to III ful tlnl tIl diall1lIl W1 1lllIlSlIlP( ill Ihl liJlI of tlw ()(llllll llliIJlll1wtl Jlgntd(s of Ill orill1tnlioll or the llIipli111 glain wilh lf(IIIl(l to IIIl lIli(roIllPll Ill- PlrtlIl[ngls of glllill l h(llld lli(1I with IhI (oll( ion ill g1l1lltd tllt f1l illdn tIl 11lI1g ur tIl lloJmuls of til Stlll( pO]ll1liliol1s This IlS jlP(inlly IIII [01 Ill UhlIJllJ1Q from lJ plnnts VOl t]( alHlmllts flOlll middotLr plnnt tllPl( 1 H tlntilllY tolIl iJltIPll-lP(l p(J((middotlltn~( of nhor ion ns (otnptU((1 wi I 11 t lIP nomll k Tit nllllllwl of ll1iclo(Vl(s ])]0(11((lt1 bv aHIIllnts Ilolll t planls 111 within (1)( JHllg chnJ(llplshyjl if of IhI 1Illlmal of till nllH (oUl inns AmOllg n IWIJtml-l from bullIJ~ pIn Ills t hos h(loll~il1g I () (I( good lIlyulp [YPl j(gtlldl(s-l of -(1111(( ltnw(d 11 slight iJIlpaSI ill llllllllWl or mifJ()(Vlps 1I (ompllJld with til( Iltlllllll1 IlJodlllpd hy (ol11palnhh- llOllllllk In gtltHrnl the sD-ndlt-d rough nllIIllllts tlvlltllld hid ill l1uml) of Ini(Io(tPfl In (lilt of till ~l(xi(llu inlllll((fatl [(Ikltiolls I~-l~J h Ublllllnts t(lng ltI 1(-l [hun hnlf ns 1llI1llY mtero[-l as llt( nOll11lt1-l uf lbnt (ollp(tion ill nIHlS tlllY nv(rn~(l abouttll( snnw

LOSS OF ClfHO10S0lES BY BERHTS

plhllol1 ll1101l wllillI (llllS to )p IH I hpJ witl(slllpnd n mong iI)(IIIH1t plnnt is lhl IOis of (hIOIllOSnJlll-l hoth TllPjO iHll) lind mitotic-nlly In oddil il))) to tIl In of a fp ChI0I11O-l0HlPH in pollett tnOtlHl ((lls through lagging It Hlll1pllllSP I 11lHI II nh HS wus ((11 ill [jJ plusmn-und 72plusmn-(hl0l1lOHOIlW plunl- pSPI(illll) (host of tIl( inllIIlldint( t)I)( 10s1 of IIlJg gtoup- of (hroIlHlSOJH(H wn OJSlIPc o((sjollnlly dUling Ilwiosisin ]lollll1 l1wtltlr (llIs of llhllItlnt plnll(S This tlnltilJHY hn-l 1)((1l 01ls(1(11 ill lol((tioll f1om Tpxns Hfi w(it as ~Ilxh() nut ill (ommlllinl stlnins nIthough it (1(111 to Ill 1ll()J( common in -l01l10 plnns (hun ill otlwllt Thnt tll spimllt IIlPlllHlliMIl mn) hI il1l1((shy(lUlltt foJ so InUI1) Chl()J1lOS01ll1S is slIgl-rlstld h) thl formntion of n mullipoln l -lpindh- (ig I () I(1( t ht HHi (hromosol1Us nre in 4

bull

bull

bull

POLYPLOIDY AND ANEUPLOIDY IN G1AYVLE 23

bull insllod of 2 gloups nt IUlnphnsl 1 FOUl gloups illsUnd of th( llslInl 2 nt sNond plopitnsc (fig 1 If) 11IP 1ll(1 j(sldt plohllbly of n Illultipolnr spindle lIlIting lhl first division For (ompIlIison till usunl 2 g-IOUpS nt this stng-l iLlt 8ho11 ill Hnothpl poJJpn lllotlwl (dl from Lhp snll1l plnnt (fig- 11) AllollH1 WIly hy whi(h til( IItUlliJtI of ChIOIllOSOlllPS enn bl d(l(Icts(l(1 is hy Ihp rOlnmlioll of Lwill polll1l Illollwl ((lls For Cxnl11pll (fig- ] J) twillpolkll lllothlt ((l1s in(oJ11plptply stpnshylnLtd (ol1tninld 107 ChlomOSOIlHs Jwt W(111l t1HIt1 10W(YPI I h(se ehlol11osomls (IP ulkluHII disllihu[pt 011( ]1lI11 hud 40 (hlollloshySOI1(s in 2 g-IOUpS of 22 nne 24 1(lsp((1 inly IHIPHS lht olh(I half had 7 ChIOJ1l0S0IHlS in 1 IOllP nnd fi4 ill th( otlWl

bull

TIl( loss or e1l1OI1l0S01lHS 110111 sonJal ic ((lIs hilS not hpl1l S((11 htlt mlly 1)(1 inllIr(d BC(fIllS( (((I(S tltlhni((llt (7) WfiS d(y(lojJ(d onl~~ l((ltllllV obspLTnliolls 011 sOIllHtiC dhisiolls ill root-lip ((lIs 111( still 111 (II( I I 10w((I it JIllS b((n ObiH(pd ill root-I ip ((lis flolll n1HIltlllt pltwlt- lhnt illln(t ((lis fro III [h(l samp plunt mny Ill in ehlomosoll1t II urn 1)(1 Tho l blHh rosses mil ()((UI SOIll( l il11l priO(

to 1IH fOlmation of poll(11 lllolh(L (Jls is sil(mn by til( fael lhllt intn(( poll(111 motllPl (plls whi(ll (onlnil1pd f(w(1 thull (11(1 tlsufLl nun11)(I of (hlOnU)SoI1lPS ha( 1gt((111 S(PI1 DilJPJ(Il((s ill til si( of [hl po)l11 mol h(L ((llis W(I( disC(IllibIP sil1(( Ihll( is tl (OIIplnUoll hp[HPIl (h(lil sit alld Iht 1l1l1l11HI of (olllnillPd ehJOIllOSOIlHS ]n tlH snnl( plnnL whieh plOdu((d (h( j10llpll moth( ((lIs whos (illoll1oshy80n1( (OlllplltS Ill( sholt11 in figtlJ( I [f and 1 olhI poIl(11 l1olh(l ((llIs whi(1t (olltnil1(d 1(IlI (hI0Il10S0111(S W(1( fOllnd On had 7S (fig 1 K) whill 11110Ih(1 had )(J (Ii 1 L) AIIOUlll poll(ll1 moth(I ((JI hnd ~ii (hlol1losolls AIlOlh(I (lXlll11pltgt is g-inll ill liguJ(1 I Jr 1](1( 11 ((Ill nL tllophns( II (ol1lnillS 0111 120 (hIOIllOAOIlltS in lILt 4 nlHICi This (pll must Iin (ontuilwd olllv liO (luoll1oS01l1(S Illtfolc 11l(liosis bpgnll AnothlI ((Ill frolll this slInip plnll[ hnd Ii (hlomoshysonws 0111(1 (h [0Il10S0111( Illl ml)(IS such flS 40 45 40 5~ 5) fiG 57 )S (iii Ii UR 0 77 RO R1 ~4 Hlld 03 W(I (oul1[pd in (XCPPshytiol1i11 poll (II mol h([ ((lIs from 14 o~h([ Itt)(ITllIlt plnn [s host (hI01l10-SOI11( llllmlWls Jllng((l flol1l 102 to 120 Ilnd whi(h (HIll( flOIll 10 difshyf(middot1e 11 t (oIke ions

Thnt slIth POIllIl1110thlJ ((lis 11111) goo through both meiolic divisions is known IwcfLllse eluomosollHS hl1n hp(Il (olllltp(l al 1(lopJHls( n One GO-chromosome (pll shollt1 only I hindpnt nl ll1elnphnp I the 1ll11n1n1ng c]uOJllOSOI1WS lwing ullivfllcmiddotnt All RO-duomosol11e e(ll flom nnoth(t plnnt showld both hhril(llls nnd 1Illhnlpnts nt I1wlllshyphusp 1 lind It 7G-ehloJ1l0AOIlW (Ill sho(d only ulliul(nts DiMlibushytion of ehlOmOSOIllPs Itt Illlllpltnsc 1 111 OtlHJ cdIs whos ehlOmosomc llumh(Js UJP lis((d in Lll( pJ(cCClillg pltIugnlph follows Chroulosomcii in pollen ChrOJllOROJllCll in POllC1lI

1110ther cell J)jlribuiQn I moth(lr celI l)iri1miOll 1

-10 IH22 MI 27230Iii 21~~211 Oii ao I 31middotHL 21215 (is a236 52_ 221 20 77 aa737

bull oiL 27 28 78 aa2middot13 50 251 30 7U 35 1 middotIa 37 213) 8 80middot12 58 27 1 30 ~H 31l 144 58 2782S l3 middot1846

l lng~lmiddotrslmlicfltNI lIy t Itl 11111011 Jlgurl in tltu srr(co of a

24 TECHNICAL BULLETIN 018 U S DEPT OF AGRICUUlUHE

Although these arc ex(cpLiolll11 cells from abelTant plants the disshytribution is not essentiully different from thnt observed in those plants with comparnble chromosome number (tablcs 1 34 and 5) bull

Reduction in (hromosollw lltunl)(t (an )re((de the dov(Iopm(nt of a tapltal c(11 For (xnmplp It tnpllnl c(11 (ontnining only 40 (hromoshysomlS wus found (fig 1 Y) on th(l snml slid( -jtll pollcmiddotn moth(ll (clls contniuing about lOR

Among nb(1Inl1ts in 54plusmn-(hromosoJ1w populntions VCty f(lw poIltn moth(tmiddot (clIs whieh ((J)ltnilwd l(ss thun the usunl 111m1gt(l1 w(rp found nlthough th(l rntio of 81 to 54is the sumo ns thnt of 108 to 72 Two cdls show(d 54 nncl 57 ehromosol11(s l(speetiYtly in un Sfi-ehromoshysoml pltmt nnd 1 show(d n (hromosollw in nIl SS-lttromosoJl1( plnllt Appnrpnn~ it is the high chlOmOSOllw lltlll1b(l (l00 Ot mol() (Il11ong al)(1nmls Lll 72plusmn -eltromosom( )opulntiolls whi(h 1(nltls to loss of chromosom(s IJw loss of Inrg( groups of C11t0I11050111(S by nbcmmt pltluts is of inl(rpst to tho plnl1t bn(iclif embryo sn(s tlr(l pIoclu(pd by chimcrns whic1t huve [CWN cl nomoHon HS Apomictic ploduclion of S((([s by such himerns (0111d knd to the cslnblishmpnt of donps hnying

-cliromosome numbcrs difllcnt from ihost 01 pilhcl the nbNTunt or tho usunl types in the (ol(gtelions to whieh tlpy bcmiddotlong

CIlHOllOSOMAL VAIUATS WITII1 THE POLYPLOID SEHIES

Onw1

One of the ehnrncl(listies of 86 +-chromosome populttt ions of gunyulp is lhltgt tyidpnt morphological diY(lsily withill the diflclent bull collcctions This is tht consequence of nmphimixis nnd cIoss-pollinushytion sinCe they n]( InlgCly Ewlf-incompntiblc (1V) In contrast to this diwlsiLy among diploi([s til( ynrious triploid and lCtrt1ploicl stnnds nre f[tidy unifolln nmong tlHInsph-ps nllltough difi(gtring from one nuother in morphologicnl ehultlcltrislics This uniformity is fl consequence of Ilpomictie reproduction Howevct Ul(l(~ ar( individun1 plnnts which dcvintc from lh( usullily)e chumcl(tislie of the 8) lind 4) stnnd5 and commpr(inl strnins ill which they OeClll rhtco typ(gts of morpho]ogicnl YHriants huYlt he(1l (listinglljshed IInploids tho so-cnlled oJftypo normnls UIlt abcrJlults iJost of th(sc plnIlts npc tho l(gtsult of umphimixis sinc( npomixis is only fncuUlltivc (10) MOlCOYOr ESllu (4) hus shown thnt 110rnml meiosis lUngt occur in plnnts which nTe ptCshydominunlly npospolOus lh(refole some eggs with the teduc(d lUllll shybct of chJo)1loson1(s fUC produc(d If tlwy d(vplop ithout f(llilizashytion (plllUUllog(lHsi) tlllt gin~ rise to hnploids It numb(l of which htwe Igt((n found in 72plusmn-chlomosome populnliolls If (ggs which huy( tht lCduce(l nUlnbcl of duomOSOllllS n1C ftrtilizN (lLluphimh-is) they giv( rig to pluuts which Jllny 01 nmy not Imyc the StlJlll number of chrOlllOSOIlllS a$ flllomicti(nlly produced plants in Ow SlLme cu1tm(gts the Yflrinhilit~~ in ~lom()s)m( ltllnbll hlting tlw ((suIt of Ow nrying numh( (oJltnllwd tIl tl1( (gg lluclltIIS ]llus t)( vtlJying l1umb(gtl brought in h)1 tIl polltll gtnin 1gtrobnbly most of th( ofrtYll( nOllnnls 1t(

mnphimi(l In IHldition ltggs hidl hiLYt the uIlI(duencl lmnbcl or chrOIllOSOIlW 1ll1lY 1)( f(Jtiliz(d lIso JPmiddotjll11tS nmiddotl( plo(lu((d in this WHY COmHCiIlPlitIy lllorphologitai YllJinnts mnv hl( till $lll10 mnn- bull btl of du()m()som(~ IlS the ul)(mi(h of the POpilllltlOns in whielt tllty OCCllI or they IUi1Y hnsc mon ()J llWeL euo111080n1lt

bull

bull

bull

lOLYPLOIDY AND ANEUPLOIDY IN GUAYULE 25

HAPLOIDS FnOll TETUAPLOIDS

Plants which result frolll the devclopnwnt of nil (lgg which hns the reduced number of chromosomes without f(lrlilizntioll hnYe been ltalled bnploids UnforLunately the word II haploi(P 111ts betll used in 2 sonses 1l1llIlely as being equaL to one-hnlf of tho diploid l1mnblr of (Ju0l110S0111eS (monoploid) nnd also ns being eq unl to onc-hnlf of the tlolllntic ntullbCl According to the first deiini Lion a haploid gUiLyule should hnvo 18 (h101110S0111(S No such plnnt hIlS been found ItS yet Accolding to th s(I(ond d(finition a series of hnploids is thcolcti(ll11y possible since thClo is n polyploid s(ri(f in gllnyulegt Regltrdless of tho sense in which the word is used howcvel a hltl)loid owes its origin to the development of nn unfo1tiliz(d egg nucleus wInch has tho reduced JHlInbl1 of e1uoU1oI)JlleS As pleviou~Jy J(potLpd (1) 11 few haploids wel(~ found III POPUltltiOIlS hieh Itayp 72 plusmn duomosoJl1(s Since then more haye been obtuilwd from seeds eoll((t(d in r(XllS nnd lllso from ltomlllc1(inl stJnins by POWOIS and Hollins (10) by RoJlins9 by Stebbins nnd Kodnni (1S) und by the writ( In genelni the plnnts w(re slow glOWLs I1J(Y floweied jlofusdy but the 110wer heads W(I( filllllll

How l1lueh lltl[upioid (xist n11l0ng lll( dipJoid collc(tions cannot bl tnt(d cxndly 1)((lw( 01 (hI ndditiollnl millinllllC duOtnOSOIllS Silw(l tlHlC is l)ut slighl ltlitr(j(lIep in Ipngth h(tween the smnll(st binlllwd ehlOmOSOI1H llnd minilltllrcs il llisolllic lIniYl1(Il might be eOllllled as a miniMUIc FortulllltIJy ill ollly 1 or lIlP (ole(tions arc mininLul(s mOlp llumerOliS tlHlIl 3 Jf the poslulation that minintulcs owe tlt(il origin to tlw brlnkngp or biurllwd chromosomes is corlpct t1WY IUt d upliCIIM portions of (tlol11osonws nlHl the plants in which tlll) O((lIl In 11IHuploid to thn (XlPIIt small ofltypeshynormal pInn L with 11IdiuWIl lnl) disk HOHt blllels WitS found to have 37 ehrolllosol11CS

Since diploid 36+-ehI01llOSOl1lC plnnts lcprod U(l IlmphimicLically thcre is It profusion of nnlllploid lliploids nmOllg their plogeni(s when tlley fl]( t1spd us lWlCi ]llllents ill e1oss(s with 54plusmn- nnd 72plusmnshychromosome plant The 111Ilgl of til( snmpk studied wns 43 Lo 56 chromosomes inclusin The plants chos(11 for cylologirnl sLudy from non(onlrollNI poUinnlioJls showNI Il INlst 50 lJllcent of the polkn nhollld nnd mostly moro (hnn 7) PPI(pnt Their chromoshysome ntlln 1)(ls lnngccl from 49 to jG [olphoIogi(~ltlly thlSC plnnts diflPled from onc nnoUHI hilL lhtil diversity did Bot pxc(ed the mOLphologicnl diYIsity ehltllletcrlstic of 86+-ehromosome popushy1I1ti(l~s Illi only small pltl1t sLudied wns II 53-chromosome plant which hnd LItick INLV(S All thQ otl1(Is WP1( vigoloUS lnd mediumshysized Ot largo in(sj)(cl i ( of (il1ol11osom( llumlwl llnd could not be distinguished from ot hll plan ts in the StUllI (uIlmes lirolll onshytrollld pollination with 54plusmn-(hromosomo plllill-S the plunts sudied ranged in ehl01110S0J1le lHllnhpr from 43 10 40 rho 48- and 49shychromosome plutlls welo of uvernglt vigor but the 43- and 45-ehromoshysome onlS grow slowly nnd were Itss vigorous

gRollins R C UnpUblished data

26 TECHNICAL BULLElIN )lS U S DEPr OF AGRICULlURE

OJIFTYPB NOn~lALS FROM TmlLOIDS AND TETHAPJOIDS

A few of tho variants classified us offtypc 1101111111s in apomictic 3x and 4x poplllntions hud (xuctiy 54 uud 2 chromosomes respectively most of th(m were nnollploids For the 54plusmn -chlomosome popuhLtions an cxample of un(uploids muy be cited As n clHck in all cxpelimcnt s(eltls obtuincd from noncontrolled pollinution of plants in colltction 4263 wme sown Among the progeny thCl( WllC 4 oltype normals which had 54 56 64 and about 66 C1l101110S0111(S llsp(etiY(~ly The 5(l-chlOmOSOl1l( plimt was lUlgN than thc otitOl 3 but nil WPI( smaller than tlH npomiets in the saull cuItme lhe plant with lIbout 66 cluomosoJ1lCs had n loose growth habi~ The 64-CI1I0111080111( plnll~ had lUlgc gLay thiekellcd leaves and ltl1ge 110w(1 ]l(uds A~ least 75 PClclnt of the pollen produced by nU 4 Wi1S abolLl(

Most of thc olltype uOlmn1s whosc elHomos0111e nUmhll wns deshyterminod OCCUlTed in culLul(s of 72plusmn-chtomosome Jexns populations and commelcinl stlnills They do not constitute a mndolll slullple Among plnnts (hnta(tptilled by slowll growth less vigol 01 utlllsual mOtphologienl chntuetelistics leples(ntativcs W01O s(lected by L POWPtS fol cytologicnl study in the hope that theil chtOl1losomo numshyber nnd 111Piotio blhfLyior would indicntc thlil modo or oligin In gelHltlI uft(11 fI yltUS glOth few of these seloct(d plnnts W(IO equal in sill( to thl apomiets in lIte sumo cultUl(s and not 111l1ny woro oyen mcdium-siz((l most of them WPlC small nlany tended towald comshypa(tlI(ss or low splonding hnbit indicativc of shottcncd intllnoti(s r1hCll w(1e vIltintions in lLnJ SiZ(I shape and thiclmoss At leust 2 of thp plants won S(Clcnial chitt1(lns 1he10 was no appalont cOlrdation b(ltween ehtomosoll1P numblt unl plant Stlll SevoraI72-chtomoso111c plants wpro litlHl smnU 01 V(ty smull whplcns one 67-(11101110S0111e plant US llwdiull1-sillNl Tho chtomosome lHtmbcts of tbo large plnnts JHnged II0m 70 lo 72 nbollt tlw snme ltlllgo as shown by thc oer)T Hmall OlHt4 Th( lnv plnnts wlli(11 hn(l 73 to 75 chromosomes W(tP smnll oj llltl1illll1-sizldj 2 had Lhick01l( leilvos

TIll IHek of ((ItJlIntion bltwlllI r1u01l10S0J1W llumlwl nnd plnnt size is no[ sUlptising Si1(l it is not chtOl1losom( llt1mber bUl itlt(1UCtions rnlOng thlt g(IllS thnt d(llltlmit1l Lhe yigol of n plant A study limited to loud (htomosonw HlIll1hlr fails Lo show how mUIlY of (neh ki11(1 of ehromosotlll IUP Pt(spnt EYltt1 a 72-(Illomosoml plnn for instnn(c mily hc iHl fltlCllploid if ins(PHd of 11Ilving 4 lflell of 18 kinds 01 lhromoshySOI11(S (llsslll11ing thnt x lqunts Un it hns 5 or one 01 more kinds and (oJtlspondingl~ 3 oj OtliPIS Tiw sltnphsl o](ln1iol1 would 1H 4 (gtiwho Hi kinds plus ii of tl1( slY(nt((nCh pitts only 3 of OJ( ligh((wnth In pinnls hnying mot( Ot hss limn 72 (hI01110S0111(S tIll 1gtnlnt1(( is fUt(hN llPSlt by tit( nbSlIHP 01 nddilion of whoh (11101110S0111(S ~[h(1 SHUll sod of ehlOlllOSomnl ttnbnlam( mn (Ixisl ill 54-hlomosome pInnts and in thosp hHill~ mOt(1 OJ hss tluwij4

Of the olltypl 110lnll1]s whosl (ltomoso11ll llllmhCt wns dp([minpd to bl 4~ 01 ntHuploicl 41 aan l(ItcNt( hnd Oil to 70 ehtOlllOS0ll1PS 3R0 )(t((]1( 70 01 71 IXH IgtPI(lI( 72 and Sii PlltlOnt73 to 7i) In nddilion (hll (IP 2 whkh hnd 133 nnd 5li to 5H lhtomosonlls 1(SI)((shytiyPly rhus il is (vid(lnt lhaL lllost of UWSl oIrlygte norl11u]s WlO

ntHllpIoich flld lhltl lhtcp-outths of (hC111 hndkss dum 72 chromoshysonHS 01(0 Ylt t tIll Chlot11osot11f11 dtfkill1(Y WHS limited to only 1

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01 2 chromosomes in the mnjoliLy of th(ls( p1nnts with l(ss lImn 72 chromosomes It is possible thnt plnnLs with mOIe thun 72 ehromoshysomes were sufficiently vigorous nnd nOlmnl iu nppcnllUJeC Lo escnpc being clnssi6ed ns ofJtype 1101mn1s However tho pltspn(e of addishytional ehlomosol1lts mny be deleterious sillec none of the hUge oIrshytype 110rmals lind mort thnll 72 clnolllosomts Analysis of luwphnso I distribuLioll of ehlomosomes of tctlnplokls (p 15) showell thnt pollen grllins produetd by 72-ehromosomt plants (onlflintd a vnlying number of ehiomosomes mostly 33 Lo 30 11 nn (quivnkllL valinshybility in the ](thwed (hlOmOSOlllt 11ll1nhN ill (gg Iluelcj is nsslllllecl i then a 41 poplllntion should PlOdll(C nmphimiels lllnging in ehlomoshyS0111e numbCL from GO to 72 This lflng( in(l1l(1(s n dtvinLioll of nt hnst 6 from 72 sin(( (Hi willjw L1H toLal procimpd by tl1( lillian 01 2 gnm(Les (neh of whi(l1 (ontnins 33 (11l01110S0l11(S TIl( lInioll of gnmctts whieit (onlnin more Lhn1l 36 ebl011l0S011ltS lach ]JlOdllt(S p]nnis wiih 1l101( ihnn 72 cillomOSOlll(S The ObS(IY(d lI1Jlgl or li5 to 75 ehlomosou1(Is nmong olltypt normnls in 4r populnliohs is in (lose nglccmtnt middotwith thl (nleulntltllflngl bused on n study of Iltlnphnse 1 diskibution of elitomosol1llS lIowPYlJ sin(l the majoJity of the ofltype normnls WPJ( ddieilllt in only 1 01 2 (illomOSOI1WR most or the (ombinntiolls which itwolYld tIl( lIllton of gnll1eLs whieh (Ollshytrdn(d less thnn 34 CillOlllOSOmls must hnyp bl(llitlyiu ble blt~n liSt of too gt(I1tly disLll1h(cl gllli( bnlnncl

Those plnnts whidl hnd th( middotIo(sl (hlomosom( ll1llnbl1 mny he mentioned s(plHn((jy Two of tlt( T(xfls 0(11)( normnls had (i5 chromosom(s This nll(uploidy mlly Ill llPllSltlt(d as 4x-7 Both plunls WlJe small hUl llnlik( nllltl iJoth WPIl pnltinlly aRynnpli( nnd showed l(sGituGion llll(hi nt llwtnpltflR( n Sixl~r-Jiy( lHtC(nt of 1Jw pol10n of one WflS nholUd nnd til( diIUllll(l of (Il( good glni1ls inshydieni((l thnt tlwy eonlnitlPd tit( unreel II ((d llum1)(1 of irlomOSOll1es Tht otlwl PJOdtl(~lld t1 scant amount 01 11Olmfll-sized glnillfgt 501wI((tlt of which were ttboltcd

Tlllle ([(12 Pxc(ptionnl plnnls llom TCSlltIH A gt() smnll plnnt hich hud 50 to 59 chromosOll1S tUllC from It llollstl(ci (I COllltioll of sc(d ~ [ciosis wns ehulne((Itiz((l by pnllinl nsynnpsis tlnd lugging chromosomes AbouL 08 pll((nt of tIl( pollt-n wus ttiJoltpd nlld the good grains probably hu(l Lhe ulll((hlc(() Ilumb(l of ChlOIllOSOIlltS rho plant flOWlIld proJus(ly TIl( otllet platlt which ns smnll nnd compnct hn(l only 53 (hlomosol1ws (BOlh it I)nltl f171-chromosome oflLypc nOlnuJ] (nnw from s(Nls (olJcoltl(d JJ()1ll fill j]](livid 1I1l1 plaut of the good gnaYlllp tyP() At l1wlnpllns( I 80nw uniYl11(tlls and many biYL]Cnts WPll obs(l((1 TIHIP W(IC Ingging (hlOmOSOllws at nnttshyphase 1 About 85 JWICpnt of thl poll(tl was aboltldilinnd thlle w(te manymi(rocyl(Is TIll (xisl(lH( of plants with suell low ehlomosome numbers nmong 72plusmn-elilOUlOSOIlll populntiol1s llClllills nn pxplnnnshytion It wns (on(1Ilt1llt1 thnt sin(c haploids bnYl hlen found muong T(sl1us (loll((LiollS by both )oliins lO and lilt wrill tlwse 2 plants which hud 53 nnd 5G to 59 ehlOlllOSOlll(S llspectiyely musL hfve come from Lite flltilizntion of tggs which had thl lPtilHNI Illlmbllshyof chromosomes by pollen from hnploids

10 Rollins R C UJPlhlishcd data

28 TECHNreAL BULLErIN 918 J U S DEPl OF AGRICULTURE

Some ofltype n01mn1s frOUl 72 plusmn -chromosome populntions showed normul meiosis nnd the pe1Celltnge of pollen aborted was not grenter thnnthnt produced by npomiets in the snme culturcs f~lthough no plnnts with less thull 68 chtomosomes were free from meiotic irregulnrishytics With a few exceptions plants showing these ilregularitirs were of small size

Most of these offtype normnls cume from seeds from non controlled pollinations However 11 came from con trolled se1pound- and crossshypollinntions mnde by Powers nncl I-tollins (10) 1he mnge of their chromosome numbers was 66 to 74 und is pmcticltlly ic1enticnl with thut of the whole glOUp of ofltypc normuls Of these plunts 5 cume from cross-pollinntions since tlley show(d both paternal nnd mnternal characteristics ther must have come from the fertilization of eggs which had the reduced number of chromosomes

It has been nssumecl that all of these offtype normals were the result of amphimixis However from a controlled cross-pollination there was 1 exceptional sm(~ll plunt which although it showed no chul1ctelistics of the pollen pnlCllt wus distinctly different from the apomicts in the same culture (10) It hud 72 cluomosomes nndlnckecl the udditional miuiuture chromosome which the seed parent had It hnd many lagging chromosomes nt nllnphuse I nllcl II and nbout 98 percent of its pollen wus nbolted Powels nud Rollins (10) sugshygested cliplospory to uccount for the origin 01 this plant Powers has furnished evidence by ll1enns of breeding tosts that diplospory is occurring in 1 of the Texus collections (8) und it can be presumed to be occurring in other collections nlso How many oiftype normals are the result of pseuclogumous cliplospory is not known It is possible thut some of the 54plusmn- and 72plusmn-chromosomc offtype normals arosc in this way It is motmiddotc probable that most of thcm resulted from normal mnphimLis ald that the morphological variances wcre the conscquence of new c)mbinations of genes brought about through amphimixis

ABEHHANTS Fnm[ THIPLOIDS AND TETHAPLOIDS

Thc morphological vflriflnt caned aberrant may bc described as follows Growth generally is slower than thut of the normals of the same cultures leftves arc thickened nnd have fin irregular pattern pollen grains (md flower hcnds are larger i und the peduncle is thicker Some plants show this syndrome of chnracteristics in morc pronounced fashion than others i some are fairly vigorous whereas others fail to livc through the fiJst yom Another characteristic is their high chromosome l11-mbcr which also is vnrinble the range extending from 81 to 122 1hc extent to which this syndrome is expressed is not necessulily cprrclated with chromosomc number For examplc of 11 aberrants obtained fronl the open pollinution of 54plusmn~ch]omosome plants belonging to collection 4263 8 had 85 to 90 chromosomes The 2 most extreme plrmts however hncl only 81 and 83 chromosomes respectively Also among offspring from u 72-chromosome plant (42163) a 99-chromosomB plflnt wuq more cxtremc in type thun 108 plusmn -chromosomc plnn ts in the same culture

As Esuu (4- 1JP 353-354-) has shown where apoIllixis is the predomishynant form of reproduction tllllmegnspole mother cell docs not form

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tetrads but directly develops into a uninucleate embryo sac with a diploid number of chromosomes Powers and Rollins (10) have shown that pollination is necessary for the production of apomictic progeny from embryo sttcs which have the unreduced number of chromosomes (pselldogamous diplospory) Aberrants which occur in apomictic stands and have a higher chromosome number than the apomicts in the sume stands are the result of the fertilization of eggs which have the umeclucecllllUnblr of chromosomes That abcrrants occur after cross- as well as self-pollination has been shown by Powers and Rollins (10 table 3) The total chrOmOSOIlll number of an ablrshyrant plantmiddot then is the SUll1 of the tllulduceel Hum r in the rgg nucleus and the numblr of clDOmOSOIlles in the pollen grttin The yariability in total CIUoIllosome Jlumblr clepenclslalgely upon the varying number of chromosomes it1 thl pollen grain

Aberrants which wrle found among 54plusmn-ehlomosol11e populations had 81 to 89 cinmiddotomosomes From non controlled pollination of 54 plusmnshychro111osome field plants at SuJ~nas aberrants were obtained whi(h had 81 to 90 clDomosomls Among 54plusmn-(hlomosoml commelcinl strains plan ts were found which hud 81 to 93 chromosomls Siner the egg nuclei before fertilization contained 5middot1 plusmn clUoIllosomes chromosome numbers between 81 anel 84 suggest flltiliziLtion hY po11l11 from 54plusmnshychromosome plants l1umhClS aboye 90 sngget fertilization by pollen from 72plusmn-chromosome plants Plants whoSQ chromosome numbers ranged from 84 to 90 (ould have resulted from fertiliziLtiotl by pollen from eithlr 54plusmn- or 72plusmn-chromosome plants

Aberrant found amollg 72 -cblomool1le populations had mostly 103 to 112 chromosonlls although infnquCntly SUell rxtnmes as 99 and12) were founel Tlltv we1( ohtaillPtl from nIl of the 72 plusmn-(h1omoshysome eoUections from Ixieo Induded also in the snmple which was examined ntologieallv were nl)(rrants froIll 12 of the Texas collecshytions from 9 of the coIllIllelcial strnins iLncl from a cross hetwern 2 of the Inst-named Both the good gunYltle aud intell1lctliltte types Wlle included umbllS nroutllt[ ob sugg(st jprtilization by pollcn from 54plusmn-ChromosoIll( plants or from 72plusmn-Chlomosome plants whi(h contained l(ss than W cillOIllOSOl1l(S

-Ianv of the al)(rrants (te ulhcl rough nbernUl t In Rollins (11 12) llllYwIl foulld in stands of the good glltlyuh tjmiddotpp as w(ll as the intermedia te type Siurl Rollins Ins shown (xpprimell tnJly that Olle way to plodtul rough uhlrnnls is through the feltiJizLtion of guayule eggs 11ieh 11ftV( the nl1ndu(lcl numbpl of elUmiddotOIllosomtS by pollltl from muriola tht rough nJlPrrunts found in stunds of the good guayuh type prohahly tU(- fL typ( of splcips hybrid produ((d by the f(rtilization by marioh polllll of gnuyu]1 eggs whieh have the ullJ((l11(ed number of (hromosomls TJw 120-chIomosome plnnt referred to was a rough nl)(tlant and probuhly wus tlll lesult of poundrshytilization by POll(Il from a OO-(Ilromosonll mlllioln wlti(h tH(orc1ing to Stebbins and Kodnni (1J) O(CUJS in the same gelwrnl region withguayue ~

Stebbins and Kodani (13 plaSl IlftTtd to ab(rnttlts as ii itlltOshytriploid with lespltt to their imnwdiutp parents They had 110 proof whrther the (gg or t11( pollPH gmin con tdbu tNl the llllrCClucecl numhrr of chlomosom(s Tlmt ttl)(gtlIllnts might br the result of fertilizltion of flU egg which Juts the leclucpltl 11111u1gtel of dUomosomes

30 TECHXICAL BCLLETIX 918 L S DEPT OF AGRICtJLTLRE

by l1 P011(ll gLl1itt which has till 11l1l(lIl(((L numlwl wns (onfidllld as a possibility by ~tlhhills nllL l(odnni (131 who itld iItdirNt (vidpll(p hy showing middottlll plpnllpllp of tlilci inslptld of (Itl(l(S of miclospol(s within POlllll 11l0thll pll TIll Wlitll w not nhlp to confirlll sueh oiJsNTtltions on hll mnwrinl ~lono(I hnlding (vidplwP dots not support this jlossihilil gthOWll hy Hollins U 1) txptrinlPntally produ(ld nhllTllIll hyhlids IPspmblpd tIll mntllLll11 pl1ILnt molP cloly thnn thl PHI PlIllll parlltt it nd fI hiO Jl1011 (losdy thnn thl hybrid which htHl Clllll from 10S hnsill thl redutld (hromosoll1P llumblr ill tlw SHIll cult II IP - -

OFFTYPE XOlUrALS FH())[ UEllItJgtTS

Informn tion (OIUlrn illg till m(thods of IlPlo( uelioll by ahlllflll t plants is limitld at tlw PlPSlllt time to Powlrs sludy of a)(rJnnts in coll(ction 421113 lSi 111( chlomooJlle numb(rs of a flw WNP detllshymil1(cl The rnng( (xtlndld from 0n to H)(l Among til( ollspring of abNlflnts th(lP PIP a iil1Ht11 numb(l of ofitypl IlQlmllls whieh iihoecL only a J(w of th( eha1llct(listicgt of a)(IIt1nts Xo 2 phUlts Wll( alike The Chromosoll1l IlUmUPlS of 12 of tlllsP pinnts from DabllTUlts were dpt(llllin((l thlY ltlnged frol1l1-plusmn lO 03 )[eioi in pol1l1l mother clll witS flnulogous tegt thnt of nl)(tLHnt haying approximntely the salll( ntll111)(1 of ehromooll1ll Tlwll wa 110 (onpLn t ion bllWlell leaf charactcristic and dllomoollt number [tii town as follows Lraf rharacteJi-ti Ch~~~~~~lIt

)fllrh thirkpllld ~lightl irrrguiar partPfll ______________ bullbull _ ________ 03 Thickrl1ld

~lightl~ ilrCgnar puttPrll _________________ bull __ bull __ bull __________ 9192 Hlglliur pattpfll ____ ________________ bull __________________ - +82

Rlightl~middot thieklnld rpgular pammiddotIlL ___________________ _____ So RO 91 ) ot t lIlCkpll((I

TrrlglllnI patterll_ _______ ____ bull __ bull __ _ __ bull ________ bull --- - - - --- 85 ~lightly irrltulnr pIH(rU __ __ __ _ ______ bull ____ -- --- - -- - IH Rlgnlar ptltttfll ________ bull _____ bull ___ bull ___________ --- _--------- Sj R8

None of lh(se plants had thickpn(d p(dwwl(s Binc( aU of the chromosome lHnnb(rs W(l( l(ss than nfgt they probably Wlr( the result of frtilizlltion of eggs which hlld till l((luc(d 111l1n)e1 of chromosomes and r(pl(slnt((l thr bcst of the g(nie llCombinalions that resulted from amphimixis

SECTORIAL CllntEIUS

Among tIll oltypP normals wltieh (Il eXitmincd cytologically at l(ltst 2 w(r( s(etolinL chimeras 1 of thes( consisted of 2 morphologshyically (lisllnet parts lwcl th( oth(r of 3 Th(se plants had 12 ehromoshySOl11(S X 0 elMr diffclfnees in (hromosome 1l1uube1 among the se(lors could be disc(lI1ed

Among abeLTllllts sedolinl chim(Ifls IUC fllirly (ommon pnlli(ulnriy after the first yCitr of glOwth Arl((tNl portions of plants of Itcognizashybie size YlllY flom s(ctors of leans to who[l bmndl( 1-0 sectorinl chim(litS llnlOng ablIIlln t plnnts hnYl b(en sllldied rytologicltlly One hnd 09 Chl0Il10S011l(S in the SIlll11 s(clor which had gle(uih malformcd tnxes and 106 in the llst of lh( pLllilt which had tllc llSlll1l gray leavcs The other plnnt probably sltlrl((l with 91 to 94 (illomosomcs At prcseuront only a small pl1lt of it hIlS this Illunb(l b((allse tll( part which has 86 chromosom(s fmel grlcillsh smooth leay(s hits oycrgrown the

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POLYPLOIDY AD A)(ECPLOIDY IX GCYlLE

91+-dlLl)lllosome part which htL gnty mnlfOllllP(l IPllWS It is possible that somC of tlte i[lCguln growth pnttCrn of alH[[ants is the result of (liffCI((lt growth rntps among ((Ill whi(h nUT in ChlOlUOSOl11e numb([ if somatic loss of (brOll10sonlls O((UlS frl(juilti)- in tisupi

Furti1PI yi(L([wC ill SIlPPOtt of It (auiial I(btion h(tw((n hirh (hrOm()~()lllf llumbpl and the O(CllIT(Il(( of chiI1lNn may hel (iucl POW(lS [8 oht ainptl hYIWrttbplntnt ofTsp[ing from tlbllTlLUts in (ol((shytion 41fj3 Of th(lsl plants onH~ of whilh luul n muny t1 1-14 ehlOIllOSOl1HS ltnd all of which p[ohuhly hall at IpHst t W lllttny olnjoush- wtrp s(ltotial chimlms In- titt tiltH th(l- wpr mOllth nitL find pnt tlll first llnw([ing tngp In llllH of tIn phut 23 or pwn 4 ditirwt pnrtiolls of YUlying sizp (oul(l ill (listingt1iIH(L _ ftw of thr Ip lxtrpnw portion of (vpml of tillS plunt WllP pxtunilwd eyto]ugindly anti fOllucl to htwl Ils than li (hroflwouW

FACTons JJf1TlG E1 PLOIDY D POLYPLOIDY Ii G1 lTLE

Bptwp lllPini in pnlllll lHntlwl (Ill of ~lhLllhi plnut i (omplltE II 4wltiy of thi normal ltlPlOti lHilavioL fltllr( nIHl qllidd) rIYtuit(l tlip pnIvplllill lun Illl of thI prip of (IIl ln1H lxumirlld The triploill HutU1 of tlu )4 -lhlllfllOSOnlt plant WL~ illlliIHtL hy thp prpPIlII of trin1pnt at I1lmplmp I ltnd titmiddot tPt mplnicl nltt mp of 72=-(middothrollooIll plant hy thl IHPStllll of If IHlllrintilIlt In gPIHnd thpL( wa tt tlllltnlv towllnl ((lmI ditlilmdoll of (IttomoshySOnltmiddot at tlllttphl1( I rhruu)dwlIt the t1i( Polllll )Illin co[ltltinpd thl lPtluI[ numher uf middothrollloorlll Tili (toil( f pollln mother cell in which to t wIv hrmllosnnH hllutviol WtL f01tunttt( l)l(tl1ls( t11(1p Stunp phmt hlginniIlg Ilt tIl( -t -(tromnome 11v(1 IUld (Imshytinuin throughOlLt tIll lmiddotritmiddot ttl) not hnw Ilormal Hwiqi in tliClr meguIJOrp motl1ll cdk In5tetLI~ tlllre is ttpmniosi- (If the type of genfllltiyl t1P0p0l I ~ bull

~(or(ling to POWl and Rollin 10) JJonw~1fgtttil1 apoIlliis is the predomimmt but not the solI method of 511(1 pto(hwtioll in plant whieh have )4 or mor( chromoome _~polllids attllully pliLY-S a dunl role It maintains a uniforlllih~ iLmong 3J ItIld -r stttLlds whieh could not exist if ampbimhis (gt1e(onun011 Prohably if it W(re common 3) euploid would be re5trietld to in(Iividllal phmts tlltel twn -r plan ts would be quite YarifLble ~Ionover by meaIlS of itpomii= the rltre fayomlllp aneuploid (ombirwtiollS that arisC through mnphimixis since apomiis is facultative or diplospol)- ]lfi e a ehallee to mrvi-e to increase in Ilumh(r of plants and thtrefon to establih what amounts to a done In efftct nonsegregating apomiis is lqui1l1ent to vegetative reproduction Beeausp most anpuploid found in 3r and 4x apomictic populations are not as -igorou as the apomicts and proshyduce fewer or inferior seeds apparlntly not many hnse giycn rise to clones and thp1epoundore to aneuploid stands or strains Furthermore because apomixis is tht predominant method of reproduction in 31 and -l popUlations in thes( apomictic stands the[e is now only limIted opportunity for the creation of ne types by means of amphishymhis For these 2 retsOlls aneuploid strains nre uncommon

In contrast to the relatie paucity of aneuploid individual in apomictic 31 and -1t stands aut commercial stmins there Wits an

32 TECHNICAL BLLLEllN 91 S U S DEPT OF AGRICULlURE

abundallce of olleuploids with volied chromosome numbers oillong the offspring of plants which reproduc(d amphimicticolly For instance 0 segregating culture was obtained Om the seeds of a Texas moriola plant which had some guaY111c chnmcteristics alld probably hod 90 chromosomes The chromosome ll11m1gt(Is of 23 of the pllLlltS in this cllUme w(tmiddotc deLelmiMd rrlHY rflllgcd from 70 to 95 No 2 of tbe plants were morphologicnll~ alike In size they varied fOm VOIY large to very s111ol1 Also by using diploi(L 36+ -chromosomc plants 11 iell ore amphimietic as sced PILlClltS allcuploids were obtained whicb 7aried in chromosoll1c numb0l from 43 to 49 when pollen from 54plusmn -ehromosomc ploots WitS used and from 40 to 56 whcn POllillfLlion wos not controllcd

OlH-lhild of thc 4rplusmn o[rtyp( nOlmnls sludicd hlLd lcss tlmn 70 chromosoll1Cs Thcs( plnn Is W(IC defkiell t for 3 Ot more ehlOl11osomps Othet e7idenee shows that (middotollsid(Inblp (hlOlllosomnJ unbtLliwcl existed among lfLbIcgt (omhinnlions FIOIll 7~plusmn-ehlOmOS()Ill( populnshytions ttIl un hlLlnn(e which nULY be rcplesetltccl itS 41-7 wns found in plfLnts which h(1(l us fe flS HI) (htOmOSOl11lS From noncontrolled pollination of )4 plusmn -chromosol1l( pInnts n04- and it (ill-chromosome plant (1( obtained From )1oneontroll(d pollinotion of 2Iplnuls iLI1(uploid 3) plunts whidl YltliNL in (hromosome ntllnbel from 40 to )() wcrc oblniu(( Th(J( wns (Y(11 glCttPt ehlomosonull unbnlnI1(c in some of thc oflspling from eloss(s bltWCt1l 30+- IUHl Mplusmn -chromosomc plnnts such as thos( whi(1t hnd 4a 4ii 4n alHl 48 eu0I110S01l(S B((iLUse such unhnlance 1gt(tw(el1 ttp 2 lnd 3t Itv(ls is possihl(the widt d(yintiolls from (lIploicl Illullipl(s of If whicmiddotit W(lt found in an(uploids with high (htOlllnSOnH I1l1l11h(I It1e not Slll[)riing and aetulllly invol-( Itss ehlol11ool1ll1 unhtlfLll(C IW(fLUSP 11l111tiplps of NL(h of thC IS (bl0l10S0nHs wcre PI(SNIt 111(1([01( if amphimictic J(pro(u(lioll WPI( moJ( (omrnon in 3t nnel lr populntions not only would 11l01C ull(Ullloid bp (X[w(middottNl but tll( dn01l10S011W nllmb(rs of lllony of tlH1ll ould d(yiat( (onsid(l1bIy from th( tllploiltL lIumb(ls

In alt(ll1pling to account fOl th( relntiYt paucity of lUlCUploid jndiyiduuLs ill flp(lIniCli( 31 fLnd 4) stnnellgt tLlHl (omin(leial slmins one is conscious of the [n(t thitt most of thosp sl udiNl rytoogicnlly WCle 1(s8 vigorous IUld hlHl n lowcL sUti 111 Iate bdoLc ltttchillg 11l1tlulity In (ontlllst most of the ttIlcuploid 3( oflspring obUtin(d from (tOSS(8 in whi(h lilt sPCd PttlPl1ts (1( lilt jLlllpilimi(lie 2t plnnts wtrc vigorous in spitp o[ the filet tbal in SOIll( of lltPlll there Wfll ((11

gleltt(L ChlOlllOS01l1nl nnbulltl1(e in proportion to tltc (middototn1 nUI1lb(l of (lt101110S01lteS iIlvolY(d than thc middotlr-1 (hlolllosollud unlmllLllN in t11( liii-fhIOIllOSOJ1H pltUits found in tr populations lh(IlfolP tto ksspn(d igo] of most of lht imNlploids found in 3( I1nd4r npomidic populations mllst 0( Itltlihntld to g(I1i( uniJnllll1(( following amphishymixis rnth(1 tluLII to YltlitliOIl in (ltronlOs0l1ll llumbtl P(l S(

forlH of illUph imixls Jlltilizn lion of Pgg wh i(it hilS Ow Ul1lld tI(middot((l nllmlHl of ehl01l10S0Il1PS kd to lil( for11lntioll of thost plnnts with high dllOIll()SOI1W Ilulllillls which (1( fOllnd ill al and middotlr PO[luitltinlls Aneuploidy WitS )llOIl (O1ll1l101l thnll euploid) IlU ellomosollll numhprs of plllnte (xnmined thus fttr (lxl(ll(in unbroken ~W(llllnCe from RO through 113 and in unbrokell sltlt11laquoI1(( feom un throup I 110 112 120 and 122 Tltn dll0ll10S01lH I1uml)(l dlt(wllded upon the number of ChlOJl)OSOll1lS in lhe lgg (54 plusmn 01 i2) plli lhl 11llll1)eL

bull

bull

bull

33 POLYPLOIDY AND AXELPLOIDY IX GlAYULE

bull brough t in by th(gt po11(gt11 CTlllil1 Sincc (IOBB- us (gt11 us B(gtlf-pollinulioll occurs the illlmb( of chromosomes (ontribut(gt(1 by th(gt pollcn grain muy be quit(gt uliublC It lIns b((n shown from brct(ling dn1l1~tllilt polleu from trlplOlds contn1l1S ubout 25 to 3a eiIlomosoU1(S and that from tetmploids ItbOll t n to a9 In a mixcd slund of iH plusmn -ilnd 72 plusmnshyeh101110S011le plants thereforl po11(11 gluins would (ontain ubout 25 to 39 clir01110S01ll(S (onslq Uln tly if the (~~ IltHI exnctly 54 ehlomosol11Ps tlip PXPP(tN lUI~( of ehlomOSOIll( Illllllb(IS of tho abeJlllnts would bl 79 to 93 and if it had (xucliy 72 tilt ran~( would be )7 to 111 Thc lxpltclNI lungp would 1)( ips8 if most of th( plants in a stnnd hnd tlit mill( (hr0111001)1( nUllllwr out of 5l plusmn-cllloillosomc stands it would be 70 to S7 if tlip Pgg hnd PXtl(Uy 54 ehrolllosomes und out of 72plusmn-(hroUlosome stHnd]l would bl 101 to 111 if the lgg hnd cXHetl y 72

bull

Onp mciy t1Jbitm1ily c(111 OO-clllomosoIlw pliLnts l)(ntltploids (5 tin1(S 18) nnd lOS-duoIllosonw plltnls tpxltploids (n tinl(s 1R) but sinee ul1tuploicly is more common than tllploidy at thts( high leYls in the polyploid Strils the tllmjnolo~y hits little lI1NLtling unci the tllrI1S ptn taploid und IHxnploid mel ieuJl only t hu t the total number of chrOlllosollWS equnJs 5 01 G tinl(s 1 whkh (qUiLls lX If chromosomc ussoeintion at llwtuphasl I W(IC llSld fiS a ([itprion fot d(tlrl11illing polyploidy tlHll tlipse pilllltS would be consid(le1 pltlti~l ldmploids at most rhus fal 110 natull11 slltuds 0[ tonuu(lC11t1 stnuns which nlC plutuploids or htxltploiltis 11aye b((11 found nIHl nom WllP lCpo[t(d by the RussifLns Chromosol11e Ilumlwrs higlHl than 72 hayo b(en foulld orJy in indh-idunl plants in 3r1tl1d 4l~ populntions nnd in their offspring

CytologiCnl studilf of plnnts with high e1l10111050l11C Humber hltye been llstrieted almost llItin-ly to plants clnssifi(d ns morphologically abtITflnt 11 nbCIlllnts whi(h exhibiltd the whole syndrOllle of chnla(tlristies hayc hnd n hi~h ehromosOllw llllmhl I A few SletOlS of Chim(ras must be ex(cptNl J but such plltlltS obyiollS1y bl~ltll with a lugh chromosome lllllnbpl frow Illany ll0I1111H[ltll1t plfLnts 11aye mo[( thnn 72 plusmn chromosomegt is not known One SO-ehrolllosomc plnllt in a 54+-chloIJ10S0111C population WllS ofrtyPl 110rmal but not ItbplTnnt

bull

That tht sYllciron1l of eharaClllislies by which ahlllnnts nrc dismiddot linguished from othPl mOlpholo~icnl typlS (ltllnot he nttributed to high chLOmOS0l111 llumbpr alolle was shown hy tillt oCltypl llormals (nonflpomiclic offspring) f1OI11 niWlrnnt plltnls ~Ilhough 11 of the 12 plants hnd 82 to 03 CliIOIllOSOIl1lS which is till SIUll( range of chrolllosome llumlwls shon hy ab(ITnnt~ flolll [)4plusmn-lhromosomc populations none could bl classrfilcl as rlal Itb(ITanls ThlSC plnnts diflplNl howlYp[ in mode of ori~ill from nl)(lLant3 in 3x populnLions 8in(( llUir eh[ornosotnt llumlHIS Wlle (onsidlrably 10([ tlutn those of theil 5(((1 pannls t1wy Cllllle from the fertilizition of ltggS which hnd the lld uecd n II mb(t of dllOmOSOl11lS TIl(~ (onsl it lll((1 only a smull [metion of thl totnl llumbt1 of ofl3pling ~ from nigtt[llullS lind lepl(s(gtntccl till best of tht [lcombinations of genlS thl1t [(sulted from nmphimixis

In nttemptillg to ullI(lSlnnd why most plants whieh hud IL high rhlomosolll( numlwr wen nbpl[tll1l it should be ((membered lhut tlt( abellilnls sLudild O((lIITNI in 31 und 4) I1pomidie populations

34 lBCHXI(L BCLLETlX 018 L ~ DET OF AGHICnLTHE

All nlH1Illnt I(ntll flolll tlw fPltilizllliol1 of nil tJg whieh of itwlf without ftllUiz11jon is ulti(l(JllIll( rOt PlOdll(inJ It viabI pInllt tuposhymid L This n(lospOlOllS ltg is (lni(ldly bnllIlJ(pd ill itsllI bullpshyptt1(ntl~ 1m t11l11l1lU(P llsillts hllI stl(h nil PJJ is fllfiliztd tltt d(JII( of 1I11bllIUl( d(1)(lldill upon til( tolal g(lli( (omphgt Thl 1l1OlC vigorous of llll lligtPIlllllls [)lohtllJy Indiltutt It tss disllllllId gtllic bnluIl((

Insoful us higlt iJlOl11(lSOlll( 1l1lmlHlS ill glllLyu1( 111( [ill1i(( to plllllH hi(11 U1( IlHIlpholoJiu 11 nl)(II1I n l I II( pst lLillisI 11 11( 11 I of I1allllul (ullds whilll huy hih (illOllloSOIU(I llumlJ(Is SlIlll 1 IlltllltLshyplois ntHt lltgtt~pl(lids will ill limit( (0 wlml(Pl SWISS HllIllllIll plttnt huyl in lslblihinJ th(ll1sllCs n lllltluni slnlHls llfol shyttHtltlll lllost ahplTIlIlls ~l() lOlC slowl than HOInut plllnls in thl RIU1)( (lilt 1lIP Iso POWS I) ohtltilHd n PIy low PplIIIIlJl of germinal iOll of s((ds plOd wPll 1) IthtITIUl h ~ rOI(OPI tli frC[lIPllt O(Ulj(Jl(( of PIlOlilll (ltill1PIlS in uldt nlHIIunl lind tIl( loss O[

elllomosoJlH o)sllwll ill indiddttul ((l1s il1diCttl( instabilit o[ (II lOmosolllt llllmlJ( alJlon~ Ih(s(I pial Is Thl (0 III billa I iOIl o(slo gt)wth f(w ollSjlljllJ am 1 illst 1 hilit of IhlOIllOSOlll1 nlllll hlr ploilnhlv (middotxphtins whY no stands of nIHIIIUlI and illtiltiPlllally of tHlllnploids nIHl IHXllpl 0 ilti htl h111 f(lllHI ill lUIUp FOl t hI t illll bping 1 t cnst l(tlnploilly S((IllS to lH Ill t1PIHIlilllil ill lltl polyploid sPli(~ fOl Ililltllltl sunlit Itnd will olltilllll III Ill nUlil [lPrlwp hY OJ illtlrnnl luljtlS1UHllt (ll [(IImiddottlIjOII nIH(h of IIhpITHlllY all (JPI(OIllP IlowshyeYp [Ol iHlliddulI plnllts (1llIlpioidy is 1101 tilt IIPlll limil sil1(p pinnb wilh hiJh 1III01l10S01111 Jllll1tiHl ltl( cOl1tilluHII lImltHl(t ltI )l()() in Ill( ~I 111111 middot11 IJOpuilltiolls IhlollJh tlIP jPltiliztltioll of l~g~ wlult hHye IIlP till Iltit1(pd IllllnlwL of dllOlllJ)SOIlHS

Srl 1 IY

TItlt sllmpip of gllllYllh planH lIs(( for lh( clll(lminlLtioll of (IUOlllOshysom( IlUIll gt(1- illciudlcl TtXtUl nllll ~J (xiclut poplliat io Il lnll of if((111hullS (OIl1Ill(ITilli -tmins flnd OfISPIllI from both (ollllollNl and nOIl(olltloll((l l)(1llinatioll of s(C((d llhml 111( llxiclLIl mu I(Iia1 inc] tlll(( I (i- -(middotIt ronwsoll1P (ltii pioid ) rimiddott plusmn -(h 1OIllOSl)IlIC (tripoiltil and I2plusmn-(hloJllosolll( ((trnploid) SltlH[- Both i-Iplusmnshynod 12 -eh l0Il10S01lt( eOlllllHJ(bl st millS (1( llHlil(L Only 12 plusmnshy(htOJllOSOllH stunds W(II found in (1( maltIinl flOIll 1(XII Both th( jut(IIll(dialp Hllti Jootl Juayull I2plusmn-(llOlllOS(JllI( IyPIS fmlll 11exit() ILnd T(xl WIIP ill(IIHkd TIlll W(IP llO sllulls with highl (11 10 Itoso 11 l( 1H111l h(r 1rOWInI illtidd lint plttn ts it It hi~dler (hlOlIHlSOIlU Iltllnlwls most m-Ialllll nhpITllut (11 fOIlIl(1 ul1long tIl( ~J 11111 4J SII~Ilc1s and stlttll1i

~l(i(Jsis iIl1)oll(Tl mo(lHl (Us WIli (ollllllp[e ill 11111(1 ill tIll polyshyploid s(ric Tn diploids hhukJlts WlLpStPIl nl nwlnphns( 1 nud the Itnnphnsp 1 distlihulioll of (hl(JJll(lSOllllS WIlS equlll In triploidraquo trimiddotlllIIlH or hhtIllls plus tlllhnl(llts W(I( S(IU III Illltuphnw 1 ill

tetrnploids (lUiHriYlllllIt trisHhllts bintipllls i111d tllliVllhlIls Wlm fouud flll IPlldll[( towtlrd (quill distribulion or elll(lll1moltl(s Itt flllltphnip 1 whitl WIts stIgpstpdin triploid was mon PlOIlOUlUld in tltrnploids 111 plnnts with hi~hll (lirolll(JsOJlW l1nmblrs (h(I( WitS an (lXetss of bhnhnls nl llIltnphnsl 1 Annphnsl 1 distribution tentlNl tOWltId ((1 liltIi t) 1I(11(( he intCfllnlily iu nnHphn~ I c1islli

bull

bull

bull

bull

bull

bull

POLYPLOIDY AXD ANETPLOlDY IX mmiddotA)TLl~ 35

buLlon umong hight polyploid~ wllsl(i~ thl1l1 til( ilHfllnlily in tripshyloid in spit( of tll( 1111gN II 1Il1l 1)(1 of lllOIlI()~Onl(~ Huploids from t(Inp]oids ghowpd SOJll(gt lgtiYI11(II(s ul lIHtn pJIIIS( J l1tlphns( I distribution WitS l(~ l(gu]nl thnlt ill diploid ~1I(h Ilwiolie ill(~~l1shyInliti(R itS nSYIHlpsis II net 1u UlI( or (middott(Jkilll~i PIp ob~pIY(d ill 11(shyltjlHlIt1y in dij)]oids Hlld in ~OIl1lt 111l(II])1oid middot11 oIIlYJll 1l()Jlllld~ ~rnh stNility dup to Iud of SPOIOg(JlOIIS t issup nnd 1(sl it lit iOIl Ilu(]ti dso lI( olJs(IYtd ItlllOllg HlllUphid tpt IUpl)id~

middotlllt(llllldilltps (1( Ihnra((lizpd (ytologilully h~ IUOt( p()l~shyYnhllls III ])lPtnpllH~( I thull UH good gllllynt tYpl 1l10lP of lhlit poll(11 Illothl (llIs show iJlg lnggillg dllOIllO~(lllll~ nl HllUplws( 1 llld ] I und by tlH ]ll(~(ll(( of mlllY llIi(IOlytp4 uIllOIl tIll pollllI gluills

Till lllUllI)I of CblOl1l0S0fl1P ill fUllct JOJlnl POUl1 gluill plodtlC((l by lI ~11 ttlld uhllllwl )lInngt was (titJlatld flolll th( IlllllJIHI4 in tilt (hlOIllOSOll1 gloups ill pollllt lIlothI (1middot11 nrtpl Htlllphns( L laquo01 tIl ~I it was 22 to W rOItlll middotlJ ~~ tolIi fllllllllTlllIls froll1 tIll lI ~s to 4 aJl(1 (ot allllIanls flOJ1l tIlt middot11 Is to ii ~ It cll((k Oil tll(s(I (~timILt(s (ulltululilllls bnsld Oll hn(dill~ IIntn sho(d thnt 101 tlIP lf plallts tllt lUll~l WIIS 2) (0 al Hnt fOl tIL 110 to ll

Instahility of (hrolllosolll( lIltllllJ(1 t111()Il~ nllIulIt tiS sll~(slaquod by tltc loss of glOllp~ of hlOlIl(lf(lIll(S in llillllll ll1othI ((11 ililough such JlI(ehllllislll~ a Illultipohu s]lindlmiddots lIlId twin (Ill Lo~s of dn()IlI()~om(s PIIyiOllS to I hI fOllllll I iOll of jlollln mol h (I ((lIs was inshyf(ITPd ftom tIll lilJdiJl~ or ((lis wldeh (olltuill(d IP(I dllOIlOSOIl1Ps a 1llollg thost whiCh 111(1111( higlt 11l1111lwl IHoduCld hy Hl)(gtlI~ll1ti in -(J~ populntions Illld Ipss fnqlllllt]Y ill 11 111( ndllnl numhpI of CI1l(1I110shy50111(s WUs Yluiahlp (onsiltipIibk ehlolll()solllal unlgtulHl(( is possibll ill gunyuhmiddot FIOIll hOlh (oJlIJoUld lIud 11011(011 [lOllp(1 pllllilln lions of 21 plants growing ul -ulil1tls WPI( Oblllilllcl olispling wllOsl dHOlllOshySOIlI( lIlUlIbll IIlllgpd flom cll to ili

B(lculIs( apondxi~ i~ t]llt l1ldo11liJltlJli f011ll of jppjoduCtioIl among tlipioid Hlld l(tluploids ~I llnd lr stands 111( fuidy 11Jlifo11ll ~ f(w of thl (OmnA reinl s[Iuill HI HllPUploid tl Hlld 11 B(IlllSl oi ]imilpd nJ1l0ullts of alllphimixis lIud I(lJ1llul JIllmiddotjOjS ill llJ(ga~p()Jl moth(middot) (pUs ho(~Yll thltp Wltj( illdhidllul vurinll(s IlnHlIlg lllt 11 nile 11 l101Hllnshyions nlltp 1IJ0lpho]ogi(nlly distin(t tYPlS ill(middotIt)(lpd IHlploids offshytYJlt n(ml1uls alld uIWllUlIls XilH huploid from TlXllS (ollp(tiolls find (OJ1ulIJlitl t Ill ills WpI st lldild hud W IUlll (i hnt H (hlollloshySOU)l~ )[ol of tIl( nlitY]ll 1100JIlnl ()p lultuploids lJ( (hlollloshySOllW lluml)(rs of [(1(0 at oHtYjlP tlolll1uls lIUlgCd flom i4 to iii Thc 11U121]HIS or t It 41 lIld H1Yliploid -11 olr~pp llolmnl lIUll(d flolll Q5 to lil lll( dISlltHIIOH hlllIg liS [ollows )LI jlll((nl hnt J(4S thnlll 0 (lUOJ)lOSOlllPS ~~l J)(I((I1I 7() 01 71 j1i ]wllpnt 72 Hilt Hii gtprshy((Ilt 7 to 7i Ah(llullt~ from lriploids hud bO to H3 chrolllosoll1es thospound flom tlllilpl(lid~jlJ 1[) 1~O

~ll alJlIJIlIls had It high Chl(J11l0S0111P nUlllh(l whi(h ((IunllPd the ullldIH(d lllllnlwl of tlw ~l(d pnrllIt pIllS tltP Humhl (oJlllilllltNl by thp ]gt011(11 trlail wItic1t (Ilplpd Jlllilizatioll Tlwy o((llLIpd uftr elthe stlf- (H ~l()s~p()lliJlnl iOll ill hOlh U Hilt -1 P(1)ullltiolls The POllPll pUJ(nt wns (iilwl gllnruhmiddot O1lllllJioln ~(d(Jlinl (hillllIHS W(IC Inid common nll10llg oleiN nl)(ITllllts ill 41llo]luhttiolls DiliPI(l1((S in cIiromOl-lOIll( llumi)(ls W(I( illYolnclill the fe (hinwlll (xtlll1iUtd

Ab(IjI1I)() stlIlwd to JJ( n~tri(lld to thost plants which nds( in 31

36 TECHNICAL BULLElIN 918 U S DE~T OF AGRICULTURE

and 4x populations throtJCYh fthnormal ftmphimixis (nonleduction folshylowed by flrtilizntion) and to their apomictically produced offspring lwlvc offLypc normnls from abcrltlnts whieh yftried in chromosome 11U1111)I1 front 74 to 93 probably nrosc thlOugh amphimixis Since bulltlllSC lattc] plnnts showed only a few of lhe phenotypic characteristics of flbelTtlnts themiddoty probably jl1dicnted more successful genic balances in spite or high chromosome number

LIT1~ltATCRE CITED (I) BlmGJm A n

H)lel OImiddotAYlrJo ILAllH W(ln row (((UO~IOSOlE ~nrnlgtfu Ekicllce 99 22middotlmiddotmiddot22[)

(2) ROCIANTIIrA Z 1 [HOJS(IIANRgVA K]H133 K IHOI()OIl lSg(gNlfA (aAllLY 1IlTIIENIl1 ARltUNlNrUM Olt lY

(slnlllN UJII)lt om 1I10J()(lJ I TlIS n IVIIO gH lA 1t11I mn nl llmrAshyTIM GIlAY) Truey Krtmiddotd-Aziatsk (josue l-ni (Acln l-niv Asia( (((11 4pr S Bol fn~e 15 13 pp iIllls

(3) DIAOWA V L 40101011 A and ~IJ~(III-A X ])35 YBIt(lImiddotItIIJgtoiI) ZYl()B~IJIlor(flIi-l(ln 1TIU~I(IIING I)lm VAHlBshy

~IXlHN Il lUTIUoitlmiddot~1 llngoiTmiddotlt~l nllAY I) 1 INIAXtTM cuw Hut (cnthl BpihcClr Aht A lorph u Physiol der lfltwzbulli3 2 )3301 iIlUl

EHA K II)H IO)I1XIS I o (A1TLI1 Xull ACllcl Sci Proc 30 352-355

(5) I(clwwA E1 H2l nH ]TW(middotlunOf(l1srllHmiddotIIrB DEH ~IXNX11CIJ-r GAIEIOlHYTBN

l1B1 PAWlfI ENIIM AIHll~NIAIlM OItA Y lXD 1 lIYSlgUOIIJOlttTS L Assol 111( lhys lllt h FIICUlt )lmcow l-nh Bul 2 145-IGO illus

(0)-shy1932 JIII~ 1llJgtOIIEXr O~ TIIB FJlArl GAIETOIIIY1pound 1 IARTlnNIU

AIIOgNlAIDI (middotOlYlLBl lour Hul de 1IHSS 17 72-)9 bullmus [In RUisinli Euglish 81ll1IUnry pp t)1-95 j English explnnation of figures pp 98-99]

(7) l[lnlm T R 1l15 lUEnxr-rG WllIr lJlAD1Clll(HOlmNZENE TO 1CI1I1ATE CHROMOshy

som STUDY llllin Techno 20 121~125 illus (8) POW1-RS L

1015 JEIIrnIZAJI0N WITIlOl1 HBDCCTJON IN GCAYULE (PARTflENIUM AIIGI~XTNIl~l OUAYl ASD A IIYlOllItJS1S AS ro rilE EYOJCrlON OF AJO~I1XJS AU lnL1PLOID1 Genetics 30 l323]-3lO ilIus

(raquo) --- nnd UAHDslm E J 1015 lHlQlgNCY OF ABOHrBD middotO[rtJN OHAINS AND ~lIcnOCYTES IN

GtAY111~ lAH1IUJNltlI AUGBNTATCll GRAY Amer Soc Agron bullJotlr 37 1IH-103

(10) ---nnd HOIIINS It r 1915 lI~rUODn~TJON AIgt POLlINATION SlUDUS ON GUAYCLB PARTIlEshy

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U s GOVERNMENT PRINTtHG OfflCtl uu bull For snle by the Supcrilltcn((middotllt or Documents U S I)ocrnment Printing Olllec

lIshington 2 D C - lrle 10 cellts

10 ~)

bull TECHNICAL BULLETIN No 919 bull August 1946

VARIABILITY OF CERTAIN SEED

SEEDLING AND YOUNGPLANT

CHARACTERS OF GUAYULE

By

WALTER T FEDEHEH

Formerly nssocillte genctieist Spccilll GUllyul(~ Reseureh Project

Bureau of Plant Industry Soils and Agricultural Engineering

AgricuIturul Hcscareh Administration

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Polyploidy and Aneuploidy in Guayule 1

By A DOROTHY BERGNER

Formerly associate cytogeneticist Special Guayule Research Project Bureau 0Plant Industry Soils and AgriculltLral Engineering Agicultura~ Research Administration

CONTENTS Page Page

Chromosomal range in gllaYllle __ 1 Loss of chromosomes by abershyrants_____________________ _Polyploid series in natural stands 22 and commercial strains______ _ 2 Chromosomal varianLs within

Euploid populations and the pol)ploid series _________ _ 24Btrains________________ _ Oligm __________________ _2 24 Aneuploid stmins_________ _ 3 Hapluids from tetraploids _ 25 Intermediate type______ _ 3 Anellploids from diploids__ _ 25

bull lIaterials and methods________ _ 4 Offtype normals from trip-Chromosome mOlphology______ _ 5 loids and tetraploids___ _ 26 Chromosome behavior in pollen Aberranls from triploids and ~lOther cells ________________ tetraploirls_____________ _7 28 Diploids_________________ _ 7 Offtype normals from abershy~ Tliploids________________ _ rants _________________ _8 30

~ Tetraploids______________ _ 12 Sectorial chimeras________ _ 30i Plants with higher chromo- Factors limiting aneuploidy and so~ nU1~rs---------- 3115 polyploidy in guayule_______ _

18 Summary___________________ _~ HapMds---O ------------34

~llen__~__________________ _ 20 Literature cited ______________ _ 36 7-J ~ ~ c1 vi ~ ~ lttROMOSOMAL RANGE IN GUAYULE

~JIn thl-fall 6f11942 LeRoYJ Powers and W B McCallum collected ~eds of gnaytJTe (Parthenium aJgentatum A Gray) and mariola (p iJcanum H B K) in Mexico and Powers and Walter T Federer Cigtllected seeds of these same species in Texas Early in 1943 many of

these seeds together with those of certain of the commercial strains developed previously through selection by McCallum and probably of Mexican origin were germinated and planted in fields at Salinas Calif Chromosome studies especially the determination of chromoshysome l1umbers were begun as soon as possible in order to facilitate genetic and breeding experiments

After a few months of exploratorystudies it was apparent that a polyploid series exists among natural populations and commercial

bull strains of gnayule Stebbins and Kodani (13)2 working indepenshydently came to the same conclusion at about the same time

1 Submitted for publication February 27 1946 Italic numbers in parentheses refer to Literature Cited p 36

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31

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bull

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bull

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31

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bull

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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




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bull

bull

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2232 1 23526

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bull

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11

bull

bull

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- 317313 L---------~-70-chroUlosoUlO ____________________ 3 3337 1 31435

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7 3438 31338shy12 3637 33331gt


bull 20 -- 342311

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bull

bull

bull

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bull

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31

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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




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bull

bull

bull

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bull

bull

bull




~_______o_o__o_~ _______0



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- 317313 L---------~-70-chroUlosoUlO ____________________ 3 3337 1 31435

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Do _________________________ bullbull_ 3530 2 3313711 L---------~- - __ --- 1 34235shy

7 3438 31338shy12 3637 33331gt


bull 20 -- 342311

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3630 34236 - 1 34335tJ 2 31 136


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bull

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bull

bull

bull

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31

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bull

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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




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bull

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- 317313 L---------~-70-chroUlosoUlO ____________________ 3 3337 1 31435

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7 3438 31338shy12 3637 33331gt


bull 20 -- 342311

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3630 34236 - 1 34335tJ 2 31 136


middot1 30a7 --- --- _-

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bull

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31

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bull

bull














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bull

bull

bull

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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




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bull

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7 3438 31338shy12 3637 33331gt


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3630 34236 - 1 34335tJ 2 31 136


middot1 30a7 --- --- _-

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- --- - 1 35131 ---_ - - _-_ 1 35236 ---_ _------ -------i---_ 1 35335 _ _--------- _- - ---- 1 36136


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19

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bull

bull






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bull

bull




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bull

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27

bull

bull

bull













bull

bull

bull

bull

bull

bull
















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bull

bull

bull

31

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bull

bull














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bull

bull



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bull

bull

bull

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bull

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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

L-0 ~ ~ ~ 0

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bull 0

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bull

bull








- 317313 L---------~-70-chroUlosoUlO ____________________ 3 3337 1 31435

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7 3438 31338shy12 3637 33331gt


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3630 34236 - 1 34335tJ 2 31 136


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- --- - 1 35131 ---_ - - _-_ 1 35236 ---_ _------ -------i---_ 1 35335 _ _--------- _- - ---- 1 36136


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bull

bull

bull

bull

bull

bull










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bull

bull

bull

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Sumber 1

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bull

bull

19

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bull

bull






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1 1 17218l~~l 1 ISl18I degi











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bull


bull

bull

bull




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4 S 11 2 10112



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bull

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bull

bull














bull

bull

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27

bull

bull

bull













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bull

bull

bull

bull

bull
















SECTORIAL CllntEIUS



bull

bull

bull

31

bull

bull

bull














bull

bull

bull



bull



bull







Srl 1 IY




bull

bull

bull

bull

bull

bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

L-0 ~ ~ ~ 0

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7 3438 31338shy12 3637 33331gt


bull 20 -- 342311

- - --_ 34835-~

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3240 1 327333339 1 33138

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3630 34236 - 1 34335tJ 2 31 136


middot1 30a7 --- --- _-

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- --- - 1 35131 ---_ - - _-_ 1 35236 ---_ _------ -------i---_ 1 35335 _ _--------- _- - ---- 1 36136


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bull

bull

bull

bull

bull

bull










------------------------ -------










il 1 3045 I 1 3S142 v 1 I 3-14 I 1 1 39240









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bull

bull

bull

bull

bull










---------------- -- --- ---_- ----shy-~---


Sumber 1

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3U5-tfi lt1-11+1



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bull

bull

bull

19

bull

bull

bull






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1 1 17218l~~l 1 ISl18I degi











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bull


bull

bull

bull




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bull

bull

bull














bull

bull

bull

27

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bull

bull













bull

bull

bull

bull

bull

bull
















SECTORIAL CllntEIUS



bull

bull

bull

31

bull

bull

bull














bull

bull

bull



bull



bull







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35 Hi2-172 iJlUll



10 ~)





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10 ~)





By

WALTER T FEDEHEH




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10 ~)





By

WALTER T FEDEHEH




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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




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L-0 ~ ~ ~ 0

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bull



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Srl 1 IY




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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




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L-0 ~ ~ ~ 0

f-I -I- tEJ

bull 0

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42-1 r

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SECTORIAL CllntEIUS



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bull



bull







Srl 1 IY




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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




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L-0 ~ ~ ~ 0

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SECTORIAL CllntEIUS



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31

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bull














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bull



bull



bull







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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

L-0 ~ ~ ~ 0

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bull 0

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bull

bull


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4 S 11 2 10112



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27

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SECTORIAL CllntEIUS



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31

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bull

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bull

bull

bull



bull



bull







Srl 1 IY




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bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

L-0 ~ ~ ~ 0

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bull 0

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27

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SECTORIAL CllntEIUS



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31

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bull

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bull



bull



bull







Srl 1 IY




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bull

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bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

L-0 ~ ~ ~ 0

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27

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SECTORIAL CllntEIUS



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31

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bull














bull

bull

bull



bull



bull







Srl 1 IY




bull

bull

bull

bull

bull

bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

L-0 ~ ~ ~ 0

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bull 0

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-




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Onw1



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27

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SECTORIAL CllntEIUS



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bull

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31

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bull

bull














bull

bull

bull



bull



bull







Srl 1 IY




bull

bull

bull

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bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




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L-0 ~ ~ ~ 0

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bull 0

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27

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SECTORIAL CllntEIUS



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31

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bull














bull

bull

bull



bull



bull







Srl 1 IY




bull

bull

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bull

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bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

L-0 ~ ~ ~ 0

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bull














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27

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bull

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bull

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bull
















SECTORIAL CllntEIUS



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bull

bull

31

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bull

bull














bull

bull

bull



bull



bull







Srl 1 IY




bull

bull

bull

bull

bull

bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

L-0 ~ ~ ~ 0

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bull 0

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bull

bull

bull

27

bull

bull

bull













bull

bull

bull

bull

bull

bull
















SECTORIAL CllntEIUS



bull

bull

bull

31

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bull

bull














bull

bull

bull



bull



bull







Srl 1 IY




bull

bull

bull

bull

bull

bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

L-0 ~ ~ ~ 0

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bull 0

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27

bull

bull

bull













bull

bull

bull

bull

bull

bull
















SECTORIAL CllntEIUS



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bull

bull

31

bull

bull

bull














bull

bull

bull



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bull







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bull

bull

bull

bull

bull

bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

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bull

bull

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bull
















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bull

bull

bull

31

bull

bull

bull














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bull

bull



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bull







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bull

bull

bull

bull

bull

bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

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SECTORIAL CllntEIUS



bull

bull

bull

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bull

bull

bull














bull

bull

bull



bull



bull







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bull

bull

bull

bull

bull

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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

L-0 ~ ~ ~ 0

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bull

bull

bull

31

bull

bull

bull














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bull

bull



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bull







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bull

bull

bull

bull

bull

bull



































35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




bull

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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




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35 Hi2-172 iJlUll



10 ~)





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WALTER T FEDEHEH




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35 Hi2-172 iJlUll



10 ~)





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WALTER T FEDEHEH




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35 Hi2-172 iJlUll



10 ~)





By

WALTER T FEDEHEH




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35 Hi2-172 iJlUll



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WALTER T FEDEHEH




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WALTER T FEDEHEH




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