M W i. f“ I x f: 4! ,1! I, N! f \ WW I W If I J (I § : _ a § f {I W w I l W A STUDY OF \I’ARIEGATION '1\‘ CHLCROPHYTUM ELATLTM Thesis for the Degree of M. S. THCHECAN STATE COLLEGE Chafies L. Weddie 3938 pfi. A STUDY OF VAEIWGAT ON I? Cl‘l'JOVOPIYTU‘jE T1. 1T3. A Thesis Submitted to the Fecultv of michigen Stcte College of Agriculture and Acclied Science in Partial Fulfillment of the Penuirenente for the Degree of Waster of Science BY Charles Leonard Veddle K LeDeTtment of Horticulture Division of fiericulture 1988 May A Z #3 k /L 7/.) THESIS Table Introduction............ Feview of Literature.... The Form end Reenahitian Types of Chimeres....... Chimeres and ficrcholoev. Chimerae enfi Cytology... Chineres and Genetics... Chimeres and Physiology. Variegetion............. Fxperimental............ Methods................. Results................. Discussion.............. suntJPTVoo0.000.000.0000. BibliOPTSDJ-‘.’. o o o c o o o o o o o 115? of Contents )1 7‘.) 7.0 O .15 .15 .17 .17 A STUDY OF CLLORQPBYTUJ TLPTUS Introfluctien The preet flejoritv of elects ere Some cherecterieticelly fievelop other veloncert of enthocvevir wirwente. V o ‘7‘ n 8779,0198 9110;; 89. L10. with little chloroohyll in certfiin Ftill other eoeciee ere leevee. dividnpls occasionellv fievoloo leaves low BTPES flevoic of chloronhvll. The 007’1‘WOT‘177 Tpcmmized FT'Q er: 1. Physiolouical. a more or lecs mottled or varieyeted urcere, r nutritional di [0 svstens. Va iecetion of this tyne is indeed caused, the narrower Sense of the ten: is veriegetion. 2. Detholoyicel. Vi ”11$ ’1 with chlorophyll doveloonvfit, (n3? .trrmrm $71!“? 375711012" Ft7fi,9tihfl, -I‘ hortions of hflrfiallv green Often the leaves qoneerence beceuee gh water table, by environmflfltfil cnnditiflns not 61909898 (‘01 ~ q *‘r 11 39 't.l( ,.L..L"1 leaves. infected TARIFGfiTIUI IN uniforfilV ETC?“- ColorS cine'to de— 5 9S 0-. .10 other LJO et- 1 ewe blunei mcv tvoicellv develon leevee their H, but in- vol- yith white 0? tvoee of vericretion of olents present Of or injured root a u l areatlv influenced, anfi oerhepe in mottled H 1‘1 "w 1 ,3 ‘_ 0 ‘ Kilt—111' 5" 3.3. l Abntilon O rith a virus w“ -2- causes a mottliny of lervee. Such infection car no trens— mittoc to heelthv olente hf yrrftinr onto them eciore of infected stoc: or by ineect crrriere of the virus. 3. Chifierel. Another tyne of veriegetefi ylent is tin? chifir121 voicfli:ie c finMJeeo.vrf'tieeuw>xfi¢ich is §WWl€- ticelly ureen enfl tiesue WLiCh is a.ueticallv vithout - ‘r r r - in ,. 7' . 'r‘ 3 ~- ‘fl ~r~ . £ 4 - clfilo;v30;czll., c:r'nlrlaer: 1?] v CPlofllS lirtdzeinie (:e went in r UJMJn K: ,7.- ‘ - '1 -. ‘ w " —. - -- . "5 . Twigs Ptinjx werxns anlJr‘.it-1'tbc Ji”ttn.”1yfb€ 01.'vrr- QidCiflfitCC. FFViPV 3? Litnfeture t3? Frau l inttwr ct to tJri renntitficfiz, the c ixfllwrist, the gnnreiologisd;, the rurnflraloeiet run} the horifixnxrtmrist Since the beninniry of the tvnntioth conturv, anc to some extent even Defore, has been the Urobleu of ChlmPTPS. A filant chimera eccircins to the ne“er eno Droeéer concept is env olent thich is c nooeefl of two or more genetically different cells, or tieenee, so combined as to form a 1 o o I S hale infiv1onel. The fiiiference in the condonente meV be little or erect, involviV3 fl difference in a single azene or'gqi EhtlTW‘(HTWYQOSQWWT(TT931PTGWH3, the litfiflz'bOinm es effinities. It follows That ‘4- 3 {Jo .Sed onlv tv :0 3'] K.) 1T- "-°. - .- . - .. ..'.(‘-.’.".. ".9 -", t ,‘.. Chinerns vhcec tiseue on»00hent9 miller little are more numerous, enfl thet thiee involvinw the greater fiifferercee hetveen heir components are nct orocuced naturellv, but are thp rssdlt of fiTfiftinx ans sfiscics or 36nd? anon southsr. 'kinklsi‘-pi‘st usefl its; teti Chiciiwaiyi Cescrfiiwiai Crfnch thst cross Mkich Vss cowsossd of the tissuas of txo fiiffsr ht 'V i. 1'.‘ .- " ‘3 ‘\ ,ru .0 “ ‘I‘ l‘ " "‘ " " ‘ ' -. ’ A Giants. $23100 timixtau.‘tsrg czs duafl LUw4l-J7 8,10T6 It- strictsd sense to infiicsts owl? the D€Tl~ spa mericlicel 0f srsftin? one varistv noon snothsr. Althouxt siait~d olsrts arc not usually 0 nsidsrod to bs csivcyas, “no“ il— fiiviéusls con?wrw ynrfnctl“ to tho VOV dsfinitioc for the term and serve to illustrsts some of the fssturss of all chimsrcs arc msny of the Uh} ologicsl problems found in a, I. an)st cifigqsrs.. .Anfl tflfl3, tine pszmxillnslq itsricljxisl amid sectorisl chimeras hsvs been kno n to arise from grrftcd nlsntg, 7“ fsct, hsvs Nash "tncucsfl sfiosriashtMllv bv Vinklcr, those work Vfis osrtlv tcsaoos bin for .— Esut's stole srfiftod two sdsciss U] chimsral hypothesis. WinVlcr '0 C‘ .', ' 1 ‘ " 1 - '2 r- ". . g ".x‘ “-11: '- ,"\ ‘ H 1 ‘L' “ ' o; QQIfiUUE RQQ Fftsr ms UUlJ“ Lao lo soc scvosac tho Pthw m “ ‘ 1 . A -‘ : --. 1 v . w "1. t -. ‘v -~ ‘ st tLe ifilgfi lELVl’f s sntll strin Ol scion tloidw. Ac- Vcttiticmuqixids LETS tknu: induced, squs of vifitnisircse QI- actlv ct tie ooint of union of stock and scion. brenctss '— , conts7nofi sows tissnss from the r J ’3 L. 0 rd 3._ i C; r—' H x '0 forging frofi 0‘ stock and some from tho scion and the srawinq ooint was interoretwd UV Saar ( 908) ss bsins a coanosite (but a [nossic or corrss iix uro Tsihor than 9 blend) of both O O a tissuss. Iarw;n tritss of grafting tvo halvss o; a hya- cinth bulb towsthcr thus ororucihr s flOTQTS Vera half one color snfi hali another. To this Ypo of comb option of tissyos gnur wave the (escriitivs n81w>trf soctorial c ststss that chifinrfil pa tfirns gst Vi‘h in VfitlT” pfrts of cleats are the “psnlt of Chi” rsl srovith hoihts, i.o. the psttern four? in tbs Hfitorc chssn is umrwt of ths batters already wressht in the gr Hing coin s. Bear's tor: 1w::;wwnnwxxith the ides of stufirihq the olfi srcft hybrio theorv uhicn has resulted from accidental ‘ \ chixermmaiflfich n5! z‘Tissn TWFYW ?“nti*H€4J‘HTtiTH{ ‘ O f ,hitsrsl hvoothcsis. DPJT'S war , Vfilfih vss «one lltfl 2 -° T’ 3 -11 n 4 r~' -° r‘ ,1 ‘3 u"! rx “fl D \‘l .7": --r 1"."1'1 '7/ r r) fww :- ,' .. T1{~""I€,-L(' -_ V'C‘f'l Q'Ca”t"‘(l 9t] (11-1“ kl}. L. {‘2 hFA ‘ .37. A 'J. glut-1:. (3 , _ \-'\ln(L that have botodc known as soctorisl morsclihs , osriclinrl find nixed cflfiivfw s shfi elucidrdywi for the f. of the ooints wkich are discussed in the fflllQVlnfl oars— ETEIHWS. The Form enC Pecosnitiou of Chineves The form which Chiceres take cedeufis noon the kind 5. enfi the Wideness of the differehces bet set the comaonent .‘ {-1 at" r‘ 1 “ ' r~ ‘ ' ‘ QOJFS soc tthT anatom_cw L-Jo t Ametowicel fiisposition of the comoonehts effects the gene- neerence and recognitisn of the 4 T) O tical behavior and the chicers. Chiuerns arise in netur, most frequently DY cistributinn i“ the indivifué somatic 3utstion, every such chenve resulting in e chimera, if the broader ownceyt holds, thaunh not alvays recogniz— “_Ja sole. These facts provide en interest n1 aspect to the problen when it is consifieted that a sometic nutrtion we? involve a change in e sinxle gene, 2 sinsle chromosome, or we? result in the doubling si a whole chromosome com- ‘olementh. iPeiut Fnui'tiwe (ifl.“X}CMUfiT¥) of Offigfillltfitufallflr affect the ehatouicel distribution of the mutant tissue. 4. - ' Plants with plastics nutetirc lets is the develoinent of the leaves uoult-Eue cherecterized by relativelv swell Pres devoic of chloroghvll, thTPDS, were the mutetion to take olece in the leaf otiflordiuw, Drecticelly the Whole of the lee? fiisht he effected; vero the chance to occur early in the Entocehy of the olent, many of the leaves would nrob- ably be affected. The first tvo tyoss voulc not be ter- Detueted since there the mutent tissue does not include 9 {ITO‘JT‘ l I... “2 point. Such late uuteting gruos are kuovn in herbitis (Iwe 1927), Polveoniun (Imni 1927). Iwei also *0 found recurrentlv outrtins streins of white—veriegetefi -5- bsrlev end of Veronica Corelline, Cessicnw, Plentssc , Lienthus, Lesoe€e7e, Pelcrccnium, Gentisrs end Helio- tronun. he clssses mutetions es suto- sné eIo-, auto- imrtetions being tlmmu>xfliich‘tske place indepencentlv of the gene connler, end exonutetions coins those vhich sre stiruletec bv other acres. Chittencen (1937) observed strains of Pelsre nine 7*nsle which snort c to white ‘brenclwns brutet33n1 eFrl“'i11 antopwvur) vitllgrreen tixncs on the stinules (reversion occurins lete in develoonent). Grafted nlsnts anc those chiperfis resulting froa lste occurring or recurrent mutcticns being excluded, there ere three tvoes of chiweres with vervirs abilities for cer- netuetinx thews~lves, viz. the sect riel, the reieclinsl, , end the nericlinnl. 7 C‘ 1 .; u L1. 1/ U) l” (n The sectoriel chimerr is one in which the two t’ are united si€e by side in the stew end the sroviny noint of the main stem or of e trench. A section of such 8 stew, Here it nossihle to stain the two tissues differentially, would soccer as a circle with one sector differinm from the other as 8 Die in vhich one or more cuts had been recieced with another kinf. In the branch in which it occurs, such an arrangement msf be comesrstivel= steule, but the lateral brcnches which cries will have the constitution of one or the other of the components. Onlv those hues erisins st the junction of the tvo conoonents rill have a chiuerel -7- structure, ehfl thev will very denenfiing deem the proscrtion of each tissue enterihs ir. In blasts having severe sec- tors of M‘ut nt t1 ssues, hfiwever, the majority of lateral branches hey have the sectorial erreniement, but Very sel- dom will the exact ccmoesition of the original stem be r:- on it hes been Pssuued that in I.Jo tested. In this 6iscuss sectoriels the mutant comacnent canerises part, but not all, of each Heristem t1 0 lever. Occasiorslly it hseoens thet in sedition t) the sectoriel structure one Oi the ca loonent tiLssues will hrve a thin strip overlenpins the region of the other tissue. Such en errenqenent, as will be seen later, often gives rise to the periclinal form. The sectnriel tyne of chimera is then cuite unstable with- n its elf end in succeed: he vegetativelv Dropageted gener— 1—10 atians. The mericlinel tvne sense 8 much as the sects -rial, the difference be ir; that the differ ring sect>r 6-We net ex— tend all the vev tn the center. The differing tissue in this tyoe consists of a skin one or more cell leyers deep, which does net reach the center of the sten, end extends only part of the wav srounfi. The mericlinsl tyne beheves in J grovth end ves tetive prouefietinn in the seas msnnrr as does the secteriel, erceyt thet brsr ches arising on the side of the stem which is overlaid vitl the sucerficiel strip of eberrent tissue, till give rise to the Deri cliual structure. Unless the striu cverlving the parent tissue -8- is severel cell levers thick, brenches pure for that type of tissue conteineu in the strip will sel on arise, the ex- plsnetion being the ot deeper lvln tissue will as drawn uo, fOTWiNH the 'core' of the Drench rrszlti “V: in e eerlclinel errsrmNWQGHt- a he oericlinel chioers is like the mericlinel, exceot thet the skin of one tyne comnlstelv surroun s end envelope e core of the other. As a rule no chen e of csttern occurs in the oroduction of lateral shoots; therefore, the peri- clinal is the nest StflJlG tvte o: chimera, since, es has been pointed out, the oth er tuo tvoes tend to resolve them- selves into bran ches sure for the c moonent tissues, or into the oericlinel srrsn elent The literature is reulete with examples of the verious tvoes of chimeras. Nenv snecies have been involved. The majoritv of instances he ve de it with “ripest d slants, obviogslv because of the ease of detecting a chimerel er- rsneenent with the green er 6 white oettern. It is now suesested (Jones 1937) that the: 1ericl ine.l tvoe arises most frecuently in neture and that many chiheres described in the liter eture es sectorisls were woDnolv in reality nericlinel arreneements. Peri— and mericlinel chiweres, eccordine to fieur's hynothesis, ere mede nossible by the fact that engiosperns grow by (livision of cells in two or gore sel‘—0er)etuet1_e veristem"tic reg; ns errensed in concentric rings or layers in the growing stem. Such an errengenent, it seens to this writer, precludes much of the nossihilitv of true secterisls being formed neturellfi. Ferns sror iron e single enicel cell and thereftre the only clinerel tvoe tossiole to them is thet in which somatic mutation occurs during develop- ment. Such cases have been known in both sporophytic end genetoohytic senere ions. (An6ersen-Kotto 192?). Greno- sperms grow from single meristenstic resions. Although there hes been much controversy concerning the point of self-oeroetuetine neristefietic levers being comoosed of different Vines of tissue, the existence of steels nericlinel chimerss with one—, t 0—, end three- leyered skins hes confirmed the existence of such arrange- ments, (Krenke 1933). It must he kept in mind thet e groving noint censistins, for example, of two levers of ootentially white tissue over 9 core of green tissue, will not renrofiuce exactly that arrangement in the leeves. And therein lies one of the mas difficult eroblems desling with chimeras, i.e. the nrohlem of essignirs the oriein of a nerticuler tissue to a certain lever of yeristen. Another problem of inportence is that concerning the number of meristens in the grovinq points of Olents. Not all chimeras are eesilv recognizes and enelvzed, as those the differences of Whose tissue cannonents are con- neretivelv greet. Ability to provuce chlorophyll is easy enough to detect in the foliage, bu 'mnossible in the'meri- stems. Henv geneticel characters such as those concerned -11")- with floral fort end color exores. \ 1 thenurlvos over a csfiphretivelv short ueriod of the life cycle and in a relatively swell oortioo J? the olsht's er tony. Such 'JI consi Ge: fit on U) (D ' I: ....J J aissize the fact thet chinerrs are olite likely much wore abundant than is reolizeo. Chi eres 9nd kao1hvlovv The study of CFj eros has vrovco nsef.;l in arnv dif- ferent lines of ifiVFPtiHPthW. The orolol“*ist hes profited inch from tho stufir of chircrss, the most V?lfl- shle in OEIBtiJ H31i.s a khovlroao of the ohrticuler lever of the floristefi concerned With the formation of .he different tissues in the mature olent. Thet the ceroels end hence tho seed in shciosoerns ere oroduced by the sub- eoidermal lever of meristew vss thought before chigeros vere understood. Feed orofiuction in many chimerul species has, however, confir1er this feet. Pelstion of diff errnt mature ls of tissues to < eiih ite reuions of the growinr ooint hes been difficult. Jones (1937) hes comoetehtlv trested this problem of chiueres in relation to morn} 3107". Son? of the more pertihent points of his discussion ore Riven here. The shoot son? 0 msists of an inner core trich con— tributes most to the growth in volume, covered by one or :norc lxrverw:nlf cellxz’“hici1;fiarc tflr> tnrinJ, erwfli lsyefl?)jooricl1hel (fl 11erss 111_Pslor1~vrwv1, treosvrnfiw: sec— tixfinct oi“ 31w :>otix>l€%: CM? vfiricli si131cwi jilst suicli e 1'ecw‘e- Si‘HY‘PC? 9137.“? D (‘10 1319,9119. TY} thPCP graph—var-\-.rljite pfifi' 1"i'JltP- .L. over-groom chimerts this str’o stares out in the contrefitinfi FI’PQT‘. Ffifi V431“? ti $337119. 0 L; o tho 1 ('7‘. r" f le1i“8 star, tioh of the —. :orcium there My orerhisetinn orfirwvrci1vi, 99""!7FICHQ hirrfva 11151111 1 7-1, +‘ \ . I tvO bitiorrvtietf born 59 tit see s, eccoru 1.1? 12:6 st :1 (j '1 i”1‘1 )J ' Of the (‘7 1ST?” 73t1")1:‘r(_§‘n , Cprt" C: J \r csse in h w :1 1 Phi" orocotsiefions. VGTK hv Pott To . r 1 4 H‘ - fi “1 ‘- ‘4" y w) . 1h91 Iixriwt u re 1— fi‘L-V- ‘ :‘ -* ‘ s -14» too toner vor- h doveloorpnt in: to Jones to he otic levers t; ,. r—I Ti 11 “a! ‘., ' 1 - -_- 3. “'11 4.03.- r134. u 'VlelC’nF} .’ ’5 ‘ ‘- is? Y'”‘ iouh1 ,nis tzw> ster (19- this with his Ftufiv o9 Pho9790o5rnh. Chittanden (19?5 so: 1927) esein confirms these T hmihws h? c stud* of t>o chirfrhss, f3r1s1111hi131 "Fiwwr: oi‘-iet1vwe" r11? Ehflirervreo 1/3?- te“sis hivslisu 131 Pelrr7nri1r1'93ror: of Evture" the stets, notioles, UGC%“Cl¢P Fro cehtvrs of the leevws ere ours vhite vith stimules, leaves snfl cslvx hevi.' ereon rqnjurinsr, iicdta tip; rqlb-Jzoi(iejris], lfWTPT‘ rtnio'nisiiflc ‘POI‘ tifé UTGOW is the merginel monqnhvll of the leeves, 8 sub- ljrrer ” V -11 » _. eoidermel 01 centers of the loevcs it ix: coruvifieTW(3 thr wreoh col 1 -5 unul ‘2 1.1. th~= AJ " EET¢lFfil mesoohvll in inc vsrintv of Nature" is perfectlv er°leeous M varietv of Chloroohvtum elstum StHC l T‘. 1 I O 23 The ovo Granules 1‘ ll (9 v In 1 L Prsihel moristcn TT‘IO KA)YI.j. t F‘- S , " o o o o t 10.6? O are rot C3nS; .. 1 .‘o (“If .0 7‘ ‘77 t 131:5" CT“. N w!- «7 (n 1.1; ( ehfi over the he ref-1.1 r1: (3 «\J is rosnohsiol for the of Pelsrwon "Fresh the medioeloihets in the present york. "fi (‘ixfi s \ 913151,? PT“.- i V T 3 f“. F? t O hr-Jires, ormelly Tl -15.. (JV-2 C _ves rise only to the eoidernis of the mature nlsnt...." Imsi mskos the eencrelizetion that nost, but not all, dicotyledons arise from three meristenstic zones end thct all monocotyledons have only t“o. he fices not consider that dicotylefions we? heve more then three. It is doubt- ful, however, that vith all the enormous fliversitv and versatilitv in the nlsnt hirgcom any such broad general- izetions csn ho mede. Chimerss end Cytology Cytology hes made 8 contribution to the stuoy of chimeres end is receiving in return answers to some nrob- lows. Cytological studv contributed much to the analysis of Winkler's Solenum chimeras. Chronosamel chimeras in creois have been stufiied by Hollirgshesd (1928), and in tobacco by Kostov (1930). The behevior of plestids in the inheritance end development of variegated olents has been a source of infornetion to the cvtolosist concerning these seeminelv inconsistent cell inclusions. hhite tissue of variegated olents .ontsin plestid differing ecnerently only in their ability to nrofluce chlorophyll. The cause of the inability of plastics to produce chloroohyll and their nronegetion and transmission hes been the concern of cvtolosists. Three theories concerning reesons for this inability of olestids heve been edvsnceo, vix., (a) an in- hibitory action of the cvtonlssm of one nsrent on the olestids of the other, (b) ineh°litv within the nlsstid H0 tself, end (0) nucleer (genetic) factors controlling chloroohvll nroduction, these usuellv beine simple re- cessives. If both tvpes of nlestids are found in the same cell, Yeshui (19?9), it is considered thet the sec- ond is the csse, whereas, if white end green plestids are never found together, the first exnlenetion is edvenced. The severel instences in which vericaeted nlents have srisen from crosses of normel green verieties or snecies, Clsusen (1929), Harland (19? ), Love and reis, Delgreen es revieved by hittenden (1927), are accounted for bv the _irst exnlenetion. Share thinks that the difference in green and white nlesti s is entirely develonwrntel, not in- herent in the nlestid itself, but C”nditioned by "some in- visible differentietion" on "one tvne of initiel bva being carried out in the cytonlesm. He does not mention the nocsibilitv of ereen end white plestids occurring in the same cell. This Writer finds only one direct mention, Veshui (1929), of such 9 condition, though Imei (1937) stronslv nostuletes it. Baur, es reviewed by Jones (1934), seems not to doubt that such a condition exists, although cvtolonicel nroof is imnossible, due to the difficulty of d'stlnsuishins between undeveloped chloronlssts end chon- driosones or leuconlests. Cvtoloeicel evidence of UlestiCs nessine into the egg along with the mole gamete is meagre, Correns (1928). Such F is indicetcd, however, by bi-psrentel, non-.endel en in- i4. heritrnce of veriweetion. The fer instances of such in- heritance tend to incicste thet in some nlents nessese of peternsl cytoolesn vith nlsstifi prinorcie into the egg occurs; but in the nejority of cssrs, es inoiceteé by the greater number of ceses of surely meternel inheritence of chlorophyll, this does not occur. The releticnshin between cytological orocesses end the orobleu of chiwcres end veriesction is thus suite eponrent. Chinnrcg ens Genetics It is the field of genetics primarily which hes con- cerned itself vith the oroblem of blent chimeres. This is exoliceble by the acts that chi eras arise from soretic mutations eni thet inheritrncc in chincres is often non- Ecndelien. An unceretrnfiine of the chimercl hvnothesis has clarified many problems concerniru bun sristions end sometirzrmrtetions, e.§u Cflxmisen ens Goocsfinmai eccoint for' the necterine soortins to neech vith the hypothesis that the nectsrine is a chimera. Theoreticslly, necterine, the recessive candition, roulfi not be likely to nutcte to beech, the dominsnt c nfiition, but here the necterine e monochlewydius chincrs tith 9 core of peach and a shin Vith tie factor for the smooth skin, the beach core woulc be exoected to break through occasionally reverting to peach, such reversion ectuslly occurring ether frequently. Since each somatic mutztion results in the plant be- -16- coming 8 chi msra, tb8 prool8ms of Jud vnriation 8nd of cni18r88 8T8 clonal? 8.88818 t7fi. Th8 ch8nC8 of 8 swmstic nut8tion LQCQ in; known at once is necessarily siinht, 108 the ch8n88 m8v occur 8nywh8ro in ths d8enly 18in U) Me ‘* tissu8 of the 018nt, or 8Ti8i.8 in tile snide? is of the nlant 8: 8V n8ro8tu8t8 its8lf for manv vngotctiV8lV propa- ?8t d gHPpr11‘te1ru]_ parent enui thrijc joint cxnrtrol with those includefl in the egg Cell of Ceveloo1ent of chloroohvll in the progeny. One of the connonly cited exemoles of the above (Ikeno 1917) proved to be en infec— tious chlorosis. Jones (1934) cites Collins (l9?$) as fiy‘ 1c ing founc nun-‘€.d.11cn Ql-OrVP“tFl inheritance in Chlorophvtum eletum, the variesptefi olent with which this study deals. Peoeetrd examination of the work, hovever, fails to disclose env such stateuent by Collins, or any grouncs for such interpretation bV Jones or Chittenoen, who also cites Collin's Clloroohvtun as an exsmole of this a tyne of 3 c4 Ht. (1 inheritance. This trite? has not hen 0p- nortunity to cxsuine the orieinsl work of the other rambles, Beur (1998) and Fevner (1924), as citec by Jones sno Chittcnc_e n of this type of inheritsnce. Vsriegetion sometimes is founfi in the offswrinc of -19- soecies crosses. Chittenfien essuwes this the re811t of the nlestics of one species not being functional in the cvto- olesm of the other. Tt is not cleer, however, vhv counlementerv Tectors for chlorophyll fieficioncv, one located in eech of the snecies, end both heinf necessgry l '- 1 1 far eyonpgslon, coulfl not he resdonsi11e in this case. ‘ I Herlsno, workinfi v1th two soocics of cotton, foun( Couhle recessives for chlorouhvll ceficient. ho cless of chiueres other then the vsriegeted slants orizec for their ornamentrl value, end bus sports, end a few others such as Cvtiesus A 211 enfi the Creteego-Iesoili, cultivated es onoities, hes es vet echieved econo 10 im— 1ortence. I‘msi states that 01.31t1'_v:atlc‘>n of ‘g1ericlinsl Torus of ?elerffinium be:eme s foo in Jeosn end thet nlsnts of neueo ericti hove sold for verv high orices. Joreensen "D D) and Crene (1927), however, have tried usin: Vinkler's grsft— . - ins methods 0 nroduce a noteto Tlth oer_clinsl skin of .. I t0m€t0 vhich would be resistant to Phytophthors. Such is theoreticsllv possible, but Jorgonsen failed to get the right combination. Asseeve (1927), hovever, found that several varieties of potatoes in cultivetion mere periclinal chimeras, both tissues in sooe cases being of well Pnovn varieties, enc that bud soorts in ootetoes Vere often the result (r9 someticz:reorren,vmumrt of ctfiwmeres. -20.. EXPERIMTNTAL TThtvahictirni . - Species and verieties of Chlorouhytud long have been grown and efinirefi by gerdehers end florists. One suecies, C. conosuu, Home (1873), we, ed ired enfi given literrrv -0 I =nc by Goethe in 1888, who wrote e letter con- (D - 1 sierific he this olent in which he praised its beeuty. The Ip—Jo 08 H n most Donuler of the chloroohvtums are the veriegeteo forms of C. eletuu ené C. cowneuq, with the former of vhich this pener deels. Two forms of eech of these suecies are ec- n hired for their ettrective green eno white leaves end their ereceful florerirg seepes. Of the two types, both of which are reoresented in eech snecies, one (the'medio-elhihete verietv) hes the White tissue in banfis dovr the certers of the leaves end the leeves of the other (the elbo-merqinete verietv) have White vergins, he green tissue being in the center of the leaf in this case. They are considereé, in fact, to be reciprocel forms of oericlinel chimores. The medio-elhinete form of C. eletum ves studied in this vork. Collins (19??) stufiied both forms of both species of this eleht enetomicellv end geneticvllv, determining their structural nature and finding that the chloroohyll produc- tion is maternally inheritcfi, inveriehlv, no metter ‘"host the source of the pollen, reprofiucing in the seedling the type of tissue which bore the ovule. The VQUPH censules when borne on norfiel tissue ere ereen and when borne on chloronhvlless tissue ere white thus urVine it oossihle to ’ deternine which tyne of tissue bore the ovule. The litersture fieeline with veriessted nlents recog— nizes the feet thet there is considerable variation in th; 0 -~. nrooortions of green end vhite in the variessted tissue. Aposrertl , however, little attention has been neid to this ssoect of the subject Vhlfih is e wetter of both scadexic 1 end precticel interest. It was therefore decided to mere a study of both the nuentitstive reletianshios of the tvo ) kinfis of tissue found in the leaves of a veriegeted slant and of the Derflnnech of such Quantitative differences from one vegetetiVe severation to another. sceous plant belonging to the I." Chlorouinftui is 8 lil tribe Asnhodelese. The soecies forus are sure green, a origin of the variegatCE tissue Define unknown. Smell vhite v‘) are borne in the siring on sleLfier terete sceses, 1...]. 0“"??? I *hich after the floverins sesson nroduce vegetstive shoots thet unéor moist cunfitions produce roots. Wet besel crovns fievelon, thus furnishin: two types of metoeiel for venetetive nronegetion. This Writer has noted towers the enfi of the flowerins seesen small Vegctstive shoots near t U) the ever of the scene erisine from Vffit enoeered at fir to be a flower, i.e. one or two petals or revels ens ner- Lens 8 stenen being nroduced at the noint thich res uro- ducing e vegetstive shoot. Thus the plsnt in some instences soneeiw:ix3'be viviosrfixn1s. Aftei'ifivs floveivzsnwaimroduced 1 the old crowns grefiuellv die 6orn end are reele‘ed UV new ones. In the nreen forms the scede is green, scsae ler**s scewo borne s “ots, ens voun; cs silos hairy {rse' elso. In the p.C. (White center) Taro the scs:e is Vhite, t1: scene leeves she the vesrtntivc shoots borne on the sccne 3' 9 k4 '— A (j thite. rm of the youn: flovu As observed Collins the viite oort ’olcnts often be00“‘ cornerstivel" 1'_‘T'C)C-h . ixw every =>L0003 1*7 9 9HP’ ’ L.us. n the orciceriLv viite w: J__| tissue " i ,1 cclorowqyi tisnvre then FT”??? to lw=*"fiet lfirw‘strow " 'virescert elhiro'. Collins nostuleted e hioito”7 ffctor “resert in tie thite tissu ‘ with ere. In View of the feet thfit ffiunf e Freeninv o"0 the vtite tissue, it woulc essnme absence of e fector to orocuce chlo tense light, but oresence of soother Vhich (tiffuse liifht. lfi7VTstrow (lfihWi) founfl ‘Jrocuctifvirif chlorothll its? ceotion o9 frctdrs for chlorooTVll oro t?“ vs the white center orttcrn of the loeveo, (In, (W ‘3 8 chtors ens the 1:) ‘7 9 I"? inns of older ‘Yowcf’*lL°utP, develoa to Plc() The Vilte (‘ I 4) Cells bleechiug 9 ‘fl? ;<:?; TY?HZIC?‘9(1 olruts elso show net better to rowtrll in ir- oroduces it in for the 5:1ct5;>r 7. cthIWVHflrtiw en cwwvr ‘ettVdn 'Phe eccV(\ *‘mwthvsirri¢"mirre out, too, hr the fact that rorvel sreeo tiosuo in intense liaht is roticeculv lirhter is 0113? then ofrtiOl shcfle. Plchts stoninw in e Poolicfi P- ‘" 7' r‘ . 5 O . 1 _ 7'. I _ orlcrter «CV9 me n, 0310? until l“fi* 1;,- “ p. pm 177311 5‘ o \" vfhflT“7 in vith ro fEN“ Fhffir?ih“tfllfifi tiwfi‘r vazltkirzzreeu “eh tflwetr color becin¢ tn frfie. filblno sn¢fil3hjc bathe by t.C. glmnts SU”VlVF-]JNWQ°T if HTJVT’lll‘th“ Shafe 3d? Evafi apvegr to be deWNiLDUirW’ C“MJ)“Otfifirll. It i}? tax) 9972:? fict, 'thmugij, to Do surn thct thov r‘ll PHTVlV‘. Tho fallHVlhr nognrvathWS v“ ttn gbhetvl Ffiatcnf of t“? 1??? of ttn V.C. DY tlli r: mfi‘. c+ ”D (lu9334 (V9115? lfi ixsszvr 52M? lcwwgr <31?fir:fi<fi¢: C‘HJfi V‘“ Brews WPWD Tmuhfi, PS VCT’ tfi“99 0? the evifier”ifl Cf the *Nitfi floveriv: shaft, to cantsih chloronlfivts. Purim: the Cuurae of ttc exjnriWfirt meets, tJtnling near a hunflrcé ‘CT? mcr1fnrtni, all of vtich wrofluced el- bivo seedlings. A f?" ¢NP53 were gatherafi frfim t0,vlly green DlFUtS, but for 93Nn rragin none gortinatpd. N0 pure white or mute {roan shunts or eaves mare voted. A wa'lpBVPF, hoxevnr, raw: “mtcd in Flieh he vhltn tiara? evtbfifiefi to the edfie of the leaf, 1.9. the ETWNH‘ edgxa\a99 tin; cont:rntdus. Collihs, as if he floutts the hywotlefiim, says little 0 ncernlnv thn chineral nntnre of his chloronbvtnn, but tlwthizps fid;IlPhfi ftffiyflut E 'Zfljrthnlc Farintic Stuptnsttinn'. ‘ .1 .2 . a r1 F 3 . “-21 ° .- .' ' ‘ 1- .. ., '- In swearlnfi Jl «Lawzt'n rnh~1n Piflhnl”lfl*b, a? sows, "in it thr;st}-e?iderw¢l lfTUYF is ETPPD inn; the 9010€T41S is albihotic. T11? HJUPFTQ ftow the fact tlfit the "uvrd I .. CD —. - 1‘ F‘ - v‘ - ‘7. I" -‘ ‘ 1‘ - '1 1 -' r‘ ,"‘ I}- ~— - "a T " coLls *bfltgwr tr Mn fr)? EU,XH the .nlt~ J? from rtswu tn .1 ET??? DSTtR of ttw 10?? are GGVfii3 nf Chlo 00L ll. tr? 0 ¢Ttr¢7“’, lv: fJLuv‘ tlw t if? t13* ffn‘é Ii th ifta Iflitc# C€vtn*r-- * fill tho cwofilinnc fiicd. O ‘ 'Q C Q . v ' - F. Pzeooloiene vor. HOClQ-VPTlOWFt7—-thfl guerc cells ere eETF‘”7 illfitlllr 1x215“? frmx1 t}fi’ PTV¢FN (3n tile VflgltE-7J?IWLS. Eenoe in C neioerction of the condition of the epic ‘ 0 ‘. .v .- ...lo .._ ,x «- ‘.- , - It tee tn}? ooeervetWAn Ilnoh led re ta «*0 ]N( the JAFTC Celle in deteii, enfl the investirrt7in if tie lfevee of 7‘ " ‘ ‘ ‘" 1 ' .».. 1w _: 1 ~ "- ’ ‘ " l" 'v. — ‘ v“ A, \ ‘ 1 CHltU"T F“t:':i we Finn 1‘tuat 2J1 fill Ef:tflhtHKL .“F h7ktb (D Lor— 51 (37)}?1f1311“71 51cf7*(:(W€> I‘i.t;i1 ED117715i_?a. " Leter, Envever, Collineth“te the at to «at tint, "In Chlo*ouh?tue the leevve D“l¢lh€ e as euoeoeeivo callers ouebeo of? es it yrvc erm U : nfl.ittin° flnuv woe ~wnietrvet7o 79T”?. After 019- O O a . cuneinfi 0e01l1er trove of venicmet on prtterne in TioCes- C‘thfl, Phineo'ie fine Teotvlie be eeve "Ho Feuht in the VBTIPJHtCH IS? F or Chlonomfivtnw i , 0r§9‘“flte the ordenl” seine etion a: to clo:r;tic Cmfldltlfl? to the Centere or cdwee J? the loevve h:vln: regejfl to leaf \riain ie not Clearlv evident." It is thue ODViduR that Collins {see not think of the €ITlFuifi13K5? fVWYrie <3i‘ CT leifiiolnxter ;=e 17e113n'5inr~ tC) tzie <30:2ior1 : «OJ-1 tVDe of‘rwxniclinfil vffiuue crwwonont ticriuwa omnwrlee 01f- ferert eelf—oeroetueting meristems. It, h reven, toes not seem PfWN33fi??V t ajwvetiletr €337 rhvttrfixz eevreretfiyni of green end albinotio celle Two» e wiiee premium fioint com- ”I eiderfix~x in tflwa ceee rc"the rfjiwé oenterwwi glant, tfiié.2teen "\ - ‘J 5- Quert cells over tlv white centrel 9T“%9 of the leef, fine - —t~ inheritance of the cherectene J lonaing to the sub-epidermal lever. Colline, too, wont? he inrovmler Olimenfis from eeee Donne on rteen qtninee flovetinw eoeflee wllon (owviwuelY beina eeotoriele «U?*vwfiiolinfle) teellved tnnmcelves into t're etehle oeriolinel enreneenent. ;1efiio—ver1erete fifn~we a? C lorookftuw elrtun ene C. oovoenw 'to 3e elfinle, rflrfijte, “etiolinal mo“CW%OIWfiWltJue (hevinn 1 O O I -. one-levered lehg) 0h1*eree. the lrevet, 1t voult seem, ere forweC, re i9 Thane to be the C??? 1“With. 3*» Home ere Certfi" .— D monoeo.vleopne b" Tenner rnf Pottivv, en Citer v? Jones (1937), ”no in Pel+nnwvvhrn “Preek of Nature" enG :yfit”n*ee Firteneie rivelis b" Ctitteneen (lcfo), tram the two outer- moet levere of the tunioe. Theee To? ere founfi in these Jo olente thet tle fiernetoven peve rise t3 5?'€#T¢ls end by \v t. 'oericli74fil fiiviei;w1'tn tlwa'nantlnel VWHMTthll of'ttve.loevee. ‘ Thus the oneen dermetouen in Chlonontytum eLetum medio- elbinete otofluoee the "?een-eofiee to the leaves. Thet the Pecond lever of he.fiorietevetio tunicr 18 not Concerned in the for :tion of the Herriral neeoshvll is shown by the feet that seefi borne bv the olent ere eldinotio, the 915- eniéetnel lever of the tunice nevinw been proved by Beur (1909), es cited bv beteson (19:0), to be reedoneible for the oroeuction of the C? pollen? leovee. Thet tie freen H tiswnxe is: onlfir DFF? celJ_ lEjW¥P tlfimfi i1: tlns tLPllCP% is :Jro\ued bv the ff: ct t}"-_:-‘t chlonon nice (' L‘Ll ”A - ‘l A --. - - lever? o’ twe geeofilvll Confited to the puere cells 0 of the flO”erinx olent. Coll l _ albinoc cells ea trrvellinr 0 1n 1 o i 19 “”0fluobé in eeverel tT’l in the P“? i? fwonh oLcfit eWirerxie in the scene f the ' -1 . - ‘0 I“'S theorv of en '510 ”1 Ppflfivfifitlufl' rill trtce rat 9 e irel benfl oeeeinm "tour: tb‘ rot? in o o '1 ' , ' .J . — . a v -~V . ~ ‘1 correletion Vita the s,irtl if er: o~if’c" oteelo.3tra fzwwr t") JTO white olerte. Such plants, lptrrol etootfi eible thet the? too correl I‘L Soirel arrenwe-ent 0f the FT” ti In the cwurre :f $39110. nlente veto fiecewiteteo et re whether efventitioae 0” “7t, ment of the white-centened a? 1 ‘ Emir C‘T‘p: :51 |( eoifiermis of‘the:fl3W, it ‘IOIfiLd INTve tween 13Tofnlcefiifrfir1 reeion. Another anon? of two nlent, ceulfi vWite etoete from root cuttin O 1_ Wlty no UCCPSR. Another wheee of this et ijvvpetiartirwr‘tnt;(InigtfiflCV internediete tieeues in lines Tn thp 3* the elem-fir (men I ‘ ' l . fiuce t? ((3 “r- if? xlf notentifflfljrtfnite tise? CS tbet see it be eccongliei MC“ in Evrnlnte Ell* vet totolly yrfifiT een Wentimnefl, never rillhrfi in origin it is I“ (I ete rith Collin'e snoooeed teeetinr eloinotic or green "- 0" C T" :Ve‘t' 1 W (21 ‘.'W m ‘.- - :1 IJ ‘_V-\ ‘_.T_'_ -.( .‘ I, .1-("- L, ](1J , ‘q V‘Jt’oL . (in ' em" fill luv—LS fOT‘JJti, WVOBUCGd the tV:icel errnnfle- ¢WHW leevaxh ]pad tiygm. bpen Le r~ec¥7nw tn the 'e white shoots .QVN uerntiterJe bufie in tlfisa ‘Uerfi cliflwel et*n1ct1rre (If tfiie eC, \Wfiulé Lm‘1rm)?ucti'n1€WT leszim99 ettaevvtec bnt @9- c.¢‘crrrnxaé my white end ereen , ijrneyntmui eecoflleleu GfiIVWWGT, w v"?lrtv. O 7'“- YW" {3 l i ; ( )I‘ Wflrflny Pits ?D19?3*h191 "I e. Srlfircl," 5n Plix}~n%riin vsriptv, (unoanliQFPfl rosxlts) haw been :70].8 to s¢dfiret “t0 c. C. D x.- rmrt Ftrzirwx1x71yW3puco a rtrwfitl . v1" . ' '-"'. "‘ ‘ ru: . 5 1 ~ 1 ' '5‘" 8 “ "" o ‘— lTl VLKLCT1 tfh, Y; Ltf‘_xfll“rn an: 1H19 ix 91 :19 \f0,- qgufruuv(7nu RPOtheT in which the flfir?ifi is gfiiTLV ride cnmnvfi:§i“c ntqfivwws hrlf'iflw= lesfk flpnrts ( “the i§11tr‘$t?fi El) .. “ ~ . -. I‘ ‘ gpggtlc rearrern" newt? in thich pitho“ thc firmed or the vhitn tisgue is ted) are Cénpfirfitivelv frequent. Such htudias hrvo thnmrrtiacl vslue, Rihce in the ‘3 O '1 .l honed (lSPR?G rc¢nstrnt Chjfiprsl notstwos O omit— (19?8), it will h“ neceqsnrv to “¢?wt?*n Rn owtimuw death to the fifin of tfimrto tEPan. To be cert'i n of rcsictcnce, the ski" W‘ll V006 to bn tvo O? MOTH cell iffvre thick the lerwv, uu wot thic“ onnu~h in tho St?" to irterfe Vith tubgr “Owwrt?fin. thfiio9 of tho {haurts of tisgue 1V37*u€w3 iyv' tEEC: 9"ir7 (znigflrvn63h139. itw \19J7i52?rvtr d . ) _, o '_ s ' V“ q -_.|_ :i. 't‘ F , .1. .1. large (nouch UHficers of nlnnts arc used, Shoulé indioat nufiher 0? lFV0TF of flwriPtCW concerned in the fawnwtian I .l‘J. leeves in a sneoipa or vmriotv. If w thin Qf tho eramhta of Ski“ tissue v—J. CD :3 d H. J , 3 \ aintrn‘ 1 Inadeé, tiv= intm=rp1VTtflti n1 Tlfii¢d 130 t}urt ti“) Ineristewlxwana inVOIVFfi 531'the forw'ticnl WT 7fionochlfivwwttus fort? twfifiwr presert; 11‘53 oi tion inflioatirf UVfiHONCO 0* bath wmno— find 2 v rietfi th {)1 ...- h .93 (D) (a .013 17 .31. F) layers of th the loavvfi, -gnfia] (Fifit? '\ w- r) 5...: y- I... "5 C 5.10 CD in .7"? e the of x} 9. only ibu- 98- forms ver; obtainpd three lovers of unriste; coulc be as- swned to be concerned. "Let hf) 6* S In JRWURTV 19?? thirty-four plants of Chlorowhytum which aonerentlv varied consicerahlv in Pnount of fireen vere selectefi from those NTOVinp in the hOTtiCultural preem- house at Past Ldrsina. four of thpse nlsnts W??? the normal green tYUP of C. elfitum, two rare C. conosun var. medio- clbinntn, and tvgntv-pinht Fern of C. oletun variptV mfidifl- q 3 nlhinnta. Latcr one nl‘nt was obscrved awonq those not selectnfi uhich vns of the alho-nnrglnatr tvoe. Heasureionts in Willirmtcrs of each tissue t’T‘JG in each loaf of suf- ficient size hero made in all oanJS. The measurements were racorflefi as an. of grppn, mm. of thitP, and mm. of intermediate tlpSUQ intermpfiiete tisruw, as inflicctcu by a lighter shade of prcnn bninfl that in which grccn did not connoso the entire thickness of the leaf. The plants were thpn propagatofl p? rpnifily PR noggihle 00th from division of thr croums and fro? the vraotfitive shoots nroCuceé on the scoops. Th9 nnrent plants vvre numhrrpd from 1 to 34 in no nPrticuler orfler With T9f970“09 to congrent vidtn of tho creer and white Stfiins. The pronunv regultina from crown divigions verc fieeignntnfl 99 1A, 1B, 2A, 25, ntc.,' thg numhor indicating the oarpnt Ulflht and thn cavitpl lgttnr Slfinifylh? 9 cro n nivisiun. Progeny resultinx fl'l fron scene offshoots "ere r"esi{nip.ted siwilerlv, e swell letter heir» used ti inoicete scape orivin. It has thought et first to cxrnwire enuunts of‘awwwnw'producefl Ln’cNfieh tvoe of shoot, but insufficient runners of olents oreventvd tfilS commerison. r). Considereble difficultv wee hen rt iirst in determin— ing the amounts of white end interrediete tissue since they in UOSt ceses mere somewhet interiixew. After severel trials, Lovevc”, it use found the tuicfest end most eccurete method MRS first to fieter*ire the c uhined ridth of the green end intermefiiate oortiors eno with the ruler still held in olece to estififitc es closely as nossiole the vidth of each of the thite end interreuiete strins. This mes fnunfl or connerison vith results obteired by seeerate, verr cerefid_rteesure1eqn3t. '9 WC, ' O 1'” l," O 175. 177. 138. Phrwnel, _A Citrn in the 70 n. Phnl1 G. 147-151 Cit? 1 . 19? 613, :39 997. 7:1;01 not. G 3116. 01;? Q—TTHTt - - . -46- o T 1101121“ 7.). ° 3 3'._3.'j,_*.,r)3~.)vp 11 SbOT‘I 1 9"3I3h . 1990. o .Lhtrk)(‘1lct n-: YOTK. J. fr. B81U‘ 1911. '7!” . (7 v r (\ r- yul. nvn.19-¢ CT‘GPt‘C “ Eu (Err Ft 8: 775-91. 0 I O ’42- 22g1T’1-1-2 , ' JOH‘. CG“( 1.. all \1 M 919150? [TIA-T: '7’: ”(I . ‘. 0 ..._LI_‘ 1. PGaTTC? V. 535*: Cn1"T"(I"(-3 C 5:?71-971. 19 Cu. ’01! 07,0731: Cr, :31) D. 11=’)T‘-7Ii 1919. 1 3‘19. (m Ifiir. d. 33:“ v 3'_ rW'pr: F‘CLQUIEW _€tw UNL ,- ‘3" l_ [- LJQO, ('0, ')§)'). ttenfien.) '9116. 'Th0 9 C 3T. . tifififhfi'- , N1 01‘13. :(-T3T'. Uon L7. 31?} t .’_) (2 TI V F“ 1‘1 1" 1 nivorc 19:0. '2 . n- ,-\ 5’35 {a (-3 I \f I- _'_ .L ‘ 1 1927. _cvwan, 3“"; ‘) r-T. .3 164, r1331 Pb‘I .LVL - O IZM). r- 911.9 FET‘J :71 :13 .— n. I“ 1948. Inn I“ <2 .1 I" i )0,” (1:3 - Cn'tcfiLOari. in 99 L (19111-1 81311. 1337 "31Pnt qcn “tjflh. T" h I 1 F11” 81:10. 2.1111." .- z. b. ( _I. 5?. P1P? -- W\T of 19? A., r:?1 191.211-. CRT1FDPTJ 150T. TQVTPVVC bufi :911.. t d . 11999 ”-31“““Tb"0tr“ve unfl 9.1961- n ‘JCTt. l‘le’rjl.) .3 Vf‘T'Tfit1 rm 815, 'Otl 1,.‘ ’1 C; 0 ,"t1 CS 0 3 (I. in 3 T |_. ‘~ JPF1011351 ynfift hybrids 3101. PCV. 5nd rwt new in ’LCGT‘ 1. U 11: u“. A 3‘9. t—I Q wrie- V. C? M, 03' n (-3111.-r-T"’pf?“091‘hf79139?r1 CITLTT, “1’1. 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