Chlorophyll varlegation in Vitis species was examined In F1 and S1 populations. A single recessive gene is proposed to account for the occurence of 25 percent variegated seedlings in both F1 and S1 populations. Variegation was not lethal and might be useful in wine grape breeding (If associated with inactive polyphenol oxldase) and for its ornamental value. This trait is one of very few seedling traits trider monogenic control that might be useful in genetic studies of grape

Relsch, B. I.

Watson, J. P.

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The Journal of Heredity 75:417-418. 1984.Inheritance of leaf variegationin Vitis speciesB. I. Relsch and J. P. WatsonResults and DiscussionMost of the variegated seedlings showed agreen and white pattern of variegation. A fewvariegated seedlings had patches of pinkish-purple and patches of green tissue in true leavesand occasionally in the cotyledons. The pink-ish-purple color eventually bleached out, leavingwhite tissue in its place. Leaves on variegatedseedlings ranged from almost entirely green toentirely white. Classification at the cotyledon tofirst true leaf stage was inaccurate; 10 days latermany seedlings classified earlier as normalshowed leaf variegation. Data collected fromseedlings showing 2-4 true leaves were subse-quently analyzed and the results are presentedhere. The symbol uar is proposed for the mutantcausing green and white leaf variegation.> ABSTRACT. Chlorophyll vartegatton in V7rts specieswas examined In F, and S t populations. A singlerecessive gene Is proposed to account for trieoccurrence of 25 percent variegated seedlings Inboth Fi and S, populations. Variegation was not. lethal and might be useful In wine grape breeding(if associated with Inactive polyphenol oxklase), and for Its ornamental value. This trait Is one ofvery few seedling traits under monogenic controlthat might be useful In genetic studies ofgrapes.LEAF VARIEGATION in Vitis has been describedamong the progeny of Aramon, an ancient Eu-. ropean grape (V. vinifera) cultivar5. A singlerecessive gene was reported to cause green andwhite variegation among selfed progeny ofAramon. However, such variegation was ex-pressed only among plants grown at 20°C, not at35°C5. Variegation also was described in a sportof cv. Sultana, but inheritance was not studied1.We have observed that, following severe coldi damage to dormant buds, small variegated shootsare a common vineyard occurrence.Well-defined mutants identifiable at an earlystage of growth are of importance in geneticstudies of plants. De Lattin3 listed 53 genesidentified in Vitis prior to 1957. Of these, fourconcerned traits identifiable among seedlings. Nofurther seedling traits have since been re-ported4.Variegated leaves appeared in seedlinggrapevine populations planted in 1982 and werestudied to identify the genetic basis for their oc-currence.Materials and MethodsCrosses and selfs among hybrid stocks (seeFigure 1) and V. oinifera grape cultivars(Riesling and Chardonnay) were made in 1981and 1982 using standard techniques4. Seed werestratified for 3 months at 1-2°C and planted inCornell-Mix2 medium in a greenhouse in mid-March of 1982 and 1983. Approximately 4 weeksafter planting, seedlings showing true leaves wereclassified as cither variegated (green and white)or normal (entirely green). The available datawere subjected to chi-square analysis to deter-mine goodness-of-fit to the Mendelian monohy-brid ratio. In June 1982 and 1983, seedlings weretransplanted to a field nursery and observed forsurvival and expression of variegation.The authors are affiliated with the Department ofHorticultural Sciences, New York State AgriculturalExperiment Station, Cornell University, Geneva, NewYork 14456. They wish to thank Dr. R. Pool for valu-able discussions.© 1984, American Genetic Association.RavatSi —Baco2-16 —- S 6905 -r s- s.*— PlnotNolr(vin)— Seedling[—— Baco 1l_NY 34217 —IAlicante_i— S. 452 —i4595 —— S.405 ——-4199-- S.85-r~U C . 132-11^IBaroque Blanc (vin)Noah (lab, rip)Folle Blanche (vin)Vitis rlparia— Pinot Blanc (vln)— Ontario (lab)Qanzin—S. 14AxR1 —L_AxR1 —(rup)(vin)AxR4 —AlicanteBouschetJaeger 70(vin)Jaeger 70(vin)Aramon (rupestrisGanzinJaeger 70(vin)i— Aramon(vln)| rupestrisGanzin(V, Aramon)[— (Unc)I— (rup)|—(Unc)I—(rup)vin)i-(linc)'-(rup)Aramon (vln)rupestrisGanzinFIGURE 1 Pedigreeof hybrid stocks used in the study of variegation in Vilii. Abbreviations for species: vin-cinifera; lab—labrusca; rip—riparia; line—lincecumii; rup—nipeslris.~ Notes September / October 1984 417Variegated plants from crosses made in 1981continued to express the trait in tissues producedin 1982, 1983, and 1984 (Figure 2). Expressionwithin each plant was irregular; some shoots wereentirely green, others were intensely variegated.Only a few seedlings classed as normal exhibitedvariegation in the field. These individuals hadapparently been misclassified at the seedlingstage. Following two seasons in the field, 78 of149 green plants in two Fi populations survived,as did 15 of 48 variegated plants. Apparently,variegation was not a lethal trait, although itappears to decrease plant survival. Survivingvariegated plants appeared to be as vigorous astheir normal sibling counterparts.It is proposed that NY 34217, Riesling,Chardonnay, Ravat 51, and Baco 2-16 are het-erozygous (+ var) for the variegated trait.Hence, hybrid and selfed progeny within thisgroup should have 25 percent variegated seed-lings. Only one of 10 populations (Table I) ana-lyzed using the chi-square test did not fit thishypothesis. In that atypical case, a self-pollinatedpopulation from Riesling included only 16 per-cent variegated seedlings. Misclassification at anearly growth stage and possible temperature ef-fects on variegation5 would help to explain a de-ficiency of seedlings classed as variegated.Albinos occur at very low frequencies (<0.5percent) in many grape seedling populations.They occurred at a frequency of 16 percentamong self-pollinated seedlings of Ravat 51(Table I) and were considered in the statisticalanalysis to be an extreme form of expression forthe variegated trait. Albinism has been consid-ered to be an extension of leaf variegation traitsin other studies as well7.In 14 progeny having one parent in commonwith those segregating for leaf variegation, allobserved seedlings were green (Table II). Thecross involved would represent + + X +oar, withthe expectation of producing all green seedlings.Self-pollinated progeny of three putative + +genotypes also produced only green seedlings(Table II).Studies on the grape cultivar Aramon sug-gested monogenic inheritance of a variegatedtrait5. Since Ravat 51 derives approximately 9.0percent of its genetic makeup from AramonFIGURE 2 Leaf variegation in a 31-year-old Vilisseedling.through four independent sources (Figure 1) itis possible that the gene investigated is identicalto that found in Aramon and now identified in thefour other grape stocks reported here.Grapevines grown in pot culture from cuttingsare being developed as an attractive floristcrop6'9. A variegated stock might have additionalornamental value.The mutant trait presently under study alsomay be of some importance in the production ofnew grape cultivars that resist oxidation in theirwine. Grapevine variegation has been found inassociation with Bruce's sport, a variant of theSultana grape, which was characterized by adecreased tendency of the fruit to brown whileTable I. Segregation data and chi-square tests for green and variegated (var) leaf traits amongVitis seedlingsCrossFi populationsNY 34217 X RieslingNY 34217 X Baco 2-16NY 34217 X Ravat 51Ravat 51 X RieslingRavat 51 X ChardonnaySi populationsNY 34217 selfedRavat 51 selfedRiesling selfedChardonnay selfedBaco 2-16 selfedtotal126220621193251421619942270No. seedlingsgreen861645296246916316833205variegated4056102379S19t31965X2(3:D3.060.022.602.070.090.860.029.470.290.12P0.05-0.200.80-0.950.05-0.200.05-0.200.5O-O.800.20-0.500.80-0.95<0.01*0.50-0.800.50-0.80drying1. A form of polyphenol oxidase charac-terized by greatly reduced activity was respon-sible for resistance to oxidation of grape phenoliccompounds sensitive to browning8. This alteredform of polyphenol oxidase may be a valuable .trait in retarding oxidation in commercial wines.The genetic relationship between variegation andaltered forms of polyphenol oxidase has not yetbeen elucidated. However, if the variegated traitpresently under study is identical to that foundin Bruce's sport and is associated with an inactiveform of polyphenol oxidase, it may be a usefultrait to manipulate for the benefit of a wine grape 'breeding program.References1. ANTCLIFF, A. J. and W. J. WEBSTER. Bruce'ssport—a mutant of the sultana. Australian J. Expt.Agr. Anim. Husb. 2:97-100. 1962.2. BOODLEY, J. W. and R. S H E L D R A K E , JR. Cornell *peatlite mixes for commercial plant growing. Inf.Bui. 43. Cornell Coop. Extension, Ithaca, NY. 1982.3. DE LATTIN, G. Zur Genetik der Reben. BisherigeErgebnisse der Faktoranalyse bei der Gattung Vilis.Vitis 1:1-8. 1957.4. ElNSET, J. and C. PRATT. Grapes. In Advances in ,Fruit Breeding. J. Janick and J. N. Moore, Eds.Purdue Univ. Press, Lafayette, IN. p, 130-153. 1975. H5. G A L Z Y , R. and P. GALZY. Action de la temperaturesur la panachure blanche de la vigne. Le ProgresAgricoleet Viticole 162:271-280. 1964.6. JANICK, J. Production of fruiting grapevines for potculture. HortScience 18:56-57. 1983.7. O R A K W U E , F. C. and L. V. CROWDER. Inheritanceof variegation in Brassica campestris. L. J. Hered. '74:65-67. 1983.8. RADLER, F. The prevention of browning during rdrying by the cold dipping treatment of sultanagrapes. J. Sd. Food Agric. 15:864-869. 1964.9. ROB1TA1LLE, H. A. and J. JANICK. Rapid pro-duction of small fruiting grapevines from softwoodcuttings. HortScience 14:161. 1979.Table II. Other crosses and sells related tosegregating populations in which no variegatedseedlings were observedCross*NY 65.444.4 X RieslingNY 65.533.13 X RieslingGW 5 X RieslingHorizon X RieslingCayuga White X RieslingS. 13047 X RieslingJ.S. 23-416 X RieslingJS. 26-627 X RieslingNY 65.444.4 X ChardonnayS. 10173 X Chardonnay111. 172-7 X ChardonnayHorizon X Baco 2-16NY 65.533.13 X Ravat 51Cayuga White X Ravat 51}S. 23-416 selfedNY 65.444.4 selfedHorizon selfedNo. seedlingsobserved9337515186751051414992473222395420054• Does not fit expected ratiot + 34 albino* Abbreviations used: NY - New York; GW =Geneva White; S. - Seibel; JS. •= Joannes-Seyve;III. - Illinois418 The Journal of Heredity Notes

Inheritance of leaf variegation In Vitis species

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