Two grapevine seedlings, V. 72181 and V. 72182, selected for extremely high methyl anthranilate (MA) and total volatile esters (TVE) content, were selfed to create families 8020 and 8021 respectively, to test the inheritance of these two components of labrusca flavour character. REYNOLDS et al. (1982) had postulated a three-gene, dominant and complementary system (M, A, F) for MA and a two-gene dominant and complementary system (T, V) for TVE. Families 8020 and 8021 segregated 3:1 for MA, indicating only one heterozygous locus for MA in the parents. This would question REYNOLDS' assignment of genotypes for the grandparents of these two families and would suggest a more complex environmentally influenced system. The TVE segregation patterns followed REYNOLDS' hypothesis and segregated 3:1 for one heterozygous locus

Fisher, K. H.

Fuleki, T.

Reynolds, A. G.

English

JKI Open Journal Systems (Julius Kühn-Institut)

Genetic improvement of grapevine form or function 159 The heritability of methyl anthranilate and total volatile esters in Vitis spp. hybrids K. H. F1sHER 1), T. FULEKI 1) and A G. REYNOLDS 2) 1) Honicultural Research Institute of Ontario, Vineland Station, Ontario, Canada 2) Agriculture Canada Research Station, Summerland, British Columbia, Canada S u m m a r y : Two grapevine seedlings, V. 72181 and V. 72182, selected for extremely high methyl anthranila.te (MA) and total volatile esters (1VE) content, were selfed to create families 8020 and 8021 respectively, to test the inheritance of these two companents of labrusca flavour character. REYNOLDS et al. (1982) bad postulated a three-gene, dominant and complementary system (M, A, F) for MA and a two-gene dominant and complementary system (T, V) for 1VE. Families 8020 and 8021 segregated 3 : 1 for MA, indicating only one heterozygous locus for MA in the parents. lbis would question REYNOLDS' assignment of genotypes for the grandparents of these two families and would suggest a more complex environmentally influenced system. The 1VE segregation patterns followed REYNOLDS' hypothesis and segregated 3 : 1 for one heterozygous locus. Key word: genetics, heritability, flavour, methyl anthranilate, volatile esters, biometry, sensory rating, breeding. Introduction Since 1913, the grape breeding programme at the Horticultural Research Institute of Ontario (HRIO) has been striving to produce wine grapes of suitable hardiness for the climatic conditions in southern Ontario, Canada. Flavour and colour requirements have changed markedly since that time and the breeding programme has been altered to meet the new requirements. With more recent demands for non·labrusca flavoured varieties, the heritability of labrusca flavour components has become quite critical. Since 1970, Dr. T. FuLEKI of this Institute has been surveying the HRIO variety collection as well as the seedlings resulting from the HRIO breeding programme for compounds thought to be instrumental in the perception of labrusca flavour, i. e. methyl and ethyl anthranilate (MA) and total volatile esters (TVE). He later used these components to develop the Vineland Grape Flavour Index (VGFI) that became the objective flavour selection criterion for the grape seedling populations at the Institute (FULEKI 1982). The analysis of grape seedling selections for the above compounds became routine procedure, but perhaps more important was the impact on the choice of parents in the breeding programme for subsequent generations. The M.\ and TVE survey values were examined to assess the potential labrusca flavour of seedlings according to the M.\{TVE/VGFI values of the parents. Based on the hypothesis that heritability could be predicted from these values, REYNOLDS studied three seedling families, 7216, 7218 and 7219, which had high and low labrusca flavoured parents. The seedling populations of these three families were examined in 1977 and 1978 and REYNOLDS postulated genotypes for MA and TVE synthesis for their parents. He also identified two seedlings, V. 72181 and V. 72182, with extraordinarily high MA and TYE values (REYNOLDS 1980). Thes~ two seedlings were maintained in the HRIO vineyard and were selfed in 1980 to create families 8020 and 8021, respectively (Fig. 1). REYNOLDS postulated that M.\ synthesis was governed by three dominant and complementary genes (M, A and F), with the F locus necessary in the dominant condition for MA synthesis to proceed and Mand A loci both dominant for excessive M.\ production. The presence of M and/or A in the recessive condition would result in lower amounts of MA, below the organoleptic threshold oflabrusca flavour influence, and the presence of the F locus in the recessive 160 Section 2 Concord x De Chaunac (Seibel 9549) V.54077 X Le General (Bertille Seyve 5563) ~ v.~81 v.~82 8020 8021 a Fisher, Ann. Rept. HRIO, 18-20, 1981. a Fig. 1: Genealogy of grape vine cultivars/ seedlings used in MA/lVE inheritance studies 1. Mm Aa Ff X MM Aa ff Tt Vv x tt Vv (excess MA) 3:5 (low or no Ma) (excess TVE) 3:5 (low TVE) Reynolds et al, Am. J. Eno!. Vitic., 33:1:14-19, 1982. Fig. 2: Segregation ratios for MA and 1VE for Family 7218 (V. 54077 x Le General) 1 condition would preclude any MA synthesis. REYNOLDS also postulated that TVE was governed by two dominant complementary genes (T and V) and that either in the recessive condition would constitute TVE production below the organoleptic level oflabrusca flavour influence (REYNOLDS et al. 1982). These hypotheses led to the genotypic assumptions presented in Fig. 2. Since seedlings V. 72181 and V. 72182 were selected for their exceptionally high M.\ and TVE, it is assumed that they would have the genotypic make-up for excess MA and TVE production. Based on REYNOLDS· genotypic assignments for the parents, the following genetic combinations could be assigned to the above seedlings, as outlined in Table 1. In order to confirm the ratios postulated in Table 1, the following study was initiated. Method Plant material Parental lines were planted in the vineyard as individuals or as populations of vegetatively propagated clones. Their genealogy is illustrated in Fig. 1. Selfed populations were created by bagging flower clusters of the appropriate parental vines priorto capfall. Seedling populations (8020, 8021) were planted as a group in the seedling block of the main vineyard. Genetic improvement of grapevine form or function 161 Table 1: Proposed genotypes and segregation ratios for MA and TVE for F 2 populations derived from V. 72181 and V. 721828 crosses Excess MA Segregation Ratio Excess TVE Segregation Ratio MM AA Ff L/NL 3:1 Tt w L/NL MM Aa Ff L/NL 9:7 Tt Vv L/NL MM aA Ff Tt vV mM AA Ff mM Aa Ff L/NL 27:37 mM aA Ff a Reynolds et al, Am. J. Enol. Vitic., 33:1:14-19, 1982 b L = labrusca, NL = non-labrusca flavour character Table 2: Threshold values for organoleptic detection oflabrusca flavour characters Methyl anthranilate Total volatile esters Harvest 0.10 ppm 12.0 ppm (Nelson et al, 1977) (Fuleki, l982) 3:1 9:7 At maturity, the total crop was harvested from each seedling vine (total crop was sub-sampled from a parent vine). Grapes were washed, removed from the stems, frozen and maintained at -30°C until analysis. Analyses for MA and TYE were carried out within 3 months of the harvest date. Data are presented forthe harvestsof1986, 1987 and 1988 in Tables 3-6. To obtain representative samples for analyses, frozen grape samples were ground without breaking the seeds. A household and a 5 HP commercial meat grinder (Butcher Boy, Model A42.50, Laser Mfg. Co. Inc., Los Angeles, CA, USA) were used for smaller and larger lots respectively. The ground frozen material was thoroughly mixed and two 50 g aliquots were removed for steam distillation. Steam distillation Samples were placed in an all-glass distillation apparatus (Cat. No. JD-2115, SGA Sci., Bloomfield, N.J., USA). The distillation was carried out on duplicate 50 g samples, collecting 100 ml in 15 min. The distillate was used to determine both the M.\ and the TYE. Glass distilled water was steam distilled under the same conditions as the grapes to produce blanks for the MA and TYE analyses. Determination of MA A highly sensitive fluordmetric method was used (CASIMIR et al. 1976). The measurements were carried out on a ZFM 4 fluorescence attachment of a Zeiss DMR21 recording spectrophotometer (C. Zeiss, Oberkochen, Germany). The exciting radiation was isolated with a 162 Section 2 Table 3: Methyl anthranilate values and equivalent la brusca/non-labrusca codes for Family 8020 ( 1986 to 1988) Methyl anthranilate (ppm) Seedling Number 1986 1987 Code 1988 Code 1 1. 776 L 7.446 L 2.776 L 2 0.136 L 3 5.096 L 4.596 L 4 0.618 L 0.560 L 5 17.218 L 14.746 L 6 0.290 L 0.482 L 0.767 L 7 0.244 L 0.676 L 2.164 L 8 8.546 L 9 0.037 NL 10 0.143 L 0.042 NL 0.052 NL 11 1. 786 L 1.950 L 12 6.446 L 11.326 L 13 0.541 L 15 0.866 L 0.039 NL 1.654 L 16 4.076 L 4.076 L 5.386 L 17 5.436 L 5.556 L 19 0.022 NL 0.079 NL 20 0.163 L 0.239 L 0.261 L 21 0.358 L 0.565 L 0.882 L 23 0.093 NL 0.479 L 1. 733 L 24 0.106 L 0.192 L 25 7.956 L 9.836 L 6.819 L 26 1. 746 L 4.876 L 2.626 L 27 0.096 NL 0.301 L 0.354 L 28 2.167 L 29 0.006 NL 0.006 NL 0.014 NL 30 0.079 NL 0.471 L 31 3.436 L 32 0.366 L 1.356 L 1.119 L 33 4.166 L 12.606 L 11.796 L 34 0.065 NL 0.070 NL 35 2.936 L 2.826 L 6.071 L 37 0.000 NL 0.011 NL 0.031 NL 38 0.992 L 39 0.013 NL 0.032 NL 0.044 NL 40 0.032 NL 0.018 NL 0.015 NL 42 o.ooo NL 0.006 NL 0.044 NL 43 0.036 NL 0.060 NL 44 3.046 L 2.583 L 45 10.096 L 15.776 L 7.826 L L labrusca, NL = non-labrusca flavour character 365 nm monochromatic filter from the line spectrum of a mercury lamp. The 4MQIII mono· chromator of the spectrophotometer was set at 425 nm to measure the fluorescence emitted by the MA. The fluorometer reading was related to the M.\ concentration by using a standard cuzve. To eliminate interference from fluorescent volatile substances other than methyl and ethyl amhranilate, the reading taken on a Vi1is vinifera L. cultivar corresponding to a few ppb M.\ was subtracted from each measurement. Genetic improvement of grapevine form or function 163 Table 4: Methyl anthranilate values and equivalent labrusca/non·labrusca codes for Family 8021 ( 1986 to 1988) Methyl anthranilate (ppm) Seedlinq Number 1986 1987 Code 1988 Code 1 0.176 L 1.526 L 2.370 L 2 0.179 L 0.816 L 0.72 L 3 0.037 NL 0.087 NL 4 2.726 L 7.516 L 2.836 L 5 0.189 L 1.436 L 0.931 L 6 0.655 L 0.530 L 8 0.386 L 0.024 NL 0.097 NL 10 0.748 L 0.511 L 11 0.023 NL 0.361 L 1.346 L 12 0.336 L 1.137 L 13 0.082 NL 0.075 NL 0.066 NL 14 0.081 NL 0.121 L 0.172 L 15 3.696 L 15.676 L 11.776 L 17 17.696 L 13.966 L 18 3.976 L 8.216 L 2.832 L 19 0.077 NL 0.221 L 20 0.136 L 0.053 NL 0.202 L 21 0.471 L 1.112 L 2.206 L 23 0.452 L 0.630 L 0.571 L 24 0.020 NL 0.026 NL 0.029 NL 25 0.034 NL 0.137 L 0.140 L 27 0.251 L 0.683 L 0.436 L 28 0.627 L 1.416 L 1. 715 L 29 1.236 L 2. 776 L 4.055 L 30 0.730 L 1.876 L 31 2.919 L 32 0.028 NL 0.384 L 0.213 L 33 0.480 L 1.424 L 0.639 L 34 1.236 L 2.546 L 2.366 L 35 0.451 L 0.752 L 0.925 L 38 1.326 L 2.705 L 2.266 L 39 0.484 L 1.276 L 1.074 L 40 2.836 L 3.766 L 5.207 L 42 2.566 L 3.586 L 6.336 L 43 0.038 NL 44 0.018 NL 3.266 L 45 0.924 L 0.920 L 46 0.559 L 1.626 L 0.694 L 47 0.051 NL 0.081 NL 48 0.159 L 0.096 NL 0.193 L 50 0.010 NL 0.063 NL 0.043 NL 54 5.206 L L labrusca, NL non-labrusca flavour character Determination of TYE A modification of THOMPsoN's method (1950) was used. The reaction of esters with hydroxylamine in aqueous alkaline solution forms hydroxamic acids, which react with the ferric ion to form red ferric hydroxamate complexes. 164 Section 2 Table 5: Total volatile esters values and equivalent labrusca/non·labrusca codes for Family 8020 (1986 to 1988) Total volatile eaters (ppm) Seedlinq Number 1986 Code a 1987 Code 1988 Code 1 56 L 66 L 59 L 2 128 L 3 90 L 456 L 4 170 L 28 L 5 81 L 197 L 6 233 L 223 L 126 L 7 3 NL 5 NL 9 NL 8 20 L 9 2 NL 10 212 L 6 NL 162 L 11 19 L 15 L 12 22 L 61 L 13 266 L 15 92 L 70 L 70 L 16 150 L 141 L 208 L 17 288 L 58 L 19 2 NL 3 NL 20 235 L 20 L 42 L 21 5 NL 8 NL 21 L 23 2 NL 2 NL 4 NL 24 62 L 297 L 25 127 L 32 L 48 L 26 239 L 75 L 134 L 27 144 L 255 L 28 36 L 29 20 L 22 L 21 L 30 3 NL 2 NL 31 116 L 32 89 L 58 L 95 L 33 52 L 51 L 181 L 34 20 L 166 L 35 45 L 39 L 66 L 37 1 NL 1 NL 1 NL 38 3 NL 39 80 L 36 L 72 L 40 4 NL 62 L 186 L 42 1 NL 2 NL 2 NL 43 4 NL 2 NL 44 40 L 152 L 45 194 L 154 L 119 L a L labrusca, NL = non-labrusca flavour character Fresh alkaline hydroxylamine solution was prepared by mixing equal volumes of 6 M hydroxylamine hydrochloride and 10.5 ~sodium hydroxide. 2 ml of this solution and 20 ml of the distillate were added to a 25 ml volumetric flask. These were mixed thoroughly and allowed to stand for 5 min. Then. 1 ml of concentrated hydrochloric acid was added, followed by 1 ml of 1.11 M ferric chloride. The flask was filled to volume with 0.046 M ferric chloride solution and the colour measured on a Zeiss DMR21 spectrophotometer at 5 00 nm. A standard curve, prepared by Genetic improvement of grapevine form or function 165 Table 6: Total volatile esters values and equivalent labrusca/non-labrusca codes for Family 8021 (1986 to 1988) Total volatile esters (ppm) Seedling Number 1986 1987 Code 1988 Code 1 139 L 63 L 38 L 2 6 NL 55 L 2 NL 3 1 NL 4 NL 4 1 NL 3 NL 3 NL 5 89 L 24 L 59 L 6 L 59 L 61 L 8 91 L 56 L 55 L 10 1 L 4 NL 11 56 L 55 L 79 L 12 120 L 26 L 13 168 L 15 L 30 L 14 4 NL l NL 1 NL 15 183 L 147 L 261 L 17 56 L 93 L 18 105 L 100 L 58 L 19 l NL 1 NL 20 118 L 28 L 100 L 21 108 L 54 L 51 L 23 174 L 82 L 154 L 24 44 L 17 L 25 L 25 1 NL 0 NL l NL 27 204 L 95 L 187 L 28 1 NL 1 NL 3 NL 29 2 NL 2 NL 2 NL 30 83 L 40 L 31 3 NL 32 54 L 16 L 128 L 33 165 L 59 L 238 L 34 51 L 12 NL 66 L 35 35 L 15 L 22 L 38 63 L 18 L 52 L 39 175 L 42 L 79 L 40 80 L 72 L 81 L 42 88 L 28 L 35 L 43 1 NL 44 0 NL 33 L 45 4 NL 23 L 46 184 L 36 L 170 L 47 0 NL 5 NL 48 56 L 28 L 52 L so l NL 0 NL 2 NL 54 73 L a L labrusca, NL = non-labrusca flavour character using various concentrations of ethyl acetate, was used to determine the concentration of the TYE in the sample. Statistical analysis Chi square· analyses were performed on the observed ratios oflabrusca/non·labrusca flavour components as they were identified by the M.\ and TYE analyses. 166 Section 2 Results The seedling families 8020 and 8021 (selfed populations of V. 72181 and V. 72182, respectively) displayed transgressive segregation in both MA and TVE distribution (Tables 3-6). All four distributions were highly skewed to the lower values of MA and TYE. To simplify the data, all M.\ and TVE values were coded as to labrusca or non-labrusca in flavour contribution, according to the threshold values in Table 2, and presented in Tables 3·6. These threshold values were also used by REYNOLDS et al. (1982). The ratios postulated by the authors using genotypic assumptions of REYNOLDS et al. (1982) were tested and are presented in Tables 7·10. Table 7: Chi square tests for MA segregation ofFamily 8020 (1986to1988) Year Predicteda Statistic Observed xa p value Significance5 1986 L/NL 3:1 18:9 0.50 o. 5o<p<o. 30 ns 1987 L/NL 3:1 24:8 0.031 0.90<p<0.70 ns 1988 L/NL 3:1 25:10 0.08 0.90(p<0.70 ns 1986 L/NL 9:7 18:9 0.80 0.50.(p(0.30 ns 1987 L/NL 9:7 24:8 3.84 p = 0.05 ** 1988 L/NL 9:7 25:10 2.67 o.1o<p<o.05 * L/NL = labrusca/non-labrusca flavour character b significance levels for rejection: p=0.10=*, p=0.05=**, p=0.001=*** Table 8: Chi square tests for MA segregation ofFamily 8021 (1986 to 1988) Year Predicteda Observed xa 1986 L/NL 3:1 26:7 0.06 1987 L/NL 3:1 28:9 0.013 1988 L/NL 3:1 33:7 0.833 1986 L/NL 9:7 26:7 5.87 1987 L/NL 9:7 28:9 4.92 1988 L/NL 9:7 33:7 10.15 L/NL = labrusca/non-labrusca flavour b significance levels for rejection: p•0.001=*** Statistic P value O. 90<p<O. 70 0.95(p<0.90 o. 5o<p<o. 30 o.os<p(o.01 O.OS(p<0.01 O.Ol<"p<0.001 character Signif icanceb ns ns ns ** ** *** 0=0.10=*, p=0.05=**, Genetic improvement of grapevine fonn or function 167 Discussion The 3 : 1 segregation ratio for both MA and TYE in both families 8020 and 8021 had the most acceptable Chi square values. This would mean that the Y. 72181 and Y. 72182 genotype for both MA and TYE must be segregating for only one heterozygous locus each. For MA, if that locus is F, then the ff recessive must allow for some production of MA because no seedlings measured in family 8020 or 8021 had a zero level for MA with any consistency over the three harvest seasons. For it to be a locus other than F would preclude the grandparent Le General and the great grandparent De Chaunac from having such low levels of MA. Perhaps the REYNOLDS hypothesis (REYNOLDS 1980) of an alternate MA synthesis pathway exists under the ff recessive condition as well as under the aa and/ or mm recessive conditions. The TYE inheritance pattern for both families 8020 and 8021 follows REYNOLDs' hypothesis (REYNOLDS 1980) since there was no condition completely precluding the synthesis ofTVE. Table 9: Chi square tests for lVE segregation ofFamily 8020 (1986 to 1988) Predicteda Statistic Year Observed x2 p value Signif icanceb 1986 L/NL 3:1 18:8 0.20 0.70<'.p<0.50 1987 L/NL 3:1 24:8 o.o p = l 1988 L/NL 3:1 27:8 0.013 o. 9o<:p<o. 10 1986 L/NL 9:7 18:8 1.32 0.3o<p<0.20 1987 L/NL 9:7 24:8 3.84 p = 0.05 1988 L/NL 9:7 27:8 5.36 0.05<p<O.Ol a L/NL = labrusca/non-labrusca flavour character b significance levels for rejection: p=0.10=*, p=0.05=**, p=0.001=*** Table 10: Chi square tests forlVE segregation of Family 8021 (1986 to 1988) Predicteda Statistic Year Observed ns ns ns ns ** ** x2 p value Significance 1986 L/NL 3:1 24:9 0.009 o. 95<p<o. 90 ns 1987 L/NL 3:1 25:12 0.73 O.So<p(0.30 ns 1988 L/NL 3:1 28:12 0.31 0.70<.p<0.50 ns 1986 L/NL 9:7 24:9 2.96 0 .10<'.p<O. 05 * 1987 L/NL 9:7 25:12 1.51 0. 30<p<O. 20 ns 1988 L/NL 9:7 28:12 2.54 o.20<pc::o.10 ns L/NL = labrusca/non-labrusca flavour character b significance levels for rejection: p=0.10=*, p=0.05=**, p=0.001=*** 168 Section 2 Acknowledgements The assistance ofR.B. P AL.>\BA Y with the analytical determinations is gratefully acknowledged. Literature cited CASIMIR, D. J.; MOYER, J. C.; MATTICK, L. R .; 1976: Fluorometric determination of methyl anthranilate in Concord grape juice. J. Assoc. Ollie. Analyt. Chem. 59, 269-272. FrsHER, K. H .; 1981: Breeding for 'vine and table grapes adapted for Ontario conditions. Ann. Rpt. Hort. Res. Inst. Ont. 18· 20. Ft:LEKI, T.; 1982: The Vineland Grape Flavour Index · a new objecrive method for the accelerated screening of grape seedlings on the basis offlavour character. Vitis 21, 111-120. NEt.soi-:, R. R.; ACREE, T. T.; LEE, C. Y.; Buns, R. M.; 1977: Methyl anthranilate as an aroma constituent of American wine. J. Food Sci. 42, 57-59. REYNOLDS, A. G.; 1980: The Inheritance of Methyl Anthranilate and Total Volatile Esters in Vitis spp. M. Sc. Thesis, Univ. ofGuelph, Guelph, Ontario, Canada. ·· ·· ; FULEKI, T.; EYAl'S, W. D.; 1982: Inheritance of methyl anthranilate and total volatile esters in Vitis spp. Amer. J. Eno!. Viticult. 33, 14-19. THOMPSO~. A. R .; 1950: A colourimetric method for determining esters. Austral. J. Sci. Res. 3 A, 128-135. Breeding grapes for basal fruitfulness R. Sr~GH and B. l'\. S. M rnrnY Di\~sion of Fruit Crops, Indian Institute of Horticultural Research, Hessaraghatta, Bangalore-560089, India Abs t r act : An attempt has been made to develop varieties with fruitful basal buds, as this character is of prime importance for any cultivar to be grown successfully on low-cost training systems like the head system. As a first step, the fruiting behaviour of several accessions from diverse geographic origin was made and certain genotypes were identified as promising sources for the desired character. Employing a microscopic bud analysis technique. bud fertility of these genotypes was measured from 2nd to 1 Jth node and various biometric parameters were worked out to detem1ine the extent of variability and inheritance of this character. The pheno\ypic and genotypic coefficients of variance were moderate to high and heritability estimate was convincing enough to begin hybridization.Extensive hybridization work was carried out for several years employing the identified parents and this has resulted in large number of hybrid progenies. The fertility analysis of these hybrids and their respective parents indicated that the varieties Bangalore Blue, Black Champa. Queen of the Vineyards and Convent Large Black are potential sources for basal fruitful buds. To incorporate this desirable character in the commercial cultivars like Anab-e-Shahi and Thompson Seedless, large number of crosses were made involving these cultivars and potential sources. From these progeny we were able to locate some promising hybrids which, besides possessing fruitful basal buds, are prolific bearers of high quality fruits. 

The heritability of methyl anthranilate and total volatile esters in Vitis spp. hybrids

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The heritability of methyl anthranilate and total volatile esters in Vitis spp. hybrids

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