tain cyanidin/peonidin should not possess a recessive phi allele. Several conclusions can be drawn from this study. 1. The inheritance of specific flower col-.
From the US National Arboretum, Floral and Nursery Plant Research, USDA, ARS, BARC-WEST, Beltsvllle, MD 20705-2350.1 would like to thank Sister M. Antonio Reneau lor her help In collecting the anthocyanin data.
Yellow-Tip: A Cytoplasmically Inherited Trait in Melon (Cucumis melo L.)
The Journal of Heredity 1996.87(3)
D. T. Ray and J. D. McCreight
A new chlorophyll-deficient mutant is the first cytoplasmically inherited trait described in melon. This mutant is characterized by yellow apices, with the leaves and stems progressively turning green in color as the branches mature. A protocol is proposed for naming and symbolizing cytoplasmic traits in melon. This mutation Grfesbach RJ, Asen S, and Leonhardt BA, 1991. Petunia is named yellow-tip and is symbolized cythybrida anthocyanlns acylated with caffelc acid. PhyYt. As a chlorophyll-deficient mutation, it is tochem 30:1729-1731. potentially useful in genetic, physiological, Hooker, J, 1837. Petunia violacear. Petunia hybrida var. and biochemical studies. Curtis. Bot Mag 64 (new series 11)^556. References Chuck G, Robblns T, NlJJar C, Ralston E, Courtney-Gutterson N, and Dooner HK, 1993. Tagging and cloning of a petunia flower color gene with the maize transposable element activator. Plant Cell 5:371-378. deVlaming P, Schram AW, and Wiering H, 1983. Genes affecting (lower color and pH of flower limb homogenates In Petunia hybrida Theor Appl Genet 66:271-278.
Reciprocal crosses were made between the chlorophyll-deficient line and Top Mark, (C. melo subsp. melo Cantalupensis Group; Kirkbride 1993) the current standard cultivar for western U.S. shippingtype cantaloupes. It is characterized by dark-green foliage and heavily netted fruit with orange-colored flesh. Crosses were performed in a greenhouse in Salinas, California, to produce F, (reciprocal), F2, and BC, families. Evaluations were done in a greenhouse at the University of Arizona. Plants were grown in a medium of equal parts by volume of peat, sand, and vermiculite in 21.6 cm high, 21.6 cm top diameter, and 17.8 cm bottom diameter containers (7.6 L volume). Greenhouse temperatures throughout the experiment (June to August 1993) ranged between
Brief Communications 2 4 5
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Jonsson LMV, Aarsmann MEG, Poulton JE, and Schram Even though all of the F2 seedlings that In melon (Cucumis melo L), 111 mutant ' AW, 1984. Properties and genetic control of four methhad the genotype Hf-Mf- contained the yltransferases Involved In the methyiatlon of anthocy- phenotypes have been described, but only anlns In flowers of Petunia hybrida Planta 160:174-179. 62 are maintained by the collection curasame anthocyanin composition, they did not have the same color. The F2 seedlings tors (Pitrat 1994). Of these mutants, nine Kondo T, Yoshlda K, Nakagawa A, Kawal T, Tamura H, and Goto T, 1992. Structural basis of blue-color develthat were red (RHS 78A) had more acidic are chlorophyll-deficient mutants, six of opment In flower petals from Commelina communis. pHs than those seedlings that were blue which are maintained (Pitrat 1994). ChloNature 358515-518. (RHS 89C). The full range of pHs (5.5-6.3) Kurkdjlan A and Guem J, 1989. Intercellular pH: mea- rophyll-deficient mutants are potentially found in the entire F2 population was not useful in genetic, physiological, and biosurement and Importance In cell activity. Annu Rev found within this Hf-Mf- subpopulation Plant Physlol Plant Mol Blol 40:271-303. chemical studies. These mutants have dis(Figure 4). None of the Hf-Mf- seedlings ex- Marshall HH, Campbell CG, and CoUicutt LM, 1983. crete phenotypes that are easily identified Breeding for anthocyanin colors In Rosa Euphytlca 32: pressed the more acidic pHs. The most and readily manipulated. Eight of the nine 205-216. acidic pHs were only found in the hfhfMfchlorophyll-deficient mutants are condiS, 1964. A survey of anthocyanlns In petuand hfhfmfmf subpopulations. This is ex- Muszynsld tioned by recessive alleles at different loci nia. Physlol Plant 17:975-979. pected since the Hfl and Phi genes are (Cox and Harding 1986; Dyutin 1979; HoffNleuwhof M, van Eljk JP, and Elkelboom W, 1989. Relaclosely linked on chromosome 1 (Wiering tion between flower color and pigment composition of man and Nugent 1973; Nugent and Hofftulip (Tulip L). Nether J Agric Scl 37365-370. and deVlaming 1984). Our data suggests man 1974; Pitrat et al. 1986,1991; Whitaker that the dominant Hfl allele is linked to 1952; Zink 1977). The ninth mutant, Pale Paris CD and Haney WJ, 1958. Genetic studies In Petuthe recessive phi allele. Flowers that con- nia I. Nine genes for flower color. Am Soc Hort Scl 72. (Pa) green foliage, is conditioned by a nu462^*72. tain cyanidin/peonidin should not possess clear semidominant allele that is lethal Rea PA and Poole RJ, 1993. Vacuolar H+ translocating a recessive phi allele. (white) when homozygous, and viable, but pyrophosphatase. Annu Rev Plant Physlol Plant Mol yellow in color, when heterozygous Blol 44:157-180. Several conclusions can be drawn from (McCreight and Bohn 1979). Stewart RN, Norrts KH, and Asen S, 1975. Mlcrospecthis study. trophotometric measurement of pH and pH effect on Dominant mutants are especially useful 1. The inheritance of specific flower col- color petal epidermal cells. Phytochem 14.937-942. in pollination studies, and now are often ors can be explained through the com- Stotz G, deVlaming P, Schram AW, and Forkmann G, used to test pollen distribution of geneti1985. Genetic and biochemical studies on flavonold-3'bined inheritance of anthocyanin pigmen- hydroxylatlon Inflowersof Petunia hybnda Theor Appl cally engineered plants (Umbeck et al. tation and vacuolar pH. Genet 70:300-305. 1991). In melon, dominant mutations have 2. The inheritance of anthocyanin pig- van Raamsdonk LWD, 1993. Flower pigment composibeen suggested for use in the screening of mentation was controlled by multiple in- tion In Tulipa Genet Res Crop Evol 40:49-54. hybrid seedlings (Foster 1968; Lee and dependent genes (///and Mf) that followed Wiering H, 1974. Genetics of flower color In Petunia hyJanick 1978). brida Genen Phaenen 17:117-134. simple Mendelian genetics. A new chlorophyll-deficient phenotype Wiering H and deVlaming P, 1977. Glycosylatlon and 3. The inheritance of vacuolar pH was methyiatlon patterns of anthocyanlns in Petunia hybri- was found in the breeding lines of Dr. R. E. more complex, being controlled by two in- da II. The genes Mil and Mf2. Z Pflanzenzuchtg 78:113Foster (University of Arizona) and given dependent codominant genes (Phi and 123. to D. T. Ray for genetic analysis. The muPhZ) and being influenced by the cellular Wiering H and deVlaming P, 1984. Inheritance and biotant line was slow-growing, with the cotychemistry of pigments. In: Petunia (Sink K, ed). New environment. ledons and growing tips (leaves, stem, York: Springer-Verlag; 49-76. 4. Linkage of the various pH and anthoand tendrils) yellow in color, but later Received August 31, 1995 cyanin genes prevented the expression of turning green (Figures 1 and 2). We report Accepted December 31, 1995 all of the potential gene combinations. It inheritance studies on this chlorophyll-deCorresponding Editor: Prem P. Jauhar was not possible to obtain seedlings exficient phenotype. pressing cyanidin/peonidin at the least acidic pHs or delphinidin/malvidin at the more acidic pHs. Materials and Methods
27°C (night) and 38°C (day) under natural lighting conditions. Results and Discussion
Figure 1. Morphology of a yellow-lip melon seedling characterized by yellow apices with leaves turning green with maturity.
Cytoplasmic inheritance was determined for the chlorophyll-deficient phenotype. All plants in the F,, F2, and BC, generations possessed the mutant phenotype when a chlorophyll-deficient mutant plant type was the female parent, and were green when the female parent was Top Mark (Table 1). The protocol for genetic nomenclature in the Cucurbitaceae was established by Robinson et al. (1976) and subsequently modified by Wehner et al. (1982). There is, however, no convention for cytoplasmic characters. The system that we used for designating cytoplasmically inherited genes is after R. G. Palmer (personal communication) and Cianzio and Palmer (1992). It has been modified for the Cucurbitaceae (Robinson et al. 1976; Wehner et al. 1982) as follows:
In the present case we have named the mutant yellow-tip and symbolized the gene as cyt-Yt. From the Department of Plant Sciences, University of Arizona, Tucson, AZ 85721 (Ray), and the U.S. Department of Agriculture, Salinas, California (McCreight). The Journal of Heredity 1996:87(3)
Figure 2. Morphology of the yellow-lip melon mutant in which the younger leaves, stems, and tendrils are yellow and turn green as they mature.
Table 1. Numbers of yellow and green melon (Cucumis melo L.) plants in a yellow-tip mutant line and Top Mark, and their F,, F,, and BC, progeny Phenotypic comparisons" Generation
BC,
Pedigree
Families
Yellow
Green
Yellow-tip Top mark Yellow-tip X Top Mark Top Mark X yellow-tip F, (selfed) Yellow-tip x F, (yellow-tip x Top Mark) Top Mark x F, (yellow-tip x Top Mark)
7 1 2 3 8 7 1
69 0 24 0 211 204 0
0 4 0 99 0 0 35
• Yellow = cotyledons and growing tips yellow in color; Green = dark green foliage.
2 4 6 The Journal of Heredity 1996:87(3)
References Cianzio SR and Palmer RG, 1992. Genetics of five cytoplasmically inherited yellow foliar mutants in soybean. J Hered 83:70-73. Cox EL and Harding KE, 1986. Linkage relationships of the light green mutant in cantaloupe. HortScience 21: 940 (abstr). Dyutin KE, 1979. Inheritance of yellow-green coloration of the young leaves in melon (in Russian). Tsitologia i genetika 13:407-408. Foster RE, 1968. F, hybrid muskmelons. J Hered 59:205207. Hoffman JC and Nugent PE, 1973. Inheritance of a virescent mutant of muskmelon. J Hered 64:311-312. Kirkbride JH Jr, 1993. Biosystematic monograph of the genus Cucumis (Cucurbitaceae). Boone, North Carolina: Parkway Publishers. Lee CW and Janick J, 1978. Inheritance of seedling bitterness in Cucumis melo L. HortScience 13:193-194. McCreight JD and Bohn GW, 1979. Descriptions, genet-
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1. The name of the trait should describe a characteristic feature of the mutant type in a minimum number of adjectives and/or nouns. 2. Genes are symbolized by one or more italicized Roman letters prefixed by cyt(also italicized). The first letter of the symbol is capitalized and the same as that of the name.
Ics and Independent assortment of red stem and pale In muskmelon (Cucumis melo L). J Am Soc Hort Scl 104: 721-723. Nugent PE and Hoffman JC, 1974. Inheritance of halo cotyledon mutant In muskmelon. J Hered 65315-316. Pltrat M, 1994. Gene list for Cucumis melo L Cucurbit Genet Coop Rep 17:135-147. Rtrat M, Ferrlere C, and Rlcard M, 1986. Flava, a chlorophyll deficient mutant In muskmelon. Cucurbit Genet Coop Rep 9:67. Pitrat M, Rlsser G, Ferrlere C, Oliver C, and Rlcard M, 1991. Two vlrescent mutants In melon (Cucumis melo L). Cucurbit Genet Coop Rep 14:45. Robinson RW, Munger HM, Whltaker TW, and Bohn GW, 1976. Genes of the Cucurbltaceae. HortSclence 11554568. limbeck PF, Barton KA, Nordhelm EV, McCarty JC, Parrott WL, and Jenkins JN, 1991. Degree of pollen dispersal by Insects from a field test of genetically engineered cotton. J Econ Entomol 84:1943-1950. Wehner TC, McCrelght JD, Henderson WR, John CA, and Robinson RW, 1982. Update of cucurbit gene list and nomenclature rules. Cucurbit Genet Coop Rep 5: 62-66. Whltaker TW, 1952. Genetic and chlorophyll studies of a yellow-green mutant In muskmelon. Plant Physlol 27: 263-268.
Unstable White Flower Color in Groundnut (Arachls hypogaea L.) S. L. Dwlvedi, A. K. Singh, and S. N. Nigam This article summarizes our observations on an unstable white flower color observed in early-generation populations of a cross between two yellow-flowered, truebreeding parents (ICGV 86694 and NC Ac 2821) in groundnut (Arachis hypogaea L). The segregation behavior of white- and chimeric-flowered plants in F2 to Fs generations of the cross did not agree with the conclusions of previous researchers that the white flower color in groundnut was controlled by one to two recessive genes. No cytological abnormality was observed in plants either with white or chimeric flowers. The probable source for this inconsistent segregation for flower color appears to be the presence of an unstable genetic element along with the alleles producing white flower phenotype. The reversion of white flower-color allele to its normal stable form—yellow—occurs at a low frequency, probably due to the excision of this element at the germinal level. When the ex-
Five distinct flower colors (white, yellow, orange, burnt orange, and amber) have been reported in groundnut (Arachis hypogaea L.) (Hayes 1933; John et al. 1954). Of these, yellow and orange are the most common. Both codominance (incomplete dominance) and complete recessiveness are reported for white flower color. Orange (Kumar and Joshi 1943) and yellow (Habib et al. 1980) flower colors in some crosses are incompletely dominant over white flower color with monogenic inheritance. Complete dominance of orange flower color over white flower color with monogenic inheritance is also reported (Hayes 1933). In some other crosses, digenic ratios are reported; 15 yellow to 1 white (Jadhav and Shinde 1979; Patil 1965), and 9 yellow to 6 pale yellow to 1 white with additive gene action (Habib et al. 1980). However, in none of these studies was any observation on the stability of white flower color made. In this article, we report our observations on the unstable white flower color observed in early generations of a cross between two yellowflowered, true-breeding parents. During the 1991 rainy reason (June-October), we observed six white-flowered plants and one plant having white, yellow, and white with yellow sector flowers (from here onward referred to as chimericflowered plant) in an F2 population of 390 plants of a cross between ICGV 86694 and NC Ac 2821. Both parents bred true for yellow flower. ICGV 86694 is a stable, interspecific derivative obtained from a cross between an A. hypogaea line and A. cardenasii. NC Ac 2821, a landrace, was obtained from the North Carolina State University. These seven plants were individually harvested and grown separately in F3 to isolate a true-breeding, white-flower line. The seeds of the chimeric-flowered plant did not germinate. The pooled data of flower color segregation in F3 and F4 generations are given in Tables 1 and 2. In F3 generation of one of the white-flowered F2 plants, only three seeds germinated. These three plants had only yellow flowers. The remaining five white-flowered F2
plants segregated for flower color in the F3 generation. Of the 18 yellow-flowered plants obtained in the F3, only five bred true for yellow flower in the F4 generation. The remaining 13 F3 plants segregated for different flower colors. Whereas the flowers of progeny of seven yellow-flowered F3 plants had all the three color patterns (yellow, white, and chimeric), the flowers of progeny of the remaining plants had only two (yellow and chimeric in the case of four plant progeny, and yellow and white in the case of two plant progeny). Except for one white-flowered plant that possibly bred true for flower color in the F< generation (only one plant), the remaining white- and chimeric-flowered F3 plants segregated for flower color patterns. In the F4, only white- and chimeric-flowered plants produced by white-Dowered F3 plants were harvested and grown individually in the F5 generation. A few of the progeny failed to germinate. Forty-nine progeny of the white-flowered plants bred true for flower color in the F5 generation and the remainder again segregated (Table 3). Among the progeny of chimeric-
Table 1. Segregation for flower color In the F, generation of the cross ICGV 86694 x NC Ac 2821 In groundnut Whiteflowered F, plant
Number of F5 plants Yellow White Chimeric flower flower flower Total
PI P2 P3 P4 P5 P6 Total
5 1 1 8 0 3 18
3 5 16 16 7 0
15 8 24 37 10 3 97
7 2 7 13
3 0 32
47
Table 2. Segregation for flower color In the F, generation of the cross ICGV 86694 x NC Ac 2821 in groundnut Num-
Flower color pattern of Fj plant White
Total Chimeric
Total Yellow
Total
Number of F
