Download Yellow-Tip: A Cytoplasmically Inherited Trait in Melon (Cucumis

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
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
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:
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.
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-
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.
Zlnk FW, 1977. Linkage of vlrescent foliage and plant
growth habit In muskmelon. J Am Soc Hort Scl 102.613615.
Received July 24, 1995
Accepted December 31, 1995
Corresponding Editor: William F. Tracy
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-
cision occurs at the somatic level, there is
a partial reversion of white-flower color allele giving rise to yellow, white, or chimeric
flowers on the same plant. Our efforts in
two subsequent generations to stabilize
white-flowered plants did not succeed.
Further studies are required to get at the
source of this unstable activity of alleles
responsible for white flower color phenotype.
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<1slants
ber
of F,
Chimeproge- Yellow White ric
flower flower flower Total
ny
31
1
13
1
1
47
23
4
3
30
7
2
4
5
18
85
2
0
1
0
88
142
22
13
177
68
21
43
72
204
326
2
95
0
1
424
100
8
0
108
17
4
0
0
21
265
0
51
1
0
317
137
0
7
144
22
0
15
0
3
676
4
146
2
1
829
379
30
20
429
107
25
58
72
262
Brief Communications 2 4 7