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Gene List for Watermelon

2007

  • by Todd C. Wehner
    • Department of Horticultural Science
    • North Carolina State University
    • Raleigh, NC 27695-7609

Abstract

Watermelon (Citrullus lanatus (Thunb.) Matsum. & Nakai) is a major vegetable crop in the world, accounting for 6.8% of the world area devoted to vegetable crops. Watermelon is a useful vegetable crop for genetic research because of its small genome size, and the many available gene mutants. The watermelon genes were originally organized and summarized in 1944, and have been expanded and updated periodically. However, the action of some watermelon genes has not been described clearly in some cases. Also, the interaction of multiple gene loci that control similar traits needs to be described more clearly. Finally, it is necessary to identify the inbred lines having each published gene mutant, for use as type lines in studies of gene action, allelism, and linkage. The objective of this work was to update the gene list, identify the cultivar or line having each gene mutant, and collect seeds of the lines for use by interested researchers. In addition, the gene descriptions were expanded and clarified, information on gene interactions was added, and errors in naming or citing previously described genes were corrected. New genes that have not previously been described (cr, Ctr, dw-3, eg, ms-2, Ti, ts and zym-FL) were added to the list, for a total of 163 watermelon gene mutants.

Introduction

Watermelon (Citrullus lanatus (Thunb.) Matsum. & Nakai) is a major cucurbit crop that accounts for 6.8% of the world area devoted to vegetable crops (FAO, 2002). Watermelon is grown for its fleshy, juicy, and sweet fruit. Mostly eaten fresh, they provide a delicious and refreshing dessert especially in hot weather. The watermelon has high lycopene content in the red-fleshed cultivars: 60% more than tomato. Lycopene has been classified as a useful in the human diet for prevention of heart attacks and certain types of cancer (Perkins-Veazie et al., 2001).

Watermelon is native to central Africa where it was domesticated as a source of water, a staple food crop, and an animal feed. It was cultivated in Africa and the Middle East for more than 4000 years, then introduced to China around 900 AD, and finally brought to the New World in the 1500s. There are 1.3 million ha of watermelon grown in the world, with China and the Middle Eastern countries the major consumers. China is the largest watermelon producer, with 68.9% of the total production. The other major watermelon producing countries are Turkey, Iran, Egypt, United States, Mexico and Korea (FAO, 2002). In the United States, watermelon is used fresh as a dessert, or in salads. U.S. production is concentrated in Florida, California, Texas, and Georgia (USDA, 2002), increasing from 1.2 M tons in 1980 to 3.9 M tons in 2002, with a farm value of $329 million (USDA, 2002).

Watermelon is a useful crop species for genetic research because of its small genome size, and the many available gene mutants. Genome size of watermelon is 424 million base pairs (Arumuganathan and Earle 1991). DNA sequence analysis revealed high conservation useful for comparative genomic analysis with other plant species, as well as within the Cucurbitaceae (Pasha 1998). Like some of the other cultivated cucurbits, watermelon has much genetic variability in seed and fruit traits. Genetic investigations have been made for some of those, including seed color, seed size, fruit shape, rind color, rind pattern, and flesh color.

This is the latest version of the gene list for watermelon. The watermelon genes were originally organized and summarized by Poole (1944). The list and updates of genes for watermelon have been expanded and published by Robinson et al. (1976), the Cucurbit Gene List Committee (1979, 1982, and 1987), Henderson (1991 and 1992), Rhodes and Zhang (1995), and Rhodes and Dane (1999). This current gene list provides an update of the known genes of watermelon, with 163 total mutants grouped into seed and seedling mutants, vine mutants, flower mutants, fruit mutants, resistance mutants, protein (isozyme) mutants, DNA (RFLP and RAPD) markers, and cloned genes.

Researchers are encouraged to send reports of new genes, as well as seed samples of lines containing the gene mutant to the watermelon gene curator (Todd C. Wehner), or to the assistant curator (Stephen R. King). Please inform us of omissions or errors in the gene list. Scientists should consult the list as well as the rules of gene nomenclature for the Cucurbitaceae (Cucurbit Gene List Committee, 1982; Robinson et al., 1976) before choosing a gene name and symbol. Please choose a gene name and symbol with the fewest characters that describes the recessive mutant, and avoid use of duplicate gene names and symbols. The rules of gene nomenclature were adopted in order to provide guidelines for naming and symbolizing genes. Scientists are urged to contact members of the gene list committee regarding rules and gene symbols. The watermelon gene curators of the Cucurbit Genetics Cooperative are collecting seeds of the type lines for use by interested researchers, and would like to receive seed samples of any of the lines listed.

This gene list has been modified from previous lists in that we have 1) added or expanded the description of the phenotypes of many of the gene mutants, 2) added descriptions for phenotypes of interacting gene loci, 3) identified the type lines that carry each form of each gene, 4) identified the gene mutant lines that are in the curator collections, and 5) added genes that have not previously been described: cr (El-Hafez et al., 1981), Ctr (Provvidenti, 1992), dw-3 (Hexun et al., 1998), eg (Gusmini et al., 2004), ms-2 (Dyutin, and Sokolov, 1990), Ti (Tanaka et al., 1995), ts (Zhang et al., 1994a), and zym-FL (Provvidenti, 1991). We had intended to include a review of gene linkage, but few reports were found except for sets of molecular markers in wide crosses of Citrullus. Finally, we attempted to correct some of the errors in gene descriptions or references from previous lists.

Previous Gene Lists

  • Poole, 1944: 15 genes total
  • Robinson et al., 1976: 10 genes added, 25 genes total
  • Robinson et al., 1979: 3 genes added, 28 genes total
  • Robinson et al., 1982: 2 genes added, 30 genes total
  • Henderson, 1987: 3 genes added, 33 genes total
  • Henderson, 1991: 3 genes added, 36 genes total (plus 52 molecular markers)
  • Rhodes and Zhang, 1995: 3 genes added, 39 genes total (plus 109 molecular markers)
  • Rhodes and Dane, 1999: 5 genes added, 44 genes total (plus 111 molecular markers)
  • Guner and Wehner, 2003: 8 genes added, 52 genes total (plus 111 molecular markers)

Gene Mutants

Seed and seedling mutants

Three major genes control seed coat color: r (Poole et al., 1941), t (McKay, 1936), and w (Poole et al., 1941) for red, tan, and white seed coat, respectively. The genes interact to produce six phenotypes: black (RR TT WW); clump (RR TT ww); tan (RR tt WW); white with tan tip (RR tt ww); red (rr tt WW); and white with pink tip (rr tt ww). A fourth gene, d was suggested by Poole et al. (1941) as a modifier, producing black dotted seed coat when dominant for r, t, and w but having no effect on other genotypes.

The genes (s) and (l) for short and long seed length (sometimes called small and large seed size) control seed size, with s epistatic to l (Poole et al., 1941). The genotype LL SS gives medium size, ll SS gives long, and LL ss or ll ss gives short seeds. The Ti gene for tiny seed was reported by Tanaka et al. (1995). Tiny seed from 'Sweet Princess' was dominant over medium-size seed and controlled by a single dominant gene. The small seed gene behaved in a manner different from Poole's medium-size seed cultivar. Tanaka et al. (1995) suggested that the Ti gene was different from the s and l genes. Unfortunately, the origin of short and long seed genes was not described in Poole's paper.

Tomato seed is shorter and narrower than the short seeded genotype, ll ss (width x length: 2.6 x 4.2 mm). It is controlled by the ts (Zhang et al., 1994a) gene, with genotype LL ss tsts. Cracked seed coat cr (El-Hafez et al., 1981) is inherited as a single gene that is recessive to noncracked seed coat. Egusi seed eg (Gusmini et al., 2004) has fleshy pericarp covering the seeds. After washing and drying, the seeds are difficult to distinguish from normal.

Vine mutants

Several genes control leaf or foliage traits of watermelon. Nonlobed leaf (nl) has the entire leaf rather than the lobed leaf type of the typical watermelon (Mohr, 1953). Seedling leaf variegation slv (Provvidenti, 1994) causes a variegation resembling virus infection on seedlings. It is linked or pleiotropic with Ctr for cool temperature resistance. The yellow leaf (Yl) gene results in yellow leaves, and is incompletely dominant to green leaves (Warid and Abd-El-Hafez, 1976). Delayed green leaf dg (Rhodes, 1986) causes pale green cotyledons and leaves for the first few nodes, with later leaves developing the normal green color. Inhibitor of delayed green leaf (i-dg) makes leaves normal green even when they have dgdg genotype (Rhodes, 1986). The juvenile albino ja (Zhang et al., 1996b) gene causes reduced chlorophyll in seedling tissues, as well as leaf margins and fruit rind when plants are grown under short day conditions. The dominant gene Sp (Poole, 1944) causes round yellow spots to form on cotyledons, leaves and fruit, resulting in the fruit pattern called moon and stars.

So far, four dwarf genes of watermelon have been identified that affect stem length and plant habit: dw-1 (Mohr, 1956) and dw-1s (Dyutin and Afanas'eva, 1987) are allelic, and dw-1, dw-2 (Liu and Loy, 1972), and dw-3 (Hexun et al., 1998) are non-allelic. Dwarf-1 plants have short internodes due to fewer and shorter cells than the normal plant type. Plants with dw-1s have vine length intermediate between normal and dwarf, and the hypocotyls were somewhat longer than normal vine and considerably longer than dwarf. The dw-1s is recessive to normal plant type. Plants with dw-2 have short internodes due to fewer cells than the normal type, and plants with dw-3 have leaves with fewer lobes than the normal leaf.

The golden yellow mutant is controlled by the single recessive gene go, where the stem and older leaves are golden yellow (Barham, 1956). The gene go was from 'Royal Golden'. One benefit of the go gene is that the fruit become golden yellow as they mature, possibly useful as a maturity indicator. The gene tl (formerly called branchless, bl) results in tendrilless branches after the 5th or 6th node (Rhodes et al., 1999 and Zhang et al., 1996a). Also, plants have half the number of branches of the normal plant type, vegetative meristems gradually become floral, tendrils and vegetative buds are replaced by flowers (with a large percentage being perfect), and growth becomes determinate.

Flower mutants

The andromonoecious gene a (Rosa, 1928) controls monoecious (AA) vs. andromonoecious (aa) sex expression in watermelon. Andromonoecious plants have both staminate and perfect flowers, and appears to be the wild type. Light green flower color is controlled by the single recessive gene, gf (Kwon and Dane, 1999).

Four genes for male sterility have been reported. Glabrous male sterile (gms) is unique, with sterility associated with glabrous foliage (Watts, 1962 and 1967). A second male sterile ms-1 (Zhang and Wang, 1990) produces plants with small, shrunken anthers and aborted pollen. A third male sterile mutant appeared simultaneously with dwarfism, and the dwarf gene was different from the three known dwarf genes. It was named male sterile dwarf (ms-dw) by Huang et al. (1998). All male sterile genes reduce female fertility as well. These mutants have been used in hybrid production, but have not been as successful as hoped, since they often have low seed yield. A new, spontaneous male sterile mutant (ms-2) with high normal seed set has been identified, and will be more useful for hybrid production (Dyutin, and Sokolov, 1990).

Fruit mutants

Considerable attention has been given to genes affecting fruit type in watermelon. A single recessive gene su (Chambliss et al., 1968) eliminates bitterness in fruit of C. lanatus, and appears to be the same as the dominant gene (Su) for bitter flavor in the fruit of the colocynth (Citrullus colocynthis).

Fruit shape is controlled by a single, incompletely dominant gene, resulting in fruit that are elongate (OO), oval (Oo), or spherical (oo) (Weetman, 1937). A single gene controls furrowed fruit surface f (Poole, 1944) that is recessive to smooth (F). Explosive rind (e) causes the fruit rind to burst or split when cut (Porter, 1937). The non-explosive genotype (EE) would be important to improve shipping ability for marketing.

Thirteen mutants have been identified that affect fruit color either in the skin or flesh. A single gene determines the intensity of green color of the fruit skin, with solid light green (g) skin color recessive to solid dark green (G) (Weetman, 1937). Green striped skin is controlled by a single gene g-s (Weetman, 1937) that is recessive to dark green, but dominant to light green. The symbol g-s for this gene was proposed, although there are narrow, medium, and wide striped patterns that were not explained by qualitative genes so far. Another type of striping of the fruit skin is where the stripes are very narrow and inconspicuous. The trait is controlled by a single recessive gene inherited independently of g-s, called pencilled lines p (Weetman, 1937). The greenish white mottling of the fruit skin is controlled by the single recessive gene m (Weetman, 1937), resulting in gray types, such as 'Iowa Belle' and 'Charleston Gray'. The gene Sp produces spotted fruit, making interesting effects as found on the cultivar 'Moon and Stars' (Poole, 1944).

Watermelon flesh color is controlled by several genes to produce red, orange, salmon yellow, canary yellow, or white. Genes conditioning flesh colors are B (Shimotsuma, 1963), C (Poole, 1944), i-C (Henderson et al., 1998), Wf (Shimotsuma, 1963), y (Porter, 1937) and y-o (Henderson, 1989 and Henderson et al. 1998). Canary yellow (C) is dominant to red flesh (c). Red flesh (Y) is dominant to salmon yellow (y). Orange flesh (y-o) is a member of multiple allelic system at that locus, where Y (red flesh) is dominant to both y-o (orange flesh) and y salmon yellow (salmon yellow), and y-o (orange flesh) is dominant to y (salmon yellow). In a separate study, two loci with epistatic interaction controlled white, yellow and red flesh. Yellow flesh (B) is dominant to red flesh. The gene Wf is epistatic to B, so genotypes WfWf BB or WfWf bb were white fleshed, wfwf BB was yellow fleshed, and wfwf bb was red fleshed. Canary yellow flesh is dominant to red, and i-C inhibitory to C, resulting in red flesh. In the absence of i-C, C is epistatic to Y.

Golden yellow was inherited as a single recessive gene go (Barham, 1956) derived from 'Royal Golden' watermelon. The immature fruit had a dark green rind which becomes more golden yellow as the fruit matures. The stem and older leaves also become golden yellow, and the flesh color changes from pink to red.

Resistance mutants

Resistance to race 1 (and 3) of anthracnose (Colletotrichum lagenarium, formerly Glomerella cingulata var. orbiculare) is controlled by a single dominant gene Ar-1 (Layton 1937). Resistance to race 2 of anthracnose is also controlled by a single dominant gene Ar-2-1 (Winstead et al., 1959). Resistance to race 1 of Fusarium oxysporum f. sp. niveum is controlled by a single dominant gene Fo-1 (Henderson et al., 1970). Gummy stem blight, caused by Didymella bryoniae (Auersw.) Rehm is inherited by a recessive gene db (Norton, 1979). Most watermelons are resistant to races of Sphaerotheca fuliginea present in the U.S., but a single recessive gene pm (Robinson et al., 1975) for susceptibility to powdery mildew was found in the plant introduction, PI 269677. A high level of resistance to Zucchini yellow mosaic virus was found in four landraces of Citrullus lanatus, but was specific to the Florida strain of the virus. Resistance was conferred by a single recessive gene zym-FL (Provvidenti, 1991).

Genes for insect resistance have been reported in watermelon. Fruit fly (Dacus cucurbitae) resistance was controlled by a single dominant gene Fwr (Khandelwal and Nath, 1978), and red pumpkin beetle (Aulacophora faveicollis) resistance was controlled by a single dominant gene Af (Vashishta and Choudhury, 1972).

Stress resistance has been found in watermelon. Seedlings grown at temperatures below 20°C often develop a foliar mottle and stunting. A persistent low temperature is conducive to more prominent foliar symptoms, malformation, and growth retardation. The single dominant gene Ctr was provided cool temperature resistance (Provvidenti, 1992).

The morphological and resistance genes of watermelon, including gene symbol, synonym, description, references, availability (y), and photograph.(z)

Symbol
Synonym
Gene description and type lines
References
Supplemental references
Availability
Photograph (click for larger image)
 a
-
andromonoecious; recessive to monoecious; a from 'Angeleno' (black seeded); A from cultivars 'Conqueror' and 'Klondike'.
Rosa, 1928
Porter, 1937; Poole, 1944
C
a
 Af
-
Aulacophora faveicollis resistance; resistance to the red pumpkin beetle; dominant to susceptibility; Af from Sl.72 and Sl.98 inbreds; af from 'Sugar Baby'.
 Vashishta and Choudhury, 1972
 -
 M
Af
 Ar-1
B, Gc
Anthracnose resistance to races 1 and 3 of Glomerella cingulata var. orbiculare (Colletotrichum lagenarium); Ar-1 from 'Africa 8'*, 'Africa 9'*, and 'Africa 13'* and 'Charleston Gray'**; ar-1 from 'Iowa Belle 476', 'Iowa Belle 487'* and N.C.9-2, N.C.11, and 'New Hampshire Midget'**.
 Layton 1937*
 Hall et al., 1960; Robinson et al., 1976; Winstead et al., 1959**
 C
 Ar-2-1
 -
Anthracnose resistance to race 2 of Colletotrichum lagenarium; Ar-2-1 from W695 citron* and PI 189225, PI 271775, PI 271779, and PI 299379**; ar-2-1 from 'Allsweet', 'Charleston Gray', and 'Florida Giant'; resistance in Citrullus colocynthis is due to other dominant factors; resistance from R309***; susceptibility from 'New Hampshire Midget'.
 Winstead et al., 1959*
 Love and Rhodes, 1988***, 1991; Sowell et al., 1980**; Suvanprakorn and Norton, 1980
 P
 B
Y
Yellow flesh; Wf is epistatic to B (Y renamed B by Henderson*); flesh color segregated into 12 white, 3 yellow and 1 red in the F2; WfWf BB or WfWf bb white fleshed; wfwf BB yellow fleshed; wfwf bb red fleshed; B from breeding line V.No.3 and b from V.No.1.
 Shimotsuma, 1963
 Henderson, 1992*
 ?
 C
 -
Canary yellow flesh; dominant to pink; i-C inhibitory to C, resulting in red flesh; in the absence of i-C, C is epistatic to Y; CC from 'Honey Cream'* and NC-517, cc from 'Dove'*; CC YY I-C I-C from 'Yellow Baby' F1** and 'Yellow Doll' F1**; cc yoyo I-C I-C from 'Tendersweet Orange Flesh'**; cc yy I-C I-C from 'Golden Honey'**; cc YY i-C i-C from 'Sweet Princess'**.
 Poole, 1944*
 Henderson et al., 1998**
 C
 cr
 
 cracked seed coat; recessive to Cr (non-cracked) seed coat; cr from 'Leeby' and Cr from 'Kaho' and 'Congo'.
 El-Hafez et al., 1981
 -
 ?
 cr
 Ctr
 
Cool temperature resistance; Ctr from line PP261-1 (a single plant selection of PI 482261 from Zimbabwe); ctr from 'New Hampshire Midget'; resistant to leaf mosaic injury when grown at air temperature below 20°C.
 Provvidenti, 1992
 Provvidenti, 2003
 P
 Ctr
 d
 -
dotted seed coat; black dotted seeds when dominant for color genes r, t, and w; d is a specific modifier of black seed coat color wherein RR TT WW DD is solid black and RR TT WW dd is dotted black seed coat; d from 'Klondike' and 'Hope Giant'; D from 'Winter Queen'.
 Poole et al., 1941
 Poole, 1944; Kanda, 1951
 C
 db 
 -
resistance to gummy stem blight caused by Didymella bryoniae; db from PI 189225; Db from 'Charleston Gray'.
 Norton, 1979
 -
 P
db 
 dg
 
delayed green; cotyledons and young leaves are initially pale green but later develop chlorophyll; first reported to be hypostatic to I-dg; more recent evidence indicates a simple recessive; dg from breeding line 'Pale 90'; Dg from 'Allsweet'.
 Rhodes, 1986
 -
 ?
 dg
 dw-1
 -
dwarf-1; short internodes, due to fewer and shorter cells than normal forms; allelic to dw-1s; dw-1 from 'Bush Desert King' (also, 'Bush Charleston Gray', 'Bush Jubilee', 'Sugar Bush'); Dw-1 from 'Sugar Baby' and 'Vine Desert King'.
 Mohr, 1956
 Liu and Loy, 1972
 C
 dw-1
 dw-1-s
 -
short vine; allelic to dw-1; vine length intermediate between normal and dwarf; hypocotyl somewhat longer than normal vine and considerably longer than dwarf; dw-1-s recessive to normal; dw-1-s from 'Somali Local' (All-Union Research Institute of Plant Growing No.4641).
 Dyutin and Afanas'eva, 1987
 -
 ?
 dw-1-s
 dw-2
 -
dwarf-2; short internodes, due to fewer cells; dw-2 from inbred line WB-2; Dw-2 from 'Sugar Baby' and 'Vine Desert King'.
 Liu and Loy, 1972
 Mohr and Sandhu, 1975
 ?
 dw-2
 dw-3
 -
dwarf-3; dwarf with fewer leaf lobes (intermediate between normal leaf and non-lobed leaf); dw-3 from 'Dwarf Male-Sterile Watermelon (DMSW)'; Dw-3 from 'Changhui', 'Fuyandagua', and 'America B'.
 Hexun et al., 1998
 -
 ?
 dw-3
 e
 t
explosive rind; thin, tender rind, bursting when cut; e from 'California Klondike'; E from 'Thurmond Gray'.
 Porter, 1937
 Poole, 1944
 ?
 eg
 -
egusi seed; immature seeds with fleshy pericarp, becoming normal at maturity; eg from PI 490383 selection NCG-529 and PI 560006; Eg from 'Calhoun Gray' and 'Charleston Gray'.
 Gusmini et al., 2003
 -
 C
 f
 -
 furrowed fruit surface; recessive to smooth; type inbreds not given; f like 'Stone Mountain' or 'Black Diamond'; F like 'Mickylee'.
 Poole, 1944
 -
 M
  f
 Fo-1
 -
Fusarium wilt resistance for race 1; dominant gene for resistance to race 1 of Fusarium oxysporum f. sp. niveum; Fo-1 from 'Calhoun Gray' and 'Summit'; fo-1 from 'New Hampshire Midget'.
 Henderson et al., 1970
 Netzer and Weintall, 1980
 C
 Fo-1
 Fwr
 -
Fruit fly resistance caused by Dacus cucurbitae; dominant to susceptibility; Fwr from breeding lines J 18-1 and J 56-1; fwr from 'New Hampshire Midget', 'Bykovski', 'Red Nectar' and breeding line 'J 20-1'.
 Khandelwal and Nath, 1978
 -
 ?
  Fwr
 g
 d
light green fruit rind pattern; light green fruit recessive to dark green (G) and striped green (g-s); g from 'Thurmond Gray' and G from 'California Klondike'.
 Weetman, 1937
 Poole, 1944; Porter, 1937
 ?
 g
 g-s
 ds
striped green fruit rind pattern; recessive to dark green but dominant to light green skin; g-s from 'Golden Honey'; G from 'California Klondike'.
 Weetman, 1937
 Poole, 1944
 C
 gf
 -
light green flower color; gf from 'KW-695' and 'Dalgona'; Gf from Korean watermelon accession 'SS-4'.
 Kwon and Dane, 1999
 -
 ?
 gf
 gms
 msg
glabrous male sterile; foliage lacking trichomes; male sterile caused by chromosome desynapsis (named glabrous male sterile by Robinson*); gms from 'Sugar Baby' irradiated with gamma rays.
 Watts, 1962, 1967
 Robinson et al., 1976*; Ray and Sherman, 1988
 ?
 gms
 go
 c
golden yellow color of older leaves and mature fruit; (named golden by Robinson*); go from 'Royal Golden'; Go from 'NC 34-9-1' and 'NC 34-2-1'.
 Barham, 1956
 Robinson et al., 1976*
 C
 i-C
 i
inhibitor of canary yellow, resulting in red flesh (renamed by Rhodes and Dane*); CC YY I-C I-C from 'Yellow Baby' F1 and 'Yellow Doll' F1; cc yoyo I-C I-C from 'Tendersweet Orange Flesh'; cc yy I-C I-C from 'Golden Honey'; cc YY i-C i-C from 'Sweet Princess'.
 Henderson et al., 1998
 Rhodes and Dane, 1999*
 C
 i-dg
 -
inhibitor of delayed green; Epistatic to dg; I-dg I-dg dgdg plants are pale green; and i-dg i-dg dgdg plants are normal; dg from breeding line Pale 90; Dg from 'Allsweet'; i-dg gene was lost when advanced inbreds were made.
Rhodes, 1986
 -
 L
  i-dg
 ins
 -
intermittent stripes; narrow dark stripes at the peduncle end of the fruit becoming irregular in the middle and nearly absent at the blossom end of the fruit; ins from 'Navajo Sweet'; Ins from 'Crimson Sweet'.
 Gusmini and Wehner, in press
 -
 C
 ja
 -
 juvenile albino; chlorophyll in seedlings, leaf margins, and fruit rind reduced when grown under short days; ja from 'Dixielee mutant' and 'G17AB' F2; Ja from 'Sweet Princess' and '20J57'.
 Zhang et al., 1996b
 -
 ?
 ja
 l
 -
long (or large) seeds; interacts with s; long recessive to medium or short; LL SS for medium, ll SS for long, and LL ss or ll ss for short seed; ll SS from 'Peerless'; LL SS from 'Klondike'; LL ss from 'Baby Delight'.
 Poole et al., 1941
 -
 ?
 m
 -
mottled skin; greenish white mottling of fruit skin; randomly-distributed, irregularly-shaped light green spots on a mostly solid dark-green rind pattern; m from 'Long Iowa Belle' (seeds not available) and 'Round Iowa Belle' (seeds not available); M from 'Japan 4' (seeds not available) and 'China 23' (seeds not available).
 Weetman, 1937
 Poole, 1944
 ?
 ms-1
 ms
male sterile; plants with small, shrunken anthers and aborted pollen; ms-1 from 'Nongmei 100'; Ms from most cultivars, e.g. 'Allsweet'.
 Zhang and Wang, 1990
 Zhang et al., 1994b
 ?
 ms-1
 ms-dw
 -
male sterile, dwarf; ms-dw from 'Dwarf Male-Sterile Watermelon (DMSW)'; Ms-dw from 'Changhui', 'Fuyandagua', and 'America B'.
 Huang et al., 1998
 -
 ?
ms-dw
 ms-2
 
male sterile with high seed productivity; ms-2 from 'Kamyzyakskii'; Ms-2 from cultivars like 'Allsweet'.
 Dyutin, and Sokolov, 1990
 -
 ?
ms-2
 nl
 -
nonlobed leaves; leaves lack the typical lobing; sinuate leaves (named nonlobed by Robinson*); leaves lack the typical lobing of most cultivars, slightly lobed with the sinus obscure; incomplete dominance; Nl is not sinuate, but pinnatifid (deeply pinnately lobed, with prominent sinuses) like most cultivars; nl from spontaneous mutant of 'Black Diamond', and probably 'Sunshade; Nl from 'Black Diamond', and most cultivars such as 'Allsweet' and 'Calhoun Gray'.
 Mohr, 1953
 Robinson et al., 1976*
 C
 O
 
Elongate fruit; incompletely dominant to spherical, so that Oo is oval; O from 'Long Iowa Belle'; o from 'Round Iowa Belle', 'China 23', 'Japan 4', and 'Japan 6'.
 Weetman, 1937
 Poole and Grimball, 1945
 ?
 p
 -
pencilled lines on skin; inconspicuous stripes; greenish-white mottling* (called pencilled by Robinson**); inconspicuous, very narrow, pencil-width stripes running the length of the fruit (originally spelled penciled by Poole); recessive to netted fruit; p from 'Japan 6' (seeds not available) and P from 'China 23' (seeds not available).
 Weetman, 1937*
 Robinson et al., 1976**
?
 pm  - powdery mildew susceptibility; susceptibility to Sphaerotheca fuliginea is recessive; pm from PI 269677; Pm from 'Sugar Baby' and most cultivars.  Robinson et al., 1975  -  P
 r   red seed coat; genes r, t and w interact to produce seeds of different colors; black from 'Klondike' (RR TT WW); clump from 'Sun Moon and Stars' (RR TT ww); tan from 'Baby Delight' (RR tt WW); white with tan tip from 'Pride of Muscatine' (RR tt ww); red from citron (rr tt WW); white with pink tip from 'Peerless' (rr tt ww).  Poole et al., 1941  -  ?
 s - short (or small) seeds; epistatic to l; long recessive to medium or short; LL SS for medium, ll SS for long, and LL ss or ll ss for short seed; ll SS from 'Peerless'; LL SS from 'Klondike'; LL ss from 'Baby Delight'.  Poole et al., 1941  - ?
 Scr
 -
Scarlet red flesh color; dark red color of the fruit flesh (darker red than the YY red color of 'Angeleno Black Seeded'); Scr from 'Dixielee' and 'Red-N-Sweet'; scr from 'Angeleno Black Seeded'.
 Gusmini and Wehner, in press
 -
 C
 slv  - seedling leaf variegation; conferred by a single recessive gene in PI 482261; linked or pleiotropic with a dominant allele for resistance to cool temperature injury (20°C for greenhouse-grown plants); slv from PI 482261 (resistant to ZYMV-FL); Slv from 'New Hampshire Midget'.  Provvidenti, 1994  -  P   slv
 Sp  - Spotted cotyledons, leaves and fruit; dominant to uniform foliage and fruit color; Sp from 'Sun, Moon and Stars'* and 'Moon and Stars'**; sp from 'Allsweet'.  Poole, 1944* Rhodes, 1986**  C
 su  Bi, suBi suppressor of bitterness; (su named by Robinson*); non-bitter fruit; su from 'Hawkesbury'; Su from bitter-fruited mutant of 'Hawkesbury'; bitterness in C. colocynthis is due to Su Su genotype.  Chambliss et al., 1968  Robinson et al., 1976*  ? su
 t  bt tan seed coat; genes r, t and w interact to produce seeds of different colors; black from 'Klondike' (RR TT WW); clump from 'Sun Moon and Stars' (RR TT ww); tan from 'Baby Delight' (RR tt WW); white with tan tip from 'Pride of Muscatine' (RR tt ww); red from citron (rr tt WW); white with pink tip from 'Peerless' (rr tt ww).  McKay, 1936  Poole et al., 1941  ?
 Ti  - Tiny seed; dominant over medium seed (ti); Ti from 'Sweet Princess'; ti from 'Fujihikari'.  Tanaka et al., 1995  -  ?
 tl  bl tendrilless (formerly called branchless*), after 4th or 5th node, vegetative axillary buds are transformed into flower buds and leaf shape is altered; tl from 'Early Branchless'; Tl from breeding lines 'G17AB', 'ASS-1', 'YF91-1-2', and S173 breeding line.  Rhodes, Zhang, Baird and Knapp, 1999; Zhang, Rhodes, Baird and Skorupska, 1996a  Lin, Tong, Wang, Zhang and Rhodes, 1992*  ?  tl
 ts  tss  tomato seed; seeds smaller than short (LLss or llss), almost the size of a tomato seed; ts from tomato seed Sugar Baby mutant; Ts from 'Gn-1'.  Zhang et al., 1994a  Zhang, 1996  C
 w   white seed coat; genes r, t and w interact to produce seeds of different colors; black from 'Klondike' (RR TT WW); clump from 'Sun Moon and Stars' (RR TT ww); tan from 'Baby Delight' (RR tt WW); white with tan tip from 'Pride of Muscatine' (RR tt ww); red from citron (rr tt WW); white with pink tip from 'Peerless' (rr tt ww).  Poole et al., 1941  -  ?
 Wf  W White flesh; (named white flesh by Robinson*); Wf is epistatic to B (Y renamed B by Henderson**); WfWf BB or WfWf bb white fleshed; wfwf BB yellow fleshed; wfwf bb red fleshed; B from breeding line V.No.3 and b from V.No.1; flesh color segregated into 12 white, 3 yellow and 1 red in the F2.  Shimotsuma, 1963  Robinson et al., 1976*; Henderson, 1992**  ?
 y  rd yellow flesh; recessive to coral (light) red flesh (Y); y from 'Golden Honey'; Y from 'Angeleno' (black seeded).  Porter, 1937  Poole, 1944; Henderson, 1989; Henderson et al., 1998  C
 y-o  - orange flesh; allelic to y; Y (red flesh) is dominant to y-o (orange flesh) and y (salmon yellow flesh); y-o (orange flesh) is dominant to y (yellow flesh); cc y-oy-o I-C I-C from 'Tendersweet Orange Flesh'; cc yy I-C I-C from 'Golden Honey'; cc YY i-C i-C from 'Sweet Princess'.  Henderson, 1989; Henderson et al., 1998  Poole, 1944; Porter, 1937 C
 Yb
 -
yellow belly; yellow colored ground spot on the fruit; Yb from 'Black Diamond Yellow Belly'; yb from 'Black Diamond'.
 Gusmini and Wehner, in press
 -
 C
 Yl  Y Yellow leaf; incompletely dominant to green leaf (yl); (Y renamed Yl by Henderson*). Yl from 'Yellow Skin'. Warid and Abd-El-Hafez, 1976  Henderson, 1991*  ?  Yl
 zym-FL  zym Resistance to zucchini yellow mosaic virus (ZYMV-FL); resistance is specific to the Florida strain; zym-FL from PI 482322, PI 482299, PI 482261, and PI 482308, Zym-FL from elite cultivars. Provvidenti, 1991  -  P

z Asterisks on cultigens and associated references indicate the source of information for each.
y C = Mutant available from Cucurbit Genetics Cooperative watermelon gene curator; M = molecular marker or isozyme; P = mutants are available as standard cultivars or accessions from the plant introduction collection; ? = availability not known; L = mutant has been lost.

 

The isozymes and molecular markers for watermelon, including gene symbol, synonym, description, references and availability.(y)

Symbol
Synonym
Gene description and type lines
References
Supplemental references
Availability
 Aco-1
 -
 Aconitase-1.
 Navot et al., 1990
 -
 M
 Aco-2
 -
 Aconitase-2.
 Navot et al., 1990
 -
 M
 Adh-1
 -
 Alcohol dehydrogenase-1; one of five codominant alleles, each regulating one band
 Navot and Zamir 1986, 1987; Zamir et al., 1984
 -
 M
 Adh-1-1
 -
Alcohol dehydrogenase-1-1; one of five codominant alleles, each regulating one band; found in C; lanatus var. citroides and C. colocynthis.
 Navot and Zamir 1986, 1987; Zamir et al., 1984
 -
 M
 Adh-1-2
 -
 Alcohol dehydrogenase-1-2; one of five codominant alleles, each regulating one band; found in C. lanatus var. citroides and C. colocynthis.
 Navot and Zamir 1986, 1987; Zamir et al., 1984
 -
 M
 Adh-1-3
 -
 Alcohol dehydrogenase-1-3; one of five codominant alleles, each regulating one band; found in Praecitrullus fistulosus.
 Navot and Zamir 1986, 1987; Zamir et al., 1984
 -
 M
 Adh-1-4
 -
 Alcohol dehydrogenase-1-4; one of five codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.
 Navot and Zamir 1986, 1987; Zamir et al., 1984
 -
 M
 Aps-1
 -
 Acid phosphase-1.
 Navot et al., 1990; Navot and Zamir 1986, 1987; Zamir et al., 1984
 -
 M
 Aps-2-1
 -
 Acid phosphatase-2-1; one of two codominant alleles, each regulating one band; found in C. lanatus and C. colocynthis.
 Navot et al., 1990; Navot and Zamir 1986, 1987
 -
 M
 Aps-2-2
 -
 Acid phosphatase-2-2; one of two codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.
 Navot et al., 1990; Navot and Zamir 1986, 1987
 -
 M
 Dia-1
 -
 Diaphorase-1
 Navot et al., 1990
 -
 M
 Est-1
 -
 Esterase-1; one of six codominant alleles, each regulating one band; found in C. lanatus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Est-1-1
 -
 Esterase-1-1; one of six codominant alleles, each regulating one band; found in C. lanatus var. citroides and C. colocynthis.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Est-1-2
 -
 Esterase-1-2; one of six codominant alleles, each regulating one band; found in C. colocynthis.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Est-1-3
 -
 Esterase-1-3; one of six codominant alleles, each regulating one band; found in Praecitrullus fistulosus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Est-1-4
 -
 Esterase-1-4; one of six codominant alleles, each regulating one band; found in C. ecirrhosus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Est-1-5
 -
 Esterase-1-5; one of six codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Est-2
 -
 Esterase-2; one of five codominant alleles, each regulating one band; found in C. lanatus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
Est-2-1 
 -
 Esterase-2-1; one of five codominant alleles, each regulating one band; found in C. colocynthis.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Est-2-2
 -
 Esterase-2-2; one of five codominant alleles, each regulating one band; found in C. colocynthis.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Est-2-3
 -
 Esterase-2-3; one of five codominant alleles, each regulating one band; found in Praecitrullus fistulosus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Est-2-4
 -
 Esterase-2-4; one of five codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.
Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Fdp-1
 -
 Fructose 1,6 diphosphatase-1.
 Navot et al., 1990; Navot and Zamir, 1986
 -
 M
 For-1
 -
 Fructose 1,6 diphosphatase-1.
 Navot et al., 1990
 -
 M
 Gdh-1
 -
 Glutamate dehydrogenase-1; isozyme located in cytosol.
 Navot and Zamir, 1986
 -
 M
 Gdh-2
 -
 Glutamate dehydrogenase-2; isozyme located in plastids.
 Navot et al., 1990; Navot and Zamir, 1986
 -
 M
 Got-1
 -
 Glutamate oxaloacetate transaminase-1; one of four codominant alleles, each regulating one band; found in C. lanatus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
Got-1-1
 -
 Glutamate oxaloacetate transaminase-1; one of four codominant alleles, each regulating one band; found in C. colocynthis and Praecitrullus fistulosus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
  Got-1-2
 -
 Glutamate oxaloacetate transaminase-1-2; one of four codominant alleles, each regulating one band; found in C. lanatus var. citroides.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Got 1-3
 -
 Glutamate oxaloacetate transaminase-13; one of four codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Got-2
 -
 Glutamate oxaloacetate transaminase-2; one of five codominant alleles, each regulating one band; found in C. lanatus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Got-2-1
 -
 Glutamate oxaloacetate transaminase-21; one of five codominant alleles, each regulating one band; found in C. colocynthis.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Got-2-2
 -
 Glutamate oxaloacetate transaminase-22; one of five codominant alleles, each regulating one band; found in C. ecirrhosus.
  Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
-
M
 Got-2-3
 -
Glutamate oxaloacetate transaminase-2-3; one of five codominant alleles, each regulating one band; found in Praecitrullus fistulosus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
Got-2-4
 -
Glutamate oxaloacetate transaminase-24; One of five codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.
Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Got-3
 -
 Glutamate oxaloacetate transaminase-3.
 Zamir et al., 1984
 -
 M
 Got-4
 -
 Glutamate oxaloacetate transaminase-4.
 Navot et al., 1990; Zamir et al., 1984
 -
 M
 hsp-70
 -
 heat shock protein 70; one gene presequence 72-kDa hsp70 is modulated differently in glyoxomes and plastids.
 Wimmer et al., 1997
 -
 M
 Idh-1
 -
 Isocitrate dehydrogenase-1
 Zamir et al., 1984
 -
 M
 Lap-1
 -
 Leucine aminopeptidase-1.
 Navot et al., 1990; Navot and Zamir, 1986
 -
 M
 Mdh-1
 -
 Malic dehydrogenase-1; one of two codominant alleles, each regulating one band; found in C. lanatus.
 Navot and Zamir, 1987; Zamir et al., 1984
 -
 M
 Mdh-1-1
 -
 Malic dehydrogenase-1-1; one of two codominant alleles, each regulating one band; found in Praecitrullus fistulosus.
 Navot and Zamir, 1987; Zamir et al., 1984
 -
 M
 Mdh-2
 -
 Malic dehydrogenase-2; one of three codominant alleles, each regulating one band; found in C. lanatus.
Navot and Zamir, 1987
 -
 M
 Mdh-2-1
 -
 Malic dehydrogenase-2-1; one of three codominant alleles, each regulating one band; found in C. colocynthis.
 Navot and Zamir, 1987
 -
 M
 Mdh-2-2
 -
 Malic dehydrogenase-2-2; one of three codominant alleles, each regulating one band; found in Praecitrullus fistulosus.
 Navot and Zamir, 1987
 -
 M
 Me-1
 -
 Malic enzyme-1; one of three codominant alleles, each regulating one band; found in C. lanatus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Me-1-1
 -
 Malic enzyme-1-1; one of three codominant alleles, each regulating one band; found in Praecitrullus fistulosus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Me-12
 -
 Malic enzyme-12; one of three codominant alleles, each regulating one band; found in C. colocynthis.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Me-2 
 -
 Malic enzyme-2.
 Zamir et al., 1984
 -
 M
 Pgd-1
 6 Pgdh-1
 6-Phosphogluconate dehydrogenase-1; one of three codominant alleles, each regulating one plastid band; found in C. lanatus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Pgd-1-1
 6 Pgdh-1-1
 6-Phosphogluconate dehydrogenase-1-1; one of three codominant alleles, each regulating one plastid band; found in Praecitrullus fistulosus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Pgd-1-2
 6 Pgdh-1-2
 6-Phosphogluconate dehydrogenase-1-2; one of three codominant alleles, each regulating one plastid band; found in Acanthosicyos naudinianus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Pgd-2
 6 Pgdh-2
 6-Phosphogluconate dehydrogenase-2; one of five codominant alleles, each regulating one cytosolic band; found in C. lanatus.
 Navot and Zamir, 1986; Zamir et al., 1984
 -
 M
 Pgd-2-1
 6 Pgdh-2-1
 6-Phosphogluconate dehydrogenase-21; one of five codominant alleles, each regulating one cytosolic band; found in C. ecirrhosus.
 Navot and Zamir, 1987; Zamir et al., 1984
 -
 M
 Pgd-2-2
 6 Pgdh-2-2
 6-Phosphogluconate dehydrogenase-2-2; one of five codominant alleles, each regulating one cytosolic band; found in Praecitrullus fistulosus.
 Navot and Zamir, 1987; Zamir et al., 1984
 -
 M
 Pgd-2-3
 6 Pgdh-2-3
 6-Phosphogluconate dehydrogenase-2-3; one of five codominant alleles, each regulating one cytosolic band; found in C. colocynthis.
 Navot and Zamir, 1987; Zamir et al., 1984
 -
 M
 Pgd-2-4
 6 Pgdh-2-4
 6-Phosphogluconate dehydrogenase-2-4; one of five codominant alleles, each regulating one cytosolic band; found in Acanthosicyos naudinianus.
 Navot and Zamir, 1987; Zamir et al., 1984
 -
 M
 Pgi-1
 -
 Phosphoglucoisomerase-1; one of three codominant alleles, each regulating one plastid band; found in C. lanatus
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Pgi-1-1
 -
 Phosphoglucoisomerase-11; one of three codominant alleles, each regulating one plastid band; found in C. colocynthis.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Pgi-1-2
 -
 Phosphoglucoisomerase-1-2; one of three codominant alleles, each regulating one plastid band; found in Acanthosicyos naudinianus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987
 -
 M
 Pgi-2
 -
 Phosphoglucoisomerase-2; one of six codominant alleles, each regulating one cytosolic band; found in C. lanatus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Pgi-2-1
 -
 Phosphoglucoisomerase-2-1; one of six codominant alleles, each regulating one cytosolic band; found in C. lanatus and C. colocynthis.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Pgi-2-2
 -
 Phosphoglucoisomerase-2-2; one of six codominant alleles, each regulating one cytosolic band; found in C. ecirrhosus.
 Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984
 -
 M
 Pgi-2-3  -  Phosphoglucoisomerase-2-3; one of six codominant alleles, each regulating one cytosolic band; found in Praecitrullus fistulosus.  Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984  -  M
 Pgi-2-4  -  Phosphoglucoisomerase-2-4; one of six codominant alleles, each regulating one cytosolic band; found in C. lanatus var. citroides.  Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984  -  M
 Pgi-2-5  -  Phosphoglucoisomerase-2-5; one of six codominant alleles, each regulating one cytosolic band; found in Acanthosicyos naudinianus.  Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984  -  M
 Pgm-1  -  Phosphoglucomutase-1; one of four codominant alleles, each regulating one plastid band; found in C. lanatus.  Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984  -  M
 Pgm-1-1  -  Phosphoglucomutase-1-1; one of four codominant alleles, each regulating one plastid band; found in C. colocynthis. Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984  -  M
 Pgm-1-2  -  Phosphoglucomutase-1-2; one of four codominant alleles, each regulating one plastid band; found in Acanthosicyos naudinianus.  Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984  -  M
 Pgm-1-3  -  Phosphoglucomutase-1-3; one of four codominant alleles, each regulating one plastid band; found in Praecitrullus fistulosus.  Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984  -  M
 Pgm-2  -  Phosphoglucomutase-2; one of four codominant alleles, each regulating one cytosolic band; found in C. lanatus.  Navot and Zamir, 1987; Zamir et al., 1984  -  M
 Pgm-2-1  -  Phosphoglucomutase-2-1; one of four codominant alleles, each regulating one cytosolic band; found in Acanthosicyos naudinianus.  Navot and Zamir, 1987; Zamir et al., 1984  -  M
 Pgm-2-2  -  Phosphoglucomutase-2-2; one of four codominant alleles, each regulating one cytosolic band; found in C. lanatus.  Navot and Zamir, 1987; Zamir et al., 1984  -  M
 Pgm-2-3  M  Phosphoglucomutase-2-3; one of four codominant alleles, each regulating one cytosolic band; found in Praecitrullus fistulosus.  Navot and Zamir, 1987; Zamir et al., 1984  -  M
 Prx-1  -  Peroxidase-1; one of seven codominant alleles, each regulating one band; found in C. lanatus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Prx-11  -  Peroxidase-11; one of seven codominant alleles, each regulating one band; found in C. colocynthis.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
Prx-12   -  Peroxidase-12; one of seven codominant alleles, each regulating one band; found in Praecitrullus fistulosus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Prx-13  -  Peroxidase-13; one of seven codominant alleles, each regulating one band; found in C. lanatus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Prx-14  -  Peroxidase-14; one of seven codominant alleles, each regulating one band; found in C. ecirrhosus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Prx-15  -  Peroxidase-15; one of seven codominant alleles, each regulating one band; found in C. lanatus and C. colocynthis.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Prx-16  -  Peroxidase-16; one of seven codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Prx-2  -  Peroxidase-2.  Navot and Zamir, 1987  -  M
 Prx-3  -  Peroxidase-3.  Navot and Zamir, 1987  -  M
 Sat  -  Serine acetyltransferase; catalyzes the formation of O-acetylserine from serine and acetyl-CoA.  Saito et al., 1997  -  M
 Skdh-1  -  Shikimic acid dehydrogenase-1.  Zamir et al., 1984  -  M
 Skdh-2  -  Shikimic acid dehydrogenase-2; one of six codominant alleles, each regulating one band.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Skdh-21  -  Shikimic acid dehydrogenase-21; one of six codominant alleles, each regulating one band; found in C. colocynthis.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Skdh-22  -  Shikimic acid dehydrogenase-22; one of six codominant alleles, each regulating one band; found in C. colocynthis.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Skdh-23  -  Shikimic acid dehydrogenase-23; one of six codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Skdh-24  -  Shikimic acid dehydrogenase-24; one of six codominant alleles, each regulating one band; found in C. ecirrhosus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Skdh-25  -  Shikimic acid dehydrogenase-25; one of six codominant alleles, each regulating one band; found in Praecitrullus fistulosus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Sod-1  -  Superoxide dismutase-1; one of three codominant alleles, each regulating one band; found in C. lanatus.  Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984  -  M
 Sod-11  -  Superoxide dismutase-11; one of three codominant alleles, each regulating one band; found in C. colocynthis.  Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984  -  M
 Sod-12  -  Superoxide dismutase-12; one of three codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.  Navot et al., 1990; Navot and Zamir, 1986, 1987; Zamir et al., 1984  -  M
 Sod-2  -  Superoxide dismutase-2; one of two codominant alleles, each regulating one band; found in C. lanatus.  Navot and Zamir, 1987  -  M
 Sod-21  -  Superoxide dismutase-21; one of two codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.  Navot and Zamir, 1987  -  M
 Sod-3  -  Superoxide dismutase-3; one of two codominant alleles, each regulating one band; found in C. lanatus.  Navot and Zamir, 1987  -  M
 Sod-31  -  Superoxide dismutase-31; one of two codominant alleles, each regulating one band; found in Praecitrullus fistulosus.  Navot and Zamir, 1987  -  M
 Spr-1  -  Seed protein-1.  Navot and Zamir, 1986  -  M
 Spr-2  -  Seed protein-2.  Navot and Zamir, 1986  -  M
 Spr-3  -  Seed protein-3.  Navot and Zamir, 1986  - M
 Spr-4  Spr-4  Seed protein-4.  Navot et al., 1990; Navot and Zamir, 1986  - M
 Spr-5  Spr-5  Seed protein-5.  Navot et al., 1990; Navot and Zamir, 1986  -  M
 Tpi-1  -  Triosephosphatase isomerase-1. one of four codominant alleles, each regulating one band; found in C. lanatus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Tpi-11  -  Triosephosphatase isomerase-11; one of four codominant alleles, each regulating one band; found in C. colocynthis.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Tpi-12  -  Triosephosphatase isomerase-12; one of four codominant alleles, each regulating one band; found in Praecitrullus fistulosus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Tpi-13  -  Triosephosphatase isomerase-13; one of four codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.  Navot et al., 1990; Navot and Zamir, 1986, 1987  -  M
 Tpi-2  -  Triosephosphatase isomerase-2; one of three codominant alleles, each regulating one band; found in C. lanatus.  Navot and Zamir, 1987  -  M
 Tpi-21  -  Triosephosphatase isomerase-21; one of three codominant alleles, each regulating one band; found in Acanthosicyos naudinianus.  Navot and Zamir, 1987  -  M
 Ure-1  -  Ureaase-1.  Navot and Zamir, 1987  -  M

y C = Mutant available from Cucurbit Genetics Cooperative watermelon gene curator; M = molecular marker or isozyme; P = mutants are available as standard cultivars or accessions from the plant introduction collection; ? = availability not known; L = mutant has been lost.

 

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