Cucurbit Genetics Cooperative Report 16:92-97 (article 35) 1993
Todd C. Wehner
Department of Horticultural Science, North Carolina State University, Raleigh, NC 27695-7609
Lists of the known genes for the Cucurbitaceae have been published previously (12, 16). Howver, in the interest of updating the information, following is a list of the 43 genes introduced or modified since thelast report (13). That makes a total (105 plus 43 minus 2 renamed) 146 reported gene mutants.
A wild type cultivar has not been proposed for cucumber as the reference for all “+” alleles. ‘Marglobe’ tomato has been chosen as the normal type for genetic studies in that crop. Perhaps ‘Wisconsin SMR 18’ should be the choice for cucumber, since it is an inbred that isued widely in genetic studies. In addition, it has many dominant and wild-type alleles.
The genes on the following list are of four categories, seedling markers, virus resistance, isozymes, and fruit mutants (Table 1). The seedling markers include a reclassification of gc as ls (25, 26) and a new group produced using gamma rays on pollen (4, 5). The fruit mutants are green mature fruit, gn (11), and palisade epidermis, Pe (2). Isozyme mutants now include 14 markers, many of which are linked (6). One virus resistance gene should be renamed according to the revised taxonomy of watermelon mosaic virus. Watermelon mosaic virus-1 was renamed papaya ringspot virus-waermelon strain (PRSV-W), so wmv-1-1 should be renamed prsv, retaining wmv-1-1 as a synonym. Watermelon mosaic virus-2 is renamed watermelon mosaic virus (WMV). Thus, the dominant gene for resistance to WMV does not change its symbol, remaining Wmv.
Isozyme variant nomenclature for this gene list follows the form according to Staub et al. (22), such that loci coding for enzymes (e.g. glutamine dehydrogenase, G2DH) are designated as abbreviations, where the first letter is capitalized (e.g.G2dh). If an enzyme system is conditioned by multiple loci, then those are designated by hyphenated numbers, which are numbered from most cathodal to most anodal and enclosed in parentheses. The most common allele of anyparticular isozyme is designated 100, and all other alleles for that enzyme are assigned a value based on their mobiility relative to that allele.
Researchers are encouraged to send reports of new genes, as well as seed samples to the cucumber gene curators (Todd C. Wehner, Jack E. Staub and Richard W. Robinson). Please let me know of any omissions or errors in the following list.
Table 1. The 43 new or revised genes of cucumber*.
Gene symbol |
Character |
References |
|
Preferred |
Synonym |
||
| al | albino cotyledons.. White cotyledoms and slightly light green hypocotyl; dying before the first true leaf stage. Wild type al+ from ‘Nishiki-suyo’; al from M2 line from pollen irradiation. | 4, 5 | |
| by | bu | bushy. Short internodes; normal seed viability. Wild type by+ from ‘Borszczagowski’; by from induced mutation of ‘Borszczagowski’. . Linked with F and gy, not with B or bi.. | 9 |
| chp | choripetalous. Small first true leaf’ choripetalous flowers; glosssy ovary; small gruits; few seeds. Wild type chp+ from ‘Borszczagowski’; chp from chemically induced mutation. | 7 | |
| cp-2 | compact-2. Short internodes; small seeds, similar to cp, but allelism not checked. Wild type cp-2+ from ‘Borszczagowski’; cp-2 from induced mutation of ‘Borszczagowski’ called W97. Not linked with B or F; interacts with by to produce super dwarf. | 10 | |
| de-2 | determinate-2. Main stem growth ceases after 3 to 10 nodes, producing flowers at the apex; smooth, fragile, dark-green leaves; similar to de, but not checked for allelism. Wild type de-2+ from ‘Borszczgowski’; de-2 from W-sk mutant induced by ethylene-imine from ‘Borszczagowski’. | 21 | |
| dm-1 | dm? | downy mildew resistance. One of three genes for resistance to downy mildew caused by Pseudoperonospora cubensis (Berk & Curt). Wild type dm-1+ from Wisconsin SMR 19; dm-1 from WI 4783. Not checked for allelism with dm. | 1 |
| dm-2 | downy mildew resistance-2. One of three genes for resistance to downy mildew caused by Pseudoperonospsora cubensis Berk & Curt). Wild type dm-2+ from Wisconsin SMR 18; dm-2 from WI 4783. Not checked for allelism with dm. | 1 | |
| dm-3 | downy mildew resistance-3. One of three genes for resistance to downy mildew caused by Pseudoperonospora cubensis (Berk & Curt). Wild type dm-3+ from Wisconsin SMR 18; dm-3 from WI 4783. Not checked for allelism with dm. | 1 | |
| dvl-2 | dl-2 | divided leaf-2. Divided leaves after the 2nd trye leaf; flower petals free; similar to dvl, but allelism not checked. Wild type dvl-2+ from ‘Borszczagowski’; dvl-2 from mutant induced by ethylene-imine from ‘Borszczagowski’. | 19 |
| dwc-1 | dwarf cotyledons-1. Small cotyledons; late germination; small first true leaf; died after 3rd trye leaf. Wild type dwc-1+ from ‘Nishiki-suyo’; dwc-1 from M2 line from pollen irradiation. | 4, 5 | |
| dwc-2 | dwarf cotyledons-2. Small cotyledons, later germination; small first true leaf. Wild type dwc-2+ from ‘Nishiki-suyo’; dwc-2 from M2 line from pollen irradiation. | 4, 5 | |
| G2dh | Glutamine dehydrogenase (E.C.# 1.1.1.29). Isozyme variant found segregating in PI 285606; 5 alleles observed. | 6 | |
| gi-2 | ginko-2.Spatulate leaf blade with reduced lobing and altered veins; recognizable at the 2nd true leaf stage; similar to gi, fertile instead of sterile. wild type gi-2+ from ‘Borszczagowski’; gi-2 from mutant in the Kubicki collection. | 19 | |
| gig | gigantism.First leaf larger than normal. Wild type gig from ‘Borszczagowski’; gig from chemicaly induced mutation. | 8 | |
| gn | green mature fruit. Green mature fruits when R+R+ gngn; creeam colored when R+R+ gn+gn+ orange when R_. Wild type gn+ from ‘Chipper’, SMR 58 and PI 165509; gn from TAMU 830397. | 11 | |
| Gpi-1 | Glucose phosphate isomerase. (E.C. # 5.3.1.9). Isozyme variant found segregating (1 and 2) in PI 176524, 200815, 249561, 422192, 432854, 436608; 3 alleles observed. | 6 | |
| Gr-1 | Glutathione reductase-1 (E.C. # 1.6.4.2). Isozyme variant found segregating in PI 109275; 5 alleles observed. | 6 | |
| hl | heart leaf. Heart shaped leaves. Wild type hl+ from Wisconsin SMR 18; hl from WI 2757. Linked with ns and ss in the linkage group with Tu-u-D-pm. | 23 | |
| hn | horn like cotyledons. Cotyledons shaped like bull horns; true leaves with round shape rather than normal lobes; circular rather than ribbed stem cross section; divided petals; spineless fruits; pollen fertile, but seed sterile. Wild type hn+ from ‘Nishiki-suyo’; hn from M2 line from pollen irradiation. | 4, 5 | |
| hsl | heart shaped leaves. Leaves heart shaped rather than lobed; tendrils branched. Wild type hsl+ from Nishiki-suyo’ hsl from M2 line from pollen irradiation. | 4. 5 | |
| Idh | Isocitrate dehydrogenase (E.C. # 1.1.1.42). Isozyme variant found segregating in PI 183967, 21589; 2 alleles observed. | 6 | |
| lg-1 | light green cotyledons-1. Light green cotyledons, turning dark green; light green true leaves, turning dark green; poorly developed stamens. Wild type lg-1+ from ‘Nishiki-suyo’; lg-1 from M2 line from pollen irradiation. | 4, 5 | |
| 1g-2 | light green cotyledons-2. Light green cotyledons,turning dark green (faster than lg-1; light green true leaves, turning dark green; normal stamens. Wild type lg-2+ from ‘Nishiki-suyo’; lg-2 from M2 line from pollen irradiation. | 4, 5 | |
| ls | gc | light sensitive. Yellow cotyledons, lethal in high light. Abstract gave cg as symbol; article that followed gave ls as symbol. Mutant ls from a selection of ‘Burpless Hybrid’. | 25, 26 |
| Mdh-1 | Malate dehydrogenase-1 (E.C. # 1.1.1.37.). Isozyme variant found segregating in PI 171613, 209064, 326594; 3 alleles observed. | 6 | |
| Mdh-2 | Malate dehydrogenase-2 (E.C.# 1.1.1.37). Isozyme variant found segregaring in PI 174164, 185690, 357835, 419214; 2 alleles observed. | 6 | |
| Mdh-3 | Maleate dehydrogenase-3 (E.C.# 1.1.1.27). Isozyme variant found segregating in PI 255236, 267942, 432854, 432887; 2 alleles observed. | 6 | |
| Mpi-2 | Mannose phosphate isomerase (E.C. # 5.3.1.8). Isozyme variant found segregating in PI 109275, 175692, 200815, 209064, 263049, 354952; 2 alleles observed. | 6 | |
| mpy | mpi | male pygmy. Dwarf plant with only staminate flowers. Wild type mpy+ from Wisconsin SMR 12′ mpy fromGnome 1, a selection of ‘Rochford’s Improved’. | 14 |
| Pe | Palisade epidermis. Epidermal cells arranged perpendicular to the fruit surface. Wild type Pe from ‘Wisconsin SMR 18’, ‘Spartan Salad’ and Gy 2 compact; pe from WI 2757. | 2 | |
| Pep-la | Peptidase with leucyl-leucine (E.C. # 3.4.13.11). Isozyme variant found segregating in PI 169380,175692, 263049, 289698, 354952, 5 alleles observed. | 6 | |
| Pep-pap | Peptidase with phenylalanyl-L-proline (E.C. # 3.4.13.11). Isozyme variant found segregating in PI 163213, 188749, 432861; 2 alleles observed. | 6 | |
| Per-4 | Peroxidase (E.C. # 1.11.1.7). Isozyme variant found segregating in PI 215589; 2 alleles observed. | 6 | |
| Pgd-1 | Phosphogluconate dehydrogenase-1 (E.C. # 1.1.1.43). Isozyme variant found segregating in PI 169380, 175692, 222782; 2 alleles observed. | 6 | |
| Pgd-2 | Phosphogluconate dehydrogenase-2 (E.C. # 1.1.1.43). Isozyme variant found segregaring in PI 171613,177364, 188749, 263049, 285606, 289698, 354952, 419214, 432858; 2 alleles observed. | 6 | |
| Pgm-1 | Phosphoglucomutase (E.C. # 5.4.2.2.). Isozyme variant found segregating in PI 171613, 177364, 188749, 263049, 264229, 285606, 289698, 354952; 2 alleles observed. | 6 | |
| prsv | wmv-1-1 | watermelon mosaic virus 1 resistance. Resistance to papaya ringspot virus (formerly watermelon mosaic virus 1). Wild type prsv+ from WI2757; prsv from ‘Surinam’. | 24 |
| sh | short hypocotyl. Hypocotyl of seedlings 2/3 the length of normal. Wild type sh+_ from ‘Borszczagowski’; sh from khp, an induced mutant from ‘Borszczagowski’. | 20 | |
| shl | shrunken leaves. First and 2nd true leaves smaller than normal; later leaves becoming normal; slow growth; often dying before fruit set. Wild type shl+ from ‘Nishiki-suyo’; shl from M2 line from pollen irradiation. | 4, 5 | |
| sp-2 | short petiole-2. Leaf petioles shorter, darker green than normal at 2-leaf stage; crinkled leaves with slow devlopment; short hypocotyl and stem; little branching. Not tested for allelism with sp. Wild type sp-2+ from ‘Borszczagowski’; sp-2 from chemically induced mutation. | 18 | |
| wi | wilty leaves. Leaves wiltingin the field, but not in shaded greenhouse; weak growth; no fruiting. Wild type wi+ from greenhouse; weak growth; no fruiting. wi from M2 line from pollen irradiation. | 4, 5 | |
| wy | wavy rimed cotyledons. Wavy rimed cotyledons, with white centers; true leaves normal. Wild type wy+ from ‘Nishiki-suyo’; wy from M2 line from pollen irradiation. | 4, 5 | |
| ys | yellow stem. Yellow cotyledons, becoming cream-colored; cream-colored stem, petiole and leaf veins; short petiole; short internode. Wild type ys+ from ‘Borszczagowski’; ys from chemically induced mutation. | 17 | |
*Isozyme nomenclature follows a modified form (22) previously described by Richmond (15) and Gottlieb (3).
Literature Cited
- Dorochowski, R.W. and E. Lakowska-Ryk. Inheritance of resistance to downy mildew (Pseudoperonospora cubensis Berk & Curt) in cucumis sativus. Proc. 5th EUCARPIA, Cucurbitaceae Symp. p. 66-69, Warsaw, Poland.
- Fanourakis, N.E. and P.W. Simon. 1987. Inheritance and linkage studies of the fruit epidermal structure in cucumber. J. Hered. 78: 369-371.
- Gottlieb, T.D. 1977. Evidence for duplication and divergence of the structural gene for phosphoglocoseisomerase in diploid species of Clarkia. Genetics 86: 289-307.
- Iida, S. and E. Amano. 1990. Pollen irradiation to obtain mutants in monoecious cucumber. Ga,,a Field Symp. 29: 95-111.
- Iida, S. and E. Amano. 1991. Mutants induced by pollen irradiation in cucumber. Cucurbit Genet. Coop. Rpt. 14:32-33.
- Knerr, L.D. and J.E. Staub. 1992. Inheritance and linkage relationships of isozyme loci in Cucumber (Cucumis sativus L.). Theor. Appl. Genet. 84:217-224.
- Kubicki, B. and A. Korzeniewska. 1984. Induced mutations in cucumber (Cucumis sativus L.) III. A mutant with choripetalous flowers. Genetica polonica 25: 53-60.
- Kubicki, B., I Goszczycka and A. Korzeniewska. 1984. Induced mutations in cucumber (Cucumis sativus L.) II. Mutant of gigantism. Genetica Polonica 25: 41-52.
- Kubicki, B., U. Soltysiak and A. Korzeniewska. 1986a. Indued mutations in cucumber (Cucumis sativus L.). IV. A mutant of the bush type of growth. Genetica Polonica 27:273-287.
- Kubicki, B., U. Soltysiak and A. Korzeniewska. 1986b. Induced mutations in cucumber (Cucumis sativus L.). V. Compact type of growth. Genetica Polonica 27: 289-298.
- Peterson, G. C. and L. M. Pike. 1992. Inheritance of green mature seed-stage fruit color in Cucumis sativus L. J. Amer. Soc. Hort. Sci. 117:643-645.
- Pierce, L.K. and T.C. Wehner. 1989. Gene list for cucumber. Cucurbit Genet. Coop. Rpt. 12:91-103.
- Pierce, L.K. and T.C. Wehner. 1990. Review of genes and linkage groups in cucumber. HortScience 25: 605-615.
- Pyzhenkov, V. and G.A. Kosareva. 1981. Description of features and their inheritance pattern in a new cucumber mutant. Biull. Vses. Inst. Rastenievod 109: 5-8 (Plant Breeding Abstracts 1983, 53: 478).
- Richmond, R.C. 1972. Enzyme variability in the Drosophilia williston group. 3. Amounts of variability in the super species D. Paulistorum. Genetics 70: 87-112.
- Robinson, R.W., H,M, Munger, T.W. Whitaker and G.W. Bohn. 1976. Genes of the Cucurbitaceae. HortScience 11:554-568.
- Rucinska, M., K. Niemirowicz-Szczytt and A. Korzeniewska. 1991. A cucumber (Cucumis sativus L.) mutant with yellow stem and leaf petioles. Cucurbit Genet. Coop. Rpt. 14: 8-9.
- Rucinska, M. K. Niemirowicz-Szczytt and A. Korzeniewska. 1992a. Cucumber (Cucumis sativus L.) induced mutations. II. A second short petiole mutant. Cucurbit Genet. Coop. Rpt. 15:33-34.
- Rucinska, M., K. Niemirowicz-Szczytt and A. Korzeniewska. 1992b. Cucumber (Cucumis sativus L.) induced mutations. III and IV. Divided and ginko leaves. Proc. 5th EUCARPIA Cucurbitaceae Symp. p. 66-69, Warsaw, Poland.
- Soltyiak, U. and B. Kubicki. 1988. Induced mutations in the cucumber (Cucumis sativus L.). VII. Short hypocotyl mutant. Genetica Polonica 29: 315-321.
- Soltysiak, U., B. Kubicki and A. Korzeniewska. 1986. Induced mutations in cucumber (Cucumis sativus L.). VI. Determinate type of growth. Genetica Polonica 27: 229-308.
- Staub, J.E., R.S. Kupper, D. Schuman, T.C. wehner and B. May. 1985. Electrophoretic variation and enzyme storage stability in cucumber. J. Amer. Soc. Hort. Sci. 110-426-431.
- Vakalounakis, D.J. 1992. Heart leaf, a recessive leaf shape marker in cucumber: linkage with disease resistance and other traits. J. Heredity 83: 217-221.
- Wang, W.J., R. Provvidenti and R.W. Robinson. 1984. Inheritance of resistance in cucumber to watermelon mosaic virus. Phytopathology 51: 423-428.
- Whelan, E. D.P. 1971. Golden cotyledon: a radiation-induced light sensitive mutant in cucumber, HortScience 6:343.
- Whelan, E.D.P. 1972. Inheritance of a radiation-induced light sensitive mutant of cucumber. J. Amer. Soc. Hort. Sci. 97-765-767.
Scientists should consult the above list as well as the rules of gene nomenclature for the Cucurbitaceae (16) before chosing a gene name and symbol. That will avoid duplication of gene names and symbols. The rules of gene nomenclature were adopted in order to provide guidelines for naming and symbolizing genes. Scienists are urged to contact members of the Gene List committee regarding rules and gene symbols.
Gene List Committee:
Cucumber – T.C. Wehner
Cucurbita spp. – R.W. Robinson, M.G. Hutton
Melon – M. Pitrat
Watermelon – W.R. Henderson
Other genera – R.W. Robinson