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Cucurbit Genetics Cooperative Report 20:66-88 (article 30) 1997
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1997 Gene List for Cucumber

Todd C. Wehner

Department of Horticultural Science, North Carolina State University, Raleigh, NC 27695--7609

Jack E. Staub

Vegetable Crops Research, USDA/ARS. Department of Horticulture, University of Wisconsin, Madison, WI 53706 U.S.A.

Complete lists and updates of genes for cucumber (Cucumis sativus L.) have been published previously (Pierce and Wehner, 1989; Robinson et al., 1982; Wehner, 1993), and this is the latest version. For the first time, this gene list includes marker loci for restriction fragment length polymorphisms (RFLPs) and random amplified DNA (RAPDs). The genes on the 1997 list are of nine categories as follows: seedling markers, stem mutants, leaf mutants, flower mutants, fruit type mutants, fruit color mutants, resistance (mostly to diseases), protein (isozyme) variants, and DNA (RFLPs and RAPDs) markers (Table 1).

Revisions to the 1997 cucumber gene list include the addition of a flower mutant, male sterile-2 pollen sterile, ms-2PS (Zhang et al., 1994), correction of the spelling of the ginko mutant to ginkgo, modification of the description of the gene for resistance to Fusarium oxysporum f. sp. cucumerinum races, Foc (Vakalounakis, 1993, 1995, 1996), and addition of 7 isozyme mutants, Ak-2, Ak-3, Fdp-1, Fdp-2, Mpi-1, Pep-gl, and Skd (Meglic and Staub, 1996). the gene list now includes a total of 269 (158 classical genes, 91 RFLPs, and 20 RAPDs) gene mutants.

Isozyme variant nomenclature for this gene list follows the form according to Staub et al. (Staub et al,, 1985), 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 my 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 any particular isozyme is designated, 100, and all other alleles for that enzyme are assigned a value based on their mobility relative to that allele. for example, an allele at locus 1 of FDP (fructose diphosphatase) which has a mobility 4 mm less that of the most common allele would be assigned the designation Fdp)(1)-96.

RFLP marker loci were identified as a result of digestion of cucumber DNA with DraI, EcoRI, EcoRV, or HindIII (Kennard et al., 1994). Partial-genomic libraries were constructed using either PstI-digested DNA from the cultivar Sable and from EcoRV-digested DNA from the inbred WI 2757. Derived clones were hybridized to genomic DNA and banding patterns were described for mapped and unlinked loci (CsC482/H3, CsP314/E1, and CsP344E1, CsC477/H3, CsP300/E1).

Clones are designated herein as CsC = cDNA, CsP = PstI-gnomic, and CsE = EcoRI-genomic. Lower case a or b represent two independently-segregating loci detected with one probe. Lower -case s denotes the slowest fragment digested out of the vector. Restriction enzymes designated as DI, IDRAI; EI, EcoRI; E5, EcoRV; and H3, HindIII. Thus, a probe identified as CsC336b/E5 is derived from a cDNA library (from 'Sable') which was restricted using the enzyme EcoRV to produce a clone designated as 336 which displayed two independently segregating loci one of which is b. Clones are available in limited supply from Jack E. Staub.

RAPD marker loci were identified using primer sequences from Operon Technologies (OP; Alameda, California, U.S.A.) and the University of British Columbia (Vancouver, BC, Canada). Loci are identified by sequence origin (OP or BC), primer group letter (e.g., A), primer group array number (1-20), and locus (a, b, c, etc.) (Kennard et al., 1994). Information regarding unlinked loci can be obtained from Jack E. Staub.

Because of their abundance, common source (two mapping populations) and the accessibility of published information on their development (Kennard et al., 1994) DNA marker loci are not included in Table 1, but are listed below.

The 60 RFLP marker loci from mapping cross Gy 14 x PI 183967 (Kennard et al., 1994): CsP129/E1, CsC032a/E1, CsP064/E1, CsP357/H3, CsC386/E1, CsC365/E1, CsP046/E1, CsP347/H3, CsC694/E5, CsC588/H3, CsC230/E1, CsC593/D1, CsP193/H3, CsP193/H3, CsP078s/H3, CsC581/E5, CsE084/E1, CsC341H3, CsPO24/E1, CsP287/H3, CsC629/H3, CsP225s/E1, CsP0501/H3, CsE051/H3, CsC366a/E5, CsC32b/E1, CsP056/H3, CsC378/E1, CsP406/E1, CsP460/E1, CsE060/E1, CsE103/E1, CsP019/E1, CsP168/D1, CsC560/H3, CsP005/E1, CsP440s/E1, CCsP221/H3, CsC625/E1, CsP475s/E1, CsP211/Eq, CsP215/H3, CsC613/E1, Csc029/H3, CsP130/E1, CsC443/H3, CsE120/H3, CsE031/H3, CsC366b/ E5, CsC082/H13, CsP094/H3, CsC362/E1, CsP441/E1, CsP280/H3, CsC558/H3, CsP037a/E1, CsP476/H3, CsP308/E1, CsP105/E1, and CsC166/E1.

The 31 RFLP marker loci from mapping cross Gy 14 x PI 432860 (Kennard et al., 1994): CsC560/D1, CsP024/E5, CsP287/H3, CsC384/E5, CsC366/E5, CsC611/D1, CsP055/D1, CsC482/H3, CsP019/E1, CsP059/D1, CsP471s/H13, CsC332/E5, CsP056/H3CsC308/E5, CsP073/E5, CsP215/H3, CsC613D1, CsP266/D1, CsC443/H3, CsE031/E1, CsE120/H3, CsE063/E1, CsP444/E1, CsC612/D1, Cs362/E1, CsP280/H3, CsC558/H3, CsP008/D1, CsP308/E1, CsC166/E1, and CsP303/H3.

The 20 RAPD marker loci from mapping cross Gy 14 x PI 432860 (Kennard et al., 1994):OPR04, OPW16, OPS17, OPE13a, OPN06, OPN12, OPP18b, BC211b, OPN04, OPA10, OPE09, OPT18, OPA14b, OPU20, BC460a, OPAB06, OPABpo5, OPH12, OPA14a, and BC211a.

Researchers are encouraged to send reports of new genes, as well as seed samples to the cucumber gene curator (Todd C. Wehner), or to the assistant curators (Jack E. Staub, and Richard W. Robinson). 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 (Robinson et al., 1976; Robinson et al., 1982) before choosing 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. Scientists are urged to contact members of the gene list committee regarding rules and gene symbols.

Table 1. the 158 (non-molecular) genes of cucumber.

  Synonym

Character

Referencesz
Supplemental referencesz
Availabley
a
-
 Androecious. Produces primarily staminate flowers if recessive for F. A from MSU 71305 and Gy 14; a from An-11 and An-314, two selections from 'E-e-szan' of China. Kubicki, 1969  
P
Ak-2
-
 Adenylate kinase (E.C. # 2.7.4.3). Isozyme variant found segregating in PI 339247, and 271754; 2 alleles observed. Meglic and Staub, 1996  
P
Ak-3
-
 Adenylate kinase (E.C.#2.7.4.3). Isozyme variant found segregating in PI 113334, 183967, and 285603; 2 alleles observed. Meglic and Staub, 1996  
P

al

-

albino cotyledons.. White cotyledons 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.

Iidea and Amano, 1990, 1991

  
?
ap
-
 apetalous. Male-sterile. Anthers become sepal-like. Ap from 'Butcher's Disease Resisting'; ap from 'Butcher's Disease Resisting Mutant'. Grimbly, 1980  
L
Ar
-
 Anthracnose resistance. One of several genes for resistance to Colletotrichum lagenarium. Ar from PI 175111, PI 175120, PI 179676, PI 183308, PI 183445; ar from 'Palmetto' and 'Santee'. Barnes and Epps. 1952.  
P
B
-
 Black or brown spines. Dominant to white spines on fruit. Strong, 1931; Tkachenko, 1935; Wellington, 1913. Cochran, 1938; Fujieda and Akiya, 1962; Hutchins, 1940; Jenkins, 1946; Youngner, 1952.
W
B-2
-
 Black spine-2. Interacts with B to produce F2 of 15 black: 1 white spine. B-02 from Wis. 9362; b-2 from PI 212233 and 'Pixie'. Shanmugasundarum et al., 1971a  
?
B-3
-
 Black spine-3. Interacts with B-4 to produce an F2 of nine black: 7 white spine. B-3 from LJ90430; b-3 from MSU 41. Cowen and Helsel, 1983.  
W
B-4
-
 Black spine-4. Interacts conversely with B-3. B-4 from LJ90430; b-4 from MSU 41. Cowan and Helsel 1983  
W
bi
-
 bitterfree. All plant parts lacking cucurbitacins. Plants with bi less preferred by cucumber beetles. Plants with Bi resistant to spider mites in most American cultivars; bi in most Dutch cultivars. Andeweg and De Bruyn, 1959 Cantliffe, 1972; Da Costa and Jones, 1971a, 1971b; Sons et al., 1973.
W
bl
t
 blind. Terminal bud lacking after temperature. Carlsson, 1961. shock. bl from 'Hunderup' nd inbred HP3.    
L
bla
-
 blunt leaf. Leaves have obtuse spices and reduced lobing and serration. bla from a mutant of 'Wis. SMR 18'. Robinson, 1987a  
W
Bt
-
 Bitter fruit. Fruit with extreme bitter flavor. Bt from PI 173889 (Wild Hanzil Medicinal Cucumber). Barham, 1953  
W
bu
-
 bush. Shortened internodes. bu from 'KapAhk 1'. Pyzenkov and Kosareva, 1981  
L
Bw
-
 Bacterial wilt resistance. Resistance to Erwinia tracheiphila. Bw from PI200818; bw from 'Marketer'. Nutall and Jasmin, 1958 Robinson and Whitaker, 1974
W

by

bu

bushy. Short internodes; normal seed viability. Wild type By from induced mutation of 'Borszczagowski'. . Linked with F and gy, not with B or bi.

Kubiki et al., 1986a

  
?
c
-
 cream mature fruit color, interaction with R is evident in the F2 ratio of 9 red (RC) : 3 orange (Rc) : 3 yellow (rC) : 1 cream (rc). Hutchins, 1940  
L
Cca
-
 Corynespora cassicola resistance. Resistance to target leaf spot; dominant to susceptibility. Cca from Royal Sluis Hybrid 72502; cca from Gy 3. Abul-Hayja et al., 1975  
W
Ccu
-
 Cladosporium cucumerinum resistance. Resistance to scab. Ccu from line 127.31, a selfed progeny of 'Longfellow'; ccu from 'Davis Perfect'. Bailey and Burgess, 1934 Abul-Hayja and Williams, 1976; Abul-Hayja et al., 1975, Andeweg, 1956.
W
cd
-
 chlorophyll deficient. Seedling normal at first, later becoming a light green; lethal unless grafted. cd from a mutant selection of backcross of MSU 713-5 x 'Midget' F1 to 'Midget'. Burnham, et al., 196  
L
chp
-
 choripetalous. Small first true leaf;' choripetalous flowers; glosssy ovary; small fruits; few seeds. Wild type Chp+ from 'Borszczagowski'; chp from chemically induced mutation.
Kubicki and Korzeniewska, 1984
  
?
cl
-
 closed flower. Staminate and pistillate flowers do not open; male-sterile (nonfertile pollen). Groff and Odland, 1963  
W
cla
-
 Colletotrichum lagenarium resistance. Resistance to race 1 of anthracnose; recessive to susceptibility. Cla from Wis. SMR 18; cla from SC 19B. Abul-Hayja et al., 1978  
W
Cm - Cornyespora melonis resistance. Resistance to C. melonis dominant to susceptibility. Cm from 'Spotvrie'; cm from 'Esvier'.    
?
Cmv
-
 Cucumber mosaic virus resistance. One of several genes for resistance to CMV. Cmv from 'Wis. SMR 12', 'Wis. SMR 15'; and 'Wis. SMR 18'; cmv gtom 'National Pickling' and 'Wis. SR 6'. Wasuwat and Walker, 1961 Shifriss et al., 1942
W
co
-
 green corolla, Green petals that turn white with age and enlarged reproductive organs; female-sterile. co from a selection of 'Extra Early Prolific'. Hutchins, 1935 Currence, 1954
L
cor-1
-
 cordate leaves-1. Leaves are cordate. cor-1 from Nezhinskii'. Gornitskaya, 1967  
L
cor-2
cor
 cordate leaves-2. Leaves re nearly round with revolute margins and no serration. Insect pollination is hindered by short calyx segments that tightly clasp the corolla, preventing full opening. cor-2 from an induced mutant of 'Lemon'. Robinson, 1987c  
?
cp
-
 compact. Reduced internode length, poorly developed tendrils, small flowers. cp from PI 308916. Kauffman and Lower, 1976  
W

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.

Kubicki et al, 1986b

  
W
cr
-
 crinkled leaf. Leaves and seed are crinkled. Odland and Groff, 1963a  
?
cs
-
 carpel splitting. Fruits develop deep longitudinal splits. cs from TAMU 1043 and TAMU 72210, which are second and fifth generation selections of MSU3249 x SC 25. Caruth, 1975; Pike and Caruth, 1977  
?
D
g
 Dull fruit skin. Dull skin of American cultivars, dominant to glossy skin of most European cultivars. Poole, 1944; Strong, 1931; Tkachenko, 1935  
W
de
I
 determinate habit. Short vine with stem terminating in flowers; modified by IN-de and other genes; degree of dominance depends on gene background. de from Penn 76.60G*, Minn 158.60*, 'Hardin's PG57', Hardin's Tree Cucumber', and S2 -1 (and inbred selection from Line 541)**. Denna, 1971*; George, Nutall and 1970**; Hutchins, 1940, Jasmin, 1958  
W

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'.

Soltysiak et al. 1986

  
?
df
-
 delayed flowering. Flowering delayed by long photoperiod; associated with dormancy. Idf from 'Baroda' (PI 212896)* and PI 215589 (hardwickii)**. Della Vecchia et al., 1982*; Shifriss and George, 1965**.  
W
dl
-
 delayed growth. Reduced growth rate; shortening of hypocotyl and first internodes. dl from 'Dwarf Marketmore' and 'Dwarf Tablegreen', both deriving dwarfness from 'Hardin's PG-57'. Miller and George, 1979  
W
dm
P
 downy mildew resistance. One of several genes for resistance to Pseudoperonospora cubensis. Dm from Sluis & Groot Line 4285; dm from 'Poinsett'. van Vliet and Meysing, 1977 Jenkins, 1946; Shimizu, 1963.
W

dm-1

dm

downy mildew resistance-1. 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.

Doruchowski and Lakowska-Ryk, 1992.

  
?

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.

Doruchowski and Lakowska-Ryk, 1992

  
?
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.
Doruchowski and Lakowska-Ryk, 1992
  
?
dvl
dl
 divided leaf-2. True leaves are partly or fully divided, often resulting in compound leaves with two to five leaflets and having incised corollas.
den Nijs and Mackiewicz, 1980
  
W
dvl-2
dl-2
 divided leaf-2. Divided leaves after the 2nd true 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". Rucinska et al., 1992b  
?
dw
-
 dwarf. Short internodes. dw from an induced mutant of 'Lemon'. Robinson and Mishanec, 1965  
?

dwc-1

-

dwarf cotyledons-1. Small cotyledons; late germination; small first true leaf; died after 3rd true leaf. Wild type Dwc-1 from 'Nishiki-suyo'; dwc-1 from M2 line from pollen irradiation.

Iida and Amano, 1990, 1991.

  
?
dwc-2
-
 dwarf cotyledons-2. Small cotyledons, late germination; small first true leaf. Wild type Dwc-2 from 'Nishiki-suyo'; dwc-2 from M2 line from pollen irradiation.
Iida and Amano, 1990, 1991.
  
?
Es-1
-
 Empty chambers-1. Carpels of fruits separated from each other, leaving a small to large cavity in the seed cell. Es-1 from PP2-75; es-1 from Gy-30-75. Kubicki and Korzeniewska, 1983  
?
Es-2
-
 Empty chambers-2. Carpels of fruits separated from each other, leaving a small to large cavity in the seed cell. Es-2 from PP-2-75; es-2 from Gy-30-75. Kubicki and Korzeniewsika, 1983.  
?
F
Acr, acrF, D, st
Female. High degree of pistillate sex expression; interacts with a and M; strongly modified by environment and gene background. F and f are from 'Japanese'. Galun, 1961; Tkachenko, 1935. Kubicki, 1965, 1969a; Poole, 1944; Shifriss, 1961
W
fa
-
 fasciated. Plants have flat stems, short internodes, and rugose leaves.fa was from a selection of 'White Lemon'. Robinson, 1987b*; Shifriss, 1950  
?
Fba
-
 Flower bud abortion. Preanthesis abortion of floral buds, ranging from 10% to 100%. fba from MSU 0612. Miller and Quisenberry, 1978  
?
Fdp-1
-
 Fructose diphosphatase (E.C. # 3.1.3.11). Isozyme variant found segregating in PI 137851, 164952, 113334 and 192940; 2 alleles observed. Meglic and Staub, 1996.  
P
Fdp-2
-
 Fructose diphosphatase (E.C. # 3.1.3.11). Isozyme variant found segregating in PI 137851, 164952, 113334 and 192940; 2 alleles observed. Meglic and Staub 1996  
P
Fl
-
 Fruit length. Expressed in additive fashion, fruit length decreases incrementally with each copy of fl (H. Munger, personal communication). Wilson, 1968  
W
Foc
Fcu-1
 Fusarium oxysporum f. sp. cucumerinum resistance. Resistance to fusarium wilt races 1 and 2; dominant to susceptibility. Foc from WIS 248; Foc from 'Shimshon'. Netzer et al., 1977; Vakalounakis, 1993, 1995, 1996  
W

G2dh

-

Glutamine dehydrogenase (E.C.# 1.1.1.29). Isozyme variant found segregating in PI 285606; 5 alleles observed.

Knerr and Staub, 1992

  
P
g
-
 golden leaves. Golden color of lower leaves. G and g are both from different selections of 'Nezhin'. Tkachenko, 1935  
?
gb
n
 gooseberry fruit. Small, oval-shaped fruit. gb from the 'Klin mutant'. Tkachenko, 1935  
?
gc
-
 golden cotyledon. Butter-colored cotyledons; seedlings die after 6 to 7 days. gc from a mutant of 'Burpless Hybrid'. Whelan, 1971  
W
gi
-
 ginkgo. Leaves reduced and distorted, resembling leaves of Ginkgo; male- and female-sterile. Complicated background: It was in a segregating population whose immediate ancestors were offspring of crosses and backcrosses involving 'National Pickling', 'Chinese Long', 'Tokyo Long Green', 'Vickery', 'Early Russian', 'Ohio 31' and an unnamed white spine slicer. John and Wilson, 1952  
L

gi-2

-

ginkgo-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.
Rucinska et al., 1992b
  
?

gig

-

gigantism. First leaf larger than normal. Wild type Gig from 'Borszczagowski'; gig from chemically induced mutation.

Kubicki et al., 1992b

  
?
gl
-
 glabrous. foliage lacking trichomes; fruit without spines. Iron-deficiency symptoms (chlorosis) induced by high temperature. gl from NCSU 75* and M834-6**. Robinson and Mishanec, 1964* Inggamer and De Ponti, 1980**; Robinson, 1987b
W
glb
-
 glabrate. Stem and petioles glabrous, laminae Whelan, 1973 slightly pubescent. glb from 'Burpless Hybrid'.    
W

gn

-

green mature fruit. Green mature fruits when rr GnGn; cream colored when rr GnGn; orange when R_ _. Wild type Gn from 'Chipper', SMR 58 and PI 165509; gn from TAMU 830397.

 

  
W
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.

Knerr and Staub, 1992

  
P
Gr-1
-
 Glutathione reductase-1 (E.C. # 1.6.4.2). Isozyme variant found segregating in PI 109275; 5 alleles observed.
Knerr and Staub, 1992
  
P
gy
-
 gynoecious. Recessive gene for high degree of pistillate sex expression. Kubicki,1974  
W
H
-
 Heavy netting of fruit. Dominant to no netting and completely linked or pleiotropic with black spines (B) and red mature fruit color (R). Hutchins, 1940; Tklaachenko, 1935  
W

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.

Vakalounakis, 1992

  
W

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.

Iida and Amano, 1990. 1991

  
?
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.
Iida and Amano, 1990, 1991
  
?
I
-
 Intensifier of P. Modifies effect of P on fruit warts in Cucumis sativus var. tuberculatus. Tkachenko, 1935  
?
Idh
-
 Isocitrate dehydrogenase (E.C. # 1.1.1.42). Isozyme variant found segregating in PI 183967, 21589; 2 alleles observed.
Knerr and Staub, 1992
  
P
In-de
In (de)
Intensifier of de. Reduces internode length and branching of de plants. In-de and in-de are from different selections(S5-1 and S5 -6, respectively) from a determinant inbred S2-1, which is a selection of line 541.      
In-F
F
 Intensifier of female sex expression. Increases degree of pistillate sex expression of F plants. In-F from monoecious line 18-1; in-F from MSU 713-5. Kubicki, 1969b  
?
l
-
 locule number. Many fruit locules and pentamerous androecium; five locules recessive to the normal number of three. Youngner, 1952.  
W
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.

Iida and Amano, 1990, 1991

  
?

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.

Iida and Amano, 1990, 1991

  
?
lh
-
 long hypocotyl. As much as a 3-fold increase in hypocotyl length. lh from a 'Lemon' mutant. Robinson and Shail, 1981  
W
ll
-
 little leaf. Normal-s9zed fruits on plants with miniature leaves and smaller stems. ll from Ark. 79-75. Goode et al., 1980; Wehner et al. 1987  
W

ls

-

light sensitive. Pale and smaller 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'.

Whelan, 1972b

  
L
m
a,g
 andromonoecious. Plants are andromonoecious if (mf); monoecious if (Mf); gynoecious if ( Mf) and hermaphroditic if (mF). m from 'Lemon'. Rosa, 1928*; Tkachenko, 1935 Shifriss, 1961; Wall, 1967; Youngner, 1952
W
m-2
h
 andromonoecious-2. Bisexual flowers with normal ovaries. Iezzoni, 1982; Kubicki, 1974.  
?

Mdh-1

-

Malate dehydrogenase-1 (E.C. # 1.1.1.37.). Isozyme variant found segregating in PI 171613, 209064, 326594; 3 alleles observed.

Knerr and Staub, 1992

  
P

Mdh-2

-

Malate dehydrogenase-2 (E.C.# 1.1.1.37). Isozyme variant found segregating in PI 174164, 185690, 357835, 419214; 2 alleles observed.

Knerr and Staub, 1992

  
P
Mdh-3
-
 Maleate dehydrogenase-3 (E.C.# 1.1.1.27). Isozyme variant found segregating in PI 255236, 267942, 432854, 432887; 2 alleles observed.
Knerr et al., 1995
  
P
Mdh-4
Mdh-3
 Maleate dehydrogenase-4 (E.C. # 1.1.1.37). Isozyme variant found segregating in PI 255236, 267942, 432854, 432887; 2 alleles observed. Knerr and Staub, 1992  
P
mp

pf+

pf1

pfp

 multi-pistillate. Several pistillate flowers per node,recessive to single pistillate flower per node. mp from MSU 604G and MSU 598G. Nandgaonkar and Baker, 1981 Fujieda et al., 1982
W

Mp-2

-

Multi-pistillate-2. Several pistillate flowers per node. Single dominant gene with several minor modifiers. Iida and Amano, 1990, 1991

Thaxon, 1974.

  
?
Mpi-1
-
 Mannose phosphate isomerase E.C. # 5.3.1.8). Isozyme variant found segregating in PI 176954, and 249562; 2 alleles observed. Meglic and Staub, 1996.  
P
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. Knerr and Staub, 1992  
P
mpy
mpi
 male pygmy. Dwarf plant with only staminate flowers. Wild type Mpy from Wisconsin SMR 12; mpy from Gnome 1, a selection of 'Rochford's Improved'.
Pyzhenkov and Kosareva, 1981
  
?
ms-1
-
 male sterile-1. Staminate flowers abort before anthesis; partially female-sterile. ms-1 from selections of 'Black Diamond' and 'A & C'. Shifriss, 1950 Robinson and Mishanec, 1967
L
ms-2
-
 male sterile-2. Male-sterile; pollen abortion occurs after first mitotic division of the pollengrain nucleus. Ms-2 from a mutant of 'Burpless Hybrid'. Whelan, 1973  
?
ms-2PS
-
 male sterile-2 pollen sterile. Male-sterile; allelic to ms-2, but not to ap. Ms-2PS from a mutant of SunSeeds 23B-X26. Whelan, 1973  
?
n
-
 negative geotropic peduncle response. Pistillate flowers grow upright; n from 'Lemon'; N produces the pendant flower position of most cultivars. Odland, 1963b  
W
ns
-
 numerous spines. Few spines on the fruit is dominant to many. Ns from Wis. 2757. Fanourakis, 1984; Fanourakis and Simon, 1987  
W
O
y
 Orange-yellow corolla. Orange-yellow dominant to light yellow. O and o are both from 'Nezhin'. Tkachenko, 1935  
?
opp
-
 opposite leaf arrangement. Opposite leaf arrangement is recessive to alternate and has incomplete penetrance. Opp from 'Lemon'. Robinson, 1987e  
W
P
-
 Prominent tubercles. Prominent on yellow rind of Cucumis sativus var. tuberculatus, incompletely dominant to brown rind without tubercles. P from 'Klin'; p from 'Nezhin'. Tkachenko, 1935  
W
Pc
P
 Parthenocarpy. Sets fruit without pollination. Pc from 'Spotvrie'; pc from MSU 713-205*. Pike and Peterson, 1969*; Wellington and Hawthorn, 1928; Whelan, 1973 de Ponti and Garretsen, 1976
?

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.

Fanourakis and Simon, 1987

  
W
Pep-gl-1
-
 Peptidase with glycyl-leucine (E.C.# 3.4.13.11). Isozyme variant found segregating in PI 113334, 212896; 2 alleles observed. Meglic and Staub 1996  
P
Pep-gl-2
-
 Peptidase with glycyl-leucine (E.C. # 3.4.13.11). Isozyme variant found segregating in PI 137851, 212896; 2 alleles observed. Meglic and Staub 1996  
P

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.

Knerr and Staub, 1992

  
P

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.
Knerr and Staub, 1992
  
P

Per-4

-

Peroxidase (E.C. # 1.11.1.7). Isozyme variant found segregating in PI 215589; 2 alleles observed.
Knerr and Staub, 1992
  
P

Pgd-1

-

Phosphogluconate dehydrogenase-1 (E.C. # 1.1.1.43). Isozyme variant found segregating in PI 169380, 175692, 222782; 2 alleles observed.
Knerr and Staub, 1992
  
P

Pgd-2

-

Phosphogluconate dehydrogenase-2 (E.C. # 1.1.1.43). Isozyme variant found segregating in PI 171613,177364, 188749, 263049, 285606, 289698, 354952, 419214, 432858; 2 alleles observed.
Knerr and Staub, 1992
  
P

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.
Knerr and Staub, 1992
  
P
pl
-
 pale lethal. Slightly smaller pale-green cotyledons; lethal after 6 to 7 days. Pl from 'Burpless Hybrid'; pl from a mutant of 'Burpless Hybrid'. Whelan, 1973  
L
pm-1
-
 powdery mildew resistance-1. Resistance to Sphaerotheca fuliginea. Pm-1 from 'Natsufushinari'. Fujieda and Akiya, 1962; Kooistra, 1971 Shanmugasundarum et al., 1972
?
pm-2
-
 powdery mildew resistance-2. Resistance to Sphaerotheca fuliginea. Pm-2 from 'Natsufushinari'. Fujieda and Akiya, 1962; Kooistra, 1971 Shanmugasundarum et al., 1972
?
pm-3
-
 Powdery mildew resistance-3. Resistance to Sphaerotheca fuliginea. Pm-3 found in PI 200815 and PI 200818. Kookiest, 1971 Shanmugasundarum et al., 1972
W
pm-h
s, pm
powdery mildew resistance expressed by the hypocotyl. resistance to powdery mildew as noted by no fungal symptoms appearing on seedling cotyledons is recessive to susceptibility. Pm-h from 'Wis. SMR 18'; pm-h from 'Gy 2 cp cp', 'Spartan Salad' and Wis. 2757. Fanourakis, 1984; Shanmugasundarum et al., 1971b  
W
pr
-
 protruding ovary. Extended carpels. Pr from 'Lemon'. Youngner, 1952  
W

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'.
Wang et al., 1984
  
W
psl
pl
 Pseudomonas lachrymans resistance. Resistance to Pseudomonas lachrymans is recessive. Psi from 'National Pickling' and 'Wis. SMR 18'; psl from MSU 9402 and Gy 14. Dessert et al., 1982  
W
R
-
 Red mature fruit. Interacts with c; linked or pleitropic with B and H. Hutchins, 1940  
W
rc
-
 revolute cotyledon. Cotyledons are short, narrow, and cupped downwards; enlarged perianth. Rc from 'Burpless Hybrid' mutant. Whelan et al. 1975  
L
ro
-
 rosetts. Short internodes,muskmelon-like leaves. Ro from 'Megurk', the result of a cross involving a mix of cucumber and muskmelon pollen. De Ruiter et al., 1980.  
W
s
f, a
spine size and frequency. Many small fruit spines, characteristic of European cultivars is recessive to the few large spines of most American cultivars. Strong, 1931; Tkachenko, 1935 Caruth, 1975; Poole, 1944
W
s-2
-
 spine-2. Acts in duplicate recessive epistatic fashion withs-3 to produce many small spines on the fruit. S-2 from Gy 14; s-2 from TAMU 72210. Caruth, 1975  
?
s-3
-
 spine-3. Acts in duplicate recessive epistatic fashion with s-2 to produce many small spines on the fruit. S-3 from Gy 14; s-3 from TAMU 72210. Caruth, 1975  
?
sa
-
 salt tolerance. Tolerance to high salt levels is attributable to a major gene in the homozygous recessive state and may be modified by several minor genes. Sa from PI 177362; sa from PI 19240.    
P
sc
cm
 stunted cotyledons. Small, concavely curved cotyledons; stunted plants with cupped leaves; abnormal flowers. Sc sc from Wis. 9594 and 9597. Shanmugasundarum and Williams, 1971; Shanmugasundarum et al., 1972  
W
Sd
-
 Sulfur dioxide resistance. Less than 20% leaf damage in growth chamber. Sd from 'National Pickling'; sd from 'Chipper'. Bressan et al., 1981  
W

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'.
Soltysiak and Kubicki 1988
  
?
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.
Iida and Amano, 19990, 1991
  
?
Skdh
-
 Shikimate dehydrogenase (E.C. #1.1.1.25). Isozyme variant found segregating in PI 302443, 390952, 487424; 2 alleles observed. Meglic and Staub, 1996  
P
sp
-
 short petiole. Leaf petioles of first nodes 20% the length of normal. Sp from Russian mutant line 1753. Den Nijs and de Ponti, 1983  
W

sp-2

-

short petiole-2. Leaf petioles shorter, darker green than normal at 2-leaf stage; crinkled leaves with slow development; short hypocotyl and stem; little branching. Not tested for allelism with sp. Wild type Sp-2 from 'Borszczagowski'; sp-2 from chemically induced mutation.

Rucinska et al., 1992a

  
?
ss
-
 small spines. Large, coarse fruit spines is dominant to small, fine fruit spines. Ss from 'Spartan Salad', 'Wis. SMR 18' and 'GY 2 cp cp'; ss from Wis. 2757. Fanourakis, 1984; Fanourakis and Simon, 1987  
W
T
-
 Tall plant. Tall incompletely dominant to short. Hutchins, 1940  
?
td
-
 tendrilless. Tendrils lacking; associated with misshapen ovaries and brittle leaves. Td from 'Southern Pickler'; td from a mutant of 'Southern Pickler'. Rowe and Bowers, 1965.  
W
te
-
 tender skin of fruit. Thin, tender skin of some European cultivars; recessive to thick tough skin of most American cultivars. Poole, 1944; Strong, 1931  
W
Tr
-
 Trimonoecious. Producing staminate, perfect, and pistillate flowers in this sequence during plant development. Tr from Tr-12, a selection of a Japanese cultivar belonging to the Fushinari group; tr from H-7-25. MOA-309, MOA-303, and AH-311-3. Kubicki, 1969d  
P
Tu
-
 Tuberculate fruit. Warty fruit characteristic of American cultivars is dominant to smooth, non-warty fruits characteristic of European cultivars. Strong, 1931; Wellington, 1913 Andeweg, 1956; Poole, 1944
W
u
M
 uniform immature fruit color. Uniform color of European cultivars recessive to mottled or stippled color of most American cultivars. Strong, 1931 Andeweg, 1956
W
ul
-
 umbrella leaf. Leaf margins turn down at low relative humidity making leaves look cupped. ul source unknown. den Nijs and de Ponti, 1983  
W
v
-
 virescent. Yellow leaves becoming green. Strong, 1931; Tkachenko, 1931  
L
vvi
-
 variegated virescent. Yellow cotyledons, becoming green; variegated leaves. Abul-Hayja and Williams, 1976  
L
w
-
 white immature fruit color. White is recessive to green.W from 'Vaughn', 'Clark's Special;', 'Florida Pickle' and 'National Pickling'; w from 'Bangalore'. Cochran, 1938  
W
wf
-
 White flesh. Intense white flesh color is recessive to dingy white; acts with yf to produce F2 of 12 white (Wf Yf and wf Yf) : 3 yellow (Wf yf) : 1 orange (wf yf). Wf from EG and G6, each being dingy white (Wf from EG and G6, each being dingy white (Wd Yf) : wf from 'NPI' which is orange (wf yf). Kooistra, 1971  
?

wi

 

-

wilty leaves. Leaves wilting in the field, but not in shaded greenhouse; weak growth; no fruiting. Wild type Wi from 'Nishiki-suyo'; wi from M2 line from pollen irradiation.

Iida and Amano, 1990, 1991

  
?
Wmv
-
 Watermelon mosaic virus resistance. Resistance to strain 2 of watermelon mosaic virus. Wmv from 'Kyoto 3 Feet'; wmv from 'Beit Alpha'. Cohen et al., 1971  
P
wmv-1-1
-
 watermelon mosaic virus-1 resistance. Resistance to strain 1 of watermelon mosaic virus by limited systematic translocation; lower leaves may show severe symptoms. Wmv-1-1 from Wis. 2757; wmv-1-1 from 'Surinam'. Wang et al., 1984  
?

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.

Iida and Amano, 1990, 1991

  
?
yc-1
-
 yellow cotyledons-1. Cotyledons yellow at first, later turning green. yc-1 from a mutant of Ohio MR 25. Aalders, 1959  
W
yc-2
-
 yellow cotyledons-2. Virescent cotyledons. Virescent cotyledons. yc-2 from a mutant of 'Burpless Hybrid'. Whelan and Chubey, 1973; Whelan et al., 1975.  
W
yf
v
 yellow flesh. Interacts with wf to produce F2 of 12 white (Wf Yf and wf Yf) : 3 yellow (Wf yf) : 1 orange (wf yf). Yf from 'Natsufushinari', which has an intense white flesh (Yf wf); yf from PI 200815 which has a yellow flesh (yf Wf). Kooistra,1971  
P
yg
gr
 yellow-green immature fruit color. Recessive to dark green and epistatic to light green. yg from 'Lemon'. Youngner, 1952  
W
yp
-
 yellow plant. Light yellow-green foliage; slow growth. Abul-Hayja and Williams, 1976  
?

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.

Rucinska et al., 1991

  
?
zymv
-
 zucchini yellow mosaic virus resistance. Inheritance is incomplete. Believed to be inherited in a recessive fashion with the source of resistance being 'TMG-1'. Provvidenti, 1985  
W

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