Cucurbit Genetics Cooperative Report 28-29: 2015
Shengping Zhang
Chinese Academy of Agricultural Sciences, Beijing, P.R. China
Todd C. Wehner
Department of Horticultural Science, North Carolina State University, Raleigh, NC 27695-7609 U.S.A.
Introduction
This is the latest version of the gene list for cucumber (Cucumis sativus L. ). In addition to morphological and resistance genes, this list includes genes that have been cloned from different plant tissues of cucumber. The genes in the list have been grouped into ten categories as follows: seedling markers, stem mutants, leaf mutants, flower mutants, fruit type mutants, fruit color mutants, resistance genes (mostly to diseases), protein (isozyme) variants, DNA (RFLPs and RAPDs) markers (Table 1), and cloned genes (Table 2). There is also a review of linkage of the morphological and resistance genes. As of 2010 the number of cloned genes of cucumber has become to vast for this article. Up to date information on cloned cucumber genes can be found at the National Center for Biotechnology Information Database (http://www.ncbi.nlm.nih.gov/). Complete lists and updates of genes for have been published previously, as follows:
Previous Gene Lists
- Robinson et al., 1976
- Robinson et al., 1982
- Pierce and Wehner, 1989
- Wehner, 1993
- Wehner and Staub, 1997
- Xie and Wehner, 2001
- Wehner, 2005
- Call and Wehner, 2010
Revisions to the 2015 cucumber gene list include the addition of ?????. Researchers are encouraged to send reports of new genes, as well as seed samples to the cucumber gene curators (Yiqun Weng and Todd C. Wehner). 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.
Seed and seedling mutants
One of the advantages of using the cucumber in genetic research is the availability of seedling markers. To date, five non-lethal color mutants [virescent (v) (Poole, 1944; Tkachenko, 1935), variegated virescence (vvi) (Abul-Hayja and Williams, 1976), yellow cotyledons-1 (yc-1) (Aalders, 1959), yellow cotyledons-2 (yc-2) (Whelan and Chubey, 1973; Whelan et al., 1975), yellow plant (yp) (Abul-Hayja and Williams, 1976)] and 4 lethal, color mutants [chlorophyll deficient (cd) (Burnham et al., 1966), golden cotyledon (gc) (Whelan, 1971), light sensitive (ls) (Whelan, 1972b), pale lethal (pl) (Whelan, 1973)] have been identified.
Six seedling traits which affect traits other than color include bitterfree (bi) (Andeweg and DeBruyn, 1959), blind, (bl) (Carlsson, 1961), delayed growth (dl) (Miller and George, 1979), long hypocotyl (lh) (Robinson and Shail., 1981), revolute cotyledons (rc) (Whelan et al., 1975) and stunted cotyledons (sc) (Shanmugasundarum and Williams, 1971; Shanmugasundarum et al., 1972).
Seedling chilling resistance is conferred by the dominant gene Ch (Kozik and Wehner, 2006, 2008).
Stem mutants
Seven genes have been identified which affect stem length: bush (bu) (Pyzenkov and Kosareva, 1981), compact (cp) (Kauffman and Lower, 1976), determinate (de) (Denna, 1971; George, 1970; Hutchins, 1940), dwarf (dw) (Robinson and Mishanec, 1965), tall height (T) (Hutchins, 1940) and In-de which behaves as an intensifier for de (George, 1970). Rosette (ro) which also affects height is characterized by muskmelon-like leaves (de Ruiter et al., 1980).
Unlike these genes, fasciated (fa) (Robinson, 1978b; Shifriss, 1950) affects stem confirmation, not length.
Leaf mutants
Several genes have been shown to control leaf or foliage characteristics. Eight in particular are responsible for leaf shape: blunt leaf apex (bla) (Robinson, 1987a), cordate leaves-1 (cor-1) (Gornitskaya, 1967), cordate leaves-2 (cor-2) (Robinson, 1987c), crinkled leaf (cr) (Odland and Groff, 1963a), divided leaf (dvl) (den Nijs and Mackiewicz, 1980), ginko leaf (gi) (John and Wilson, 1952), little leaf (ll), (Goode et al., 1980; Wehner et al., 1987) and umbrella leaf (ul) (den Nijs and de Ponti 1983). Note that ginko leaf is a misspelling of the genus Ginkgo.
The original cordate leaf gene identified by Gornitskaya (1967) differs from cor proposed by (Robinson, 1987c) which also had calyx segments which tightly clasp the corolla, hindering flower opening and insect pollination. Therefore, we propose that the first gene identified by Gornitskaya be labeled cor-1 and the second identified by Robinson be labeled cor-2. It should be noted that plants with stunted cotyledon may look similar to those with ginko at the younger stages but the cotyledons of sc mutants are irregular and gi mutants are sterile.
Opposite leaf arrangement (opp) is inherited as a single recessive gene with linkages to m and l. Unfortunately, incomplete penetrance makes the opposite leaf arrangement difficult to distinguish from normal plants with alternate leaf arrangement (Robinson, 1987e).
Five mutants which affect color or anatomical features of the foliage are golden leaves (g) (Tkachenko, 1935), glabrous (gl) (Inggamer and de Ponti, 1980; Robinson and Mishanec, 1964), glabrate (glb) (Whelan, 1973), short petiole (sp) (den Nijs and Boukema, 1985) and tendrilless (td) (Rowe and Bowers, 1965).
Flower mutants
Sex expression in cucumber is affected by several single-gene mutants. The F locus affects gynoecy (femaleness), but is modified by other genes and the environment, and interacts with a and m (androecious and andromonoecious, respectively) (Galun, 1961; Kubicki, 1969; Rosa, 1928; Shifriss, 1961; Tkachenko, 1935; Wall, 1967). Androecious plants are produced if aa and ff occur in combination, otherwise plants are hermaphroditic if mm FF, andromonoecious if mm ff, gynoecious if MM FF and monoecious if MM ff. The gene F may also be modified by an intensifier gene In-F which increases the femaleness (Kubicki, 1969b). Other genes that affect sex expression are gy for gynoecious (Kubicki, 1974), m-2for andromonoecious (Kubicki, 1974) and Tr for trimonoecious expression (Kubicki, 1969d).
Cucumbers, typically considered day-neutral plants, have occasionally been shown to express sensitivity to long days. Della Vecchia et al. (1982) and Shifriss and George (1965) demonstrated that a single gene for delayed flowering (df) is responsible for this short-day response.
Another gene which may give the impression of eliciting daylength sensitivity by causing a delay in flowering is Fba. In reality, Fba triggers flower bud abortion prior to anthesis in 10 to 100% of the buds (Miller and Quisenberry, 1978).
Three separate groups have reported single genes for multiple pistillate flowers per node. Nandgaonkar and Baker (1981) found that a single recessive gene mp was responsible for multiple pistillate flowering. This may be the same gene which Fujieda et al. (1982) later labeled as pf for plural pistillate flowering. However, they indicated that 3 different alleles were responsible, with single pistillate being incompletely dominant over multiple pistillate: pf + for single pistillate, pf d for double pistillate and pf m for multiple pistillate (more than 2 flowers per node).
Thaxton (1974), reported that clustering of pistillate flowers is conditioned by a single dominant gene (we propose the symbol, Mp-2), and that modifier genes influence the amount of clustering. Thaxton (1974) also determined that clustering of perfect flowers is controlled by genes different from clustering of gynoecious flowers.
Several genes for male sterility have been reported for cucumber, but because of the ease of changing sex expression with growth regulators, little commercial use has been made of them. Five genes, ms-1, ms-2, ap, cl and gi have been identified. The genes ms-1 and ms-2 cause sterility by pollen abortion before anthesis; ms-1 plants are also partially female sterile (Robinson and Mishanec, 1965; Shanmugasundarum and Williams, 1971; Whelan, 1972a). Apetalous mutants (ap) on the other hand have infertile anthers which appear to have been transformed into sepal-like structures (Grimbly, 1980). Ginko (gi), mentioned earlier as a leaf mutant, also causes male sterility (John and Wilson, 1952).
One of these male steriles may be of little use except as a genetic marker. Closed flower (cl) mutants are both male and female sterile, so seed production must be through the heterozygotes only (Groff and Odland, 1963). With this mutant, the pollen is inaccessible to bees because the buds remain closed.
Three genes alter floral characteristics: green corolla (co) (Currence, 1954; Hutchins, 1935), orange-yellow corolla (O) (Tkachenko 1935), negative geotropic peduncle response (n) (Odland and Groff, 1963b). Green corolla (co), named because of its green petals, has enlarged but sterile pistils (Currence, 1954; Hutchins, 1935), and has potential for use as a female sterile in hybrid production.
Fruit mutants
Because the fruit is the most important part of the cucumber economically, considerable attention has been given to genes affecting it. One such gene is Bitter fruit, Bt, (Barham, 1953) which alters fruit flavor by controlling cucurbitacin levels. The gene Bt is different from bi because it consistently alters only the fruit cucurbitacin levels compared to bi which affects the whole plant.
Five genes conditioning skin texture are Tu (Strong, 1931; Wellington, 1913), te (Poole, 1944; Strong, 1931), P (Tkachenko, 1935), I (Tkachenko, 1935) and H (Hutchins, 1940; Tkachenko, 1935). Smooth (Tu) and tender (te) skin are usually associated with European types, while American types are generally warty and thick skinned (Poole, 1944; Strong 1931). Heavy netting, H, which occurs when fruit reach maturity may be tightly linked or pleiotropic with R and B (discussed later).
In Cucumis sativus var. tuberculatus, Tkachenko (1935) found that gene P, causing fruit with yellow rind and tubercles, was modified by gene I, an intensifier which increases the prominence of the tubercles (Tkachenko, 1935).
There are 3 genes which affect internal fruit quality, each identified by viewing transections of fruits; Empty chambers-1 (Es-1), Empty chambers-2 (Es-2) (Kubicki and Korzeniewska, 1983) and locule number (l) (Youngner, 1952).
Hutchins (1940) proposed that 2 genes controlled spine characteristics, with f producing many spines and being tightly linked with s which produced small spines. Poole (1944) used the data of Hutchins (1940) to suggest that s and f were the same gene and proposed the joint symbol s for a high density of small spines. Tkachenko (1935) who used the same symbol for control of less dense spines, did not look at spine size, and the same gene might have been involved. However, Fanourakis (1984) and Fanourakis and Simon (1987) reported 2 separate genes involved, and named them ss and ns for small spines and numerous spines, respectively.
These may differ from those that led Carruth (1975) to conclude that 2 genes act in a double recessive epistatic fashion to produce the dense, small spine habit. We propose that these genes be labeled s-2 and s-3 and s1 be used instead of sproposed by Poole (1944).
Carruth (1975) and Pike and Carruth (1977) also suggested that carpel rupture along the sutures was inherited as a single recessive gene that was tightly linked with round, fine-spined fruits. This may be similar to what Tkachenko (1935) noted in the ‘Klin mutant’ as occasional deep-splitting flesh. We suggest the symbol cs for carpel splitting, but note that because penetrance of the trait may be lower under certain environmental conditions (Carruth, 1975) this trait may be related to the gooseberry (gb) fruit reported by Tkachenko (1935). Another character not found in commercial cultivars was protruding ovary (pr) reported by Youngner (1952).
There is dispute over the inheritance of parthenocarpy, a trait found in many European cucumbers (Wellington and Hawthorn, 1928). Pike and Peterson (1969) suggested an incompletely dominant gene, Pc, affected by numerous modifiers, was responsible. In contrast, de Ponti and Garretsen (1976) explained the inheritance by 3 major isomeric genes with additive action.
A modifier of fruit length, Fl, was identified by its linkage with scab resistance (Cca) (Henry Munger, personal communication; Wilson, 1968). Expressed in an additive fashion, fruit length decreases incrementally from heterozygote to homozygote (fl fl).
A gene for inheritance of twin fused fruit, tf, was discovered in gynoecicous inbred B 5263 (Klosinska et al., 2006). The trait is characterized by pairs of two separate pistallate flowers with partially joined ovaries on a single peduncle at a node developing into twin fused fruit during development. Twin fused fruit were only observed on gynoecious plants, indicating epistatis.
Fruit Color
Twelve mutants have been identified which affect fruit color either in the spines, skin, or flesh and a few of these appear to act pleiotropically. For example, R for red mature fruit color is very closely linked or pleiotropic to B for black or brown spines and H for heavy netting (Hutchins, 1935; Tkachenko, 1935; Wellington, 1913). It also interacts with c for cream colored mature fruit in such a way that plants which are (RR CC), (RR cc), (rr CC) and (rr cc) have red, orange, yellow and cream colored fruits, respectively (Hutchins, 1940).
The B gene produces black or brown spines and is pleiotropic to or linked with R and H (Wellington, 1913). The homozygous recessive plant is white spined with cream colored mature fruit and lacks netting. Other spine color genes are B-2, B-3 and B-4 (Cowen and Helsel, 1983; Shanmugasundarum et al., 1971a).
White immature skin color (w) is recessive to the normal green (Cochran, 1938), and yellow green (yg) is recessive to dark green and epistatic with light green (Youngner, 1952). Skin color may also be dull or glossy (D) (Strong, 1931; Tkachenko, 1935) and uniform or mottled (u) (Andeweg, 1956; Strong, 1931).
Kooistra (1971) reported 2 genes that affect fruit mesocarp color. White flesh (wf) and yellow flesh (yf) gene loci interact to produce either white (WfWf YfYf or wfwf YfYf), yellow (WfWf yfyf), or orange (wfwf yfyf) flesh color.
Insect Resistance
Bitterfree, bi (Andeweg and DeBruyn, 1959), is responsible for resistance to spotted and banded cucumber beetles (Diabrotica spp. ) (Chambliss, 1978; Da Costa & Jones, 1971a; Da Costa & Jones, 1971b) and two-spotted spider mites (Tetranychus urticae Koch. ) (Da Costa & Jones, 1971a; Soans et al., 1973). However, this gene works inversely for the 2 species. The dominant allele which conditions higher foliage cucurbitacin levels incites resistance to spider mites by an antibiotic affect of the cucurbitacin. The homozygous recessive results in resistance to cucumber beetles because cucurbitacins are attractants.
In the 1989 Cucurbit Genetics Cooperative Report the authors labeled the gene for resistance to Diabrotica spp. di, but wish to retract it in light of recent evidence.
Disease Resistance
Currently there are 15 genes known to control disease resistance in C. sativus. Three of these condition virus resistance. Wasuwat and Walker (1961) found a single dominant gene, Cmv, for resistance to cucumber mosaic virus. However, others have reported more complex inheritance (Shifriss et al., 1942). Two genes condition resistance to watermelon mosaic virus, Wmv (Cohen et al, 1971) and wmv-1-1 (Wang et al., 1984). Most recently, resistance to zucchini yellow mosaic virus (zymv) has been identified (Provvidenti, 1985).
Both resistance to scab, caused by Cladosporium cucumerinum Ell. & Arth., and resistance to bacterial wilt caused by Erwinia tracheiphila (E. F. Smith) Holland are dominant and controlled by Ccu (Abul-Hayja et al., 1978; Andeweg, 1956; Bailey and Burgess, 1934) and Bw (Nuttall and Jasmin, 1958; Robinson and Whitaker, 1974), respectively. Other dominant genes providing resistance are: Cca for resistance to target leaf spot (Corynespora cassiicola) (Abul-Hayja et al., 1978), Cm for resistance to Corynespora blight (Corynespora melonis) (van Es, 1958), Foc for resistance to Fusarium wilt (Fusarium oxysporum f. sp. cucumerinum) (Netzer et al., 1977) and Ar for resistance to anthracnose [Colletotrichum lagenarium (Pars. ) Ellis & Halst. ] (Barnes and Epps, 1952). In contrast, resistance to Colletotrichum lagenarium race 1 (Abul-Hayja et al., 1978) and angular leaf spot (Pseudomonas lachrymans) (Dessert et al., 1982) are conditioned by the recessive genes cla and psl, respectively.
Several reports have indicated that more than one gene controls resistance to powdery mildew [Sphaerotheca fuliginea (Schlecht) Poll. ] with interactions occurring among loci (Hujieda and Akiya, 1962; Kooistra, 1968; Shanmugasundarum et al., 1971b). The resistance genes pm-1 and pm-2 were first reported by Hujieda and Akiya (1962) in a cultivar which they developed and named ‘Natsufushinari’. Kooistra (1968) using this same cultivar, later confirmed their findings and identified one additional gene (pm-3) from USDA accessions PI200815 and PI200818. Shimizu et al. (1963) also supported 3 recessive genes which are responsible for resistance of ‘Aojihai’ over ‘Sagamihan’.
Several genes with specific effects have been identified more recently (Shanmugasundarum et al., 1971b) but unfortunately, direct comparisons were not made to see if the genes were identical with pm-1, pm-2 and pm-3. Fanourakis (1984) considered a powdery mildew resistance gene in an extensive linkage study and proposed that it was the same gene used by Shanmugasundarum et al. (1971b) which also produces resistance on the seedling hypocotyl. Because expression is identified easily and since it is frequently labeled in the literature as ‘pm‘ we believe that this gene should be added to the list as pm-h with the understanding that this may be the same as pm-1, pm-2 or pm-3.
There are several proposed inheritance patterns for resistance to downy mildew. They range from three recessive genes (Doruchowski and Lakowska-Ryk, 1992; Shimizu et al., 1963) to three partially dominant genes (Pershin et al., 1988) to an interaction between dominant susceptible and recessive resistance genes (Badr and Mohamed, 1998; El-Hafaz et al., 1990) to one or two incompletely dominant genes (Petrov et al., 2000) to a single recessive gene (Angelov, 1994; Fanourakis and Simon, 1987; Van Vliet and Meysing, 1974; 1976). Doruchowski and Lakowska-Ryk (1992) had evidence that downy mildew resistance was controlled by three recessive genes (dm-1, dm-2 and dm-3), where dm-3 and either dm-1 or dm-2 had to be homozygous recessive for maximum resistance. However, there was discrepancy in the F2 results, which did not agree with their model. They argued that this resulted from testing too narrow a population. The three genes were included in previous cucumber gene lists but have been removed from the current list as none of the genes were identified and no type lines are available to use in studies of separate genes.
One gene, dm (dm-1), has been identified which confers resistance to downy mildew [Pseudoperonospora cubensis (Berk. & Curt. ) Rostow] (van Vliet and Meysing, 1974). Inherited as a single recessive gene, it also appeared to be linked with pm (van Vliet, 1977). The dm gene traces its resistance to PI 197087, a wild accession collected in India. There are, however, indications that more than one gene may be involved (Jenkins, 1946). Angelov (1994) reported that PI 197088 resistance was due to two recessive genes. PI 197088 was collected from the same region and at the same time as PI 197087. PI 197088 was recently reported as highly resistant to the current downy mildew in the southeastern U.S., while cultigens tracing resistance to PI 197087 are no longer highly resistant (Call, 2010). Interestingly, it appears the 2004 change of the downy mildew population resulted in a change in rank of resistant and moderately resistant cultigens. Previously highly resistant cultignes are now moderate, while cultigenes that are now highly resistant were only moderate prior to 2004 (Call and Wehner, 2010). The first reported source of resistance was from a Chinese cultigen tested at the Puerto Rico Agricultural Experiment Station in 1933. This cultigen was crossed with the best available commercial cultivars to combine the resistance with good horticultural traits. This eventually led to seven resistant lines having good characteristics. Of these, PR 37, PR 39, and PR 40 had high quality fruit and yield superior to commercial checks. These were used in further development of resistant breeding lines and cultivars (Barnes et. al. 1946; Barnes, 1955; Barnes and Epps, 1955). It appears that there are at least three genes for resistance to downy mildew in cucumber: one from the Chinese cultivar used in developing the PR lines, one from PI 197087, and one from PI 197088 (assuming that PI 197087 and PI 197088 share one resistance gene, dm-1).
Multiple genes for virus resistance have been reported, including genes conferring resistance to CMV, MWM, PRSV, WMV, and ZYMV. The resistance gene cmv was identified in ‘National Pickling’ and ‘Wis. SR 6’ for resistance cucumber mosaic virus was reported by Wasuwat and Walker (1961). Kabelka and Grumet (1997) reported Moroccan watermelon mosaic virus resistance as a single recessive gene, mwm, from Chinese cucumber cultivar ‘TMG-1’. Resistance to papaya ringspot virus (formerly watermelon mosaic virus 1) was identified in the cultivar ‘Surinam’ by Wang et al. (1984). Multiple genes for resistance to watermelon mosaic virus (WMV) have been reported. Cohen et al. (1971) identified a dominant gene, Wmv, in the cultivar ‘Kyoto 3 Feet’, resistant to strain 2 of WMV. The gene wmv-1-1 from the cultivar ‘Surinam’ was reported by Wang et al. (1984) to give resistance to strain 1 of WMV by means of limiting systemic translocation. Cultigens having wmv-1-1 may show severe symptoms on lower leaves, while symptoms on the rest of the plant are limited. Wai et al. (1997) reported three genes for watermelon mosaic virus resistance from the cultigen TMG-1: wmv-2, wmv-3, and wmv-4. Expression of wmv-3 and wmv-4 is limited to true leaves only, while wmv-2 is expressed throughout the plant. Two genes have been reported conferring resistance to zucchini yellow mosaic virus: zym-Dina from the cultigen Dina-1 and zym-TMG1 from the cultigen TMG-1. Inheritance of zym-TMG1 is incomplete but typically recessive.
Environmental Stress Resistance
Two genes have been identified for stress resistance, resistance to sulfur dioxide air pollution conditioned by Sd (Bressan et al., 1981) and increased tolerance to high salt levels conditioned by the gene, sa, Jones (1984).
Other Traits
The dominant allele, Psm, induces paternal sorting of mitochondria, where Psm is from MSC 16 and psm is from PI 401734 (Havey et al., 2004).
Molecular and Protein Markers
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 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 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 EcoRVdigested 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 CsP344/E1, CsC477/H3, CsP300/E1).
Clones are designated herein as CsC = cDNA, CsP = PstI-genomic, 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, DraI; 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, CsP078s/H3, CsC581/E5, CsE084/E1, CsC341/H3, CsP024/E1, CsP287/H3, CsC629/H3, CsP225s/E1, CsP303/H3, CsE051/H3, CsC366a/E5, CsC032b/E1, CsP056/H3, CsC378/E1, CsP406/E1, CsP460/E1, CsE060/E1, CsE103/E1, CsP019/E1, CsP168/D1, CsC560/H3, CsP005/E1, CsP440s/E1, CsP221/H3, CsC625/E1, CsP475s/E1, CsP211/E1, 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, CsC137/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/H3, CsC308/E5, CsP073/E5, CsP215/H3, CsC613/D1, 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, OPAB05, OPH12, OPA14a, and BC211a.
In addition to the isozymes, RFLPs and RAPDs, nearly 100 cloned genes are listed here (Table 2).
Possible Allelic or Identical Genes
Several of the genes listed may be either pleiotropic, closely linked, or allelic. Additional research is needed to compare the sources of the various similar genes to ensure that they are not duplicates. In some instances this may be difficult because many of the earlier publications did not list the source of the genes or the methods used to measure the traits, and many of these authors are deceased.
An example of this is the two-locus model (R c) for fruit color. We have been unable to locate any plants with red or yellow colored mature fruits. All plants evaluated in other studies have color inherited as a single gene. Hutchins may have separated fruit with cream color into 2 groups, yellow and cream, and fruits with orange color into two groups, orange and red. However, those distinctions are difficult to make using available germplasm. Situations like these may be impossible to resolve. Red mature fruit may be the same as the brownish-red cracked or netted exocarp found in some accessions such as PI 330628. Orange fruit occur in black spined accessions such as Wis. SMR 18.
In the future, researchers should use the marker lines listed here, or describe and release the marker lines used so that allelism can be checked by others. Currently, groups of similar genes that need to be checked to determine how they are related include the following: the chlorophyll deficiency mutants (cd, g, ls, pl, v, vvi, yc-1, yc-2, and yp), the stem mutants (bu, de, dw, In-de, and T), the leaf shape mutants (rc and ul), the sex expression mutants (a, F, gy, In-F, m, m-2, and Tr), the male sterility genes (ap, cl, ms-1, and ms-2), the flowering stage mutants (df and Fba), the flower color mutants (co and O), the powdery mildew resistance mutants (pm-1, pm-2, pm-3 and pm-h), the fruit spine color mutants (B-1, B-2, B-3, and B-4), the fruit skin color mutants (c, R, and w), the spine size and density mutants (s, s-2, and s-3) and the seed cell mutants (cs and gb).
Two groups of associated traits, one from ‘Lemon’ cucumber (m, pr, and s) and the other involving fruit skin color, surface texture, and spine type (R, H, and B), need to be checked using large populations to determine whether they are linked or pleiotropic. Recent gains have been made in this area by Robinson (1978a) who demonstrated that the m gene is pleiotropic for fruit shape and flower type, producing both perfect flowers and round fruits, and Abul-Hayja et al. (1975) and Whelan (1973) who determined that gl and glb are independent genes.
New information indicates that comparisons also need to be made between resistance to scab (Ccu) and Fusarium wilt (Foc) and between resistance to target leaf spot (Cca) and Ulocladium cucurbitae leafspot. Mary Palmer (personal communication) found a fairly consistent association between resistances to scab and Fusarium wilt, which suggests that they might be linked or using the same mechanism for defense against the pathogen.
Similar defense mechanisms might also be responsible for similarities in resistance to target leaf spot (Cca) and Ulocladium cucurbitae leafspot (Henry Munger, personal communication).
Genetic Linkage
Since cucumber has just 7 chromosome pairs and over 100 known genes, it would seem that linkage maps would be fairly complete by now. Unfortunately, we know of few references reporting linkages of more than 2 gene loci, and this is the first review to summarize the literature for linkages and attempt to describe different linkage groups.
Many difficulties were encountered and should be considered when reading this review. First, a portion of the nomenclature is still unclear and some of the genes may be duplicates of others since common parents were not compared. This problem was discussed in the previous section. Secondly, some of the linkage relationships analyzed in previous studies did not involve specific genes. Linkages in several reports were discussed for plant traits that might have been inherited in multigenic fashion, or if a single gene were involved, it was not specifically identified.
Therefore, in this review linkages for traits without genes will be omitted and a ‘?’ will follow each gene which has a questionable origin. Six linkage groups could be determined from the current literature (Fig. 1). The order in which the genes were expressed in each group does not necessarily represent the order in which they may be found on the chromosome.
Linkage Group A
The largest linkage group in cucumber has 14 genes, composed of wmv-1-1, wmv-2, Prsv-2, gy, gl, dl, dvl, de, F, ms-2, glb, bi, df and B-3 or B-4. In contributing to this grouping, Whelan (1974) noted that ms-2 is linked with glb (rf=.215+.029) and de (rf=.335+.042) while being independent of bi, gl, yc-1, yc-2, and cr. Gene de is linked with F (Odland and Groff, 1963b; Owens and Peterson, 1982) which in turn is linked with B-3 or B-4 (Cowen and Helsel, 1983), gy (rf=.04) (Kubicki, 1974), bi (rf=.375) and df (rf=34.7) (Fanourakis, 1984; Fanourakis and Simon, 1987). Gene de is also weakly linked with dl (Miller and George, 1979), strongly linked with dvl (Netherlands, 1982), and independent of cp (Kauffman and Lower, 1976). Gene wmv-1-1 is linked with bitterfree (bi) but independent of Ccu, B, F or pm (Wang et al., 1987). Wai et al. (1997) showed linkage between Prsv-2 and bi (rf=.28), bi and F (rf=.34), and F and wmv-2 (rf=.33).
Two reports show that dvl is weakly linked with gl (rf=.40) and independent of bi and Ccu (Netherlands, 1982; den Nijs and Boukema, 1983), while Robinson (1978f) originally indicated that gl was linked with yc and independent of B, m, l, and yg as well as bi (Netherlands, 1982) and sp (den Nijs and Boukema, 1985), but more recently he indicated that gl was independent of yc (Robinson, 1987d).
Completing linkage group A, Cowen and Helsel (1983) demonstrated that the spine color genes (B-3 and B-4) were independent of the genes for bitterness, and Whelan (1973) found that pl was independent of glb and bi, while glb was independent of gl, bi, ls, yc, and cr. The last clarifies that gl and glb must indeed be separate loci.
Linkage Group B
Group B is composed of 9 genes (n, pr, l, m, opp, m-2, Bw, s? and ms?) unless s? (Robinson, 1978) is the same as s from Hutchins (1940) and Poole (1944). If these are the same, then linkage groups II and III will be joined for a total of 12 genes. Of the first 7, two pairs have been defined with recombination values. Youngner (1952) determined that m and l were linked with a recombination frequency of .326 + .014 and Robinson determined that opp was linked to both (Robinson, 1987e). Iezzoni and Peterson (1979, 1980) found that m and Bw were separated by only one map unit (rf=.011+.003). Iezzoni et al. (1982) also determined that m-2 was closely linked with both m and Bw, and that Bw was independent of Ffrom linkage group I (Iezzoni and Peterson, 1980).
Robinson (1978c, 1978d), and Youngner (1952) found that linkages existed between m, l, n, pr and spine number (s?) with the possibility of pleiotropy being responsible for the m/pr relationship. They also demonstrated that B, yg, and pm? were independent of the same genes (Robinson, 1978c; Youngner, 1952).
Rounding out the linkage group is one of the male sterility genes (ms?). Robinson (1978d) found that it was linked with both m and l, but did not identify which male sterile gene it was.
Linkage Group C
Group C is the oldest and most mystifying linkage group. It is currently composed of R for red or orange mature fruit color, H for heavy netting, B for black or brown spine color, c for cream mature fruit color and s for spine frequency and size (Hutchins, 1940; Poole, 1944; Strong, 1931; Tkachenko, 1935). However, there is speculation on the nature of this linkage group. Since very few recombinants of the R, H, B and c, h, b linkage groups have been reported, it is also felt that these characteristics may be the response of 2 alleles at a single pleiotropic gene. There is also speculation that R and c are different alleles located at the same locus (see earlier discussion).
Hutchins (1940) found that s was independent of B and H while s was linked with R and c. If he was correct, then pleiotropy of H and B with R and c is ruled out. His report also indicated that B and s were independent of de as was de of R, c and H.
A possibility exists that this linkage group may be a continuation of group II through the s gene. Poole (1944) used the data of Hutchins (1940) to determine that c and s are linked with a recombination frequency of .163 + .065. The question that remains is whether s (Hutchins, 1940; Poole, 1944) is the same as the gene for spine number in the findings of Robinson (1978c). If Cowen and Helsel (1983) are correct in their finding that a linkage exists between F and B then groups I and III may be on the same chromosome. However, in this text they will remain separated based on conclusions of Fanourakis (1984) which indicate that errors may be common when attempting to distinguish linkages with F since classification of F is difficult. This difficulty may also explain many conflicting reports.
Linkage Group D
Twelve genes (ns, ss, Tu, Pc, D, U, te, cp, dm, Ar, coca and pm? or pm-h) are in group D, but the identity of the specific gene for powdery mildew resistance is elusive. Van Vliet and Meysing (1947, 1977) demonstrated that the gene for resistance to downy mildew (dm) was either linked or identical with a gene for resistance to powdery mildew (pm?), but because the linkage between pm? and D was broken while that of dm and D was not, pm? and dm must be separate genes. The problem lies in the lack of identity of pm? because Kooistra (1971) also found that a gene for powdery mildew resistance (pm?) was linked to D.
Further complicating the identity of pm, Fanourakis (1984) found that pm-h was linked to te and dm, yet cp, which must be located at approximately the same locus, was independent of te. He suggested that there were either 2 linkage groups, ns, ss, Tu, Pc, D, U, te and cp, dm, Ar, located at distal ends of the same chromosome with pm-h at the center, or the 2 groups are located on different chromosomes with a translocation being responsible for apparent cross linkages. However, evidence for the latter which suggested that F was associated with the 7-gene segment is not probable since there are few other supportive linkages between genes of this segment and linkage group I. A more likely explanation is the occurrence of 2 or more genes conditioning resistance to powdery mildew being found on this chromosome.
More recently Lane and Munger (1985) and Munger and Lane (1987) determined that a gene for resistance to powdery mildew (pm?) was also linked with coca for susceptibility to target leaf spot but that linkage, though fairly tight, was breakable.
The last 4 genes in this group are Tu, D, te and u (Strong, 1931). Until recently it was believed that each in the recessive form were pleiotropic and consistent with European type cucumbers and each in the dominant form were pleiotropic and consistent with American type cucumbers. Fanourakis (1984) and Fanourakis and Simon (1987) reported that crossing over (R=23.7) occurred between te and the other 3 genes which still appeared to be associated. However, using triple backcrosses they demonstrated that there is a definite order for Tu, D and u within their chromosome segment and that the Tu end is associated with the ns and ss end.
Linkage Group E
Group E is currently composed of 3 genes lh, sp and ul. The gene sp was strongly linked with lh and weakly linked with ul (Zijlstra and den Nijs, 1986). However Zijlstra and den Nijs (1986) expressed concern for the accuracy of the sp and ul linkage data since it was difficult to distinguish ul under their growing conditions.
Linkage Group F
Group F is comprised of 2 genes, Fl and Ccu which appear to be tightly associated. Wilson (1968) concluded that pleiotropy existed between scab resistance and fruit length because backcrossing scab resistance into commercial varieties consistently resulted in reduced fruit length. However, Munger and Wilkinson (1975) were able to break this linkage producing varieties with scab resistance and longer fruit (Tablegreen 65 and 66, Marketmore 70 and Poinsett 76). Now when these varieties are used to introduce scab resistance long fruit length is consistently associated.
Unaffiliated Genes
Independent assortment data are as important in developing linkage maps as direct linkage data and several researchers have made additional contributions in this area. One of the most extensive studies, based on the number of genes involved, is by Fanourakis (1984). He indicated that Ar was independent of df, F, ns, B, u, mc, pm, Tu, and D; dm was independent of bi, df, F, ns, ss, B, te, u, mc, Tu and D; bi was independent of cp, df, B, pm-h, te, u, mc and Tu; cp was independent of df, F, ns, ss, te, u, Tu, and D; F was independent of sf, B, pm-h, te, u, mc, Tu and D; df was independent of te, u, Tu, and D; ns was independent of B, pm-h and mc; ss was independent of B and mc; and B is independent of pm-h, te, u, Tu and D.
Two other extensive studies indicated that yc-2 was not linked with rc, yc-1, de, bi, cr, glb, gl, and m, (Whelan et al., 1975) and both Ccu and Bw were independent of bi, gl, glb, ls, rc, sc, cr, mc, gy-1 and gy-2 (Abul-Hayja et al., 1975). Meanwhile, white immature fruit color (w) was inherited independently of black spines (B), and locule number (l) (Cochran, 1938; Youngner, 1952).
Whelan (1973) found that light sensitive (ls) was not linked with nonbitter (bi?) but did not indicate which bitter gene he used. Zijlstra (1987) also determined that bi was independent of cp, gl is independent of lh and ccu is independent of lh, roand cp.
Powdery mildew has been the subject of several linkage studies. Robinson (1978e) indicated that resistance in ‘Ashley’ which contains 3 recessive factors was independent of B, l, pr, yg, fa, s, and H. Kooistra (1971) found that powdery mildew resistance was not linked with yf or wf and Barham (1953) determined that the resistance genes in USDA PI 173889 were independent of Bt.
Like linkage data, independent assortment data may be very valuable in developing gene maps, but care must be taken when utilizing them. For example, resistance to powdery mildew was demonstrated in the previous paragraph but none of the researchers were able to identify the particular gene involved.
Table 1. The qualitative genes of cucumber.
| Gene | Synonym | Character | Referencesz | Supplemental referencesz | Availabley | Photograph |
| a | – | androecious. Produces primarily staminate flowers if recessive for F. A from MSU 713-5 and Gy 14; a from An-11 and An-314, two selections from ‘E-e-szan’ of China. | Kubicki, 1969 | P | 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 | 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 | P | |
| al | – | albino cotyledons. White cotyledons and slightly light green hypocotyl; dying before first true leaf stage. Wild type Al from ‘Nishiki-suyo’; al from M2 line from pollen irradiation. | Iida 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 | 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 | P | |
| B-1 | B | Black or brown spine-1. Dominant to white spines on fruit. Completely linked or pleiotropic with heavy netting of fruit (H) and red mature fruit color (R). B from ‘Wis. SMR 18’; b from Gy 14 and most elite cultivars. | Strong, 1931; Tkachenko, 1935; Wellington, 1913 | Cochran, 1938; Fujieda and Akiya, 1962; Hutchins, 1940; Jenkins, 1946; Youngner, 1952 | W | W |
| B-2 | – | Black spine-2. Interacts with B to produce F2 of 15 black: 1 white spine. B-2 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 | W | |
| B-4 | – | Black spine-4. Interacts conversely with B-3. B-4 from LJ90430; b-4 from MSU 41. | Cowen and Helsel, 1983 | W | 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 DeBruyn, 1959 | Cantliffe, 1972; Da Costa and Jones, 1971a, 1971b; Soans et al., 1973 |
W | W |
| bi-2 | – | bitterfree-2. Leaves lacking cucurbitacins; bi-2 from NCG-093 (short petiole mutant). | Wehner et al., 1998a | W | W | |
| bl | t | blind. Terminal bud lacking after temperature shock. bl from ‘Hunderup’ and inbred HP3. | Carlsson, 1961. | L | L | |
| bla | – | blunt leaf. Leaves have obtuse apices and reduced lobing and serration. bla from a mutant of ‘Wis. SMR 18’. | Robinson, 1987a | W | W | |
| Bt | – | Bitter fruit. Fruit with extremely bitter flavor. Bt from PI 173889 (Wild Hanzil Medicinal Cucumber). | Barham, 1953 | W | W | |
| Bt-2 | – | Bitter fruit-2. Fruit with extremely bitter flavor. Bt-2 from LJ 90430. | Walters et al., 2001 | W | W | |
| Bu | – | bush. Shortened internodes. bu from ‘KapAhk 1’. | Pyzenkov and Kosareva, 1981 | L | L | |
| Bw | – | Bacterial wilt resistance. Resistance to Erwinia tracheiphila. Bw from PI 200818; bw from ‘Marketer’. | Nutall and Jasmin, 1958 | Robinson and Whitaker, 1974 | W | W |
| 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. |
Kubicki 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 | L | |
| Cca | – | Corynespora cassiicola resistance. Resistance to target leaf spot; dominant to susceptibility. Cca from Royal Sluis Hybrid 72502; ccafrom Gy 3. | Abul-Hayja et al., 1975 | W | 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; |
W | 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., 1966 | L | L | |
| Ch | Seedling chilling resistance. Ch from inbred NC-76. Originally from PI 246930 | Kozik and Wehner, 2006, 2008 | W | W | ||
| chp | – | choripetalous. Small first true leaf; choripetalous flowers; glossy 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 | 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 | W | |
| Cm | – | Corynespora melonis resistance. Resistance to C. melonis dominant to susceptibility. Cm from ‘Spotvrie’; cm from ‘Esvier’. | van Es, 1958 | ? | ? | |
| 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 from ‘National Pickling’ and ‘Wis. SR 6’. | Wasuwat and Walker, 1961 | Shifriss et al., 1942 | W | 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 | L |
| cor-1 | – | cordate leaves-1. Leaves are cordate. cor-1 from ‘Nezhinskii’. | Gornitskaya, 1967 | L | L | |
| cor-2 | cor | cordate leaves-2. Leaves are 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 | Ando et al., 2007 | W | 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 | ? | ? | |
| 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 MSU 3249 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 | 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, 1970**; Hutchins, 1940 | W | 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 ‘Borszczagowski’; 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. df from ‘Baroda’ (PI 212896)* and PI 215589 (hardwickii)**. | Della Vecchia et al., 1982*; Shifriss and George, 1965**. | W | W | |
| dl | – | delayed growth. Reduced growth rate; shortening of hypocotyl and first internodes. dl from ‘Dwarf Marketmore’ and ‘Dwarf Tablegreen’, both eriving dwarfness from ‘Hardin’s PG-57’. | Miller and George, 1979 | W | W | |
| dm-1 | P | downy mildew resistance. One of several genes for resistance to Pseudoperonospora cubensis. Dm-1 from Sluis & Groot Line 4285; dm-1 from ‘Poinsett’. | van Vliet and Meysing, 1974 | Jenkins, 1946; Shimizu, 1963 | W | W |
| dvl | dl | divided leaf. 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 | 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 PP-2-75; es-1from 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-2from Gy-30-75. | Kubicki and Korzeniewska, 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’. Plants are andromonoecious if (mm ff); monoecious if (MM ff); gynoecious if (MM FF) and hermaphroditic if (mm FF). | Galun, 1961; Tkachenko, 1935 | Kubicki, 1965, 1969a; Poole, 1944; Shifriss, 1961 |
W | 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 192940, 169383 and 169398; 2 alleles observed. | Meglic and Staub, 1996 | P | 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 | P | |
| Fl | – | Fruit length. Expressed in an additive fashion, fruit length decreases incrementally with each copy of fl (H. Munger, personal communication). | Wilson, 1968 | W | 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 | 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 | 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 | 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 | 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., 1984 | ? | ? | |
| gl | – | glabrous. Foliage lacking trichomes; fruit without spines. Iron-deficiency symptoms (chlorosis) induced by high temperature. gl from NCSU 75* and M834-6**. | Inggamer and de Ponti, 1980**;Robinson and Mishanec, 1964* | Robinson, 1987b | W | W |
| glb | – | glabrate. Stem and petioles glabrous, laminae slightly pubescent. glb from ‘Burpless Hybrid’. | Whelan, 1973 | W | 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. | Peterson and Pike, 1992 | W | 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 | 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 | P | |
| gy | – | gynoecious. Recessive gene for high degree of pistillate sex expression. | Kubicki, 1974 | W | W | |
| H | – | Heavy netting of fruit. Dominant to no netting; completely linked or pleiotropic with black spines (B) and red mature fruit color (R). | Hutchins, 1940; Tkachenko, 1935 | W | 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 | 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, 215589; 2 alleles observed. | Knerr and Staub, 1992 | P | 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. | George, 1970 | ? | ? | |
| 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 | 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 | ? | ? | |
| lg-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 | W | |
| ll | – | little leaf. Normal-sized fruits on plants with miniature leaves and smaller stems. ll from Ark. 79-75. | Goode et al., 1980; Wehner et al., 1987 | W | W | |
| ls | – | light sensitive. Pale and smaller cotyledons, lethal at high light intensity. ls from a mutant of ‘Burpless Hybrid’. | Whelan, 1972b | L | L | |
| m | a, g | andromonoecious. Plants are andromonoecious if (mm ff); monoecious if (MM ff); gynoecious if (MM FF) and hermaphroditic if (mm FF). m from ‘Lemon’*. | Rosa, 1928*; Tkachenko, 1935 | Shifriss, 1961; Wall, 1967; Youngner, 1952 | W | W |
| m-2 | h | andromonoecious-2. Bisexual flowers with normal ovaries. | Kubicki, 1974 | Iezzoni, 1982 | ? | ? |
| 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 | 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 | P | |
| Mdh-3 | – | Malate dehydrogenase-3 (E.C. # 1.1.1.37). | Knerr et al., 1995 | P | P | |
| Mdh-4 | Mdh-3 | Malate 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 | P | |
| mj | – | A single recessive gene for resistance to the root-knot nematode (Meloidogyne javanica) from Cucumis sativus var. hardwickii; mj from NC-42 (LJ 90430). | Walters et al., 1996; 1997 | Walters and Wehner, 1998 | W | W |
| mp | pf +,pf d, pf p | 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 | W |
| Mp-2 | – | Multi-pistillate-2. Several pistillate flowers per node. Single dominant gene with several minor modifiers. Mp-2 from MSU 3091-1. | Thaxton, 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 | 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 | 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 | L |
| ms-2 | – | male sterile-2. Male-sterile; pollen abortion occurs after first mitotic division of the pollen grain nucleus. ms-2 from a mutant of ‘Burpless Hybrid’. | Whelan, 1973 | ? | ? | |
| ms-2(PS) | – | male sterile-2 pollen sterile. Male-sterile; allelic to ms-2, but not to ap. ms-2(PS) from a mutant of Sunseeds 23B-X26. | Zhang et al., 1994 | ? | ? | |
| mwm | – | Moroccan watermelon mosaic virus resistance single recessive gene from Chinese cucumber cultivar ‘TMG-1’ | Kabelka and Grumet, 1997 | W | W | |
| n | – | negative geotropic peduncle response. Pistillate flowers grow upright; n from ‘Lemon’; N produces the pendant flower position of most cultivars. | Odland and Groff, 1963b | W | W | |
| ns | – | numerous spines. Few spines on the fruit is dominant to many. ns from Wis. 2757. | Fanourakis, 1984; Fanourakis and Simon, 1987 | W | 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 | 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 | 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 | 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 | 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 | 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 | 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 | 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 | 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 | 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 | 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 | 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 | L | |
| pm-1 | – | powdery mildew resistance-1. Resistance to Sphaerotheca fuliginia. pm-1 from ‘Natsufushinari’. | Fujieda and Akiya, 1962; Kooistra, 1971 | Shanmugasunda rum et al., 1972 | ? | ? |
| pm-2 | – | powdery mildew resistance-2. Resistance to Sphaerotheca fuliginia. pm-2 from ‘Natsufushinari’. | Fujieda and Akiya, 1962; Kooistra, 1971 | Shanmugasunda rum et al., 1972 | ? | ? |
| pm-3 | – | powdery mildew resistance-3. Resistance to Sphaerotheca fuliginia. pm-3 found in PI 200815 and PI 200818. | Kooistra, 1971 | Shanmugasunda rum et al., 1972 | W | 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 | W | |
| pr | – | protruding ovary. Exerted carpels. pr from ‘Lemon’. | Youngner, 1952. | W | W | |
| prsv | wmv-1-1 | watermelon mosaic virus 1 resistance. Resistance to papaya ringspot virus (formerly watermelon mosaic virus 1). Wild type Prsv from WI 2757; prsv from ‘Surinam’. | Wang et al., 1984 | ? | ? | |
| Prsv-2 | – | Resistance to papaya ringspot virus; Prsv-2 from TMG-1. | Wai and Grumet, 1995 | Wai et al., 1997 | W | W |
| psl | pl | Pseudomonas lachrymans resistance. Resistance to Pseudomonas lachrymans is recessive. Psl from ‘National Pickling’ and ‘Wis. SMR 18’; psl from MSU 9402 and Gy 14. | Dessert et al., 1982 | W | W | |
| Psm | – | Paternal sorting of mitochondria. Mitochondria sorting induced by dominant gene Psm, found in MSC 16; psm from PI 401734. | Havey et al., 2004. | W | W | |
| R | – | Red mature fruit. Interaction with c is evident in the F2 ratio of 9 red (R-C-) : 3 orange (R-cc) : 3 yellow (rrC-) : 1 cream (rrcc); completely linked or pleiotropic with black spines (B) and heavy netting of fruit (H). | Hutchins, 1940 | W | W | |
| rc | – | revolute cotyledon. Cotyledons are short, narrow, and cupped downwards; enlarged perianth. rc from ‘Burpless Hybrid’ mutant. | Whelan et al., 1975 | L | L | |
| rc-2 | – | recessive gene for revolute cotyledons; rc-2 from NCG-0093 (short petiole mutant) | Wehner et al., 1998b | W | W | |
| ro | – | rosette. 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 | 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 | W |
| s-2 | – | spine-2. Acts in duplicate recessive epistatic fashion with s-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 192940. | Jones, 1984 | P | 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 | 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 | 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, 1990, 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 | 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, 1985 | W | 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’; ssfrom Wis. 2757. | Fanourakis, 1984; Fanourakis and Simon, 1987 | W | 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 | 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 | W | |
| tf | twin fused fruit. Two fruit fused into single unit. Type line: B 5263 | Klosinska et al., 2006 | W | 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 | 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 | 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 | 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 | W | |
| v | – | virescent. Yellow leaves becoming green. | Poole, 1944; Tkachenko, 1935 | L | L | |
| vvi | – | variegated virescent. Yellow cotyledons, becoming green; variegated leaves. | Abul-Hayja and Williams, 1976 | L | L | |
| w | – | white immature fruit color. White is recessive to green. W from ‘Vaughan’, ‘Clark’s Special’, ‘Florida Pickle’ and ‘National Pickling’; wfrom ‘Bangalore’. | Cochran, 1938 | W | W | |
| wf | – | White flesh. Intense white flesh color is recessive to dingy white; acts with yf to produce F2 of 12 white (WfWf YfYf or wfwf YfYf) : 3 yellow (WfWf yfyf) : 1 orange (wfwf yfyf). Wf from EG and G6, each being dingy white (WfWf YfYf ): wf from ‘NPI ‘ which is orange (wfwf yfyf). | 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’; wifrom 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 | P | |
| wmv-1-1 | – | watermelon mosaic virus-1 resistance. Resistance to strain 1 of watermelon mosaic virus by limited systemic translocation; lower leaves may show severe symptoms. Wmv-1-1 from Wis. 2757; wmv-1-1 from ‘Surinam’. | Wang et al., 1984 | Provvidenti, 1985 | ? | ? |
| wmv-2 | – | watermelon mosaic virus resistance. Expressed in the cotyledon and throughout the plant; wmv-2 from TMG-1. | Wai et al., 1997 | W | W | |
| wmv-3 | – | watermelon mosaic virus resistance. Expressed only in true leaves; wmv-3 from TMG-1. | Wai et al., 1997 | W | W | |
| wmv-4 | – | watermelon mosaic virus resistance. Expressed only in true leaves; wmv-4 from TMG-1. | Wai et al., 1997 | W | W | |
| wy | – | wavy rimed cotyledons. Wavy rimed cotyledons, with white centers; true leaves normal. Wild type Wy from ‘Nishiki-suyo’; wy from M2line 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 | W | |
| yc-2 | – | yellow cotyledons-2. Virescent cotyledons. yc-2 from a mutant of ‘Burpless Hybrid’. | Whelan and Chubey, 1973; Whelan et al., 1975 | W | 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 | P | |
| yg | gr | yellow-green immature fruit color. Recessive to dark green and epistatic to light green. yg from ‘Lemon’. | Youngner, 1952 | W | 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 | ? | ? | |
| zym-Dina | – | zucchini yellow mosaic virus resistance; zym- Dina from Dina-1. | Kabelka et al., 1997 | Wai et al., 1997 | P | P |
| zym-TMG1 | zymv | zucchini yellow mosaic virus resistance. Inheritance is incomplete, but usually inherited in a recessive fashion; source of resistance is ‘TMG-1’. | Provvidenti, 1987; Kabelka et al., 1997 | Wai et al., 1997 | W | W |
zAsterisks on cultigens and associated references indicate the source of information for each. y W = Mutant available through T.C. Wehner, cucumber gene curator for the Cucurbit Genetics
Cooperative; P = mutants are available as standard cultivars or accessions from the Plant
Introduction Collection; ? = availability not known; L = mutant has been lost.
* Isozyme nomenclature follows a modified form of Staub et al. (1985) previously described by
Richmond (1972) and Gottlieb (1977).
Table 2. The cloned genes of cucumber and their function.z
| Gene accession | Tissue source | Function | Clone type | Reference |
|---|---|---|---|---|
| Genes involved in seed germination or seedling development | ||||
| X85013 | Cotyledon cDNA library | Encoding a T-complex protein | cDNA | Ahnert et al., 1996 |
| AJ13371 | Cotyledon cDNA library | Encoding a matrix metalloproteinases | cDNA | Delorme et al., 2000 |
| X15425 | Cotyledon cDNA library | Glyoxysomal enzyme malate synthase | Genomic DNA fragment | Graham et al., 1989; 1990 |
| X92890 | Cotyledon cDNA library | Encoding a lipid body lipoxygenase | cDNA | Höhne et al., 1996 |
| L31899 | Senescing cucumber cotyledon cDNA library | Encoding an ATP-dependent phosphoenolpyruvate carboxykinase (an enzyme of the gluconeogenic pathway) | cDNA | Kim and Smith, 1994a |
| L31900 | Cotyledon cDNA library | Encoding microbody NAD(+)- dependent malate dehydrogenase (MDH) | cDNA | Kim and Smith, 1994b |
| L44134 | Senescing cucumber cDNA library | Encoding a putative SPF1-type DNA binding protein | cDNA | Kim et al., 1997 |
| U25058 | Cotyledons | Encoding a lipoxygenase-1 enzyme | cDNA | Matsui et al., 1995; 1999 |
| Y12793 | Cotyledon cDNA library | Encoding a patatin like protein | cDNA | May et al., 1998 |
| X67696 | Cotyledon cDNA library | Encoding the 48539 Da precursor of thiolase | cDNA | Preisig-Muller and Kindl, 1993a |
| X67695 | Cotyledon cDNA library | Encoding homologous to the bacterial dnaJ protein | cDNA | Preisig-Muller and Kindl, 1993b |
| X79365 | Seedling cDNA library | Encoding glyoxysomal tetrafunctional protein | cDNA | Preisig-Muller et al., 1994 |
| X79366 | Seedling cDNA library | Encoding glyoxysomal tetrafunctional protein | cDNA | Preisig-Muller et al., 1994 |
| Z35499 | Genomic library | Encoding the glyoxylate cycle enzyme isocitrate lyase | Genomic gene | Reynolds and Smith, 1995 |
| M59858 | Cotyledon cDNA library | Encoding a stearoyl-acyl-carrier- protein (ACP) desaturase | cDNA | Shanklin and Somerville, 1991 |
| M16219 | Cotyledon cDNA library | Encoding glyoxysomal malate synthase | cDNA | Smith and Leaver, 1986 |
| Genes involved in photosynthesis and photorespiration activities | ||||
| M16056 | Cotyledon cDNA library | Encoding ribulose bisphosphate carboxylase/oxygenase | cDNA | Greenland et al., 1987 |
| M16057 | Cotyledon cDNA library | Encoding chlorophyll a/b-binding protein | cDNA | Greenland et al., 1987 |
| M16058 | Cotyledon cDNA library | Encoding chlorophyll a/b-binding protein | cDNA | Greenland et al., 1987 |
| X14609 | cotyledon cDNA library | Encoding a NADH-dependent hydroxypyruvate reductase (HPR) | cDNA | Greenler et al., 1989 |
| Y09444 | Chloroplast genomic library | tRNA gene | Chloroplast DNA fragment | Hande and Jayabaskaran, 1997 |
| X75799 | Chloroplast genomic library | Chloroplast tRNA (Leu) (cAA) gene | Genomic DNA fragment | Hande et al., 1996 |
| D50456 | Cotyledon cDNA library | Encoding 17.5-kDa polypeptide of cucumber photosystem I | cDNA | Iwasaki et al., 1995 |
| S69988 | Hypocotyls | Cytoplasmic tRNA (Phe) | cytoplasmic DNA fragment | Jayabaskaran and Puttaraju, 1993 |
| S78381 | Cotyledon cDNA library | Encoding NADPH- protochlorophyllide oxidoreductase | cDNA | Kuroda et al., 1995 |
| D26106 | Cotyledon cDNA library | Encoding ferrochelatase | cDNA | Miyamoto et al., 1994 |
| U65511 | Green peelings cDNA library | Encoding the 182 amino acid long precursor stellacyanin | cDNA | Nersissian et al., 1996 |
| AF099501 | Petal cDNA library | Encoding the carotenoid-associated protein | cDNA | Ovadis et al., 1998 |
| X67674 | Cotyledon cDNA library | Encoding ribulosebisphosphate carboxylase/oxygenase activase | cDNA | Preisig-Muller and Kindl, 1992 |
| X58542 | Cucumber genomic library | Encoding NADH-dependent hydroxypyruvate reductase | Genomic DNA fragment | Schwartz et al., 1991 |
| U62622 | Seedling cDNA library | Encoding monogalacto- syldiacylglycerol synthase | cDNA | Shimojima et al., 1997 |
| D50407 | Cotyledon cDNA library | Encoding glutamyl-tRNA reductase proteins | cDNA | Tanaka et al., 1996 |
| D67088 | Cotyledon cDNA library | Encoding glutamyl-tRNA reductase proteins | cDNA | Tanaka et al., 1996 |
| D83007 | Cotyledon cDNA library | Encoding a subunit XI (psi-L) of photosystem I | cDNA | Toyama et al., 1996 |
| Genes expressed mainly in roots | ||||
| AB025717 | Root RNA | Lectin-like xylem sap protein | cDNA | Masuda et al., 1999 |
| U36339 | Root cDNA library | Encoding root lipoxygenase | cDNA | Matsui et al., 1998 |
| AB015173 | Root cDNA library | Encoding glycine-rich protein-1 | cDNA | Sakuta et al., 1998 |
| AB015174 | Root cDNA library | Encoding glycine-rich protein-1 | cDNA | Sakuta et al., 1998 |
| Flower genes | ||||
| AF035438 | Female flower cDNA library | MADS box protein CUM1 | cDNA | Kater et al., 1998 |
| AF035439 | Female flower cDNA library | MADS box protein CUM10 | cDNA | Kater et al., 1998 |
| D89732 | Seedlings | Encoding 1-aminocyclo-propane- 1-carboxylate synthase | cDNA | Kamachi et al., 1997 |
| AB003683 | seedlings | Encoding 1-aminocyclo-propane- 1-carboxylate synthase | cDNA | Kamachi et al., 1997 |
| AB003684 | Seedlings | Encoding 1-aminocyclo-propane- 1-carboxylate synthase | cDNA | Kamachi et al., 1997 |
| AB035890 | Fruit RNA | Encoding polygalacturonase | cDNA | Kubo et al., 2000 |
| AF022377 | Floral buds | Encoding agamous-like putative transcription factor (CAG1) mRNA | cDNA | Perl-Treves et al., 1998 |
| AF022378 | Floral buds | Encoding agamous like putative transcription factor (CAG2) mRNA | cDNA | Perl-Treves et al., 1998 |
| AF022379 | Floral buds | Encoding agamous-like putative transcription factor (CAG3) mRNA | cDNA | Perl-Treves et al., 1998 |
| U59813 | Genomic DNA | Encoding 1-aminocyclo-propane- 1-carboxylate synthase | Genomic DNA fragment | Trebitsh et al., 1997 |
| X95593 | Corolla cDNA library | Encoding carotenoid-associated protein | cDNA | Vishnevetsky et al., 1996 |
| AB026498 | Shoot apex RNA | Ethylene-receptor-related gene | cDNA | Yamasaki et al., 2000 |
| Genes involved in fruit development and maturation | ||||
| AB010922 | Fruit cDNA library | Encoding the ACC synthase | cDNA | Mathooko et al., 1999 |
| J04494 | Fruit cDNA library | Encoding an ascorbate oxidase | cDNA | Ohkawa et al., 1989; 1990 |
| AB006803 | Fruit cDNA library | Encoding ACC synthase | cDNA | Shiomi et al., 1998 |
| AB006804 | Fruit cDNA library | Encoding ACC synthase | cDNA | Shiomi et al., 1998 |
| AB006805 | Fruit cDNA library | Encoding ACC synthase | cDNA | Shiomi et al., 1998 |
| AB006806 | Fruit cDNA library | Encoding ACC oxidase | cDNA | Shiomi et al., 1998 |
| AB006807 | Fruit cDNA library | Encoding ACC oxidase | cDNA | Shiomi et al., 1998 |
| AB008846 | Pollinated fruit cDNA library | Corresponding genes preferentially expressed in the pollinated fruit | cDNA | Suyama et al., 1999 |
| AB008847 | Pollinated fruit cDNA library | Corresponding genes preferentially expressed in the pollinated fruit | cDNA | Suyama et al., 1999 |
| AB008848 | Pollinated fruit cDNA library | Corresponding genes preferentially expressed in the pollinated fruit | cDNA | Suyama et al., 1999 |
| Genes involved in cell wall loosening and cell enlargement | ||||
| AB001586 | Hypocotyl RNA | Encoding homologous to serine/threonine protein kinases (for CsPK1.1) | cDNA | Chono et al., 1999 |
| AB001587 | Hypocotyl RNA | Encoding homologous to serine/threonine protein kinases (for CsPK1.2) | cDNA | Chono et al., 1999 |
| AB001588 | Hypocotyl RNA | Encoding homologous to serine/threonine protein kinases (for CsPK2.1) | cDNA | Chono et al., 1999 |
| AB001589 | Hypocotyl RNA | Encoding homologous to serine/threonine protein kinases (for CsPK2.2) | cDNA | Chono et al., 1999 |
| AB001590 | Hypocotyl RNA | Encoding homologous to serine/threonine protein kinases (for CsPK3) | cDNA | Chono et al., 1999 |
| AB001591 | Hypocotyl RNA | Encoding homologous to serine/threonine protein kinases (for CsPK4.1) | cDNA | Chono et al., 1999 |
| AB001592 | Hypocotyl RNA | Encoding homologous to serine/threonine protein kinases (for CsPK4.2) | cDNA | Chono et al., 1999 |
| AB001593 | Hypocotyl RNA | Encoding homologous to serine/threonine protein kinases (for CsPK5) | cDNA | Chono et al., 1999 |
| U30382 | Hypocotyl cDNA library | Encoding expansins | cDNA | Shcherban et al., 1995 |
| U30460 | Hypocotyl cDNA library | Encoding expansins | cDNA | Shcherban et al., 1995 |
| Genes induced or repressed by plant hormones | ||||
| D49413 | Hypocotyl cDNA library | Corresponding to a gibberellin- responsive gene encoding an extremely hydrophobic protein | cDNA | Chono et al., 1996 |
| AB026821 | Seedling RNA | Encoding IAA induced nuclear proteins | cDNA | Fujii et al., 2000 |
| AB026822 | Seedling RNA | Encoding IAA induced nuclear proteins | cDNA | Fujii et al., 2000 |
| AB026823 | Seedling RNA | Encoding IAA induced nuclear proteins | cDNA | Fujii et al., 2000 |
| M32742 | Cotyledon cDNA library | Encoding ethylene-induced putative peroxidases | cDNA | Morgens et al., 1990 |
| D29684 | Cotyledon cDNA library | Cytokinin-repressed gene | cDNA | Teramoto et al., 1994 |
| D79217 | Genomic library | Cytokinin-repressed gene | Genomic DNA fragment | Teramoto et al., 1996 |
| D63451 | Cotyledon cDNA library | Homologous to Arabidopsis cDNA clone 3003 | cDNA | Toyama et al., 1995 |
| D63384 | Cotyledon cDNA library | Encoding catalase | cDNA | Toyama et al., 1995 |
| D63385 | Cotyledon cDNA library | Encoding catalase | cDNA | Toyama et al., 1995 |
| D63386 | Cotyledon cDNA library | Encoding catalase | cDNA | Toyama et al., 1995 |
| D63387 | Cotyledon cDNA library | Encoding lectin | cDNA | Toyama et al., 1995 |
| D63388 | Cotyledon cDNA library | Encoding 3-hydroxy-3- methylglutaryl CoA reductase | cDNA | Toyama et al., 1995 |
| D63389 | Cotyledon cDNA library | Encoding 3-hydroxy-3- methylglutaryl CoA reductase | cDNA | Toyama et al., 1995 |
| D63388 | Cotyledon cDNA library | Encoding a basic region/helix- loop-helix protein | cDNA | Toyama et al., 1999 |
| Resistance genes | ||||
| M84214 | Genomic library | Encoding the acidic class III chitinase | cDNA | Lawton et al., 1994 |
| M24365 | Leave cDNA library | Encoding a chitinase | cDNA | Metraux et al., 1989 |
| D26392 | Seedling cDNA library | Encoding FAD-Enzyme monodehydroascorbate (MDA) reductase | cDNA | Sano and Asada, 1994 |
| Somatic embryo gene | ||||
| X97801 | Embryogenic callus cDNA library | MADS-box gene | cDNA | Filipecki et al., 1997 |
| Repeated DNA sequences | ||||
| X03768 | Genomic DNA | Satellite type I | Genomic DNA fragment | Ganal et al., 1986 |
| X03769 | Genomic DNA | Satellite type II | Genomic DNA fragment | Ganal et al., 1986 |
| X03770 | Genomic DNA | Satellite type III | Genomic DNA fragment | Ganal et al., 1986 |
| X69163 | Genomic DNA | Satellite type IV | Genomic DNA fragment | Ganal et al., 1988a |
| X07991 | rDNA | Ribosomal DNA intergenic spacer | Genomic DNA fragment | Ganal et al., 1988b |
| X51542 | Cotyledons | Ribosomal DNA intergenic spacer | Genomic DNA fragment | Zentgraf et al., 1990 |
zTable listing includes only the sequences published in journals as well as the genebank database through the year 2001.
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