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Cucurbit Genetics Cooperative Report 28-29:105-141 (article 26) 2005-2006

Cucurbit Genetics Cooperative

Gene List 2005 for Cucumber

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. 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

Revisions to the 2005 cucumber gene list include the addition of Psm for paternal sorting of mitochondria (Havey et al., 2004). Researchers are encouraged to send reports of new genes, as well as seed samples to the cucumber gene curator (Nischit V. Shetty), or to the assistant curator (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.

Gene Mutants

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).

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-2 for 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 s proposed 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).

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.

Currently, one gene, dm, 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). There are, however, indications that more than one gene may be involved (Jenkins, 1946).

Environmental Stress Resistance

Presently, only 2 genes have been identified in this category; resistance to sulfur dioxide air pollution conditioned by Sd (Bressan et al., 1981) and increased tolerance to high salt levels conditioned by major 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.

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, m2, 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, B-2, B-3, and B-4), the fruit skin color mutants (c, R, and w), the spine size and density mutants (s, s2, 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 12 genes, composed of wmv-1-1, 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).

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 I, 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 II 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 F from 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 III 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 IV, 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 V 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 VI 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, ro and 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

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

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 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

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-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

B-4

-

Black spine-4. Interacts conversely with B-3. B-4 from LJ90430; b-4 from MSU 41.

Cowen 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 DeBruyn, 1959

Cantliffe, 1972; Da Costa and
Jones, 1971a,
1971b; Soans et
al., 1973

W

bi-2

-

bitterfree-2. Leaves lacking cucurbitacins; bi-2 from NCG-093 (short petiole mutant).

Wehner et al., 1998a

 

W

bl

t

blind. Terminal bud lacking after temperature shock. bl from 'Hunderup' and inbred HP3.

Carlsson, 1961.

 

L

bla

-

blunt leaf. Leaves have obtuse apices 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 PI 200818; bw from 'Marketer'.

Nutall and Jasmin, 1958

Robinson and
Whitaker, 1974

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

Cca

-

Corynespora cassiicola 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;

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

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

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

-

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

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 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

 

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

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

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

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

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 18; 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 Pseudoperonospora 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. 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 PP-2-75; 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 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'.

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 192940, 169383 and 169398; 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 an 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., 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

glb

-

glabrate. Stem and petioles glabrous, laminae slightly pubescent. glb from 'Burpless Hybrid'.

Whelan, 1973

 

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

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; Tkachenko, 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, 215589; 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.

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

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

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

ls

-

light sensitive. Pale and smaller cotyledons, lethal at high light intensity. ls from a mutant 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.

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

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

-

Malate dehydrogenase-3 (E.C. # 1.1.1.37).

Knerr et al., 1995

 

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

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

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

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

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 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

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

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 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

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. Exerted 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 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

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

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

R

-

Red mature fruit. Interacts with c; linked or pleiotropic 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

rc-2

-

recessive gene for revolute cotyledons; rc-2 from NCG-0093 (short petiole mutant)

Wehner et al., 1998b

 

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

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 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

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, 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

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

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.

Poole, 1944; Tkachenko, 1935

 

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 'Vaughan', '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 (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'; 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 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

wmv-3

-

watermelon mosaic virus resistance. Expressed only in true leaves; wmv-3 from TMG-1.

Wai et al., 1997

 

W

wmv-4

-

watermelon mosaic virus resistance. Expressed only in true leaves; wmv-4 from TMG-1.

Wai et al., 1997

 

W

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. 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

 

?

zym-Dina

-

zucchini yellow mosaic virus resistance; zym- Dina from Dina-1.

Kabelka et al., 1997

Wai et al., 1997

P

zym-TMGI

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

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

zOnly the sequences published in both journals and the genebank database are listed.

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