New Cucumis Rootstocks for Melon: ‘UPV-FA’ and ‘UPV-FMy’

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  • 1 Instituto de Conservación y Mejora de la Agrodiversidad Valenciana (COMAV), Universitat Politècnica de València (UPV), Camino de Vera 14, 46022 Valencia, Spain
  • 2 Instituto Agroforestal Mediterráneo (IAM-Universitat Politècnica de València)
  • 3 Instituto de Conservación y Mejora de la Agrodiversidad Valenciana (COMAV), Universitat Politècnica de València (UPV), Camino de Vera 14, 46022 Valencia, Spain

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Grafting plants onto resistant rootstocks is a cultural practice spread worldwide to cope with biotic and abiotic stresses, such as fungal diseases, nematodes, drought, salinity, and extreme temperatures. For Cucurbitaceae crops, the most common rootstocks are interspecific hybrids between Cucurbita maxima Duchesne and Cucurbita moschata Duchesne (Davis et al., 2008). Besides their tolerance to biotic and abiotic stresses, hybrid Cucurbita rootstocks are preferred because they show good emergence performance and develop long and thick hypocotyls that facilitate grafting. These hybrids have however some important shortcomings. Their excess of vigor can cause a delay in the flowering and ripening processes in grafted plants (Soteriou et al., 2016), they are not resistant to nematodes (Cohen et al., 2014; Özarslandan et al., 2011), and often they have negative impacts on fruit quality (Guan et al., 2015; Rouphael et al., 2010; Soteriou et al., 2014). These effects on quality are dependent on the rootstock–scion interactions. In melon (Cucumis melo L.), for example, an increase of fruit size and seed cavity (Verzera et al., 2014) or modifications of flesh firmness (Colla et al., 2006; Zhao et al., 2011) as a result of grafting are common. Flesh color variations (Colla et al., 2006), vitrescence (Jang et al., 2014), and changes in pH and soluble solids content (Colla et al., 2006; Verzera et al., 2014) have also been reported.

In part because of these quality problems, grafting is less common in melon than in watermelon [Citrullus lanatus (Thunb.) Mansf.] and cucumber (Cucumis sativus L.). However, in recent years, the withdrawal of methyl bromide (a broad spectrum pesticide highly efficient against soilborne pathogens), and the increment of global temperature, have been favoring the spread of nematodes and fungi highly damaging for melons, such as Fusarium spp., Monosporascus spp., Macrophomina phaseolina (Tassi) Goidanich and Podosphaera xanthii (Castagne) Braun & Shishkoff. The global warming has also a negative impact on soil salinization and drought. Consequently, the use of grafted melon plants has gained increased attention. The use of rootstocks belonging to the same species/genus as the scions could minimize the quality problems observed when using Cucurbita hybrids as rootstocks. Even though some promising melon accessions have been selected and used to develop Fusarium oxysporum Schlechtend. Fr. f. sp. melonis (Leach & Currence) Snyder & Hans spp. and Monosporascus cannonballus Pollack & Uecker resistant rootstocks (Condurso et al., 2012; Fredes et al., 2016; Verzera et al., 2014), the assessment of a new germplasm that may provide a wider range of resistances without decreasing the fruit quality is a new challenge for melon grafting.

Apart from intraspecific variation, intrageneric variation can also be exploited in melon grafting. The genus Cucumis includes a large number of African, Asian, and Australian wild species (Den Nijs and Visser, 1985; Renner et al., 2007; Sebastian et al., 2010; Singh and Yadava, 1984). Strong reproductive barriers prevent the use of these resources for melon breeding (Chen and Adelberg, 2000), but the genus is potentially a good source of new rootstocks. Resistance to nematodes and to soilborne and aerial fungi has been reported in some Cucumis species. To date, two species have been assayed as rootstocks: Cucumis metuliferus Naudin (Gisbert et al., 2017; Guan et al., 2014; Kokalis-Burelle and Rosskopf, 2011; Nisini et al., 2002; Sigüenza et al., 2005) and Cucumis pustulatus Naudin ex Hook.f. (Liu et al., 2015). Despite their effectiveness in the management of soil pathogens, lower soluble solids content and softer flesh have been observed in fruits from plants grafted onto C. metuliferus (Guan et al., 2014; Nisini et al., 2002). Other Cucumis species useful as sources of resistance to nematodes, Fusarium spp., or both are C. ficifolius A.Rich., C. zeyheri Sond., C. africanus L.f., C. anguria L., and C. myriocarpus Naudin (Den Nijs and Custers, 1990; Nisini et al., 2002). One of the main difficulties when using these wild species is the reduced size and vigor of the seedlings. Differences in hypocotyl diameters between the rootstock and the scion at the grafting union, either at the time of grafting or later during the development of the grafted plants (Pofu and Mashela, 2011; Pofu et al., 2013), can result in physiological collapse of melon plants under field conditions. Vigor of wild rootstocks can be increased by using interspecific hybrids, which have the additional advantage of combining resistances or other suitable traits from both parental lines. Although hybridization between different Cucumis species is extremely difficult, some successful crosses are possible (Kho et al., 1980; Matsumoto et al., 2012; Walters and Wehner, 2002), especially between some wild related African species, which are phylogenetically relatively close (Den Nijs and Visser, 1985; Sebastian et al., 2010).

The research groups of “Cucurbits” and “Grafting Breeding” at COMAV-UPV (Centro de Conservación y Mejora de la Agrodiversidad Valenciana-Universitat Politècnica de València) are working together in the development and characterization of new experimental rootstocks for cucurbit crops (Fredes et al., 2017; Gisbert et al., 2016). Germplasm characterization and crossability studies have been performed within the Cucumis genus to generate vigorous rootstocks suitable for melon. Two new interspecific Cucumis hybrids with good characteristics to be used as melon rootstocks have been developed: ‘UPV-FA’ (C. ficifolius × C. anguria) and ‘UPV-FMy’ (C. ficifolius × C. myriocarpus). Both hybrids have good compatibility with cultivars of the two main melon market classes (muskmelon and Piel de Sapo), are resistant to some of the main fungi that are pathogenic to melon, and do not negatively modify melon fruit quality.

Origin

The two new interspecific hybrids ‘UPV-FA’ and ‘UPV-FMy’ were obtained by crossing three wild African species: Cucumis ficifolius (F) (accession BGV012786) as female parent and the two other wild species as male parents, Cucumis anguria var. longipes (A) (accession BGV12795) and Cucumis myriocarpus (My) (accession BGV008535). The three accessions were held at the COMAV’s Genebank. Hybridizations were conducted in a greenhouse at the UPV, under controlled conditions during the spring–summer season of 2015. Direct and reciprocal crosses were performed in all cross combinations, but fruit set was only obtained using C. ficifolius as female parent. Results are in accordance with previous studies, reporting that these three species are in the same compatibility group (Singh and Yadava, 1984), and with the occurrence of some form of unilateral incongruity between C. ficifolius and other African species such as C. anguria, C. myriocarpus, C. zeyheri, C. dipsaceus Ehrenb. ex Spach., and C. figarei A.Rich. (Den Nijs and Visser, 1985). The fruit set percentage was around 70% in both hybrids, quite similar to the value of 80% found in the self-pollinations of the parentals. Also, the average number (± se) of viable seeds per fruit found in the hybrid fruits (227.3 ± 9.9 and 132.7 ± 17.1 for ‘UPV-FMy’ and ‘UPV-FA’, respectively) was similar or even higher than the number found in fruits derived from selfings (164.3 ± 32.8, 64.3 ± 14.5, and 129.3 ± 5.2 for F, My, and A, respectively). The germination capacity of the hybrid seeds was tested under different experimental conditions, reaching 100% in treatments with cold stratification (Cáceres et al., 2016a). The hybrid seeds germinated earlier in comparison with their respective parentals (Cáceres et al., 2016a). Hypocotyl diameters were measured in the germinated seedlings at 21 d after sowing. The diameters of ‘UPV-FMy’ (0.28 ± 0.0062 cm) and ‘UPV-FA’ (0.25 ± 0.0075 cm) were higher than those of the wild parents (0.22 ± 0.0068, 0.24 ± 0.0033, 0.24 ± 0.0061 cm for F, My, and A, respectively). Seedlings were transplanted to the greenhouse, and the hybrid nature was confirmed using discriminant taxonomic traits between the parentals, such as the presence of aculei in stems and petioles, the ratio between the hyaline and opaque parts of aculei of ovaries, and the shape of ovaries and fruits (Kirkbride, 1993). Intermediate phenotypes between those of the parentals were observed for these characters, which support their hybrid nature (Fig. 1; Cáceres et al., 2016b).

Fig. 1.
Fig. 1.

Discriminant morphological characters among Cucumis ficifolius (F), C. anguria (A), C. myriocarpus (My), and their hybrids ‘UPV-FMy’ and ‘UPV-FA’. From top to bottom: leaves, stems, aculei from ovaries with an inferior opaque part and a superior hyaline part, and fruits. The morphology was evaluated on greenhouse growing plants.

Citation: HortScience horts 52, 5; 10.21273/HORTSCI11791-17

The good cross compatibility of these two interspecific crosses, along with the higher germination rates and wider hypocotyls found in the hybrids compared with their parents, confirms the advantage of using interspecific hybrids instead of single wild Cucumis genotypes for grafting purposes.

Responses to Biotic and Abiotic Stress

Wild Cucumis species have been reported to possess resistance to several fungal diseases (Matsumoto et al., 2011; Thomas and More, 1990). Among these diseases, fusarium wilt, caused by F. oxysporum f. sp. melonis, is one of the most severe for melons worldwide. Both hybrids and their respective parents were confirmed as resistant to fusarium wilt by artificial inoculation of F. oxysporum f. sp. melonis race 1.2 that was performed in a growth chamber following the protocol described by Perchepied and Pitrat (2004). At 20 d after inoculation, plants of ‘UPV-FA’ and ‘UPV-FMy’ showed no symptoms of infection, whereas wilting of leaves and browning of the stem were observed in 50% of melon muskmelon plants (C. melo var. cantalupensis Naudin), used as susceptible control. All muskmelon plants finally died (Fig. 2A–C), whereas ‘UPV-FA’ and ‘UPV-FMy’ plants continued to grow, remaining symptomless at the end of the assay (40 d). These results are in agreement with those of previous reports that describe resistance to several races of F. oxysporum in C. ficifolius, C. myriocarpus, and C. anguria (Liu et al., 2015; Matsumoto et al., 2011; Nisini et al., 2002).

Fig. 2.
Fig. 2.

Response of the two hybrids, ‘UPV-FA’ (C. ficifolius × C. anguria) and ‘UPV-FMy’ (C. ficifolius × C. myriocarpus), compared with a melon susceptible control against Fusarium oxysporum f sp. melonis: plants of cantaloupe (A), ‘UPV-FA’ (B), and “UPV-FMy” (C); against Monosporascus cannoballus: roots extracted from naturally infested soil of cv. Piñonet Piel de Sapo (D), ‘UPV-FA’ (E), and ‘UPV-FMy’ (F); against powdery mildew: plants cultivated in an open field of cv. Piñonet Piel de Sapo (G), ‘UPV-FA’ (H), and ‘UPV-FMy’ (I).

Citation: HortScience horts 52, 5; 10.21273/HORTSCI11791-17

Other soilborne pathogens with increasing economic importance for melon cultivation are nematodes (mainly Meloidogyne ssp.) and the fungus Monosporascus cannonballus, responsible for the melon vine decline disease (Pivonia et al., 2010). Nematode resistance has been described in some C. anguria accessions (Den Nijs and Custers, 1990), and nematicide activity was also reported in C. myriocarpus fruit extracts (Mashela et al., 2008). Nematode resistance in ‘UPV-FMy’ and ‘UPV-FA’ is under evaluation. Both hybrids were included in a screening assay for evaluating their response to M. cannoballus in a naturally infested field. In comparison with the Piel de Sapo Piñonet cultivar, used as susceptible control, both hybrids showed different levels of tolerance, having moderate to mild root damage and no wilted plants. Roots of each genotype extracted from the infested field are shown in Fig. 2D–F. ‘UPV-FMy’ was moderately resistant (with a root damage score of 3 ± 0.50, in a scale of 0 = no symptoms to 5 = roots severely affected by the fungus), whereas ‘UPV-FA’ was highly resistant (root score 1 ± 0). Roots of both hybrids had also a more vigorous and branched root structure than the Piñonet cultivar. In this field assay, the response to powdery mildew was also observed in both, rootstocks and scion plants. The Piñonet cultivar was highly susceptible (symptom score of 4 ± 0.23), whereas ‘UPV-FMy’ and ‘UPV-FA’ were highly (0.2 ± 0.31) and moderately (1.02 ± 0.31) resistant (Fig. 2E–G).

The two new putative rootstocks were also evaluated against osmotic stress using clones from plants maintained in vitro. These clones were grown on basal medium (4.4 mg·L−1 Murashige and Skoog salts, including vitamins, 2% sucrose, and 0.75% plant agar), containing sorbitol at four different concentrations: 0, 0.05, 0.1, and 0.2 M. The addition of compounds such as sorbitol to culture medium reduces the water potential, so it is harder for plants to uptake water, simulating water-deficient conditions in soil (Verslues et al., 2006). The Piel de Sapo Piñonet cultivar showed great growth inhibition in a previous assay in medium with sorbitol after 45 d of culture (the fresh weight of plants was reduced 49% and 92% in media with sorbitol at 0.05 and 0.1 M, respectively). Cucumis metuliferus, which is another wild species used as putative rootstock for melon (Gisbert et al., 2017; Guan et al., 2014; Kokalis-Burelle and Rosskopf, 2011; Nisini et al., 2002; Sigüenza et al., 2005), was also included in the assay conducted to evaluate ‘UPV-FMy’ and ‘UPV-FA’. Differences in vine growth and rooting inhibition were observed after 30 and 45 d (Table 1). After these periods, plant fresh and dry weights were noted. Root characteristics (length, number of tips, and root diameter) were measured using the WinRhizo’s software. At the lowest sorbitol concentration (0.05 M), vine growth and root development were inhibited in C. metuliferus after 30 d of culture, whereas both hybrids ‘UPV-FMy’ and ‘UPV-FA’ only showed a mild vine growth inhibition. This behavior was maintained after 45 d of culture, even at the intermediate sorbitol concentration (0.1 M) (Table 1). Rooting was not observed in any genotype in culture medium with 0.2 M sorbitol. This result was expected as no growth was also observed in other works using this very high concentration (Claeys et al., 2014). Of the two hybrids, ‘UPV-FA’ was more tolerant than ‘UPV-FMy’ (Table 1), with sorbitol causing less root length and branching reduction. A good root development is an important trait for putative rootstocks.

Table 1.

Plant fresh and dry weights and root parameters of ‘UPV-FA’ (C. ficifolius × C. anguria), ‘UPV-My’ (C. ficifolius × C. myriocarpus), and C. metuliferus after 45 d of culture on BM medium with sorbitol (0, 0.05, 0.1, and 0.2 M).

Table 1.

Performance as Rootstocks

The performance of ‘UPV-FA’ and ‘UPV-FMy’ as rootstocks was evaluated with two different melon scions. A first evaluation was performed in a greenhouse under hydroponic conditions, using a muskmelon Charentais as scion. Plants were grafted using the cleft procedure (Lee et al., 2010), and high rates of success were obtained (>90%). The responses of muskmelon plants grafted onto ‘UPV-FA’ and ‘UPV-FMy’ were compared with cantaloupe nongrafted (NG) plants. Similar vine growth was observed in grafted and NG plants. Grafting onto ‘UPV-FA’ and ‘UPV-FMy’ did not delay female flowering or fruit ripening and did not alter the number of flowers and fruit set. On average, the first female flower opened from 35.3 to 37.3 d after transplanting (DAT), and fruits needed from 40.0 to 41.3 d after pollination to reach full maturity. To evaluate the impact of grafting on fruit quality, fruits were characterized for the following traits: weight (grams), length and width (centimeters), shape (length/width), cavity width (centimeters), rind and flesh firmness (kg·cm−2) (measured using a digital Penetrometer (8 mm) FHT-803; Melrose, MA), total soluble solids (quantified through a digital Rephractometer Atago; Tokyo, Japan), pH (measured using pH-indicator paper pH 1–14 Merck, Darmstadt, Germany), and flesh color (measured with a colorimeter, Minolta CR-400; NJ, using the color parameters Hunter L, a, and b). No significant differences were found in any of these traits between fruits from plants grafted onto ‘UPV-FA’ and fruits from NG plants (Table 2). The fruits from plants grafted onto ‘UPV-FMy’ only differed from those of NG plants in the fruit shape, being slightly more flattened, in the sugars content (°Brix 13.3 vs. 12.3), and in the flesh color (significantly higher value of the parameter a that measures color from red to green) (Table 2; Fig. 3). Therefore, quality parameters were not modified or only slightly positively modified by these rootstocks, which did not cause negative impact on commercial traits.

Table 2.

Characteristics of muskmelon fruits harvested from nongrafted plants (NG) and plants grafted onto the experimental hybrids ‘UPV-FMy’ (C. ficifolius × C. myriocarpus) and ‘UPV-FA’ (C. ficifolius × C. anguria).

Table 2.
Fig. 3.
Fig. 3.

Fruit of muskmelons cv. Vedrantais (Charentais type) from ungrafted plants (A) and plants grafted onto ‘UPV-FMy’ (B) and ‘UPV-FA’ (C).

Citation: HortScience horts 52, 5; 10.21273/HORTSCI11791-17

A second evaluation of the experimental hybrids ‘UPV-FA’ and ‘UPV-FMy’ was performed under field conditions using the commercial melon ‘Finura’ (Rijk ZWaan) (C. melo var. inodorus H.Jacq. market class ‘Piel de Sapo’) as scion. In this assay, both experimental rootstocks were also compared with a commercial rootstock of melon (64-376RZ, Rijk Zwaan) and one C. maxima × C. moschata rootstock (the commercial hybrid Cobalt, Rijk Zwaan). In addition, NG and self-grafted (SG) plants of ‘Finura’ were used as controls. All grafted plants were produced by a commercial nursery “Viveros la Sala” (San Pedro del Pinatar, Spain) and were transplanted to an experimental field located in Picassent (Valencia, Spain). Plant vigor was evaluated during the growing period (at 10, 30, and 60 DAT) in a scale of 0 (very low), 1 (low), 2 (intermediate), 3 (high), and 4 (very high), and the appearance of the first female flowers was recorded. Both commercial rootstocks, ‘Cobalt’ and ‘64-376RZ’, conferred high early vigor, similar to the SG melons (average scores, ± se, at 10 DAT from 3.25 ± 0.32 to 3.75 ± 0.14), which was associated to an earlier appearance of the first female flower, whereas the two interspecific hybrids resulted in less vigorous plants, similar to the NG melons (scores from 2.63 ± 0.24 to 2.75 ± 0.14). However, no significant differences in plant vigor were found among rootstocks at 30 and 60 DAT (scores ranging from 3.25 ± 0.25 to 4 ± 0 in plants grafted onto the different rootstocks). In fact, at 60 DAT, when most fruits were considered to have reached the commercial ripening stage, the average number of fruits and marketable yield per plant were similar in all the combinations (3.04 fruit/plant and 6.74 kg/plant in average).

Similar to the previous assay with muskmelons, fruit quality traits (including rind and flesh thickness) were measured in 16 representative fruits per combination (4 of each plot). External quality traits, such as fruit shape and rind thickness, were similar in fruits of cv. Finura harvested in plants grafted onto the two interspecific Cucumis hybrids (‘UPV-FMy’ and ‘UPV-FA’) and onto the commercial melon rootstock ‘64-376RZ’ (Table 3). No differences in fruit size and shape were found among melons developed on these Cucumis rootstocks and those developed on NG and SG plants, whereas the Cucurbita hybrid ‘Cobalt’ resulted in significantly larger fruits (average, ± se, fruit weight 2960 ± 168.15 g vs. 2502 ± 60.94 g and 2578 ± 100.30 g in NG and SG melons, respectively) with a wider seed cavity (7.22 ± 2.25 cm vs. 5.76 ± 1.71 and 5.77 ± 2.67 cm) (Table 3). These effects are commonly observed when Cucurbita hybrids are used for grafting melon or watermelon (Colla et al., 2006; Verzera et al., 2014). The ‘UPV-FA’ rootstocks also caused a slight increase of fruit seed cavity (6.47 ± 2.35 cm). Figure 4 shows fruits from the different rootstocks in which the higher fruit size and seed cavity can be appreciated.

Table 3.

Marketable yield and quality traits of melon fruits cv. Finura (Piel de Sapo) harvested from nongrafted plants (NG), self-grafted plants (SG), and plants grafted onto the experimental Cucurbita (C. maxima × C. moschata) hybrid F1, the commercial Cucurbita hybrid ‘Cobalt’, the commercial Cucumis melo rootstock ‘64-376RZ’, and two new experimental Cucumis F1: ‘UPV-FMy’ (C. ficifolius × C. myriocarpus) and ‘UPV-FA’ (C. ficifolius × C. anguria).

Table 3.
Fig. 4.
Fig. 4.

Fruit of Piel de Sapo melons cv. Finura from ungrafted plants (A) and plants grafted onto C. maxima × C. moschata hybrid ‘Cobalt’ (B), the Cucumis melo ‘64-376RZ’ (C), “UPV-FMy” (D), and “UPV-FA” (E).

Citation: HortScience horts 52, 5; 10.21273/HORTSCI11791-17

Regarding flesh properties, we found significantly firmer flesh in fruits from ‘UPV-FA’ (3.26 ± 0.19 vs. 2.15 ± 0.16 and 2.41 ± 0.26 in NG and SG plants, respectively) and flesh with significantly higher Brix degree in those fruits from the Cucurbita rootstocks (13.59 ± 0.28 vs. 11.93 ± 0.45 and 12.13 ± 0.39 °Brix) (Table 3). These two effects do not negatively affect fruit quality. An effect on flesh color was also observed in fruits derived from the commercial Cucurbita and melon rootstocks. The former showed a significantly higher value of the a parameter (−1.87 ± 0.26), which means less greenish flesh, and the latter caused a significant increase of the b parameter value (11.56 ± 0.63), which means a more yellowish flesh. The wild Cucumis hybrids did not alter flesh color in comparison with fruits of NG and SG plants (−2.57 ± 0.19 to −2.67 ± 0.12 and 9.24 ± 0.32 to 9.75 ± 0.22 for the a and b parameter values, respectively, in all combinations).

Use

The results obtained in the characterization of ‘UPV-FA’ and ‘UPV-FMy’ indicated that both hybrids can be used as melon rootstocks. The good cross compatibility between the parents facilitates hybrid seed generation, and the high germination rates along with the wider hypocotyls will be essential for grafting management. Their resistance/tolerance to F. oxysporum and M. cannonballus and their good agronomical performance with two commonly used melon scions make both hybrids comparable with other rootstocks used for grafting melon. In comparison with Cucurbita commercial rootstocks, ‘UPV-FA’ and ‘UPV-FMy’ have less early vigor and do not anticipate female flowering. However, no differences in vigor and fruit set are found at the end of the growing cycle. These Cucumis hybrids do not cause fruit size increase and only ‘UPV-FA’ slightly increased cavity thickness, both effects commonly observed with Cucurbita hybrids rootstocks. An additional value of these rootstocks is their good tolerance to osmotic stress, especially in ‘UPV-FA’. Also, their tolerance to powdery mildew can facilitate their management during rootstock seed production, and as it has been reported in other crops (Guan et al., 2012), it can contribute to improve the scion response to this disease.

Availability

Small trial seed samples of all the breeding lines are available for research purposes (please contact authors).

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  • Liu, B., Ren, J., Zhang, Y., An, J., Chen, M., Chen, H., Xu, Ch. & Ren, H. 2015 A new grafted rootstock against root-knot nematode for cucumber, melon, and watermelon Agron. Sustain. Dev. 35 1 251 259

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    • Export Citation
  • Mashela, P.W., Shimelis, H.A. & Mudau, F.N. 2008 Comparison of the efficacy of ground wild cucumber fruits, aldicarb and fenamiphos on doses against root-knot nematode, Meloidogyne incognita on tomato Ind. J. Nematol. 35 38 41

    • Search Google Scholar
    • Export Citation
  • Matsumoto, Y., Miyagi, M., Watanabe, N. & Kuboyama, T. 2012 Temperature-dependent enhancement of pollen tube growth observed in interspecific crosses between wild Cucumis spp. and melon (C. melo L.) Sci. Hort. 138 144 150

    • Search Google Scholar
    • Export Citation
  • Matsumoto, Y., Ogawara, T., Miyagi, M., Watanabe, N. & Kuboyama, T. 2011 Response of wild Cucumis to inoculation with Fusarium oxysporum f. sp. melonis race 1, 2y J. Jpn. Soc. Hort. Sci. 80 4 414 419

    • Search Google Scholar
    • Export Citation
  • Nisini, P.T., Colla, G., Granati, E., Temperini, O., Crinò, P. & Saccardo, F. 2002 Rootstock resistance to fusarium wilt and effect on fruit yield and quality of two muskmelon cultivars Sci. Hort. 93 3–4 281 288

    • Search Google Scholar
    • Export Citation
  • Özarslandan, A., Söğüt, M.A., Yetişir, H. & Elekçioğlu, I.H. 2011 Screening of bottle gourds (Lagenaria siceraria (Molina) Standley) genotypes with rootstock potential for watermelon production for resistance against Meloidogyne incognita (Kofoid & White, 1919) Chitwood and Meloidogyne javanica (Treub, 1885) Chitwood. Turkish J. Entomol. 35 4 687 697

    • Search Google Scholar
    • Export Citation
  • Perchepied, L. & Pitrat, M. 2004 Polygenic inheritance of partial resistance to Fusarium oxysporum f. sp. melonis race 1.2 in melon Phytopathology 94 1331 1336

    • Search Google Scholar
    • Export Citation
  • Pivonia, S., Gerstl, Z., Maduel, A., Levita, R. & Cohen, R. 2010 Management of Monosporascus sudden wilt of melon by soil application of fungicides Eur. J. Plant Pathol. 128 2 201 209 doi: 10.1007/s10658-010-9644-7

    • Search Google Scholar
    • Export Citation
  • Pofu, K.M. & Mashela, P.W. 2011 Improving survival of inter-generic grafts for suppressing Meloidogyne species in Citrullus cultivars and indigenous Cucumis species Acta Agr. Scand. 62 4 383 386

    • Search Google Scholar
    • Export Citation
  • Pofu, K.M., Mashela, P.W. & Mafeo, T.P. 2013 Optimising stem diameters of watermelon cultivars and indigenous Cucumis species for improving compatibility of inter-generic grafts Acta Hort. 1007 807 812

    • Search Google Scholar
    • Export Citation
  • Renner, S.S., Schaefer, H. & Kocyan, A. 2007 Phylogenetics of Cucumis (Cucurbitaceae): Cucumber (C. sativus) belongs in an Asian/Australian clade far from melon (C. melo) BMC Evol. Biol. 7 58

    • Search Google Scholar
    • Export Citation
  • Rouphael, Y., Schwarz, D., Krumbein, A. & Colla, G. 2010 Impact of grafting on product quality of fruit vegetables Sci. Hort. 127 2 172 179

  • Sebastian, P., Schaefer, H., Telford, I.R.H. & Renner, S.S. 2010 Cucumber (Cucumis sativus) and melon (C. melo) have their wild progenitors in India, and the sister species of Cucumis melo is from Australia Proc. Natl. Acad. Sci. USA 107 32 14269 14273

    • Search Google Scholar
    • Export Citation
  • Sigüenza, C., Schochow, M., Turini, T. & Ploeg, A. 2005 Use of Cucumis metuliferus as a rootstock for melon to manage Meloidogyne incognita J. Nematol. 37 3 276 280

    • Search Google Scholar
    • Export Citation
  • Singh, A.K. & Yadava, K.S. 1984 An analysis of interspecific hybrids and phylogenetic implications in Cucumis (Cucurbitaceae) Plant Syst. Evol. 147 3–4 237 252

    • Search Google Scholar
    • Export Citation
  • Soteriou, G.A., Kyriacou, M.C., Siomos, A.S. & Gerasopoulos, D. 2014 Evolution of watermelon fruit physicochemical and phytochemical composition during ripening as affected by grafting Food Chem. 165 282 289

    • Search Google Scholar
    • Export Citation
  • Soteriou, G.A., Papayiannis, L.C. & Kyriacou, M.C. 2016 Indexing melon physiological decline to fruit quality and vine morphometric parameters Sci. Hort. 203 207 215

    • Search Google Scholar
    • Export Citation
  • Thomas, P. & More, T.A. 1990 Screening wild Cucumis sp. in the field and with artificial inoculation against Fusarium oxysporum f.sp. melonis Cucurbit Genet. Coop. Rpt. 13 18 19

    • Search Google Scholar
    • Export Citation
  • Verslues, P.E., Agarwal, M., Katiyar-Agarwal, S., Zhu, J. & Zhu, J.K. 2006 Methods and concepts in quantifying resistance to drought, salt and freezing, abiotic stresses that affect plant water status Plant J. 45 4 523 539

    • Search Google Scholar
    • Export Citation
  • Verzera, A., Dima, G., Tripodi, G., Condurso, C., Crinò, P., Romano, D., Mazzaglia, A., Lanza, C.M., Restuccia, C. & Paratore, A. 2014 Aroma and sensory quality of honeydew melon fruits (Cucumis melo L. subsp. melo var. inodorus H. Jacq.) in relation to different rootstocks Sci. Hort. 169 118 124

    • Search Google Scholar
    • Export Citation
  • Walters, S.A. & Wehner, T.C. 2002 Incompatibility in diploid and tetraploid crosses of Cucumis sativus and Cucumis metuliferus Euphytica 128 3 371 374

    • Search Google Scholar
    • Export Citation
  • Zhao, X., Guo, Y., Huber, D.J. & Lee, J. 2011 Grafting effects on postharvest ripening and quality of 1-methylcyclopropene-treated muskmelon fruit Sci. Hort. 130 3 581 587

    • Search Google Scholar
    • Export Citation

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Contributor Notes

C. Gisbert and B. Picó thank the Programa Hispano–Brasileño de Cooperación Universitaria PHBP14/00021. The authors also thank the MINECO projects AGL2013-49040-C2-1-R and AGL2014-53398-C2-2-R, cofunded with FEDER funds. We also thank the germplasm bank of COMAV for supplying Cucumis spp. seeds and Rijk Zwaan for providing seeds of commercial scions and rootstocks and open field cultivation facilities.

Corresponding author. E-mail: cgisbert@btc.upv.es.

  • View in gallery

    Discriminant morphological characters among Cucumis ficifolius (F), C. anguria (A), C. myriocarpus (My), and their hybrids ‘UPV-FMy’ and ‘UPV-FA’. From top to bottom: leaves, stems, aculei from ovaries with an inferior opaque part and a superior hyaline part, and fruits. The morphology was evaluated on greenhouse growing plants.

  • View in gallery

    Response of the two hybrids, ‘UPV-FA’ (C. ficifolius × C. anguria) and ‘UPV-FMy’ (C. ficifolius × C. myriocarpus), compared with a melon susceptible control against Fusarium oxysporum f sp. melonis: plants of cantaloupe (A), ‘UPV-FA’ (B), and “UPV-FMy” (C); against Monosporascus cannoballus: roots extracted from naturally infested soil of cv. Piñonet Piel de Sapo (D), ‘UPV-FA’ (E), and ‘UPV-FMy’ (F); against powdery mildew: plants cultivated in an open field of cv. Piñonet Piel de Sapo (G), ‘UPV-FA’ (H), and ‘UPV-FMy’ (I).

  • View in gallery

    Fruit of muskmelons cv. Vedrantais (Charentais type) from ungrafted plants (A) and plants grafted onto ‘UPV-FMy’ (B) and ‘UPV-FA’ (C).

  • View in gallery

    Fruit of Piel de Sapo melons cv. Finura from ungrafted plants (A) and plants grafted onto C. maxima × C. moschata hybrid ‘Cobalt’ (B), the Cucumis melo ‘64-376RZ’ (C), “UPV-FMy” (D), and “UPV-FA” (E).

  • Cáceres, A., Ferriol, M., Gisbert, C. & Picó, B. 2016b Use of wild Cucumis as potential new rootstocks for melons. Proc. XI EUCARPIA Mtg. Cucurbit Genet. and Breeding, Warsaw, Poland, 24–28 July 2016. p. 300–304

  • Cáceres, A., Picó, B. & Gisbert, C. 2016a Germination of wild Cucumis species and interspecific hybrids. Proc. XI EUCARPIA Mtg. Cucurbit Genet. Breeding, Warsaw, Poland, 24–28 July 2016. p. 227–230

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  • Cohen, R., Tyutyunik, J., Fallik, E., Oka, Y., Tadmor, Y. & Edelstein, M. 2014 Phytopatological evaluation of exotic watermelon germplasm as a basis for rootstock breeding Sci. Hort. 165 203 210

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  • Colla, G., Rouphael, Y., Cardarelli, M., Massa, D., Salerno, A. & Rea, E. 2006 Yield, fruit quality and mineral composition of grafted melon plants grown under saline conditions J. Hort. Sci. Biotechnol. 81 146 152

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  • Condurso, C., Verzera, A., Dima, G., Tripodi, G., Crinò, P., Paratore, A. & Romano, D. 2012 Effects of different rootstocks on aroma volatile compounds and carotenoid content of melon fruits Sci. Hort. 148 9 16

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  • Gisbert, C., Gammoudi, N., Munera, M., Giné, A., Porcurull, M., Sorribas, F.J. & Picó, M.B. 2017 Evaluation of two potential Cucumis spp. resources for grafting melons Acta Hort. 1151 157 162

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  • Gisbert, C., Perpiñà, G., Cáceres, A., Segarra, G., Pérez de Castro, A. & Picó, B. 2016 El injerto como técnica para la recuperación de variedades tradicionales: “el meló d’Or de Ontinyent”, un caso de estudio Phytoma 282 40 46

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  • Jang, Y., Huh, Y.C., Park, D.K., Mun, B., Lee, S. & Um, Y. 2014 Greenhouse evaluation of melon rootstock resistance to Monosporascus root rot and vine decline as well as of yield and fruit quality in grafted “Inodorus” melons Kor. J. Soc. Hort. Sci. 32 5 614 622

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  • Kokalis-Burelle, N. & Rosskopf, E.N. 2011 Microplot evaluation of rootstocks for control of Meloidogyne incognita on grafted tomato, muskmelon, and watermelon J. Nematol. 43 3–4 166 171

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  • Lee, J.M., Kubota, C., Tsao, S.J., Bie, Z., Echevarria, P.H., Morra, L. & Oda, M. 2010 Current status of grafting: Diffusion, grafting techniques, automation Sci. Hort. 127 93 105

    • Search Google Scholar
    • Export Citation
  • Liu, B., Ren, J., Zhang, Y., An, J., Chen, M., Chen, H., Xu, Ch. & Ren, H. 2015 A new grafted rootstock against root-knot nematode for cucumber, melon, and watermelon Agron. Sustain. Dev. 35 1 251 259

    • Search Google Scholar
    • Export Citation
  • Mashela, P.W., Shimelis, H.A. & Mudau, F.N. 2008 Comparison of the efficacy of ground wild cucumber fruits, aldicarb and fenamiphos on doses against root-knot nematode, Meloidogyne incognita on tomato Ind. J. Nematol. 35 38 41

    • Search Google Scholar
    • Export Citation
  • Matsumoto, Y., Miyagi, M., Watanabe, N. & Kuboyama, T. 2012 Temperature-dependent enhancement of pollen tube growth observed in interspecific crosses between wild Cucumis spp. and melon (C. melo L.) Sci. Hort. 138 144 150

    • Search Google Scholar
    • Export Citation
  • Matsumoto, Y., Ogawara, T., Miyagi, M., Watanabe, N. & Kuboyama, T. 2011 Response of wild Cucumis to inoculation with Fusarium oxysporum f. sp. melonis race 1, 2y J. Jpn. Soc. Hort. Sci. 80 4 414 419

    • Search Google Scholar
    • Export Citation
  • Nisini, P.T., Colla, G., Granati, E., Temperini, O., Crinò, P. & Saccardo, F. 2002 Rootstock resistance to fusarium wilt and effect on fruit yield and quality of two muskmelon cultivars Sci. Hort. 93 3–4 281 288

    • Search Google Scholar
    • Export Citation
  • Özarslandan, A., Söğüt, M.A., Yetişir, H. & Elekçioğlu, I.H. 2011 Screening of bottle gourds (Lagenaria siceraria (Molina) Standley) genotypes with rootstock potential for watermelon production for resistance against Meloidogyne incognita (Kofoid & White, 1919) Chitwood and Meloidogyne javanica (Treub, 1885) Chitwood. Turkish J. Entomol. 35 4 687 697

    • Search Google Scholar
    • Export Citation
  • Perchepied, L. & Pitrat, M. 2004 Polygenic inheritance of partial resistance to Fusarium oxysporum f. sp. melonis race 1.2 in melon Phytopathology 94 1331 1336

    • Search Google Scholar
    • Export Citation
  • Pivonia, S., Gerstl, Z., Maduel, A., Levita, R. & Cohen, R. 2010 Management of Monosporascus sudden wilt of melon by soil application of fungicides Eur. J. Plant Pathol. 128 2 201 209 doi: 10.1007/s10658-010-9644-7

    • Search Google Scholar
    • Export Citation
  • Pofu, K.M. & Mashela, P.W. 2011 Improving survival of inter-generic grafts for suppressing Meloidogyne species in Citrullus cultivars and indigenous Cucumis species Acta Agr. Scand. 62 4 383 386

    • Search Google Scholar
    • Export Citation
  • Pofu, K.M., Mashela, P.W. & Mafeo, T.P. 2013 Optimising stem diameters of watermelon cultivars and indigenous Cucumis species for improving compatibility of inter-generic grafts Acta Hort. 1007 807 812

    • Search Google Scholar
    • Export Citation
  • Renner, S.S., Schaefer, H. & Kocyan, A. 2007 Phylogenetics of Cucumis (Cucurbitaceae): Cucumber (C. sativus) belongs in an Asian/Australian clade far from melon (C. melo) BMC Evol. Biol. 7 58

    • Search Google Scholar
    • Export Citation
  • Rouphael, Y., Schwarz, D., Krumbein, A. & Colla, G. 2010 Impact of grafting on product quality of fruit vegetables Sci. Hort. 127 2 172 179

  • Sebastian, P., Schaefer, H., Telford, I.R.H. & Renner, S.S. 2010 Cucumber (Cucumis sativus) and melon (C. melo) have their wild progenitors in India, and the sister species of Cucumis melo is from Australia Proc. Natl. Acad. Sci. USA 107 32 14269 14273

    • Search Google Scholar
    • Export Citation
  • Sigüenza, C., Schochow, M., Turini, T. & Ploeg, A. 2005 Use of Cucumis metuliferus as a rootstock for melon to manage Meloidogyne incognita J. Nematol. 37 3 276 280

    • Search Google Scholar
    • Export Citation
  • Singh, A.K. & Yadava, K.S. 1984 An analysis of interspecific hybrids and phylogenetic implications in Cucumis (Cucurbitaceae) Plant Syst. Evol. 147 3–4 237 252

    • Search Google Scholar
    • Export Citation
  • Soteriou, G.A., Kyriacou, M.C., Siomos, A.S. & Gerasopoulos, D. 2014 Evolution of watermelon fruit physicochemical and phytochemical composition during ripening as affected by grafting Food Chem. 165 282 289

    • Search Google Scholar
    • Export Citation
  • Soteriou, G.A., Papayiannis, L.C. & Kyriacou, M.C. 2016 Indexing melon physiological decline to fruit quality and vine morphometric parameters Sci. Hort. 203 207 215

    • Search Google Scholar
    • Export Citation
  • Thomas, P. & More, T.A. 1990 Screening wild Cucumis sp. in the field and with artificial inoculation against Fusarium oxysporum f.sp. melonis Cucurbit Genet. Coop. Rpt. 13 18 19

    • Search Google Scholar
    • Export Citation
  • Verslues, P.E., Agarwal, M., Katiyar-Agarwal, S., Zhu, J. & Zhu, J.K. 2006 Methods and concepts in quantifying resistance to drought, salt and freezing, abiotic stresses that affect plant water status Plant J. 45 4 523 539

    • Search Google Scholar
    • Export Citation
  • Verzera, A., Dima, G., Tripodi, G., Condurso, C., Crinò, P., Romano, D., Mazzaglia, A., Lanza, C.M., Restuccia, C. & Paratore, A. 2014 Aroma and sensory quality of honeydew melon fruits (Cucumis melo L. subsp. melo var. inodorus H. Jacq.) in relation to different rootstocks Sci. Hort. 169 118 124

    • Search Google Scholar
    • Export Citation
  • Walters, S.A. & Wehner, T.C. 2002 Incompatibility in diploid and tetraploid crosses of Cucumis sativus and Cucumis metuliferus Euphytica 128 3 371 374

    • Search Google Scholar
    • Export Citation
  • Zhao, X., Guo, Y., Huber, D.J. & Lee, J. 2011 Grafting effects on postharvest ripening and quality of 1-methylcyclopropene-treated muskmelon fruit Sci. Hort. 130 3 581 587

    • Search Google Scholar
    • Export Citation
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