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Article

Evaluation of Figleaf Gourd and White-Seeded Pumpkin Genotypes as Promising Rootstocks for Cucumber Grafting

by
Gengyun Li
,
Jiamei Zou
,
Tianrui Gong
,
Xuejiao Li
,
Jing Meng
,
Jie Zhang
,
Bin Xu
and
Shuilian He
*
Key Laboratory of Vegetable Biology of Yunnan Province, College of Landscape and Horticulture, Yunnan Agricultural University, Kunming 650201, China
*
Author to whom correspondence should be addressed.
Horticulturae 2025, 11(7), 778; https://doi.org/10.3390/horticulturae11070778
Submission received: 1 June 2025 / Revised: 16 June 2025 / Accepted: 26 June 2025 / Published: 3 July 2025
(This article belongs to the Special Issue Genomics and Genetic Diversity in Vegetable Crops)
Browse Figures
Versions Notes

Abstract

Rootstocks are vital in cucumber production. Although figleaf gourd (Cucurbita ficifolia) is among the species used, its application remains limited due to the perception that white-seeded pumpkin (C. maxima × C. moschata) offers superior commercial traits. This perception is partly due to the insufficient collection and evaluation of local figleaf gourd germplasm, which has obscured its potential as a rootstock. Based on prior screening, four wild figleaf gourd genotypes from Yunnan Province were selected and compared with seven commercial white-seeded pumpkin rootstocks. Scions grafted onto figleaf gourd exhibited vegetative growth (stem diameter, plant height, and leaf area) and fruit morphology (length, diameter, biomass, and surface bloom) comparable to the top-performing white-seeded pumpkin genotypes. Fruits from figleaf gourd rootstocks also displayed comparable or significantly higher nutritional quality, including vitamin C, total soluble solids, soluble sugars, and proteins. Notably, figleaf gourd itself showed significantly greater intrinsic resistance to Fusarium wilt than white-seeded pumpkin. When used as a rootstock, it protected the scion from pathogen stress by triggering a stronger antioxidant response (higher SOD and POD activity) and mitigating cellular damage (lower MDA levels and electrolyte leakage). These results provide evidence that these figleaf gourd genotypes are not merely viable alternatives but are high-performing rootstocks, particularly in enhancing nutritional value and providing elite disease resistance.

1. Introduction

Grafting is a horticultural technique that joins a scion and a rootstock through tissue fusion, resulting in a single, unified plant. Among major vegetable crops, grafting is primarily applied to species in the Solanaceae and Cucurbitaceae families. In cucurbitaceous vegetables, grafting enhances resistance to abiotic stresses and soil-borne diseases, improves yield, and reduces the excessive use of fertilizers and pesticides [1,2,3]. As one of the most economically important cucurbits, cucumber (Cucumis sativus) is widely produced using grafting techniques. Over the past decade, grafted cucumber seedlings have accounted for approximately 75% of total cucumber production in Japan and Korea and nearly 100% of cucumbers grown in solar greenhouses in northern China [4,5,6], making the availability of high-quality rootstocks critical for the industry’s sustainable development.
In cucumber production, one of the most favored rootstocks is the white-seeded pumpkin (Cucurbita maxima × C. moschata), celebrated for its ability to promote vigorous scion growth, its broad environmental adaptability, and its capacity to produce fruits with an attractive appearance and excellent flavor [5,7,8,9]. In contrast, figleaf gourd (C. ficifolia) is considered a niche rootstock, primarily utilized for its strong cold tolerance in winter cultivation [10,11]. However, the broader application of figleaf gourd is hindered by a prevailing perception of its inferiority. It has been reported that cucumbers grafted onto figleaf gourd may exhibit slower growth, lower fruit sugar content, and increased surface bloom, which diminishes their commercial value compared to those on white-seeded pumpkin rootstocks [12,13,14].
Crucially, this negative perception may not reflect the true potential of the species but rather stem from a significant research gap: the historical lack of systematic collection and evaluation of its diverse germplasm. The performance evaluations to date have likely been based on a limited and non-representative selection of figleaf gourd accessions [9,15]. It is well-established that significant intraspecific variation exists in rootstock performance [13,16,17], yet little is known about how different—particularly wild—genotypes of figleaf gourd affect cucumber quality. This knowledge gap has obscured the potential of elite figleaf gourd genotypes that could rival or even surpass commercial rootstocks in key agronomic traits.
To begin addressing this gap, we turned to the rich, untapped genetic resources of wild figleaf gourd. Native to Mexico, this species has naturalized in China, with wild populations thriving in Yunnan Province. Our previous field investigations revealed that these populations exhibit strong adaptability, vigorous growth, and remarkable phenotypic diversity [18]. This suggests that within this wild gene pool lie genotypes with superior rootstock potential. Based on this prior work, where we identified numerous genetic variations (SNPs and InDels), we selected four representative genotypes exhibiting robust growth and adaptability. This led us to hypothesize that these elite wild genotypes, when used as rootstocks, could challenge the conventional view and demonstrate performance comparable or superior to the dominant white-seeded pumpkin rootstocks.
To test this hypothesis, we conducted a comparative study. The four selected wild figleaf gourd genotypes were benchmarked against seven leading commercial white-seeded pumpkin rootstocks. We evaluated their effects on cucumber scion growth, fruit morphology, nutritional quality, and, critically, their ability to confer resistance to Fusarium wilt. This study aims to provide a rigorous, evidence-based assessment of these novel figleaf gourd rootstocks, offering new insights into their value for cucumber production and a scientific foundation for developing high-performance, disease-resistant cucurbit germplasm.

2. Materials and Methods

2.1. Plant Material

The cucumber cultivar ‘Jin Early Round’, obtained from Kerun Agricultural Technology Co., Ltd. (Tianjin, China), was used as the self-rooted control (SR). This specific cultivar was selected for three primary reasons: (1) its significant commercial relevance as a widely cultivated variety in China; (2) its susceptibility to Fusarium wilt, making it an ideal indicator to evaluate the disease resistance conferred by different rootstocks; and (3) to provide a uniform genetic background, ensuring that any observed differences in scion performance could be directly attributed to the influence of the rootstock genotype. For white-seeded pumpkin, seven commercial varieties were selected: The ‘Diamond’ strain was obtained from Shouguang Fanggeng Agricultural Technology Co., Ltd., Shouguang, China. The other six strains—‘Gold Armor Rootstock’ (GAR), ‘Phoenix No.3’ (P3), ‘Japanese Cedar’ (JC), ‘Korean Gold Rootstock’ (KGR), ‘Root King 99’ (RK99), and ‘Black Dominant No. 1’ (BD1)—were provided by Qingdao Yucheng Rootstock Technology Co., Ltd., Qingdao, China. For figleaf gourd, four genetically distinct genotypes (NH-G1, LQ-H2, SP-G5, and HZ-H3) were selected from 233 local accessions previously collected in Yunnan Province. Their selection was based on superior performance in preliminary screenings for growth and grafting suitability. These genotypes originate from Nanhua, Luquan, Shiping, and Huize Counties, respectively, and their genetic and phenotypic diversity has been previously confirmed [18]. The formal identification of the figleaf gourd material was performed by Yongjie Guo (Kunming Institute of Botany) based on morphological characteristics, and voucher specimens were deposited at the Herbarium of the Institute of Botany, Chinese Academy of Sciences (voucher no. KUN 1580438). All experiments were conducted during the summer in a greenhouse at Yunnan Agricultural University (25°13′ N, 102°74′ E; 1890 m elevation). Plants were grown under natural light with an approximate photoperiod of 12 h/day. Average daytime and nighttime temperatures were approximately 28 °C and 17 °C, respectively.

2.2. Experimental Design

The study was designed to evaluate three key performance areas: (1) Vegetative growth: Starting one week after transplanting, scion stem diameter, plant height, and leaf area were measured weekly to monitor growth dynamics until the first fruit harvest. (2) Fruit traits and quality: At commercial maturity, harvested fruits were assessed for morphological traits (fruit length, maximum diameter, fresh biomass, and surface bloom content) and nutritional quality (contents of total soluble solids, soluble protein, soluble sugars, and vitamin C). (3) Fusarium wilt resistance: Resistance to Fusarium oxysporum f. sp. cucumerinum (FOC) was evaluated in two parallel experiments. First, the intrinsic resistance of the rootstocks themselves was assessed by inoculating seedlings of a representative figleaf gourd (SP-G5), a white-seeded pumpkin (RK99), and self-rooted cucumber (SR), followed by observation of disease symptoms. Second, to evaluate the protective effect on the scion, grafted plants (on SP-G5 and RK99) and SR plants were inoculated. Physiological stress responses were then measured in the scion leaves, including antioxidant enzyme activities—specifically superoxide dismutase (SOD) and peroxidase (POD)—as well as malondialdehyde (MDA) content and electrolyte leakage (EL).

2.3. Measurement of Vegetative Growth Dynamics

Rootstock and scion seeds were sown simultaneously. Grafting was performed using the cleft grafting method once the scion cotyledons had fully expanded. After grafting, seedlings with uniform growth were selected for the experiment and carefully managed during the seedling stage. To ensure consistent measurements across replicates, a specific internode and its corresponding leaf at the same developmental stage were marked on each scion. To track growth dynamics, measurements were taken weekly for eight consecutive weeks. The following parameters were recorded: stem diameter, measured at the midpoint of the marked internode using digital calipers; plant height, measured from the soil surface to the apical growing point; and the area of the marked leaf, which was determined from digital images using ImageJ software (v1.4.3) [19]. All measurements were performed on three biological replicates for each grafting combination. Significant differences among treatments at each time point were determined using Tukey’s HSD test (p < 0.05).

2.4. Assessment of Cucumber Fruit Phenotypes

Two months post-grafting, commercially mature fruits were harvested for phenotypic analysis. For each rootstock treatment, three representative fruits of average size, each from a different plant, were selected as biological replicates. The following morphological traits were measured: (1) Fruit length and diameter: The maximum length and the diameter at the widest point of each fruit were measured. (2) Fruit biomass: The fresh weight of each individual fruit was recorded. (3) Surface bloom content: The amount of bloom (the white, waxy powder on the fruit surface) was quantified using a colorimetric method based on previous studies [20,21]. Briefly, the brightness (L-value) of a piece of transparent tape on a black background was first measured using a precision colorimeter (Model HP-200, Hanpu Optoelectronics Technology Co., Ltd., Shanghai, China) to establish a baseline. The same piece of tape was then gently pressed onto the central surface of the fruit to lift the bloom, transferred back to the black background, and its brightness was measured again. The difference in brightness (ΔL) between the two readings was used as a quantitative measure of the bloom content.

2.5. Determination of Cucumber Fruit Nutritional Quality

The same fruits used for phenotypic assessment were also analyzed for nutritional quality. For each biological replicate, equal-weight tissue samples were obtained from the middle section of the fruit. The following physiological parameters were measured: (1) Total soluble solids (TSS): The TSS content was measured using an automatic temperature-compensated digital refractometer (Atago Palette PR101, Atago Co., Tokyo, Japan) [22]. (2) Soluble protein: The concentration of soluble protein was determined by the Coomassie Brilliant Blue G-250 method [23]. (3) Soluble sugar: The soluble sugar content was quantified by the anthrone colorimetric method [24]. (4) Vitamin C (ascorbic acid): The vitamin C content was measured via the the 2,6-dichloroindophenol titration method [25]. Three biological replicates were performed for each trait. Differences among rootstock grafting treatments were analyzed using Tukey’s HSD test.

2.6. FOC Inoculation and Plant Response Assessment

The FOC strain, provided by the Department of Plant Nutrition at Nanjing Agricultural University, was cultured on potato dextrose agar (PDA) medium for six days at 25 °C. To prepare the inoculum, fungal cultures were flooded with sterile distilled water and gently scraped. The resulting suspension was filtered through four layers of sterile cheesecloth to remove mycelial fragments. The spore concentration was then adjusted to 1 × 108 spores/mL using a hemocytometer. For inoculation, seedlings were carefully uprooted at the third-true-leaf stage. Their roots were washed and then submerged in 10 mL of the spore suspension for 30 min (root-dip inoculation). Following inoculation, the seedlings were repotted in their original pots, and the remaining spore suspension was poured into the soil to ensure successful infection. The intrinsic resistance to FOC was evaluated on non-grafted seedlings of a representative figleaf gourd (SP-G5), a white-seeded pumpkin (RK99), and SR. Disease symptoms and plant phenotypes were visually assessed and photographically documented at 0 (pre-inoculation), 7, and 14 days post-inoculation (dpi). To assess the protective effect of rootstocks, the same FOC inoculation procedure was applied to grafted plants (cucumber scions on SP-G5 and RK99) and the SR control. Leaf samples from the scions were collected at 0, 2, 4, and 6 dpi to measure key physiological stress markers. For physiological measurements, 0.1 g of fresh leaf tissue was homogenized in 1 mL of the specific extraction buffer provided in the commercial assay kits (Grace Biotechnology, Suzhou, China). The homogenates were centrifuged at 8000× g for 10 min at 4 °C, and the resulting supernatants were collected for enzyme activity and lipid peroxidation assays. SOD activity was determined based on its ability to inhibit the photochemical reduction of nitroblue tetrazolium (NBT) by superoxide anions under light exposure, with absorbance measured at 560 nm. POD activity was measured by monitoring the increase in absorbance at 470 nm caused by the oxidation of guaiacol in the presence of hydrogen peroxide. MDA content was quantified using the thiobarbituric acid (TBA) method. After reaction under acidic and high-temperature conditions, the absorbance of the resulting MDA–TBA adduct was measured at 532 nm. To correct for interference from soluble sugars and other non-specific compounds, absorbance readings were also taken at 450 nm and 600 nm, and these values were subtracted from the 532 nm reading according to the kit’s formula. All measurements were conducted according to the manufacturer’s instructions provided with the kits. Electrolyte leakage was measured following the method described by Liu et al. (2022) [26]. Briefly, cucumber pieces with a diameter of 10 mm and a thickness of 1 mm were rinsed three times with deionized water, then submerged in 25 mL of deionized water at 25 °C for 2 h. The initial conductivity (C0, μS/cm) was measured using a conductivity meter. Subsequently, the samples were boiled for 10 min, cooled to room temperature, and the total conductivity (C1, μS/cm) was recorded. Electrolyte leakage (EL%) was calculated using the formula: EL% = (C0 × 100)/C1. Three biological replicates were conducted for each trait. Statistical differences among treatments were analyzed using Tukey’s HSD test.

3. Results

3.1. Cucumber Scions Grafted onto Figleaf Gourd Rootstocks Exhibit Comparable Vegetative Growth Rates to Those on White-Seeded Pumpkin Strains

Periodic monitoring of cucumber vegetative growth was initiated seven days after grafting onto the two types of rootstocks. Although scions with similar initial vigor and size were selected, significant differences in all measured traits emerged one week after grafting, and these differences grew more pronounced over time (Table 1). Among the white-seeded pumpkin strains, scions grafted onto GAR exhibited a consistently high growth rate, achieving significantly greater stem diameter and leaf area than most other genotypes throughout the observation period (Figure 1A,B). Across all tested rootstocks, Diamond, KGR, and BD1 exhibited the highest growth rates in stem diameter and plant height (Figure 1A,C). Among the figleaf gourd rootstocks, LQ-H2 showed a notable growth rate in leaf area, resulting in a final leaf area that was significantly greater than that of most other rootstocks and comparable to that of GAR (Figure 1B). Regarding plant height, NH-G1 and LQ-H2 also grew rapidly, with rates comparable to the fastest-growing white-seeded pumpkin strains (Diamond, KGR, and BD1) (Figure 1C). Overall, cucumber scions grafted onto figleaf gourd rootstocks grew at rates similar to those on white-seeded pumpkin rootstocks. Additionally, all grafted plants ultimately surpassed self-rooted cucumber seedlings in final vegetative growth (Table 1).

3.2. Figleaf Gourd Rootstocks Produce Cucumber Fruits with Morphological Traits Comparable to Those from White-Seeded Pumpkin Rootstocks

Phenotypic measurements of fruit length revealed the strong performance of figleaf gourd rootstocks. The resulting fruits showed no significant difference in length from those of the top-performing white-seeded pumpkin strains (e.g., KGR, Diamond, and RK99), yet they were significantly longer than those from the lowest-performing strain, BD1, and the SR control (Figure 2A). A similar trend was observed for fruit diameter (Figure 2B). In terms of fruit biomass, while KGR and RK99 produced the heaviest fruits, the values were not significantly different from those of the figleaf gourd rootstocks (Figure 2C). Fruit surface bloom, quantified by brightness difference, where a smaller value indicated less bloom, also showed few significant variations. Only fruits from SP-G5, P3, and KGR had significantly less bloom (a lower brightness difference) than SR, while no significant differences were found among the other rootstocks. These results indicate that figleaf gourd rootstocks yield cucumber fruits with a length, diameter, biomass, and surface bloom content comparable to those from commercial white-seeded pumpkin strains, with some rootstocks even outperforming self-rooted seedlings and white-seeded pumpkin strains.

3.3. Figleaf Gourd Rootstocks Confer Comparable or Superior Nutritional Quality to Cucumber Fruits Relative to White-Seeded Pumpkin Rootstocks

The figleaf gourd rootstocks NH-G1 and HZ-H3 produced fruits with significantly higher vitamin C (VC) levels compared to SR seedlings and several white-seeded pumpkin strains (Diamond, JC, and KGR), while showing no significant difference from strains P3 and RK99 (Figure 3A). The figleaf gourd genotypes NH-G1, SP-G5, and HZ-H3 also yielded fruits with significantly higher total soluble solids contents than SR and white-seeded pumpkin strains KGR and RK99 (Figure 3B). The soluble sugar content in cucumber fruits from figleaf gourd rootstocks was generally higher, with HZ-H3, NH-G1, and LQ-H2 showing the highest levels among all tested plants, significantly exceeding those of SR seedlings and all white-seeded pumpkin strains (Figure 3C). For soluble protein, only minor differences were observed between figleaf gourd and white-seeded pumpkin rootstocks. The highest values were observed in SP-G5, LQ-H2, Diamond, HZ-H3, P3, and JC strains, all of which were significantly higher than in SR fruits. Notably, genotypes like HZ-H3 and NH-G1 significantly enhanced key metrics such as soluble sugar and vitamin C content, surpassing most tested white-seeded pumpkin rootstocks.

3.4. Figleaf Gourd Demonstrates Superior Resistance to Fusarium Wilt Compared to White-Seeded Pumpkin and Cucumber

To compare the intrinsic resistance to Fusarium wilt, seedlings of figleaf gourd (SP-G5), white-seeded pumpkin (RK99), and cucumber (SR) were inoculated with FOC. At seven days post-inoculation, SR seedlings exhibited severe yellowing and wilting. In contrast, the RK99 seedlings were more tolerant, with only a single leaf showing mild symptoms, while the SP-G5 seedlings remained asymptomatic (Figure 4). By 14 days post-inoculation, the infection had progressed dramatically in the susceptible plants: SR seedlings were necrotic and near death, while RK99 plants showed significant yellowing and wilting, primarily on older leaves. In contrast, SP-G5 plants remained largely healthy, with only slight chlorosis observed on a few lower leaves (Figure 4). These results align with field observations and confirm that figleaf gourd exhibits significantly higher resistance to Fusarium wilt than both white-seeded pumpkin and cucumber.

3.5. Cucumber Scions Grafted onto Figleaf Gourd Rootstocks Exhibit Higher Antioxidant Enzyme Activity and Lower Cellular Damage Under Fusarium Wilt Infection

Following FOC infection, the protective effect of the rootstock was evident. Scions grafted onto figleaf gourd (SP-G5) remained asymptomatic, whereas those on white-seeded pumpkin (RK99) exhibited leaf wilting and marginal necrosis. In comparison, SR plants displayed severe yellowing and wilting by 7 days post-inoculation and were near death (Supplementary Figure S1). Measurements of antioxidant enzyme activity indicated that FOC infection triggered a substantial increase in SOD activity in scions grafted onto both SP-G5 and RK99, while SOD activity in SR scions remained largely unchanged. Notably, at 4 days post-inoculation, SOD activity in SP-G5 scions was significantly higher than in RK99 scions (Figure 5A). Similarly, POD activity increased significantly in all treatments following inoculation, but the induction was markedly greater in SP-G5 scions than in RK99 or SR scions (Figure 5B). Conversely, markers for cellular damage showed the opposite trend. MDA content, an indicator of lipid peroxidation, was highest in SR. Crucially, from 4 dpi onward, MDA levels in SP-G5 scions were significantly lower than in RK99 scions (Figure 5C). A similar trend was observed for electrolyte leakage (EL), which reflects cell membrane damage (Figure 5D). Collectively, the lower MDA and EL levels in SP-G5, coupled with its heightened antioxidant enzyme activity, indicate that the figleaf gourd rootstock confers superior protection to cucumber scions against FOC-induced oxidative stress compared to the white-seeded pumpkin rootstock.

4. Discussion

Grafting is a vital technology in modern cucurbit production, as appropriate rootstocks can improve scion resistance to biotic and abiotic stresses, enhance nutrient uptake, and increase yield [2]. In China, white-seeded pumpkin is one of the most widely used rootstocks in cucumber cultivation, valued for conferring favorable traits such as improved fruit quality and disease resistance [9,27,28]. In contrast, figleaf gourd, primarily found in regions like Yunnan Province, remains an underutilized genetic resource. Its potential has been largely overlooked, not due to inherent inferiority, but due to a history of insufficient collection and systematic evaluation—a situation that is common for crop wild relatives [29,30]. Much of its genetic diversity, which is substantial due to its semi-wild status [9,18], remains untapped. The four genotypes in this study, selected from our extensive collections in Yunnan, represent elite germplasm with demonstrated ecological adaptability and preliminary resistance to Fusarium wilt. This study was therefore conducted to systematically evaluate these promising figleaf gourd genotypes alongside widely used commercial white-seeded pumpkin rootstocks, with the aim of exploring their potential value in cucumber grafting systems.
Several studies have already established the excellence of figleaf gourd as a rootstock for conferring tolerance to abiotic stresses. Its most noted benefit is enhancing the cold tolerance of cucumber scions [11], a critical trait for extending the growing season. The mechanism often involves the upregulation of antioxidant enzyme activities (e.g., SOD and POD) and improved photosynthetic efficiency under low-temperature stress [11,31]. Similarly, grafting onto figleaf gourd markedly improves cucumber resistance to salinity [32,33]. These findings provide a strong precedent for its stress-mitigating capabilities. Building on this, our results show that these figleaf gourd genotypes also possess elite resistance against Fusarium wilt, a devastating soil-borne disease. Fusarium wilt is a major threat to cucumber production. The pathogen penetrates roots and invades the plant’s vascular tissues via xylem vessels, ultimately causing plant death [34,35,36] and resulting in severe economic losses [37,38,39]. Our results confirm that figleaf gourd possesses significantly higher intrinsic resistance than white-seeded pumpkin (Figure 4), consistent with previous reports [40,41,42]. More importantly, we showed that this resistance is effectively transferred to the susceptible scion. Scions grafted onto figleaf gourd exhibited a powerful and rapid antioxidant defense response (higher SOD and POD activity) upon pathogen challenge, which successfully mitigated cellular damage (lower MDA and EL levels) (Figure 5). This proactive defense priming appears to be a key feature contributing to effective resistance, whereby the rootstock may help the scion restrict pathogen invasion before substantial damage occurs [43].
Despite its clear advantages in stress resistance, the broader adoption of figleaf gourd has been hampered by perceptions of inferior fruit quality—perceptions that our findings suggest are based on limited genetic sampling. For instance, some studies reported that cucumbers grafted onto figleaf gourd had lower soluble solids and fructose contents [12,44]. While those findings are likely valid for the specific genotypes evaluated in those studies, our results reveal significant variation among different figleaf gourd lines. Genotypes like NH-G1, SP-G5, and HZ-H3 produced fruits with nutritional quality comparable or superior to many commercial white-seeded pumpkin rootstocks (Figure 3). Furthermore, even in cases where the sugar content might be lower, this is not necessarily a negative trait; such fruits may be well-suited for specialized markets, such as for consumers managing diabetes. Indeed, figleaf gourd itself has been reported to help stabilize blood glucose levels [45,46,47], suggesting that cucumbers with reduced sugars from these rootstocks could be a healthier option for specific consumer groups.
This principle of genotype-dependency also extends to critical aesthetic traits like fruit bloom. Figleaf gourd is often associated with producing fruits with heavy bloom, a trait considered undesirable in some markets that prefer a cleaner appearance and longer shelf life [20,48]. Bloom formation is known to be influenced by various environmental factors, including UV radiation, planting season, air pollution, and humidity [13,14]. In addition to these external factors, the rootstock genotype also plays a critical role, with experimental evidence demonstrating significant variation in bloom production even among different genotypes of white-seeded pumpkin [9,14,49]. Our results showed that figleaf gourd genotype SP-G5 produced cucumbers with bloom levels as low as those of the best-performing white-seeded pumpkin genotypes (KGR and P3). Ultimately, our findings for both nutritional and aesthetic traits align with the broader consensus that rootstock effects on scion performance are highly genotype-specific [50,51,52]. This underscores that previous reports of drawbacks likely reflect the limited germplasm evaluated and that systematic screening, like the one presented here, is crucial for identifying elite figleaf gourd genotypes that overcome these perceived commercial limitations.
A significant finding of this study is the profound impact of the rootstock’s genetic identity on scion performance. The wide-ranging variations we observed—from vegetative growth and fruit morphology to nutritional quality—all occurred on a genetically uniform cucumber scion, strongly indicating that the rootstock genotype is a primary driver of these outcomes. This principle is clearly illustrated by the significant performance differences not only between the figleaf gourd and white-seeded pumpkin groups but also among the distinct genotypes within each group. Our use of a single scion cultivar in this context allowed for a direct comparison of the rootstocks’ effects, minimizing confounding variables from the scion. While this approach effectively highlighted the genotype-specific influence of the rootstocks, we also recognize its inherent limitations. Rootstock–scion interactions can be highly specific [51,52], and the superior performance we observed may not be universally applicable to all cucumber cultivars. Therefore, several crucial next steps are needed to fully capitalize on the potential of figleaf gourd as a high-value rootstock. First, it is essential to evaluate these elite genotypes with a broader range of commercial scions to validate the general applicability of their benefits. Second, large-scale field trials are necessary to confirm their performance and commercial viability under real-world agricultural conditions. Finally, future research should also address fundamental biological barriers, such as seed dormancy and pollination behavior, which currently hinder the efficient breeding and multiplication of this promising species. Such a multi-faceted approach will be essential for translating these promising findings into tangible benefits for modern agriculture.

5. Conclusions

The four evaluated figleaf gourd genotypes proved to be high-performing cucumber rootstocks. Scions grafted onto them exhibited growth rates comparable to, and in some cases exceeding, those on commercial white-seeded pumpkin rootstocks. The resulting fruits maintained similar key morphological traits and nutritional quality, with some genotypes showing superior levels of beneficial compounds. Most critically, figleaf gourd rootstock conferred significantly enhanced resistance to Fusarium wilt, a major threat to cucumber production, by activating higher antioxidant enzyme activity in the scions and mitigating pathogen-induced damage. These findings challenge the conventional perception of figleaf gourd’s inferiority and highlight its substantial potential as a high-value, disease-resistant rootstock for cucumber breeding programs and commercial application.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/horticulturae11070778/s1, Figure S1: The phenotypes of the scions of cucumbers grafted onto figleafgourd (SP-G5), white-seeded pumpkin (RK99), and self-rooted seedlings at 0 days (before FOC inoculation), and at 3 days and 7 days after inoculation; Table S1: p-values for Tukey’s HSD test of SOD, POD, MDA, and Electrolyte leakage rate (EL) of Figure 5.

Author Contributions

Conceptualization, S.H.; methodology, G.L. and S.H.; validation, formal analysis, and investigation, J.Z. (Jiamei Zou) and T.G.; data curation, X.L. and J.M.; writing—original draft preparation, G.L., J.M., and S.H.; writing—review and editing, G.L., J.Z. (Jie Zhang), B.X., and X.L.; supervision, S.H.; funding acquisition, G.L., S.H., B.X., and J.Z. (Jie Zhang). All authors have read and agreed to the published version of the manuscript.

Funding

The project was supported by the Yunnan Fundamental Research Projects (202501BD070001-056, 202402AE090012, 202301BD070001-027, and 202301BD070001-168), the Yunnan Provincial Science and Technology Talent and Platform Project (202205AF150017), and the Young Talents of Yunnan Xingdian (XDYC-QNRC-2022-0233).

Data Availability Statement

The raw data generated in this study are publicly available in the Zenodo repository at https://doi.org/10.5281/zenodo.15674596 (accessed on 16 June 2025).

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Vegetative growth dynamics of cucumber scions grafted onto different rootstocks. (A) Internode diameter, (B) leaf area, and (C) plant height were measured every 7 days for 8 weeks after grafting. Data are presented as means ± SEs (n = 3).
Figure 1. Vegetative growth dynamics of cucumber scions grafted onto different rootstocks. (A) Internode diameter, (B) leaf area, and (C) plant height were measured every 7 days for 8 weeks after grafting. Data are presented as means ± SEs (n = 3).
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Figure 2. Morphological traits of cucumber fruits grafted onto different rootstocks. (A) Fruit length. (B) Fruit diameter. (C) Fruit biomass. (D) Brightness difference (ΔL). Different letters indicate significant differences (p < 0.05) evaluated using Tukey’s HSD test (n = 3).
Figure 2. Morphological traits of cucumber fruits grafted onto different rootstocks. (A) Fruit length. (B) Fruit diameter. (C) Fruit biomass. (D) Brightness difference (ΔL). Different letters indicate significant differences (p < 0.05) evaluated using Tukey’s HSD test (n = 3).
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Figure 3. Nutritional quality of cucumber fruits grafted onto different rootstocks. (A) Vitamin C (VC) content. (B) Total soluble solids. (C) Soluble sugar. (D) Soluble protein. Data represent means ± SEs of three biological replicates. Different letters indicate statistically significant differences according to Tukey’s HSD test (p < 0.05).
Figure 3. Nutritional quality of cucumber fruits grafted onto different rootstocks. (A) Vitamin C (VC) content. (B) Total soluble solids. (C) Soluble sugar. (D) Soluble protein. Data represent means ± SEs of three biological replicates. Different letters indicate statistically significant differences according to Tukey’s HSD test (p < 0.05).
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Figure 4. Disease symptoms of Fusarium wilt in figleaf gourd (SP-G5), white-seeded pumpkin (RK99), and cucumber (SR) at 0 day (before FOC inoculation) and at 7 and 14 days post-inoculation.
Figure 4. Disease symptoms of Fusarium wilt in figleaf gourd (SP-G5), white-seeded pumpkin (RK99), and cucumber (SR) at 0 day (before FOC inoculation) and at 7 and 14 days post-inoculation.
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Figure 5. Physiological responses in cucumber scions grafted onto figleaf gourd (SP-G5, blue line) and white-seeded pumpkin (RK99, red line) rootstocks following Fusarium oxysporum f. sp. cucumerinum (FOC) inoculation, with self-rooted seedlings (SR, grey line) as the control. (A) Superoxide dismutase (SOD) activity. (B) Peroxidase (POD) activity. (C) Malondialdehyde (MDA) content. (D) Electrolyte leakage (EL). Data are presented as means ± SDs (n = 3). Different letters indicate significant differences among treatments at each time point according to Tukey’s HSD test (p < 0.05). Exact p-values for all comparisons are provided in Supplementary Table S1.
Figure 5. Physiological responses in cucumber scions grafted onto figleaf gourd (SP-G5, blue line) and white-seeded pumpkin (RK99, red line) rootstocks following Fusarium oxysporum f. sp. cucumerinum (FOC) inoculation, with self-rooted seedlings (SR, grey line) as the control. (A) Superoxide dismutase (SOD) activity. (B) Peroxidase (POD) activity. (C) Malondialdehyde (MDA) content. (D) Electrolyte leakage (EL). Data are presented as means ± SDs (n = 3). Different letters indicate significant differences among treatments at each time point according to Tukey’s HSD test (p < 0.05). Exact p-values for all comparisons are provided in Supplementary Table S1.
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Table 1. Dynamic growth of cucumber scions grafted onto different rootstocks from 1 week to 8 weeks after grafting.
Table 1. Dynamic growth of cucumber scions grafted onto different rootstocks from 1 week to 8 weeks after grafting.
Stock7 d14 d21 d28 d35 d42 d49 d56 d
Stem diameter (mm)
NH-G16.48 ± 0.19 abc7.50 ± 0.34 a8.11 ± 0.42 a8.56 ± 0.31 a9.34 ± 0.33 ab10.11 ± 0.36 a10.38 ± 0.28 ab10.48 ± 0.25 bc
LQ-H26.95 ± 0.25 a7.30 ± 0.25 ab7.65 ± 0.30 abcd8.26 ± 0.36 ab8.53 ± 0.36 bcde9.26 ± 0.34 bc9.81 ± 0.27 bcd9.86 ± 0.16 cd
SP-G55.85 ± 0.19 cd6.35 ± 0.22 def6.64 ± 0.22 ef6.96 ± 0.25 d7.23 ± 0.23 f8.68 ± 0.36 c9.26 ± 0.28 cde9.67 ± 0.20 cd
HZ-H36.76 ± 0.18 ab7.08 ± 0.17 abcd7.32 ± 0.16 abcde7.47 ± 0.18 bcd8.10 ± 0.18 e8.59 ± 0.23 c9.38 ± 0.28 cde9.64 ± 0.21 d
P35.36 ± 0.20 d5.76 ± 0.22 f6.32 ± 0.18 f6.62 ± 0.14 d7.78 ± 0.23 ef8.56 ± 0.25 c9.05 ± 0.21 de9.56 ± 0.16 d
Diamond6.08 ± 0.12 bc6.43 ± 0.14 cdef6.82 ± 0.17 def7.11 ± 0.20 cd8.23 ± 0.14 de9.66 ± 0.17 ab10.94 ± 0.22 a11.85 ± 0.28 a
GAR6.34 ± 0.23 abc6.62 ± 0.25 bcde6.93 ± 0.31 cdef7.22 ± 0.27 cd8.52 ± 0.29 bcde9.78 ± 0.20 ab10.38 ± 0.26 ab11.23 ± 0.28 ab
JC6.42 ± 0.23 abc7.04 ± 0.21 abcd7.75 ± 0.22 abc8.60 ± 0.28 a9.54 ± 0.23 a9.65 ± 0.18 ab9.85 ± 0.25 bcd10.13 ± 0.29 cd
KGR6.65 ± 0.31 ab7.24 ± 0.32 abc7.83 ± 0.28 ab8.18 ± 0.29 ab9.13 ± 0.20 abc9.96 ± 0.24 ab10.40 ± 0.21 ab10.50 ± 0.21 bc
RK996.24 ± 0.23 abc6.76 ± 0.21 abcde7.22 ± 0.22 bcde7.83 ± 0.22 abc8.38 ± 0.23 cde8.64 ± 0.18 c8.75 ± 0.19 e8.80 ± 0.23 e
BD16.25 ± 0.22 abc6.94 ± 0.27 abcd7.53 ± 0.28 abcd8.61 ± 0.33 a8.99 ± 0.29 abcd9.74 ± 0.24 ab9.94 ± 0.31 bc10.14 ± 0.35 cd
SR5.33 ± 0.28 d6.05 ± 0.27 ef6.54 ± 0.28 ef6.88 ± 0.28 d7.21 ± 0.29 f7.55 ± 0.24 d7.78 ± 0.31 f7.86 ± 0.35 f
Leaf area (mm2)
NH-G1349.67 ± 9.22 a410.40 ± 10.42 a482.46 ± 9.35 b606.64 ± 8.07 b656.92 ± 12.36 cd675.35 ± 12.32 c689.28 ± 15.23 e693.83 ± 14.16 d
LQ-H2339.14 ± 10.24 a440.52 ± 10.81 a592.42 ± 10.81 a689.53 ± 14.61 a759.12 ± 7.66 a789.93 ± 15.15 a815.12 ± 13.40 b827.08 ± 12.82 a
SP-G5238.86 ± 6.19 d279.15 ± 7.63 e341.71 ± 11.67 g462.30 ± 6.32 e500.58 ± 12.54 e518.31 ± 12.84 def530.18 ± 11.70 g537.91 ± 5.89 f
HZ-H3232.49 ± 8.79 d280.79 ± 9.65 e341.64 ± 8.21 g451.52 ± 8.49 e530.69 ± 7.06 e568.91 ± 10.23 d583.06 ± 9.65 f601.4 ± 10.80 e
P3286.14 ± 8.79 b321.99 ± 9.65 d364.82 ± 8.21 fg472.96 ± 8.49 e514.21 ± 6.76 e551.71 ± 10.23 de582.99 ± 8.79 f587.42 ± 10.23 e
Diamond287.21 ± 9.36 b327.11 ± 10.80 cd396.65 ± 12.83 def461.40 ± 9.65 e526.48 ± 10.80 e566.02 ± 11.38 d573.24 ± 9.94 f575.34 ± 11.38 ef
GAR261.24 ± 11.99 bcd343.35 ± 15.71 bcd457.43 ± 15.14 bc595.60 ± 10.80 bc701.25 ± 16.07 b816.00 ± 21.49 a856.25 ± 19.28 a865.42 ± 18.03 a
JC253.85 ± 8.15 cd332.81 ± 8.15 cd431.19 ± 2.95 cd558.94 ± 5.27 d654.31 ± 11.73 cd702.65 ± 7.33 bc722.13 ± 4.40 cde747.22 ± 11.72 bc
KGR336.42 ± 9.88 a376.90 ± 10.74 b451.88 ± 12.47 bc560.64 ± 14.50 d631.30 ± 11.05 d676.58 ± 11.33 c702.88 ± 9.88 de709.07 ± 18.25 cd
RK99271.75 ± 11.84 bc332.15 ± 14.46 cd421.48 ± 20.50 cde617.69 ± 12.13 b670.72 ± 19.65 bc696.97 ± 20.21 bc736.26 ± 12.71 cd754.38 ± 15.31 b
BD1284.35 ± 7.22 b351.95 ± 8.08 bcd451.43 ± 9.82 bc574.88 ± 7.52 cd699.87 ± 10.40 b731.55 ± 10.19 b744.63 ± 6.41 c744.06 ± 14.43 bc
SR322.96 ± 8.96 a357.55 ± 8.72 bc393.48 ± 11.26 ef412.29 ± 8.67 f429.68 ± 9.82 f432.90 ± 10.11 e438.48 ± 9.84 h441.35 ± 12.44 g
Plant height (cm)
NH-G172.00 ± 8.08 a100.00 ± 9.64 a138.00 ± 9.54 a183.67 ± 6.64 abc253.00 ± 13.58 ab297.00 ± 6.66 ab329.33 ± 9.82 ab347.00 ± 6.93 a
LQ-H259.67 ± 4.63 ab89.33 ± 4.91 a113.00 ± 4.62 abc160.33 ± 4.91 cde195.67 ± 5.21 de244.00 ± 4.62 c316.00 ± 4.04 abc345.33 ± 4.33 ab
SP-G576.00 ± 7.23 a95.67 ± 3.18 a122.67 ± 4.98 abc172.00 ± 9.54 bcd214.67 ± 8.09 cd238.67 ± 8.95 c260.67 ± 10.68 e280.00 ± 7.23 c
HZ-H366.33 ± 5.21 ab83.67 ± 6.06 a106.33 ± 4.63 bc150.33 ± 5.78 de196.33 ± 6.94 de239.67 ± 7.80 c275.00 ± 6.35 de285.33 ± 6.94 c
P363.33 ± 6.94 ab88.33 ± 7.22 a113.00 ± 8.39 abc173.00 ± 6.93 bcd230.00 ± 9.24 bc252.33 ± 8.09 c295.67 ± 7.22 cd315.00 ± 9.82 b
Diamond78.33 ± 7.51 a102.33 ± 7.80 a131.00 ± 8.96 ab186.00 ± 7.51 ab231.00 ± 9.82 bc283.33 ± 8.67 b329.67 ± 7.80 ab355.00 ± 10.39 a
GAR67.67 ± 4.33 ab90.33 ± 5.81 a115.00 ± 8.96 abc150.00 ± 7.51 de191.00 ± 9.82 de251.33 ± 8.67 c312.67 ± 7.80 bc342.00 ± 10.39 ab
JC62.33 ± 4.91 ab86.00 ± 2.89 a109.00 ± 2.89 bc158.33 ± 6.06 cde215.00 ± 2.89 cd251.67 ± 8.09 c261.33 ± 7.54 e273.67 ± 7.80 c
KGR75.67 ± 10.11 a87.33 ± 9.26 a118.67 ± 10.41 abc200.33 ± 8.37 a263.33 ± 10.68 a317.67 ± 10.14 a340.00 ± 8.66 a358.33 ± 11.26 a
RK9965.33 ± 8.95 ab85.00 ± 8.96 a111.33 ± 11.55 abc147.67 ± 9.82 de191.33 ± 11.84 de229.67 ± 11.84 c264.00 ± 10.69 e278.33 ± 12.72 c
BD177.33 ± 6.94 a102.00 ± 7.51 a139.33 ± 10.14 a181.00 ± 9.29 abc253.00 ± 13.28 ab307.00 ± 11.55 ab340.67 ± 9.26 a364.00 ± 14.80 a
SR48.33 ± 6.64 b80.33 ± 9.26 a101.00 ± 10.39 c136.00 ± 8.96 e173.33 ± 11.55 e189.33 ± 9.82 d203.67 ± 7.54 f219.33 ± 12.14 d
Different letters indicate significant differences (p < 0.05) among grafting treatments on the same date according to Tukey’s HSD test (n = 3).
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Li, G.; Zou, J.; Gong, T.; Li, X.; Meng, J.; Zhang, J.; Xu, B.; He, S. Evaluation of Figleaf Gourd and White-Seeded Pumpkin Genotypes as Promising Rootstocks for Cucumber Grafting. Horticulturae 2025, 11, 778. https://doi.org/10.3390/horticulturae11070778

AMA Style

Li G, Zou J, Gong T, Li X, Meng J, Zhang J, Xu B, He S. Evaluation of Figleaf Gourd and White-Seeded Pumpkin Genotypes as Promising Rootstocks for Cucumber Grafting. Horticulturae. 2025; 11(7):778. https://doi.org/10.3390/horticulturae11070778

Chicago/Turabian Style

Li, Gengyun, Jiamei Zou, Tianrui Gong, Xuejiao Li, Jing Meng, Jie Zhang, Bin Xu, and Shuilian He. 2025. "Evaluation of Figleaf Gourd and White-Seeded Pumpkin Genotypes as Promising Rootstocks for Cucumber Grafting" Horticulturae 11, no. 7: 778. https://doi.org/10.3390/horticulturae11070778

APA Style

Li, G., Zou, J., Gong, T., Li, X., Meng, J., Zhang, J., Xu, B., & He, S. (2025). Evaluation of Figleaf Gourd and White-Seeded Pumpkin Genotypes as Promising Rootstocks for Cucumber Grafting. Horticulturae, 11(7), 778. https://doi.org/10.3390/horticulturae11070778

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