ABSTRACT
While drought negatively affects plant production, vermicompost and grafted seedlings offer a promising strategy to reduce these effects. This study, conducted in Ankara (greenhouse) and Antalya (field) provinces in 2020–2021, investigated the effects of grafting and vermicompost on morphophysiological traits, yield, and water efficiency of eggplant under drought stress. The greenhouse experiment was conducted in a factorial design (n = 3), and the field experiment was conducted using a split-split plot design (n = 3). In the greenhouse, grafted and non-grafted plants were subjected to three irrigation levels—100%, 70%: moderate stress (MS), 30%: severe stress (SS)—and vermicompost doses (0, 1, 2, 3%). Under MS conditions, grafted plants supplemented with 2% vermicompost showed significant improvements: shoot fresh weight increased by 48.81%, dry weight by 42.92%, shoot length by 49.52%, and stomatal conductance by 88.89%, while yield losses were reduced by 96% and irrigation water productivity improved by 62.79%. Field experiments applied the same irrigation levels (SS = 0.30 Epan, MS = 0.70 Epan, control = 1.00 Epan), and the optimal vermicompost dose (2%) identified in the greenhouse trials. Under MS, shoot fresh and dry weights increased by 121.20 and 162.18%, respectively, with total productivity rising by 101.13%. The innovative contribution of this study lies in its synergistic integration of grafting and vermicompost, two sustainable practices previously untested in eggplant cultivation to mitigate drought effects across both fields. The results demonstrated that this strategy reduces yield losses by 96% under severe stress and doubles productivity under moderate drought, offering a sustainable and environmentally friendly solution for water-limited agriculture.
Key words
Solanum melongena L.; water stress; grafted plant; organic material; water productivity
INTRODUCTION
Drought is a major abiotic stressor that significantly reduces crop productivity, including in eggplant. As is the case with many other cultivated crops (for example, corn, wheat, and tomatoes), insufficient rainfall and irrigation have been demonstrated to result in considerable yield and quality losses in eggplant cultivation (Cantürk et al. 2023). Although eggplant exhibits moderate drought tolerance, its genetic diversity leads to variable tolerance among cultivars (Díaz-Pérez and Eaton 2015). While wild relatives offer important genetic resources for drought tolerance (Plazas et al. 2022, Gramazio et al. 2023), breeding of drought-tolerant cultivars continues to be constrained by the polygenic nature of stress tolerance and significant costs (Toppino et al. 2022). High variation in yield and biotic stress tolerance has also been reported among genotypes (Goutam et al. 2025). Grafting onto drought-tolerant rootstocks has emerged as a practical solution that increases water productivity, nutrient uptake, and physiological tolerance (Schwarz et al. 2010, Kıran et al. 2017, Kıran et al. 2018, Zhang et al. 2020). However, its efficacy decreases under prolonged or severe drought and requires complementary strategies.
Combining grafting with bio-organic soil amendments such as vermicompost provides a novel approach to enhance plant resilience in water deficient conditions. Vermicompost is a natural soil conditioner obtained by decomposing organic waste with the help of worms. The high microbial activity, nutrients, and phytohormones in it promote plant growth. It also enhances plant growth by improving soil structure, water retention, and nutrient availability (Demir 2019, Wang et al. 2021), and strengthens drought tolerance by increasing shoot growth, stomatal conductance, and photosynthetic efficiency under drought conditions (Alharbi et al. 2023, Naeemi Golzard et al. 2023, Ahmad et al. 2024). When integrating with grafting, these treatments synergistically target both plant physiology (e.g., non-enzymatic/enzymatic antioxidant systems, root hydraulic conductance) and soil health (e.g., organic matter, porosity) and holistically address drought-induced limitations. Despite previous studies examining grafting or vermicompost in isolation, their combined effects on eggplant under drought stress are still understudied, and this study addresses this gap.
This study hypothesized that grafting and vermicompost application synergistically improves eggplant growth, physiology, and water productivity under water-limited conditions. The objectives were to evaluate their combined effects on growth and yield, physiological reactions, and soil physicochemical properties in both greenhouse and field environments. This study differentially evaluated scalability and practical applicability by combining controlled experiments with real-world validation. The findings aimed to advance sustainable agricultural practices by combining biological and organic strategies and provide a cost-effective solution for water-scarce regions.
MATERIALS AND METHODS
Research materials
In both stages of the research, ‘Aydın Siyahı’ eggplant variety seedlings, grafted and ungrafted (grown on their own roots) on ‘Köksal’, a commercial eggplant rootstock with high drought and salt tolerance (Kıran et al. 2018, Kıran et al. 2019), were used. Grafted seedlings were obtained by Antalya Tarım Hybrid Seedling Company (Antalya, Türkiye) using the tube grafting technique on seedlings with two true leaves (April 10, 2020). Ekosol Farming & Livestock Company (Manisa, Türkiye) supplied the vermicompost used in the study. Selected properties of the vermicompost used in this study are presented in Kıran et al. (2025).
Greenhouse experiment
Experimental design and planting
The research was conducted in the glass greenhouse of the Soil Fertilizer and Water Resources Central Research Institute (Ankara, Türkiye), where temperature (25–27°C) and humidity (50–55%) control is provided automatically. The research was organized using a factorial experimental design in randomized plots with three replications (n = 3). The experiment comprised three factors:
-
Factor 1: grafting (grafted plants—G, nongrafted—NG);
-
Factor 2: four different levels of vermicompost (V0 = 0%—control, V1 = 1%, V2 = 2%, V3 = 3%);
-
Factor 3: different levels of drought-irrigation level—control, 100% field capacity (FC), moderate stress (MS; 70% FC), severe stress (SS; 30% FC).
In the greenhouse experiment, the soil was passed through a 4-mm sieve and placed equally in 35 L volume and 39 × 35-cm dimensions PE pots. Before planting seedlings, vermicompost in solid form was weighed as 1, 2, and 3% (w/w) of the weight of the soil in the pot and mixed into a 10–15-cm part of the potting soil. The 1, 2, and 3% vermicompost doses used are based on previous studies by Demir (2019) and Kıran (2019), which found positive effects on plants and soil with similar dosages. Fifteen days after the vermicompost application, grafted and ungrafted seedlings with four or five true leaves were planted in the pots (May 11, 2020). Based on the soil analysis results (Kıran et al. 2025), the fertilizer requirement of the plants was fulfilled with 100 kg P2O5·ha-1 and 70 kg N·ha-1.
Drought stress treatment
All pots were irrigated until the stress treatments were started to ensure that the available moisture was brought to FC. After the plants had developed for two weeks under these conditions, drought stress applications were started (May 25, 2020). To determine the FC (pot capacity), after weighing two randomly selected pots from each subject, they were placed in separate basins, watered from above, and allowed to reach the saturation point. The pots were covered and left for 48 hours. After it was confirmed that no drainage was coming out of the bottoms of the pots, they were weighed again. The measured weights constituted the pots’ FC (pot capacity) values. During the trial period, water was given to the pots belonging to the control (100% irrigation) subject until the FC was reached. For drought stress, pots for moderate stress were given 70% of the water provided to control pots and 30% for severe stress. At the end of 98 days of stress treatments (September 1, 2020), all plants were examined for morphophysiological characteristics.
Open experiment
Experimental area and growth conditions
The current research was conducted at the study field (at a latitude of 36’96”N, a longitude of 30’81”E, and an altitude of 45 m) of the Antalya Tarım Hybrid Seeds Company in Antalya, Türkiye. The climatic parameters for the current study year (2021), long-term means (1954–2020) for April-August and the chemical and physical parameters of experimental soil and applied irrigation water are given in Kıran et al. (2025).
Statistical design, treatments, and planting
The current study arranged three replications (n = 3) according to the experimental design of the split-split plot. Eggplant plants (non-grafted and grafted), two vermicompost levels (non-application—0%—and 2%), and three irrigation levels (SS = 0.30 Epan; MS = 0.70 Epan; control = 1.00 Epan) were examined in the study. The ‘Aydın Siyahı’ eggplant variety was used as the plant material, either non-grafted or grafted onto the Köksal rootstock. The most effective 2% vermicompost dose obtained in the greenhouse experiment was applied on the row in the relevant plots according to the experimental design and mixed to a depth of 0.15 m of the soil depth (April 5, 2021). Irrigation applications were carried out to take the evaporation data (Epan, mm) obtained from a class A pan placed next to the study plots (Doorenbos and Pruitt 1977). The class A pan data were recorded at seven-day intervals. Fertilizer applications were based on soil analysis results, and all the plots received the same amount of total fertilizer. The eggplant seedlings were planted on April 20, 2021, with 0.50 × 1.20-m spacing. Each plot was created 5-m wide and 10-m long, and 120 plants were used. There are five plant rows in each experimental plot. A gap of 1 m was left between experimental plots with different irrigation treatments. Deficit irrigation practices lasted 72 days (from May 31 to August 12, 2021).
Irrigation method and management
The surface drip irrigation system was used to irrigate experimental plots. A 16-mm diameter polyethylene (PE) drip line was placed in the plant rows. The drippers have a flow rate of 2 L·h-1 and are spaced 40 cm apart. The amount of water applied to the control = 1.00 Epan was calculated with the help of Eq. 1 by using the evaporation values obtained by reading from the class A pan at one-week intervals (Tarı and Sapmaz 2017).
where: I: applied irrigation water (L); A: experimental area (m2); Epan: pan evaporation at seven-day intervals (mm); Kpc: plant-pan coefficient ranging from 0.30 to 1.00 for irrigation treatments (KcpControl = 100% of pan evaporation, KcpMS = 70% of pan evaporation, KcpSS = 30% of pan evaporation, Kcp0.50 = 50% of pan evaporation); P: percentage of the wetted area (%).
The percentage of the wetted area was taken as 35% since the percent cover was below 35% when the first irrigation was started. When the percent cover values exceeded 35%, the actual measured values were used (Keller and Bliesner 1990). Percent cover values were calculated from measurements on the same five pre-selected plants before both irrigations (Çetin and Üzen 2018). For this, plant crown development was divided by the planting spacing (plant row spacing, 1.20 m).
Evapotranspiration, water productivity, irrigation water productivity, and yield response factor
Evapotranspiration (ET) was determined using the soil water balance method described by Doorenbos and Pruitt (1977) (Eq. 2).
where: ET: evapotranspiration (mm); I: applied irrigation water (mm); R: rainfall (mm); D: deep percolation (mm); ΔW: change in soil water storage (mm).
Since the amount of irrigation water was controlled, deep percolation was assumed to be negligible.
Water productivity and irrigation water productivity were calculated using the Eqs. 3 and 4, given by Howell (2001).
where: WP: water productivity (kg·m-3), IWP: irrigation water productivity (kg·m-3); ET: evapotranspiration (mm); I: applied irrigation water (mm), Y: yield (kg·ha-1).
The yield response factor (Ky) was determined in the study by Stewart et al. (1977) and represents the effect of a decrease in crop water use on yield loss (Eq. 5).
where: Yx: the maximum yield; Ya: the actual yield; ETx: the maximum ET; ETa: the actual ET; Ky: the yield response factor.
Morphological and physiological measurements
The shoots of the harvested plants were weighed on a 1/1,000 precision balance to record their fresh weight and dried at 65°C to determine their dry weight. The shoot length of the plants was measured from the root collar to the apical part of the plant. The shoot diameter was measured with a caliper, and the mean shoot diameter was determined by averaging the measurements taken from a point just above the point of grafting, half the height of the plant, and the uppermost terminal node. Leaf area was measured on the same leaf (third and fourth) of each plant and determined using Licor LI-3000A. Chlorophyll SPAD measurements were made on the third leaf of each plant using a Minolta SPAD-502 instrument. Stomatal conductance (gs) was measured on the fifth leaf from the top of each plant between 1 and 2 p.m. using a model SC-1 porometer, and readings were taken three times. Leaf relative water content was determined from fresh leaf samples. Fresh weight (FW) was determined by weighing the leaf sections; turgor weight (TW) was recorded after soaking in pure water for 4 hours and weighing; dry weight (DW) was recorded after drying at 60°C for 24 hours. The relative water content (RWC) was calculated using Eq. 6:
Cumulative yield was calculated by weighing the fruit between the first and last harvest, and total yield (kg·ha-1) was obtained.
Soil analysis
Soil pH values were measured with a pH meter, and electrical conductivity (EC) values were measured with an EC meter (Richards 1954). Scheibler calcimeter was used to determine soil lime contents (Tüzüner 1990). The modified Walkley-Black method was used to determine soil organic matter (SOM) content (Rowell 1996). Total N was determined by the Kjeldahl method (Rowell 1996). Soil available P, K, Ca, and Mg contents were determined through extraction with 0.5 M NaHCO3 (pH = 8.5) (Rowell 1996). Cation exchange capacity (CEC) values were determined as described by Bower et al. (1952). Micronutrients (Fe, Cu, Zn, Mn) were determined in a spectrophotometer using DTPA extraction (Rowell 1996). Extractable Na, K, Ca, and Mg were determined with 1 N NH4OAc according to Richards (1954). Soluble anions and cations were determined according to Richards (1954).
Statistical analysis
Statistical Package for the Social Sciences 11.0 software was used for data analysis. The normality of the data was tested using the Shapiro-Wilk test (p > 0.05), and the results indicated that the data followed a normal distribution. The homogeneity of variance was tested using Levene’s test (p > 0.05), which showed that the variances were homogeneous. The data obtained from analysis of variance (ANOVA) were evaluated using Duncan’s multiple comparison test (p < 0.05). Duncan’s multiple comparison test (p < 0.05) was applied to assess treatment differences. To visualize correlations between the measured variables, OriginPro 2024 software (OriginLab Corporation, United States of America) was used to generate a polar heatmap for hierarchical clustering.
RESULTS
Greenhouse experiment
Changes in plant morpho-physiological characteristics
Under drought stress conditions, grafted plants grown in vermicompost pots showed better growth than those that were not grafted and without vermicompost (Fig. 1). It was observed that as the severity of drought stress increased, the values of growth characteristics gradually decreased and reached the lowest values in the SS environment. The statistical significance of the findings of all examined parameters was considered and interpreted as the interaction of ‘grafted (G) × vermicompost (V) × drought stress (DS)’. Accordingly, the effect of the ‘G × V × DS’ interaction was found to be statistically significant on shoot fresh (p < 0.01), dry weights (p < 0.05), and shoot length (p < 0.01) (Table 1). It was observed that plants under MS and SS were in the same group in terms of growth characteristics. In terms of shoot fresh biomass, the combinations of ‘G × V2 × MS’, ‘G × V3 × MS’ (48.81 and 45.39%), and ‘G × V2 × SS’, ‘G × V3 × SS’ (180.25 and 176.54%) gave the highest values with the lowest losses compared to the control in MS and SS. In dry biomass, ‘G × V2 × MS’, ‘G × V3 × MS’ (42.92 and 39.43%), ‘G × V2 × SS’, and ‘G × V3 × SS’ (62.05 and 53.19%) stood out. In shoot length, the lowest reduction rates were achieved in the interactions of ‘G × V3 × MS’, ‘G × V2 × MS’ (49.52 and 42.86%), and ‘G × V3 × SS’, ‘G × V2 × SS’ (48.21 and 44.64%) (Table 2). The impacts of three factors on gs and relative leaf water content, which are considered important physiological indicators in determining the tolerance of plants to drought stress, were found to be significant (p < 0.01) (Table 1). It was observed that gs gradually decreased as the severity of drought stress increased, but it was noted that the interaction of vermicompost and grafting reduced gs losses. Combinations ‘G × V2 × MS’, ‘G × V3 × MS’ (88.89 and 82.33%) and ‘G × V2 × MS’, ‘G × V3 × SS’ (116.05 and 91.67%) had the least decrease rates in MS and SS environments (Table 2). Statistical analysis showed that the effect of the triple combination for RWC was similar to gs. The combination of factors yielded the lowest losses at 13.21 and 9.08%, as well as 12.72 and 0.22%. These outcomes were associated with the interactions denoted as ‘G × V2 × MS’, ‘G × V3 × MS’, and ‘G × V2 × SS’.
Morphological characteristics of grafted and non-grafted eggplant plants subjected to different levels of vermicompost under moderate and severe drought stress in the greenhouse.
Variance analysis results of the investigation of the characteristics of eggplant grown in greenhouse conditions.
Effects of applications on irrigation water productivity
The irrigation water productivity (IWP) was determined by considering the irrigation water amounts and eggplant yields of the trial subjects, and the variance analysis table for the yield is given in Table 1. According to these results, the effect of the ‘G × V × DS’ interaction on yield and IWP was significant (p < 0.05). The highest efficiency values with the lowest loss rates in MS environments were in the same statistical group and appeared in the interactions of ‘G × V2 × MS’ and ‘G × V3 × MS’ (96 and 91.37%, 44,010 and 43,130 kg·ha-1). The findings showed that the use of grafted seedlings and vermicompost application significantly increased the IWP values of plants under drought stress. These increases were found to be most effective, especially at high doses of vermicompost.
Effects of applications on soil properties
The results of the ANOVA between the experimental subjects and the chemical properties of soils under greenhouse conditions are given in Table 3, and the Duncan grouping is given in Table 4. The effects on the chemical properties of soils were statistically insignificant in the triple interaction ‘G × V × DS’ (p > 0.05). The impact of ‘V × DS’ on the chemical properties of soils, except CEC and available Mn values, was found significant (p < 0.05). Under drought stress, the highest combination in terms of organic matter was ‘V3 × control’ (0.92%), and the lowest combination was ‘V1 × SS’ (0.37%). The effects of vermicompost, grafting, irrigation levels, and ‘V × DS’ interactions on available P2O5 and Ca concentrations were significant. The highest available P2O5 value was obtained from the ‘V3 × control’ interaction (4.55 kg·da-1), while the lowest available P2O5 value was determined in the ‘V1 × SS’ interaction (0.92 kg·da-1).
Field experiment
Effects of applications on plant morpho-physiological characteristics
Field evaluations confirmed that combining 2% vermicompost with grafting under MS and SS conditions yielded statistically comparable results to greenhouse findings (Fig. 2). The findings showed that the ‘G × V × DS’ interaction provided significant increases in shoot fresh and dry weights and plant height (p < 0.01) (Table 5). These findings were parallel to the findings obtained under greenhouse conditions. The interaction effect appeared more clearly in plants grown under SS. The highest increase rates for shoot fresh-dry weights and plant height were observed in ‘G × V2 × MS’, ‘G × V2 × SS’ (121.20 and 76.28%). ‘G × V2 × MS’, ‘G × V2 × SS’ (162.18 and 88.28%) had the interactions ‘G × V2 × SS’, ‘G × V2 × MS’ (42.72 and 40.83%), respectively (Table 5).
Effects of applications on water productivity and irrigation water productivity
In terms of yield, the interaction of the three factors was statistically significant (p < 0.01). Variance analysis results of yields are given in Table 5. Decreasing irrigation water amounts caused significant losses in yield. The combined use of grafted plants and vermicompost was found to be more effective in reducing yield losses under SS. The lowest yield loss was encountered in ‘G × V2 × MS’ (104.13%), while the highest yield value was reached (5,641.7 kg·ha-1) (Table 5). However, the interaction of the three factors caused significant differences (p < 0.01) for WP calculated considering yield and plant water consumption, and IWP calculated in the greenhouse environment (Table 5). WP and IWP had significant increase rates, especially in the SS environment and the interactions of ‘G × V2 × SS’, ‘G × V2 × MS’, with 112.20 and 104.76% for WP; and ‘G × V2 × SS’, ‘G × V2 × MS’, with 107.14 and 101.35% for IWP.
Grafted and non-grafted eggplant plants grown under different drought levels and with or without vermicompost in soil under field conditions.
Analysis of variance and the effect of ‘G × V × DS’ interaction on biomass, physiological traits, yield, and irrigation water productivity of eggplant under field conditions***.
Effects of applications on soil properties
The results of ANOVA of chemical characteristics of soils of vermicompost applications under different irrigation levels under field conditions are given in Table 6, and Duncan grouping is given in Table 7. Except for pH, CEC, and available Mn, the effect of EC, organic matter, available P2O5, available K2O, available Fe, Cu, and Zn on vermicompost × irrigation levels interaction was found statistically significant (p < 0.01). In this study, except pH, EC, CEC, P, K, organic matter, Ca and Mg values increased with vermicompost application. The highest electrical conductivity value of the soils was determined as 1.688 dS m-1 in the ‘V2 × SS’ interaction, and the lowest EC value was determined as 0.557 dS m-1 in the ‘V0 × control’ interaction. The highest organic matter content was determined as 1.82% in the ‘V2 × control’ interaction, while the lowest organic matter content was determined as 0.48% in the ‘V0 × SS’ interaction.
Effects of ‘V × DS’ interaction on soil chemical properties, effects of vermicompost on available Mn, and effects drought stress on cation exchange capacity*.
Correlation matrix
Strong positive correlations were observed between morpho-physiological traits, chlorophyll content, gs, and RWC under both greenhouse and field conditions (Figs. 3a and 3b). Growth traits under greenhouse conditions were observed to be positively correlated with chlorophyll, gs, and RWC. Yield also showed a strong positive correlation with all morphophysiological traits examined. However, IWP had positive but relatively low correlations with growth and physiological parameters (except leaf area, root fresh weight, and relative water content). Growth parameters under field conditions showed positive and strong correlations with yield. Strong correlations were observed between yield and shoot fresh weight, shoot dry weight, and shoot diameter, while IWP and WP did not show any correlation with growth traits. However, positive correlations were determined between WP with yield and IWP.
Correlation analysis of drought stress, grafting, and vermicompost treatments on the studied properties of eggplant. (a) Greenhouse conditions; (b) open field conditions. The correlation matrix illustrates the significance of relationships using asterisks: *p < 0.05.
Polar heatmap with dendrogram
More differences were observed in grafted and vermicomposted eggplant plants grown in the greenhouse in terms of growth parameters, physiological parameters, and yield under severe and moderate drought stress (Fig. 4a). Dendrogram showed indirect clustering between plant growth parameters and physiological parameters, and yield. Also, direct clustering was observed between IWP and root dry weight. Differences were also observed between grafted and vermicompost treatments in terms of the investigated traits under stress in field-grown eggplants (Fig. 4b). While an indirect clustering occurred between growth characteristics and yield and chlorophyll, a direct clustering was observed between IWP and WP.
Polar heatmap for the effects of drought stress, grafting and vermicompost treatments on the studied traits of eggplants. (a) Greenhouse conditions; (b) open field conditions.
DISCUSSION
Drought stress, one of the major suppressions of physiological functions obstacle to agricultural sustainability, causing yield losses, morphological deterioration, and the suppression of physiological functions, particularly in water-sensitive plants such as eggplant (Solanum melongena). This study demonstrated that combining grafting and vermicompost applications enhances drought resilience in eggplant under both greenhouse and field conditions. These treatments improved WP, photosynthesis, and defense mechanisms, thereby promoting tolerance under both moderate and severe drought stress.
The findings on the morphological and physiological characteristics, as well as the yield of eggplant plants in greenhouse and field conditions, showed that the combined application of grafting and vermicompost significantly mitigates the adverse effects of moderate and severe drought. These findings underscore the importance of grafting onto suitable rootstocks and also the use of vermicompost in the improvement of eggplant plants against drought-induced stress. The synergistic effect of these two approaches was evident not only in the morphological characteristics of the plants (e.g., fresh and dry shoot weight, shoot length) but also in physiological traits such as root structure, hydraulic conductivity, and nutrient uptake capacity. Grafting onto suitable rootstocks and vermicompost application together enhanced shoot biomass, shoot length, and water productivity. Improved root architecture, increased hydraulic conductivity, and enhanced nutrient uptake were among the key physiological benefits. Cytokinin synthesis, promoted by grafting, likely supported photosynthesis and growth even under stress (Groppa et al. 2012, Li et al. 2020, Jin et al. 2023).
The effects of this combination were further supported by the Köksal rootstock, which had previously been identified as effective under drought stress conditions (Kıran et al. 2018, Kıran et al. 2019). In comparative trials involving various eggplant rootstocks, Köksal-F1 demonstrated superior performance in terms of morphophysiological traits and yield. Therefore, it was selected for grafting in this study to better observe the physiological and yield responses when combined with vermicompost under drought conditions. However, reliance on a single rootstock may limit the generalizability of the findings; future studies should include genetically diverse rootstocks to enhance broader applicability. The Köksal rootstock enhanced water and nutrient uptake due to its vigorous root system and high root biomass. Simultaneously, the grafted scion benefited from this robust root system, reducing the plant’s overall water demand and increasing WP (Argento et al. 2023).
The use of compatible rootstocks enhanced growth by increasing cytokinin synthesis and facilitating the transport of metabolites, hormones, and signaling molecules, thereby promoting cell proliferation and stress-resilient development (Sorce et al. 2002, Albacete et al. 2015).
In contrast, vermicompost application is distinguished by its direct impact on improving soil conditions. Vermicompost increased soil fertility by increasing the availability of macronutrients such as P, K, Ca, and Mg; improved nutrient uptake by plants by accelerating the mineralization processes of various microorganism species in the compost; and consequently improved the physiological performance of plants (Demir and Kıran 2020; Feizabadi et al. 2021, Benaffari et al. 2022). Additionally, vermicompost improved macro- and micronutrient availability and soil water-holding capacity, creating a favorable rhizosphere for root growth under low water potential. This effect, coupled with the robust root structure induced by grafting application, contributed to the increased root biomass in root biomass of grafted plants.
Increased root volume in grafted plants likely enhanced the secretion of organic acids and amino acids, supporting rhizobacterial functions such as nitrogen fixation, nutrient mobilization, and hormone production (Ahemad and Kibret 2014). This symbiotic relationship, through the combined effect of inoculum and vermicompost applications, contributed to promoted plant growth, regulate metabolic processes, and ultimately increase yield under drought stress. Beneficial microorganisms in vermicompost extracts, biofertilizers, and biocontrol agents—rich in nutrients and vitamins—can produce subtilin and phytohormones such as auxin, gibberellin, cytokinin, and kinetin, thereby synergistically supporting plant growth during vermicompost formation (Sulaiman and Mohamad 2020, Rehman et al. 2023, Farfour et al. 2025). These processes may have favored the growth and development of grafted plants under drought stress.
Within the scope of physiological and biochemical processes triggered by vermicompost, the possible accumulation of certain osmolytes (such as proline and sugars), enzymatic antioxidants (superoxide dismutase, catalase, glutathione reductase), and non-enzymatic antioxidants (phenolics, glutathione, ascorbic acid, etc.) may have contributed to the plant’s capacity to cope with oxidative stress under limited water conditions (Kıran 2019, Benazzouk et al. 2020). These compounds likely supported the maintenance of cell turgor by increasing intracellular water retention, thereby enhancing leaf RWC and facilitating stomatal opening. Increased CO2 diffusion into stomatal guard cells with preserved turgor promoted gas exchange and led to higher gs, which may have indirectly contributed to increased photosynthetic rates and improved plant growth. Moreover, grafting-induced improvements in root system architecture (e.g., root length, density, and number of root hairs) enhanced water uptake from soil layers with low water potential. Simultaneously, increased potassium (K+) uptake in grafted plants may have supported stomatal opening and maintained continuous gas exchange. Previous studies have reported a positive relationship between K+ levels and gs, with potassium known to enhance photosynthetic performance by maintaining osmotic balance (Hassan et al. 2017, Miranda et al. 2021). Vermicompost’s porosity and water-holding capacity contributed to soil moisture retention and sustained gas exchange (Castellini et al. 2024). Consequently, CO2 uptake continued, potentially increasing photosynthetic carbon assimilation and thereby supporting plant development.
Despite limited water availability, the enhanced growth and development observed in vermicompost-treated grafted plants translated into increased yield, consistently across both greenhouse and field conditions. This effect was largely attributed to the improved morphophysiology of grafted plants and the positive impact of vermicompost on soil properties, including increased organic matter, CEC, and water retention. Although vermicompost slightly increased soil EC under drought, values remained below salinity thresholds and did not adversely affect yield. Additionally, vermicompost raised soil pH, enhancing nutrient availability. Notably, it compensated for drought-induced phosphorus depletion by enriching plant-available phosphorus in the soil, as it is a phosphorus-rich amendment owing to its phosphorus-rich composition and high bioavailability (Nieto-Cantero et al. 2025). WP and IWP increased due to enhanced soil properties and water retention. Polysaccharides secreted by microbes may have improved soil structure and fine particle content (Habib et al. 2016).
CONCLUSION
This study revealed that vermicompost application combined with grafting significantly enhanced drought tolerance in eggplant under both greenhouse and field conditions. The integration of these treatments increased the physiological tolerance of eggplant plants under moderate and severe drought conditions, resulting in improvements in traits such as water productivity and growth performance. This combination increased yields under moderate and severe stress conditions, resulting in an average yield increase of 100% in the field and 122% in the greenhouse. Vermicomposting enriched soil fertility by increasing nutrient availability, organic matter, and porosity, while grafting increased root productivity and optimized water use. Together, these approaches not only maintained productivity with insufficient irrigation but also highlighted a sustainable and environmentally friendly strategy for water-deficient agricultural systems. This study evaluated the efficacy of vermicompost combined with grafted plants under drought conditions in both greenhouse and single-season field environments. However, multi-year and multi-location trials involving a broader range of vermicompost doses and cost-benefit analyses are required to determine economically viable application rates and to assess the broader potential of this strategy under various abiotic stresses. Furthermore, integrating transcriptomic and proteomic approaches will be essential for a more comprehensive understanding of the underlying molecular mechanisms.
ACKNOWLEDGMENTS
We thank Antalya Tarım Hybrid Seeds Co. for its support in the second phase of the trial and for the additional financial assistance.
-
How to cite:
Kiran, S., Demir, Z., Boyacı, H. F., Aydınşakir, K., Kuşvuran, S., Zengin, S. and Ellialtıoğlu, S. S. (2025). Improving drought tolerance in eggplant through grafting and vermicompost: yield and water productivity. Bragantia, 84, e20250034. https://doi.org/10.1590/1678-4499.20250034
-
FUNDING
General Directorate of Agricultural Research and PoliciesGrant No.: TAGEM/TSKAD/B/19/A9/P3/888
DATA AVAILABILITY STATEMENT
Supporting datasets have been archived in Zenodo and will become publicly accessible upon publication (https://doi.org/10.5281/zenodo.17190590).
REFERENCES
-
Ahemad, M. and Kibret, M. (2014). Mechanisms and applications of plant growth promoting rhizobacteria: Current perspective. Journal of King Saud University — Science, 26, 1-20. https://doi.org/10.1016/j.jksus.2013.05.001
» https://doi.org/10.1016/j.jksus.2013.05.001 -
Ahmad, A., Aslam, Z., Abbas, N. R., Bellitürk, K., Hussain, S., Hussain, S., Ahmad, M., Zulfiqar, U., Moussa, I. M. and Elshikh, M. S. (2024). Enhancing wheat crop resilience to drought stress through cellulolytic microbe-enriched cow dung vermicompost. ACS Omega, 9, 2123-2133. https://doi.org/10.1021/acsomega.3c04402
» https://doi.org/10.1021/acsomega.3c04402 -
Albacete, A., Martinez-Andujar, C., Martinez-Perez, A., Thompson, A., Dodd, I. C. and Perez-Alfocea, F. (2015). Unravelling rootstock × scion interactions to improve food security. Journal of Experimental Botany, 66, 2211-2226. https://doi.org/10.1093/jxb/erv027
» https://doi.org/10.1093/jxb/erv027 -
Alharbi, K., Hafez, E. M., Omara, A. E. D. and Osman, H. S. (2023). Mitigating osmotic stress and enhancing developmental productivity processes in cotton through integrative use of vermicompost and cyanobacteria. Plants, 12, 1872. https://doi.org/10.3390/plants12091872
» https://doi.org/10.3390/plants12091872 -
Argento, S., Treccarichi, S., Arena, D., Rizzo, G. F. and Branca, F. (2023). Exploitation of a grafting technique for improving the water use efficiency of eggplant (Solanum melongena L.) grown in a cold greenhouse in Mediterranean climatic conditions. Agronomy, 13, 2705. https://doi.org/10.3390/agronomy13112705
» https://doi.org/10.3390/agronomy13112705 -
Benaffari, W., Boutasknit, A., Anli, M., Ait-El-Mokhtar, M., Ait-Rahou, Y., Raja Ben-Bem-Laouane, R., Ahmed, H. B., Mitsui, T., Baslam, M. and Meddich, A. (2022). The native arbuscular mycorrhizal fungi and vermicompost-based organic amendments enhance soil fertility, growth performance, and the drought stress tolerance of quinoa. Plants, 11, 393. https://doi.org/10.3390/plants11030393
» https://doi.org/10.3390/plants11030393 -
Benazzouk, S., Dobrev, P. I., Djazouli, Z. E., Motyka, V. and Lutts, S. (2020). Positive impact of vermicompost leachate on salt stress tolerance in tomato (Solanum lycopersicum L.) at the seedling stage: A phytohormonal approach. Plant and Soil, 446, 145-162. https://doi.org/10.1007/s11104-019-04361-x
» https://doi.org/10.1007/s11104-019-04361-x - Bower, C. A., Reitemeier, R. F. and Fireman, M. (1952). Exchangeable cation analysis of saline and alkali soils. Soil Science, 73, 251-261.
-
Cantürk, A., Cemek, B., Taşan, M. and Taşan, S. (2023). Effect of deficit irrigation on yield, water productivity, energy indices and economic productivity in eggplant cultivation. Gesunde Pflanzen, 75, 1579-1589. https://doi.org/10.1007/s10343-022-00814-z
» https://doi.org/10.1007/s10343-022-00814-z -
Castellini, M., Bondì, C., Giglio, L. and Iovino, M. (2024). Impact of vermicompost addition on water availability of differently textured soils. Heliyon, 10, 15. https://doi.org/10.1016/j.heliyon.2024.e35699
» https://doi.org/10.1016/j.heliyon.2024.e35699 -
Çetin, Ö. and Üzen, N. (2018). Effects of surface and subsurface drip irrigation on soil water moisture variation and soil-water tension. Harran Journal of Agricultural and Food Sciences, 22, 461-470. https://doi.org/10.29050/harranziraat.442314
» https://doi.org/10.29050/harranziraat.442314 -
Demir, Z. (2019). Effects of vermicompost on soil physicochemical properties and lettuce (Lactuca Sativa var. crispa) yield in greenhouse under different soil water regimes. Communications in Soil Science and Plant Analysis, 50, 2151-2168. https://doi.org/10.1080/00103624.2019.1654508
» https://doi.org/10.1080/00103624.2019.1654508 -
Demir, Z. and Kıran, S. (2020). Effect of vermicompost on macro and micro nutrients of lettuce (Lactuca Sativa var. crispa) under salt stress conditions. Kahramanmaraş Sütçü İmam University (KSU) Journal of Agricultural and Natural Sciences, 23, 33-43. https://doi.org/10.18016/ksutarimdoga.vi.579695
» https://doi.org/10.18016/ksutarimdoga.vi.579695 -
Díaz-Pérez, J. C. and Eaton, T. E. (2015). Eggplant (Solanum melongena L.) plant growth and fruit yield as affected by drip irrigation rate. HortScience, 50, 1709-1714. https://doi.org/10.21273/HORTSCI.50.11.1709
» https://doi.org/10.21273/HORTSCI.50.11.1709 - Doorenbos, J. and Pruitt, W. O. (1977). Guidelines for predicting crop water requirements. FAO Irrigation and Drainage Paper, 24, 107 p.
-
Farfour, S. A., Mohamed, M. A., Mousa, I. E. and Salem, M. F. (2025). Biological control of charcoal rot caused by macrophomina phaseolina through the suppressive role of bioactive vermicompost compared with chemical control. Journal of Current Veterinary Research, 7, 216-233. https://doi.org/10.21608/jcvr.2025.421857
» https://doi.org/10.21608/jcvr.2025.421857 -
Feizabadi, A., Noormohammadi, G. and Fatehi, F. (2021). Changes in growth, physiology, and fatty acid profile of rapeseed cultivars treated with vermicompost under drought stress. Journal of Soil Science and Plant Nutrition, 21, 200-208. https://doi.org/10.1007/s42729-020-00353-4
» https://doi.org/10.1007/s42729-020-00353-4 -
Goutam, E., Tripathi, V., Kumar, A., Bharti, Mehandi, S., Kumar, L. and Kumar, A. (2025). Assessment of genetic variability in eggplant (Solanum melongena L.) genotypes for disease resistance and yield parameters. Plant Science Today, 12, 1-7. https://doi.org/10.14719/pst.4818
» https://doi.org/10.14719/pst.4818 -
Gramazio, P., Alonso, D., Arrones, A., Villanueva, G., Plazas, M., Toppino, L., Barchi, L., Portis, E., Ferrante, P., Lanteri, S., Rotino, G. L., Giuliano, G., Vilanova, S. and Prohens, J. (2023). Conventional and new genetic resources for an eggplant breeding revolution. Journal of Experimental Botany, 74, 6285-6305. https://doi.org/10.1093/jxb/erad260
» https://doi.org/10.1093/jxb/erad260 -
Groppa, M. D., Benavides, M. P. and Zawoznik, M. S. (2012). Root hydraulic conductance, aquaporins and plant growth promoting microorganisms: A revision. Applied Soil Ecology, 61, 247-254. https://doi.org/10.1016/j.apsoil.2011.11.013
» https://doi.org/10.1016/j.apsoil.2011.11.013 -
Habib, S. H., Kausar, H. and Saud, H. M. (2016). Plant growth-promoting rhizobacteria enhance salinity stress tolerance in okra through ROS-scavenging enzymes. BioMed Research International, 6284547. https://doi.org/10.1155/2016/6284547
» https://doi.org/10.1155/2016/6284547 -
Hassan, M. U., Aamer, M., Chattha, M. U., Ullah, M. A., Sulaman, S., Nawaz, M., Zhigiang, W., Yangin, M. and Guogin, H. (2017). The role of potassium in plants under drought stress: Mini review. Journal of Basic and Applied Sciences, 13, 268-271. https://doi.org/10.6000/1927-5129.2017.13.44
» https://doi.org/10.6000/1927-5129.2017.13.44 -
Howell, T. A. (2001). Enhancing water use efficiency in irrigated agriculture. Agronomy Journal, 93, 281-289. https://doi.org/10.2134/agronj2001.932281x
» https://doi.org/10.2134/agronj2001.932281x -
Jin, M., Liu, Y., Shi, B. and Yuan, H. (2023). Exogenous IAA improves the seedling growth of Syringa villosa via regulating the endogenous hormones and enhancing the photosynthesis. Scientia Horticulturae, 308, 111585. https://doi.org/10.1016/j.scienta.2022.111585
» https://doi.org/10.1016/j.scienta.2022.111585 - Keller, J. and Bliesner, R. D. (1990). Sprinkler and Trickle Irrigation. New York: Van Nostrand Reinhold.
-
Kıran, S. (2019). Effects of vermicompost on some morphological, physiological and biochemical parameters of lettuce (Lactuca sativa var. crispa) under drought stress. Notulae Botanicae Horti Agrobotanici Cluj-Napoca, 47, 352-358. https://doi.org/10.15835/nbha47111260
» https://doi.org/10.15835/nbha47111260 -
Kıran, S., Ateş, Ç., Kuşvuran, Ş., Talhouni, M. and Ellialtioglu, Ş. Ş. (2019). Antioxidative response of grafted and non-grafted eggplant seedlings under drought and salt stresses. Agrochimica, LXIII, 123-137. https://doi.org/10.12871/00021857201922
» https://doi.org/10.12871/00021857201922 - Kıran, S., Kuşvuran, Ş., Çağla, A., and Ellialtioglu, Ş. Ş. (2017). Some physiological properties and analysis of yield parameters of grafted and non-grafted eggplants under waterless conditions. Soil and Water Journal, 6, 18-25.
-
Kıran, S., Kuşvuran, Ş. Ş., Ateş, Ç. and Ellialtioglu, Ş. Ş. (2018). The changes of fruit quality parameters at using of different eggplant rootstock/scion combinations which growing under salt and drought stress. Derim, 35, 111-120. https://doi.org/10.16882/derim.2018.427095
» https://doi.org/10.16882/derim.2018.427095 -
Kıran, S., Zeynep, D., Hatice Filiz, B., Köksal, A., Şebnem, K., Sinan, Z., and Şeküre Şebnem, E. (2025). Suppl. Mat.1, 2, 3.Zenodo. https://doi.org/10.5281/zenodo.17190590
» https://doi.org/10.5281/zenodo.17190590 -
Li, W. J., Mao, J. L., Wu, Y. H., Tang, K., Pan, F. F., Fang, F., Cai, M. Z. and Liu, P. (2020). Effects of IAA on physiological response to aluminum stress and DNA damage of Trichosanthes kirilowii. Ying Yong Sheng Tai Xue Bao, 31, 4235-4242. https://doi.org/10.13287/j.1001-9332.202012.009
» https://doi.org/10.13287/j.1001-9332.202012.009 -
Miranda, M. T., Da Silva, S. F., Silveira, N. M., Pereira, L., Machado, D. C. and Ribeiro, R. V. (2021). Root osmotic adjustment and stomatal control of leaf gas exchange are dependent on citrus rootstocks under water deficit. Journal of Plant Growth Regulation, 40, 11-19. https://doi.org/10.1007/s00344-020-10069-5
» https://doi.org/10.1007/s00344-020-10069-5 - Naeemi Golzard, M., Ghanbari Jahromi, M. and Kalateh Jari, S. (2023). Effect of biochar and vermicompost on growth parameters and physiological characteristics of feverfew (Tanacetum parthenium L.) under drought stress. Journal of Ornamental Plants, 13, 109-120.
-
Nieto-Cantero, J., García-López, A.M., Recena, R., Quntero, J. M. and Degado, A. (2025). Recycling manure as vermicompost: assessing phosphorus fertilizer efficiency and effects on soil health under different soil management. Journal of Soil Science and Plant Nutrition, 25, 5046-5061. https://doi.org/10.1007/s42729-025-02446-4
» https://doi.org/10.1007/s42729-025-02446-4 -
Plazas, M., Gonzalez-Orenga, S., Nguyen, H. T., Morar, I. M., Fita, A., Boscaiu, M., Prohens, J. and Vicente, O. (2022). Growth and antioxidant responses triggered by water stress in wild relatives of eggplant. Scientia Horticulturae, 293, 110685. https://doi.org/10.1016/j.scienta.2021.110685
» https://doi.org/10.1016/j.scienta.2021.110685 -
Rehman, S. U., De Castro, F., Aprile, A., Benedetti, M. and Fanizzi, F. P. (2023). Vermicompost: Enhancing plant growth and combating abiotic and biotic stress. Agronomy, 13, 1134. https://doi.org/10.3390/agronomy13041134
» https://doi.org/10.3390/agronomy13041134 - Richards, L. A. (1954). Diagnosis and ımprovement of saline and alkaline soils. United States: United States Salinity Laboratory Staff, United States Department of Agriculture, 60, 160 p.
- Rowell, D. L. (1996). Soil science: Methods and applications. Harlow: Longman Scientific & Technical, 350 p.
-
Schwarz, D., Rouphael, Y., Colla, G, and Venema, J. H. G. (2010). Grafting as a tool to improve tolerance of vegetables to abiotic stresses: Thermal stress, water stress and organic pollutants. Scientia Horticulturae, 127, 162-171. https://doi.org/10.1016/j.scienta.2010.09.016
» https://doi.org/10.1016/j.scienta.2010.09.016 -
Sorce, C., Massai, R., Picciarelli, P. and Lorenzi, R. (2002). Hormonal relationships in xylem sap of grafted and ungrafted Prunus rootstocks. Scientia Horticulturae, 93, 333-342. https://doi.org/10.1016/S0304-4238(01)00338-7
» https://doi.org/10.1016/S0304-4238(01)00338-7 - Stewart, J. L., Hanks, R. J., Danielson, R. E., Jackson, E. B., Pruitt, W. O., Franklin, W. T. and Hanks, R. J. (1977). Optimizing crop production through control of water and salinity levels in the soil. Logan: Utah Water Research Laboratory Report PRWG151-1, Utah State University.
-
Sulaiman, I. S. C. and Mohamad, A. (2020). The use of vermiwash and vermicompost extract in plant disease and pest control. In Natural remedies for pest, disease and weed control (p. 187-201). Elsevier. https://doi.org/10.1016/B978-0-12-819304-4.00016-6
» https://doi.org/10.1016/B978-0-12-819304-4.00016-6 -
Tarı, A. F. and Sapmaz, M. (2017). The effect of different ırrigation levels on the yield and quality of tomatoes in greenhouse. Soil Water Journal, 6, 11-17. https://doi.org/10.21657/topraksu.339821
» https://doi.org/10.21657/topraksu.339821 -
Toppino, L., Barchi, L. and Rotino, G. L. (2022). Next generation breeding for abiotic stress resistance in eggplant. In C. Kole (Ed.). Genomic designing of climate-smart vegetable crops (p. 115-151). Cham: Springer. https://doi.org/10.1007/978-3-031-03964-5_4
» https://doi.org/10.1007/978-3-031-03964-5_4 - Tüzüner, A. (1990). Soil and water analysis laboratory manual. Ankara: Ministry of Agriculture, Forestryand Rural Affairs, General Directorate of Rural Services.
-
Wang, F., Wang, X. and Song, N. (2021). Biochar and vermicompost improve the soil properties and the yield and quality of cucumber (Cucumis sativus L.) grown in plastic shed soil continuously cropped for different years. Agriculture, Ecosystems & Environment, 315, 107425. https://doi.org/10.1016/j.agee.2021.107425
» https://doi.org/10.1016/j.agee.2021.107425 -
Zhang, Z., Liu, Y., Cao, B., Chen, Z., and Xu, K. (2020). The effectiveness of grafting to improve drought tolerance in tomato. Plant Growth Regulation, 91, 157-167. https://doi.org/10.1007/s10725-020-00596-2
» https://doi.org/10.1007/s10725-020-00596-2
Edited by
-
Section Editor:
Alberto Cargnelutti Filho https://orcid.org/0000-0002-8608-9960
Publication Dates
-
Publication in this collection
03 Nov 2025 -
Date of issue
2025
History
-
Received
08 Feb 2025 -
Accepted
18 Aug 2025





NG: nongrafted plants; V: vermicompost; C: control; G: grafted plants; MS: moderate stress; SS: severe stress.
NG: nongrafted plants; V: vermicompost; C: control; G: grafted plants; MS: moderate stress; SS: severe stress.
SFW: shoot fresh weight; SDW: shoot dry weight; RFW: root fresh weight; RDW: root dry weight; SL: shoot length; SD: shoot diameter; LA: leaf area; gs: stomatal conductance; RWC: relative water content; SPAD: chlorophyll SPAD value; WP: water productivity; IWP: irrigation water productivity.
SFW: shoot fresh weight; SDW: shoot dry weight; RFW: root fresh weight; RDW: root dry weight; SL: shoot length; SD: shoot diameter; LA: leaf area; gs: stomatal conductance; RWC: relative water content; SPAD: chlorophyll SPAD value; WP: water productivity; IWP: irrigation water productivity.