1 Introduction

Sapindus mukorossi is a subtropically and tropically widespread ecologically and economically useful tree species with widespread ecological services that include soil fixation, carbon sequestration, and wildlife support, while its seeds and saponin-yielding fruits are used in soap, cosmetics, and traditional medicine, hugely contributing to the economy at the local level. The dual ecological and economic importance of S. mukorossi focuses on the need to ensure healthy and strong seedling production in order to achieve sustainable industry growth (Liu et al., 2024b).

Seedling production is the foundation of S. mukorossi plantation establishment and long-term productivity. High-quality seedlings with good growth and stress tolerance are essential for achieving stable survival percentages, maximum growth, and sustained yields in plantations. Efficient nursery operations and propagation techniques have direct bearings on the success of reforestation, commercial yield, and ecological rehabilitation programs (Liu et al., 2022).

Despite its adaptability, S. mukorossi seedlings are prone to abiotic and biotic stresses. Drought and salinity may undermine water uptake and growth, low temperature may cause tissue damage and retarded growth, and pest and disease pressure may reduce survival rates and seedling performance. All these problems limit the development of productive plantations and lower general productivity, making stress-resistance research a priority (Zhao et al., 2019).

This study provides comprehensive details on the molecular mechanisms of stress tolerance in S. mukorossi seedlings and associated cultivation practices that optimally enhance seedling performance under stress. Through integrating physiological, molecular, and multi-omics data with realistic cultivation strategies, the study highlights the need to blend basic research with practical methodology and providing paths toward developing stress-tolerant seedlings and environmentally friendly industry operations.

2 Research Status of Stress Tolerance in S. mukorossi Seedlings

2.1 Physiological and biochemical responses related to stress tolerance

Sapindus mukorossi seedlings have a range of physiological and biochemical mechanisms to survive environmental stress conditions such as drought and heavy metal exposure. An increase in the osmotic regulators proline and soluble protein content under water stress conditions maintains cellular water balance. Indicators of membrane stability and stress damage, i.e., malondialdehyde (MDA) and relative electrical conductivity (REC), also rise as water stress increases. The activity of antioxidant enzymes—peroxidase (POD), superoxide dismutase (SOD), and catalase (CAT)—is enhanced in the beginning to resist oxidative damage but decreases in extreme or prolonged stress. The adaptation is controlled to maintain homeostasis and drought tolerance, making S. mukorossi a candidate for afforestation in semi-arid regions. Furthermore, leaf water physiological status is maintained by collaboration between osmoregulation substances and protective enzymes

2.2 Advances and limitations in current research on stress tolerance

Recent studies have enhanced knowledge on S. mukorossi stress responses, particularly towards drought and heavy metal tolerance. Drought stress not only induces physiological defense but also compounds allelopathic effects and can contribute to habitat expansion. S. mukorossi has shown significant tolerance to lead (Pb) and phytoremediation potential with seedlings sustaining growth and Pb accumulation in roots and leaves and no visible toxicity on long durations. Fertilization management has also been marked as critical, with optimal levels of nitrogen, phosphorus, and potassium significantly increasing soil fertility, leaf physiological traits, and production. Overfertilization, however, has detrimental effects on soil health as well as on plant physiology. Studies are still lagging in many aspects: the majority of work addresses drought and heavy metals, fewer address low temperature, salt stress, and biotic stresses. The molecular mechanism for these reactions remains unknown, and field experiments of long term are scarce (Sahito et al., 2023; Zhong et al., 2023).

2.3 Comparative insights from studies on other woody or economic forest seedlings

Comparative studies show that mechanisms of stress tolerance in S. mukorossi, e.g., osmotic adjustment, antioxidant enzyme activity, and allelopathy, are similar to those in other woody and economic tree species. For instance, moderate fertilization enhances physiological characteristics and yield in S. mukorossi and other crops such as blueberries and mung beans, whereas high levels of fertilization may deteriorate development and soil health. Even the tolerance and remediation of heavy metal-contaminated soils occur to other tree species that are fast-growing in nature, which are involved in urban forestry and ecosystem restoration. Convergence shows that insight from general forestry science can be utilized in designing more resilient strategies in the cultivation of S. mukorossi (Liu et al., 2024a).

3 Molecular Basis of Stress Tolerance in S. mukorossi Seedlings

3.1 Signal perception and transduction mechanisms

Recent genomics and transcriptomics in Sapindus mukorossi have identified a number of genes for perception and transduction of stress signals. Candidate genes such as SmPP2C (abscisic acid signaling), SmAHP (cytokinin signaling), and SmLRR-RKs (leucine-rich repeat receptor kinases) suggest that the pathways of hormone signaling, calcium signaling, and ROS (reactive oxygen species) signaling pathways play crucial roles to mediate abiotic stresses (Xue et al., 2022). These kinds of mechanisms are consistent with evidence in other crop and woody plants, where hormone and ROS signaling plays a critical role in stress acclimation (Wang et al., 2021).

3.2 Roles of key transcription factor families in stress regulation

Genome-scale analyses have shown the presence and selection of functionally diverse transcription factor families in S. mukorossi, including WRKY (e.g., SmWRKY6, SmWRKY26, SmWRKY33), bHLH (e.g., SmbHLH1), and others. These transcription factors regulate downstream stress-responsive gene expression, enabling the modulation of drought, salinity, and other stress physiological and biochemical reactions (Xue et al., 2022). Comparative studies in other plants, such as Lycium ruthenicum and rice, highlight conserved functions of the MYB, WRKY, NAC, and bZIP families in regulating stress tolerance through gene regulatory networks (Wang et al., 2018).

3.3 Stress-responsive genes and functional proteins

Selective sweep and association mapping in S. mukorossi have also identified genes that encode functional proteins such as SmPCBP2 and SmCSLD1 involved in stress adaptation and important agronomic traits (Xue et al., 2022). Direct evidence from research on LEA proteins, heat shock proteins (HSPs), and antioxidant enzyme genes in S. mukorossi are under restriction, the presence of these gene families in the genome and well-studied functions in other species suggests their likely function in tolerance to stresses (Wang et al., 2018; Yang et al., 2023). For example, the genes for antioxidant enzymes are upregulated in salt and drought stress in S. mukorossi and other model plants.

3.4 Epigenetic regulation

Despite the fact that direct research on epigenetic regulation in M. pomifera is not yet published, having a chromosome-scale genome assembly provides a platform for epigenetic research on DNA methylation, histone modification, and regulation by small RNAs in later research. All of these epigenetic mechanisms are known to control gene expression under stress in the majority of plant species and are likely also to play an important role in S. mukorossi (Xue et al., 2022; Song et al., 2023).

4 Multi-Omics Analysis of Stress Tolerance Mechanisms in Sapindus mukorossi

4.1 Applications of transcriptomics in stress response studies

Transcriptomics enables systematic description of abiotic stress-related gene expression modification in S. mukorossi, with the identification of significant regulatory genes and pathways implicated in stress adaptation. High-throughput RNA sequencing identifies abiotic stress-responsive differentially expressed genes in hormone signaling, antioxidant defense, and osmotic adjustment, providing targets for breeding and genetic engineering. Transcriptome data also allow new stress-responsive genes to be identified and the comprehension of intricate networks of stress-tolerant genes (Gupta et al., 2023; Singh et al., 2023).

4.2 Metabolomics insights into secondary metabolites associated with stress tolerance

Metabolomics demonstrates the dynamic change of primary and secondary metabolites in response to stress, such as the accumulation of osmoprotectants, antioxidants, and some secondary metabolites that provide tolerance to stress. In S. mukorossi and other woody plants, metabolomic investigations have identified molecules like proline, flavonoids, and phenolics with protective roles against drought, salinity, and heat stress (Sochacki and Vogt, 2022). These results help link metabolic pathways to physiological responses noticed and identify biomarkers for stress tolerance (Razzaq et al., 2021; Mondal et al., 2024) (Figure 1).

Figure 1 Chemical structure description of the pulp of Sapindus mukorossi (Adopted from Sochacki and Vogt, 2022)

4.3 Proteomics and functional protein network analysis

Proteomics provides direct quantification of stress response-related proteins and post-translational modifications. With the use of S. mukorossi, proteomic analysis can detect differentially abundant proteins such as heat shock proteins, LEA proteins, and antioxidant enzymes that have significant functions in cell protection and repair during stress. Functional protein network analysis describes the protein interaction further with the identification of significant hubs and pathways controlling stress response (Roychowdhury et al., 2023; Singh et al., 2023).

4.4 Multi-omics integration revealing regulatory networks of stress tolerance

Integration of transcriptomics, metabolomics, and proteomics offers a complete view of the regulatory networks that govern stress tolerance in S. mukorossi. Multi-omics enables the identification of candidate gene, protein, and metabolite and their elucidation in coordinated regulation under stress. This integration identifies sophisticated cascades of signaling, metabolic adaptation, and protein interaction, providing a systems-level understanding that can be applied to precision breeding and genetic engineering for enhanced stress tolerance (Gupta et al., 2023; Wang et al., 2023; Sarfraz et al., 2025).

5 Stress-Resistant Improvement Methods in S. mukorossi Seedling Cultivation

5.1 Traditional nursery management practices

Enhancement of substrate composition, regulation of water and fertilizer supply, and application of exogenous hormones are the basis for enhancing stress resistance in S. mukorossi seedlings. Nutrient fertilization with nitrogen, phosphorus, and potassium at optimum levels significantly improves soil quality, root fineness, and leaf physico-chemical properties, leading to enhanced yield as well as stress acclimation. However, over-fertilization can have negative impacts on soil structure and plant physiology, underpinning the necessity of precise management of nutrients (Figure 2). Techniques such as seed dormancy-breaking treatments (e.g., sulfuric acid scarification) also increase seed germination percentages and seedling vigor, improving strong establishment under stress (Gao et al., 2023; Kheloufi et al., 2024; Liu et al., 2024a).

Figure 2 Redundancy analysis (RDA) between soil properties and leaf physiological traits. Soil factors are indicated by solid arrows. Leaf properties are indicated by dashed lines. The first (horizontal) and second (vertical) axes explain 20.07% and 15.23% of the variation. ** means that correlation is significant at the 0.01 level (Adopted from Liu et al., 2024a)

5.2 Molecular breeding and utilization of stress-resistance gene resources

Molecular breeding employs genetic variation and stress-resistance gene pools to produce improved S. mukorossi varieties. Genome sequencing and population genetic research conducted in recent years have identified candidate genes for stress tolerance, such as hormone signaling, antioxidant defense, and root development-related genes. These resources enable marker-assisted selection and accelerate breeding for stress-resistant varieties (Li et al., 2021; Sochacki and Vogt, 2022).

5.3 Potential of genetic engineering and genome editing technologies

Genome editing (e.g., CRISPR/Cas9) and genetic engineering offer powerful means of targeted improvement of stress tolerance in S. mukorossi. Direct applications to the species remain to be developed, but availability of a high-quality genome and identification of genes relevant to stress response lay the foundation for future gene editing uses. These technologies hold the promise of being able to introduce or enhance traits such as drought tolerance, disease resistance, and nutrient use efficiency (Li et al., 2021; Attri, 2023).

5.4 Role of microbial symbiosis in enhancing stress tolerance

The symbiotic association with mycorrhizal fungi and endophytes has been reported to enhance stress tolerance in woody crops such as S. mukorossi significantly. These stress-tolerant microorganisms improve nutrient uptake, induce root growth, and exhibit systemic resistance against abiotic and biotic stresses. Introduction of microbial inoculants into the nursery stage is one of the possible ways for sustainable stress management, yet more studies on S. mukorossi must be conducted (Shi et al., 2022).

6 Studies on Cultivation Methods of S. mukorossi Seedlings

6.1 Rapid screening and selection techniques for stress-resistant seedlings

Recent research emphasizes the importance of effective dormancy-breaking and pre-sowing treatments for maximum germination and early vigor, being critical in selection for vigorous, stress-resistant S. mukorossi seedlings. Sulfuric acid scarification (2 hours) and heat treatment have been found to play a crucial role in high germination percentages as well as seedling growth, providing a convenient means for rapid screening of high-vigor individuals for stressful conditions (Kheloufi et al., 2024). Seed size grouping and mechanical scarification even improve the effectiveness of selection by identifying seedlings with more potential for growth (Attri, 2023).

6.2 Construction and promotion of efficient seedling cultivation systems

Successful cultivation systems for S. mukorossi integrate substrate improvement, rational fertilization, and enhanced propagation methods. NPK fertilizer application in balanced proportions not only increases yield but also improves soil fertility as well as leaf physiological properties, allowing for vigorous seedling development (Liu et al., 2024b). Grafting methods—such as cleft, side, and tongue grafting—on suitable rootstocks (e.g., Dodonaea viscosa) have been developed to enhance survival, shorten the juvenile phase, and develop resistance to stress, enabling mass production of high-quality seedlings on a large scale (Wen-Rong, 2007; Begum et al., 2019). Micropropagation and tissue culture techniques also seem to have potential for effective multiplication of superior genotypes.

6.3 Integration of stress-resistant seedling cultivation with ecological restoration and industrial applications

Production of stress-resistant S. mukorossi seedlings increasingly combines ecological restoration and industrial uses. Improved seedling vigor and survival facilitate reforestation and soil stabilization, especially on degraded or marginal soils. Fruiting of the species with high saponin content has extensive use in pharmaceuticals, cosmetics, and natural washing powders, directly relating the cultivation of seedlings to economic and ecological benefits. Improved transplanting and spacing are among the effective propagation and planting practices that also maximize restoration efficacy and industrial productivity (Chen et al., 2021; Kheloufi et al., 2024).

7 Linking Stress Tolerance Research with Practical Applications in Sapindus mukorossi

7.1 Bridging differences between greenhouse and field trials

Translation from controlled greenhouse to field environments is required for practical application. Field studies in S. mukorossi have indicated that environmental factors such as soil fertility, water status, and climatic variation have significant roles to play in seedling performance and stress response, at times leading to contrasting results with the greenhouse experiments. For example, field-oriented rational fertilization practices (e.g., specific NPK ratios) were shown to improve yield, soil fertility, and physiological traits, revealing the importance of validating laboratory findings in the real environment. Furthermore, field studies on drought stress and allelopathy illustrate that stress responses can be intensified or interact differently in the field compared to controlled environments (Zhong et al., 2023).

7.2 Establishment of evaluation systems for stress tolerance

Developing robust evaluation systems is necessary to ascertain stress tolerance in S. mukorossi. Contemporary research emphasizes the use of multiple combined indices—i.e., seedling vigor, root and shoot biomass, physiological characteristics, and tolerance indices—to evaluate performance collectively under various stressors, such as drought and heavy metals (Selvaraj et al., 2020; Sahito et al., 2023). Factor analysis and redundancy analysis (RDA) were employed to filter out the key soil and plant characteristics that are correlated with stress tolerance, providing a scientific basis for standardization evaluation protocols (Liu et al., 2024a).

7.3 Optimization of seedling quality standards and cultivation systems

Seedling quality criteria and cultivation systems can be streamlined through the translation of research into practice recommendations. High-vigor seedling selection, balanced fertilization, and substrate optimization have been recommended to achieve maximum stress tolerance and productivity (Liu et al., 2022). Furthermore, habitat modeling and climate adaptation studies offer the foundation for developing region-specific cultivation strategies, ensuring that S. mukorossi seedlings are best suited for ecological restoration and industrial applications under environmental changes (Liu et al., 2022a).

8 Concluding Remarks

The latest studies on Sapindus mukorossi seedlings have pointed out the pivotal developments in their stress tolerance mechanism. Research has indicated the pivotal physiological and biochemical mechanisms, including osmotic adjustment, antioxidant defense, enzyme protection, as well as molecular regulatory networks implicating stress signal perception, hormone cross-talk, transcription factor activity (e.g., MYB, WRKY, NAC, bZIP), functional genes and proteins (e.g., LEA proteins, HSPs, antioxidant enzymes), and epigenetic regulation. Multi-omics approaches, together with transcriptomics, metabolomics, and proteomics, have also further clarified the complex interactions underlying stress responses and identified potential targets for seedling tolerance enhancement.

Integration of molecular information with good cultivation practices is of utmost importance for improving S. mukorossi seedling performance. Traditional nursery practices, molecular breeding, gene editing, and microbial symbiosis technologies can be used best for optimization in seedling production with increased drought, low-temperature, salinity, and pest/disease stress resistance. The combined approach links fundamental research and applied methods so that stress-resistant seedlings have excellent survival, satisfactory growth, and plantation establishment over a prolonged period.

Following molecular and multi-omics understanding of stress tolerance, the subsequent efforts are needed towards the areas of molecular breeding programs for the development of elite stress-tolerant S. mukorossi cultivars, optimization of nursery and cultivation protocols, and ecological restoration and commercial culturing. Synergistic application of precision molecular tools and applied culture technologies is anticipated to enhance seedling quality, industrial yields, and underpin sustainable development in the S. mukorossi industry.

Acknowledgments

The authors sincerely thank the research team for their meticulous support and active assistance in data collection and literature organization, which provided a solid foundation for the successful completion of this study. The authors also extend heartfelt gratitude to the two anonymous studyers for their valuable comments and constructive suggestions.

Conflict of Interest Disclosure

The authors affirm that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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