1 Introduction

Tea (Camellia sinensis) is the world's second most popular non-alcoholic beverage after water. It is cultivated in more than 60 countries in the tropics and subtropics, supporting the livelihoods of more than a million smallholder growers and providing foreign exchange earnings for numerous Asian and African economies. Other than its financial worth, tea is valued for its cultural worth and variety of health-promoting bioactive chemicals, including catechins, theaflavins, and amino acids, making it a functional food. That tea is a farm business commodity and a product with a relationship to health offers evidence of its global significance in farming, business, and human wellness (Wang et al., 2025).

Tea is a perennial crop that is subjected to a range of abiotic and biotic stresses during its long life cycle. Abiotic stress by way of drought, cold, and salinity tends to cause injury to the growth of tea and reduces yield and leaf quality. Drought and temperature stress are particularly relevant under climate change scenarios, leading to impaired photosynthesis, oxidative stress, and reduced biomass accumulation. Similarly, low temperature limits tea geographic extent and directly affects winter survival overwintering and spring bud flush. Soil salinity, though less well-researched, indicates future hardship in certain tea-producing areas by way of soil deterioration and irrigation control. In addition to abiotic stresses, tea plants are also vulnerable to biotic stresses such as fungal diseases (Exobasidium vexans, Colletotrichum spp.), bacterial blights, and insect pests such as tea geometrid (Ectropis obliqua) and tea green leafhopper (Empoasca onukii), which collectively threaten sustainable production (Wang et al., 2022).

Not only does stress reduce tea yields but also quality of leaves, which is immediately translated into flavor, aroma, and the health-contributing constituents of finished tea products. As global demand for quality tea is on the rise, stress-resistant cultivars are now indispensable in order to gain yield stability as well as consistency of quality under changing environmental conditions. Understanding of the genetic basis of stress tolerance will provide molecular tools for improvement through marker-assisted selection, genomic selection, and transgenic or genome editing technologies. Furthermore, understanding of stress-responsive genes and regulation networks will also be useful in developing sustainable cultivation strategies that are in conformity with climatic resilience and ecological balance (Li et al., 2025).

This study presents a comprehensive overview of recent advances in the dissection of the genetics underlying tea plant stress tolerance. We provide molecular underpinnings of primary abiotic (drought, cold, salinity) and biotic (pathogens and pests) stress responses, describe key stress-responsive genes, transcription factors, and hormonal signaling pathways, and outline advances achieved through multi-omics approaches. Apart from this, future prospects of integrative strategies such as systems biology, genome editing, and molecular breeding are evaluated in terms of improving tea stress resistance. Opportunities and challenges are also shown, keeping in mind the aim of providing visions on future research directions and applications for sustainable tea production and industry growth.

2 Major Stress Factors and Physiological Responses in Tea Plants

2.1 Drought stress and osmotic adjustment mechanisms

Drought is the most restrictive factor in tea production, with decreased water content, altered photosynthesis, and increased oxidative damage. The tea plants respond by synthesizing osmoprotectants such as proline and soluble carbohydrates, inducing antioxidant enzyme activity (e.g., superoxide dismutase, catalase, peroxidase), and activating abscisic acid (ABA) signaling and flavonoid biosynthesis genes. These activities help to maintain cell turgor, protect cellular constituents, and detoxify ROS under drought stress. Exogenous treatments like fulvic acid, ABA, and potassium also enhance drought tolerance by modulating these activities and enhancing antioxidant defense (Chaeikar et al., 2020; Gu et al., 2020; Sun et al., 2020; Zhang et al., 2020).

2.2 Cold Stress and adaptation to low temperature

Cold stress suppresses photosynthetic process and induces ROS accumulation, resulting in cell injury. Tea plants react through the activation of antioxidant mechanisms, compatible solute accumulation, and induction of cold-inducible genes such as CBFs, ICE1, and dehydrins. Volatile organic compounds, including eugenol and (Z)-3-hexenol, also play signaling roles, inducing cold tolerance through ABA homeostasis and ROS scavenging. Such characteristics provide stability to membranes and retard leaf senescence under cold stress (Samarina et al., 2020; 2023) (Figure 1).

Figure 1 Relative expression levels of the studied genes in response to cold and drought (Adopted from Samarina et al., 2020)

2.3 Salt stress and ionic homeostasis regulation

Salt stress triggers ion imbalance and oxidative stress in tea leaves. Ionic homeostasis, especially by selective uptake and compartmentalization of Na+ and K+ ions, is necessary. Antioxidant defense and osmolyte accumulation (such as proline and sugars) are also activated to prevent salt-mediated injury. Plant growth-promoting bacteria also enhance salt tolerance by regulating ion transport and stress signaling (Li et al., 2019; Gamalero and Glick, 2022).

2.4 Biotic stress: disease, pest resistance, and plant immune responses

Tea plants are attacked by various pathogens and insects, which trigger immunological responses by the induction of pathogenesis-related proteins, secondary metabolites (flavonoids and lignin), and hormone signaling mechanisms (salicylic acid and jasmonic acid). Drought stress and salt stress could increase biotic vulnerability but cross-protection can be elicited by beneficial microbes and the augmentation of antioxidant defense (Gamalero and Glick, 2022).

2.5 Combined effects of multiple stresses and interactive mechanisms

Tea plants are typically exposed to combined stresses, e.g., cold and drought, triggering specific and general molecular responses. Transcriptome profiling indicates that combined stresses lead to unique patterns of gene expression, with crosstalk between ABA-dependent and -independent signaling pathways, and regulation of antioxidant and osmoprotectant systems. Unstable signals and some transcription factors (e.g., bHLH, WRKY, NAC) consolidate responses to multiple stresses, achieving maximum energy efficiency and prolonging leaf lifespan (Zhang et al., 2017; Ni et al., 2024).

3 Genetic Basis of Stress Resistance in Tea Plants

3.1 Whole-genome sequencing and population genetic diversity

High-quality reference genomes and resequencing of diverse tea accessions have revealed the extensive genetic diversity and population structure of tea plants. The study identified three populations, domestication signatures, and hundreds of allelic variations for quality and stress resistance traits. Gene family expansions for stress responses such as terpene synthases have been involved in adaptation and domestication (Xia et al., 2020). Phased genome assemblies also uncover allelic imbalance and structural variations that are accountable for stress tolerance.

3.2 Stress-related gene families

Certain gene families play significant roles in abiotic and biotic stresses of the tea plant. WRKY, NAC, MYB, bZIP, HD-Zip, and HSP transcription factors are differentially regulated during abiotic and biotic stresses and regulate downstream defense and adaptation processes (Shen et al., 2022). For example, WRKY48 induces cold and pest resistance, while HD-Zip and BZR1 families are implicated in hormone and stress signaling (Shen et al., 2019). The NB-ARC domain-containing genes also play roles in disease and cold resistance (Li et al., 2025).

3.3 Signaling pathways and regulatory networks

Tea plants also use complex signaling networks such as abscisic acid (ABA), salicylic acid (SA), jasmonic acid (JA), and reactive oxygen species (ROS) to regulate stress responses. Transcriptomics indicate ABA, ethylene, and JA pathway genes to be induced in drought and cold stress with adaptation being controlled by hormone and redox cross-talk (Shen et al., 2019). Regulation miRNAs and transcription factors further tune these networks (Zhou et al., 2019).

3.4 Key structural and functional genes

Genes coding for osmoprotectants, antioxidant enzymes (e.g., SODs, peroxidases), transporters (e.g., CsGAT1 for GABA, SAT for cysteine), and lignin and flavonoid biosynthetic enzymes are major contributors to stress tolerance. CsSOD genes, for instance, are induced by cold-drought, and CsGAT1 enhances cold tolerance by controlling GABA (Zhou et al., 2019; Li et al., 2024). The genes CsHCT and laccase control lignin synthesis and impart abiotic and biotic stress resistance. Regulation by alternative splicing and miRNA further confers complexity on gene performance under stress (Zhou et al., 2019).

3.5 Advances in genome-wide association studies (GWAS), QTL mapping, and genomic selection (GS)

Next-generation sequencing and new genomic tools have enabled GWAS, QTL mapping, and genomic selection in tea. They have also been employed to discover candidate genes and markers related to resistance to stress for marker-assisted and genomic selection for breeding stress-resistant cultivars (Xia et al., 2020). SSR marker construction and multi-omics integration also accelerate genetic improvement.

4 Advances in Multi-Omics Research on Stress Resistance in Tea Plants

4.1 Transcriptomic insights into stress-responsive networks

Transcriptome analysis revealed that tea plants are capable of activating massive gene networks in response to abiotic and biotic stress. Differentially expressed genes tend to be enriched within flavonoid biosynthesis, hormone signaling, and disease resistance pathways. Transcriptome analysis of pathogen and cold stress identified key NB-ARC domain genes and highlighted the central role played by the flavonoid pathway for stress adaptation (Li et al., 2025). During drought stress, transcriptomics has identified genes involved in photosynthesis, transmembrane transport, phytohormone metabolism, and secondary metabolite biosynthesis, among a number of transcription factors and protein kinases (Figure 2) (Samarina et al., 2023; Yue et al., 2023).

Figure 2 UPGMA clustering of tea transcriptome samples (A); Heat map of stress response correlation in tea plants based on transcriptomic profiles (B) (Adopted from Samarina et al., 2023)

4.2 Proteomics and metabolomics applications in stress responses

Proteomic and metabolomic analysis complements transcriptomics to identify responsive proteins and metabolites under stress. Metabolomic profiling has identified the buildup of catechins and flavonoids under cold and disease stress, which agrees with defense gene induction (Li et al., 2025). Proteomics has identified protein abundance fluctuation and post-translational modification in response to drought and pathogen infection and proved the dynamic regulation of stress responses (Li et al., 2022).

4.3 Epigenetic regulation in stress resistance

Epigenetic control by DNA methylation, microRNAs (miRNAs), and long non-coding RNAs (lncRNAs) is crucial in controlling gene expression during stress response. System-level omics strategies have identified stress-inducible miRNAs that regulate defense genes post-transcriptionally, such as miR530b and miRn211, which regulate ROS-related genes during infection. DNA methylation modifications have also been shown to be implicated in the regulation of stress-inducible genes and with preservation of stress resistance phenotypes (Li, 2024).

4.4 Integration of multi-omics data and systems biology approaches

Integration of multi-omics data facilitates the construction of global regulatory networks and the discovery of candidate genes, proteins, and metabolites in stress tolerance. Co-expression network analysis and machine learning approaches in systems biology allow for the discovery of biomarkers and regulation modules for breeding stress-tolerant tea cultivars (Yue et al., 2023; Kajrolkar, 2025). Integrated study is driving climate-resilient tea plant and precision breeding.

5 Molecular Breeding and Strategies for Stress Resistance Improvement in Tea Plants

5.1 Marker-assisted selection (MAS) in stress-resistant breeding

Marker-assisted selection (MAS) has become a valuable technique for breeding tea plants with enhanced abiotic and biotic stress tolerance. Reference genomes of high quality and molecular markers with versatility facilitate the identification and selection of major genes and quantitative trait loci (QTLs) for stress tolerance, such as those involved in cold, drought tolerance, and insect resistance (Xia et al., 2020; Joshi et al., 2023). MAS speeds up breeding through early and precise selection of the targeted traits, and is being integrated more and more with multi-omics data for enhanced efficiency.

5.2 Potential of gene editing (CRISPR/Cas) in tea stress resistance

Gene editing technology, particularly the CRISPR/Cas systems, holds high promise for precision editing of stress-resistance genes in tea plants. Though successful implementation in tea is yet to arrive in the spotlight due to technical challenges, CRISPR/Cas has been extensively employed in other crops for the engineering of drought, salinity, and heat stress tolerance through regulation gene and transcription factor targeting (Nascimento et al., 2023). The development of efficient transformation systems and candidate gene identification in tea will also enable the use of gene editing for the enhancement of stress tolerance in a rapid manner.

5.3 Germplasm innovation and identification of stress-tolerant resources

Their analysis and use are the foundation for breeding stress-tolerant cultivars. Hybridization, extensive crossing, and identification of trace alleles in less studied Camellia species enhanced the genetic basis for resistance breeding. CsAFS2 and CsWRKY48 gene activities have unveiled their activities for cold, drought, and insect resistance enhancement and provided valuable targets for molecular breeding (Wang et al., 2024). Germplasm innovation is supported by multi-omics techniques that reveal allelic variation and adaptation characteristics (Xia et al., 2020).

5.4 Balancing stress resistance with tea quality improvement

One of the main challenges of molecular breeding is to enhance tolerance to stress while preserving tea quality. Combined breeding strategies highlight selection for resistance characteristics and for quality-characteristic metabolites such as catechins and theanine. Experiments have proven that exogenous treatment (e.g., methyl jasmonate) and transcription factor manipulation (e.g., MYB, WRKY) increase not just tolerance to stress but also quality-related metabolite content (Li et al., 2023). Genomics and metabolomics enable breeders to monitor and control such characteristics in selection (Han et al., 2022).

6 Challenges and Limitations in Tea Stress Resistance Research

6.1 Complexity of the tea genome and difficulties in genetic studies

The tea plant genome is very big and complex with a high proportion of repetitive sequences and high genetic heterogeneity. It complicates genome assembly, gene annotation, and identification of functional stress-resistance related genes. High gene duplication events and occurrence of numerous gene families complicate further dissection of stress-related pathways and the development of useful molecular markers for breeding (Xia et al., 2020). Functional genomic studies are further hampered by the evergreen nature and prolonged generation time of tea plants, which make genetic transformation and validation of candidate genes time-consuming.

6.2 Insufficient understanding of cross-talk among stress factors

While single-stress responses (e.g., to drought, cold, or salinity) have been studied, cross-talk and interactive effects of two or more stress factors are unclear. Experiments are mostly carried out under single-stress conditions, but tea plants in the field are exposed to simultaneous or sequential stresses. Molecular processes of the integration of multisensorial signals and of their combined action on growth, yield, and quality remain unknown, which is a significant research gap.

6.3 Gaps between laboratory research and field application

A majority of gains in stress resistance have their origins in laboratory or controlled-environment experiments that may not always be true in the field. Environmental variation, soil variations, and biotic interactions within tea estates can potentially alter stress responses and the effectiveness of resistance traits. There is a need for increased validation of candidate genes, molecular markers, and adaptive traits at the field level to verify practical application in breeding and production.

6.4 Integration of genetic improvement with traditional cultivation practices

Dissemination of new genetic improvement technology to traditional cultivation and management practices remains challenging. Socioeconomic factors, farmer acceptability, and agroclimatic culture of the region may discourage new cultivar growth with stress tolerance. Stress tolerance vs. maintenance of tea quality and acclimatization to local environments must be maintained through multidisciplinary approaches combining molecular, ecological, and socioeconomic research (Ramakrishnan et al., 2023).

7 Integrative Perspectives and Practical Implications of Tea Stress Resistance Research

7.1 Complementarity of different research approaches

Molecular biology, genetics, and multi-omics (transcriptomics, metabolomics, and proteomics) present complementary and unique perspectives of tea plant stress responses. Molecular and genetic studies identify key genes and regulatory networks involved in stress tolerance, whereas omics approaches identify global changes in gene expression, metabolite accumulation, and protein function under stress conditions. For example, transcriptomics and metabolomics have enlightened the critical role of flavonoid metabolites in redox homeostasis and the regulation of tea quality under stressed conditions. Combination of these approaches gives a general idea of the stress resistance mechanisms that include physiology and molecules, and thus breeding improved tea varieties is possible (Zhang et al., 2019).

7.2 Contribution of stress resistance research to sustainable tea production

Stress resistance research is ensuring sustainable tea production via the prospect of developing cultivars capable of resisting abiotic stresses such as cold, drought, and salinity, and biotic stress. Increased stress tolerance guarantees yield and quality maintenance, reduced economic loss, and chemical inputs. Stress-activated genes and metabolites, such as flavonoids, also direct cultivation methods and exogenous treatments (e.g., ABA, MeJA, melatonin) that can trigger field-level resistance in plants (Wang et al., 2023). This research supports long-term sustainability and profitability for tea farming.

7.3 Relevance of stress resistance studies under global climate change

With increasing intensity and frequency of environmental stresses due to global climate change, research on stress resistance in tea plants is more relevant than before. Understanding the processes at the molecular and physiological level underlies stress adaptation permits rapid breeding and sharing of climatically resilient tea varieties. This is essential for safeguarding tea yields, quality, and farmer incomes against irregular weather conditions and climatic extremes. In addition, stress resistance research is supportive of producing cultivation strategies that can provide a buffer against climate change impacts on tea production systems (Li et al., 2023).

8 Concluding Remarks

The past several years have witnessed significant progress in deciphering the genetic and molecular mechanisms of stress tolerance in tea (Camellia sinensis). Multi-omics approaches, i.e., genomics, transcriptomics, proteomics, and metabolomics, have revealed key stress-responding genes, transcription factors, and regulatory networks that regulate abiotic stresses such as drought, cold, and salinity and biotic stresses caused by pathogens and pests. Candidate genes for reactive oxygen species (ROS) scavenging, osmolyte biosynthesis, hormonal signaling, and secondary metabolite production have also been located, which are responsible for the mechanistic aspects of environmental stress perception and adaptation of tea plants. Functional characterization of the NAC, WRKY, and MYB transcription factor families has then highlighted their key roles as master regulators of stress-response pathways. Collectively, they form a solid foundation for rational improvement of tea stress tolerance using molecular breeding and biotechnology methods.

Genetics and molecular research is a platform to drive tea breeding for tolerance towards stress. The discovery of genes and factors regulating responses towards stress can make it easier to utilize marker-assisted selection, genomic selection, and targeted genome editing approaches, which can shorten the length of breeding cycles and improve selection efficiency. In addition, the integration of genetic data with phenotypic information makes it possible to develop predictive models guiding cultivar selection under different environmental conditions. With this description of heritability of stress adaptation, genetic research provides valuable tools for breeding improvement as well as theoretical insight into plant-environment interactions and hence reconciles basic science and applied breeding.

The integrated genetic basis of tea stress tolerance identifies clear paths for enhancing crop resilience, maximizing yield stability, and maintaining leaf quality under stress. Translation of the discovery into high-throughput phenotyping, multi-omics, and functional confirmation will further clarify breeding strategy. Translation of research gains into industrial yield—some of which are stress-tolerant cultivars and climate-resilient management practices development—a few of them are going to ensure long-term sustainability of tea production and economic stability. Lastly, structured genetics research not only progresses plant stress biology knowledge but also contributes to sustainable development and competitiveness of the global tea industry.

Acknowledgments

The author sincerely appreciate the research team members for their valuable contributions and support in the collection of relevant materials and literature organization during the study of tea plants. The authors also thank the two anonymous reviewers for their valuable comments, which played an important role in improving and refining the manuscript.

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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