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Review

Multi-Scale Drought Resilience in Terrestrial Plants: From Molecular Mechanisms to Ecosystem Sustainability

by
Weiwei Lu
1,2,
Bo Wu
1,2,*,
Lili Wang
1 and
Ying Gao
1,2
1
Institute of Ecological Conservation and Restoration, Chinese Academy of Forestry, Beijing 100091, China
2
Key Laboratory of Desert Ecosystem and Global Change, State Administration of Forestry and Grassland, Beijing 100091, China
*
Author to whom correspondence should be addressed.
Water 2025, 17(17), 2516; https://doi.org/10.3390/w17172516
Submission received: 14 July 2025 / Revised: 16 August 2025 / Accepted: 21 August 2025 / Published: 23 August 2025
(This article belongs to the Special Issue Wetland Conservation and Ecological Restoration)
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Abstract

Global climate change has intensified the frequency, intensity, and spatial heterogeneity of drought events, posing severe threats to the stability of terrestrial ecosystems. Plant drought resilience, which encompasses a plant’s capacity for drought resistance, post-stress recovery, and long-term adaptation and transformation to sustain ecosystem functionality, has emerged as a central focus in botanical and ecological research. This review synthesizes the conceptual evolution of plant drought resilience, from early emphasis on resistance and recovery to the current multi-dimensional framework integrating adaptation and transformation, and synthesizes advances in understanding multi-scale drought resilience in terrestrial plants—spanning molecular, physiological, individual, community, and ecosystem levels. Key mechanisms include molecular/physiological adaptations (osmotic adjustment, antioxidant defense, hydraulic regulation, carbon–water reallocation via gene networks and aquaporins), morpho-anatomical traits (root architectural plasticity, leaf structural modifications, and hydraulic vulnerability segmentation), community/ecosystem drivers (biodiversity effects, microbial symbioses, and soil–plant–feedback dynamics). We critically evaluate quantitative metrics and expose critical gaps, including neglect of stress legacy effects, oversimplified spatiotemporal heterogeneity, and limited integration of concurrent stressors. Future research should prioritize multi-scale and multi-dimensional integrated analysis, long-term multi-scenario simulations with field validation, and harnessing plant–microbe interactions to enhance drought resilience, providing a theoretical basis for ecosystem sustainability and agricultural production under climate change.

1. Introduction

Global climate is undergoing nonlinear evolution characterized predominantly by warming, exhibiting complex features including multi-dimensionality, nonlinearity, and threshold-crossing dynamics [1]. Observational data indicate a 1.09 °C rise in global surface temperature between 1880 and 2022, with the 2013–2022 decade witnessing an accelerated warming rate of 0.38 °C/decade [2]. This warming trajectory intensifies the hydrological cycle, triggering marked increases in the frequency, intensity, and spatial heterogeneity of extreme climate events—manifested as a 56% expansion in regional dry-wet oscillation amplitude (1979–2022) and a 3.2-fold rise in compound extreme event probability [3]. Global climate change intensifies drought dynamics. CMIP6 model simulations project that a global temperature rise of 1.5–4.0 °C will increase potential evapotranspiration across most regions. In East Asia (EAS), for instance, potential evapotranspiration rises by 11.28% under 4.0 °C warming [4]. Drought intensity, coverage area, and duration exhibit significant increases under warming scenarios. In China, anthropogenic climate change has amplified drought frequency by approximately 23%, intensified drought severity by 15%, and expanded affected areas by over 10% [5]. This nonlinear response is particularly pronounced in the Asian monsoon region, where severe drought frequency surges by 30% and duration extends by 20–40% under 2.0 °C warming [6]. At the ecohydrological scale, spatiotemporal reconfiguration of precipitation patterns (e.g., monsoon advancement/delay, rainbelt migration) exacerbates water allocation imbalances, with 34% of global terrestrial ecosystems now experiencing persistent water deficits [7]. Drought emerges as a pivotal threat to terrestrial ecosystem stability under climate change. China, a global drought-stress hotspot, confronts multifaceted challenges of water scarcity and ecological security, directly contributing to a 42% interannual coefficient of variation in degradation rates of northern shelterbelts [8]. Late 20th-century intensive human activities drove vegetation degradation in arid zones exceeding 0.8%/a, catalyzing regional desertification. Implementation of ecological water diversion and Grain-for-Green programs elevated the Normalized Difference Vegetation Index (NDVI) by 0.12–0.18 (2000–2020), significantly restoring ecosystem services [9]. However, synergistic effects between warming-driven glacial retreat acceleration and atmospheric aridification are reducing ecological carrying capacity thresholds and water-use efficiency in arid regions [8].
Drought threatens terrestrial ecosystems through a dual-threat mechanism: directly inducing plant hydraulic failure and carbon starvation, while indirectly elevating pest outbreak risks by 2.7–4.5-fold via altered plant secondary metabolite profiles that compromise chemical defenses [10,11]. Plant drought resilience has emerged as a central focus in botanical and ecological research under climate change, encompassing a plant’s capacity for drought resistance, post-stress recovery, and long-term adaptation and transformation to sustain ecosystem functionality and productivity [12,13]. Under global change, vegetation drought resilience emerges as the pivotal regulator of ecosystem sustainability, necessitating elucidation of multi-scale adaptive mechanisms. Current research predominantly focuses on acute drought stress effects (e.g., photosynthetic inhibition, hydraulic dysfunction) and vegetation decline modeled through linear response frameworks [14,15]. Critical knowledge gaps persist; however, including neglect of post-stress recovery dynamics, oversimplification of spatiotemporal heterogeneity in drought–vegetation interactions, and absence of long-term adaptive capacity quantification [16]. These limitations impede holistic understanding of ecological resilience’s multidimensional hierarchy, hindering comprehensive assessment of vegetation responses to episodic droughts versus secular climate change.
This review systematically synthesizes recent advances in terrestrial plant drought resilience research to establish a unified conceptual framework spanning molecular, physiological, individual, community, and ecosystem scales. We clarify the conceptual evolution from traditional resistance–recovery paradigms toward multidimensional frameworks integrating adaptation and transformation capacities; comprehensively elucidate key response mechanisms across scales—encompassing molecular, physiological/morphological, and community/ecosystem processes; critically evaluate strengths and limitations of existing quantitative metrics while identifying critical knowledge gaps in spatiotemporal heterogeneity, stress legacy effects, and multi-factor interactions; and propose priority research directions including multi-scale integrative analyses, long-term multi-scenario simulations with field validation, and resilience-enhancing strategies leveraging plant–microbe interactions. This integrated perspective aims to advance theoretical foundations for climate-resilient ecosystem management and sustainable agriculture.

2. Methodology

This review employed a purpose-driven narrative synthesis to trace the evolution, current trends, and future directions of drought resilience research. Literature was primarily sourced from major academic databases (Web of Science, Scopus, Google Scholar) using core search terms (“drought resilience”) and related keywords (e.g., “drought adaptation,” “drought resistance”), combined with scale-specific terms (e.g., “ micro-scale,” “individual scale,” “ecosystem,” “community,” “global”). We prioritized peer-reviewed journal articles (original research, reviews) but included influential meta-analyses, books, and reports from organizations (e.g., IPCC (Intergovernmental Panel on Climate Change)) where critical. Selection emphasized: (a) Recency: Focus on seminal works (pre-2010) and high-impact studies from the last decade (2014–2024), particularly within the last 5 years; (b) Scholarly impact: Citation metrics identified field-defining and widely recognized literature; (c) Spatial scale representation: Studies were systematically categorized into scales to contrast research themes, methods, and interventions; (d) Source rigor: Preference for primary research and systematic reviews. Analyses addressed historical concept/method evolution, scale-specific challenges/strategies, key knowledge gaps, and emerging research frontiers. As a narrative review, this approach allows integrative, cross-scale insights but acknowledges limitations in exhaustive coverage versus systematic methods.

3. Advancing the Conceptualization of Plant Drought Resilience

3.1. Conceptualization and Dimensions of Drought Resilience

The concept of resilience has a longstanding history in ecology, originally describing an ecosystem’s capacity to return to its original state following disturbances (Table 1). With intensifying global climate change driving increased frequency and severity of drought events, researchers have progressively incorporated resilience theory into studies of plant drought responses [17]. Initial research primarily centered on resistance (a plant’s ability to maintain functionality during drought) and recovery (its capacity to restore functionality after drought cessation) [12,18,19]. Holling [20] established ecological resilience as an ecosystem’s capacity to maintain functional and structural stability post-disturbance, introducing two fundamental dimensions: absorptive capacity (threshold of disturbance absorption without structural change, e.g., photosynthetic tolerance under drought) and reorganizational capacity (self-organization into new stable states beyond tipping points, e.g., post-desertification succession). This conceptual foundation, contrasting with engineering resilience (recovery speed), revealed the nonlinear dynamics underpinning drought responses. From the 1990s, Holling’s framework evolved into discipline-specific constructs: crop drought resilience and ecological drought resilience [21]. Tilman subsequently decoupled resilience into quantifiable metrics via grassland experiments [22,23]: resistance (functionality maintenance during stress) and recovery (post-stress restoration rate), demonstrating their critical trade-off—a principle guiding drought-tolerant crop breeding. A widely adopted resilience framework conceptualizes resistance and recovery as complementary components of ecosystem adaptability. This concept with its calculation of three resilience indices was proposed in 2011 by Lloret et al. [24]. However, an exclusive focus on resistance and recovery proves insufficient for comprehensively characterizing plants’ complex responses to prolonged and recurrent droughts. Schwarz et al. [19] critically evaluated conventional resilience metrics, emphasizing the necessity for more robust and standardized methodologies to assess tree drought responses (Figure 1). Through analyses of divergent drought-response scenarios—including delayed drought responses, pre-drought perturbations, and protracted recovery phases—they demonstrated significant variations in how different metrics capture authentic plant physiological behaviors. Subsequent studies have identified limitations in the Lloret indices, which may yield biased results or misinterpretations under specific conditions. Schwarz et al. [19] examining three common detrended growth scenarios in dendrochronology (Figure 1) revealed deviations from the theoretical assumptions underlying Lloret’s original resilience framework. Case analyses demonstrate: (i) drought-induced growth minima occurring after climatic events due to delayed responses (Figure 1a); (ii) violated pre-drought growth assumptions caused by non-drought-related suppression in reference periods (Figure 1b); and (iii) protracted recovery from sequential droughts necessitating both multi-year drought data aggregation and post-drought reference period re-calibration (Figure 1c). Subsequently, drought resilience has emerged as a comprehensive conceptual framework that extends beyond resistance and recovery to encompass a plant’s capacity for adaptive adjustment and structural transformation in response to persistent drought stress, thereby ensuring sustained ecological functionality [12,25]. This conceptual evolution reflects the research community’s deepening understanding of the intricate dynamics in plant–environment interactions. Post-2000, accelerated by climate change, drought resilience has been systematically defined with progressively enriched dimensions. In this section, where applicable, authors are required to disclose details of how generative artificial intelligence (GenAI) has been used in this paper (e.g., to generate text, data, or graphics, or to assist in study design, data collection, analysis, or interpretation). The use of GenAI for superficial text editing (e.g., grammar, spelling, punctuation, and formatting) does not need to be declared.

3.2. Drought Resilience Indicators

Drought resilience indicators serve as critical tools for assessing the ability of ecosystems, agricultural systems, and even socio-economic systems to maintain functionality and recover from drought disturbances. These indicators span multiple dimensions—from plant physiological levels to regional scales—and are designed to quantify a system’s resistance to drought, recovery capacity, and overall adaptation capabilities [26,27]. An in-depth understanding of these indicators is crucial for developing effective drought response strategies and ensuring regional sustainable development [2,28]. Resistance refers to a system’s ability to withstand negative impacts and preserve structure/function during drought [26,29]. For instance, highly resistant systems exhibit minimal initial impact, such as drought-tolerant maize varieties maintaining high germination vigor, germination rates, and growth rates of coleoptiles and roots under water stress during seedling stages [30]. Quantitatively, plant resistance can be measured by reduced growth or productivity decline during drought, often expressed as the ratio of growth under drought to growth under favorable conditions [31]. Recovery denotes the ability and speed of a system to return to its pre-drought state after drought cessation [25,26,29]. This is typically assessed by monitoring post-drought recovery trajectories of ecosystem functions (e.g., vegetation productivity via satellite remote sensing), where prolonged recovery may predict increased future mortality risks [25,29,32].
Beyond resistance and recovery, drought resilience includes adaptation—a system’s capacity to adjust its structure or function for improved future drought response—and transformation, which involves fundamental system changes to address long-term or extreme climate shifts, moving beyond mere recovery toward more sustainable states [19,33] (Figure 2). Figure 2 conceptualizes drought resilience dimensions through comparative vegetation dynamics. The hierarchy progresses from overarching Adaptation (evolutionary/physiological adjustments) to core Drought Resilience (integrated stress response capacity), which bifurcates into Resistance (stress-withstanding ability via osmotic regulation, hydraulic control) and Recovery (post-stress function restoration via metabolic reprogramming). Trees/Ecosystems A and B exemplify divergent resilience strategies: A demonstrates high resistance but limited recovery (e.g., deep-rooted species maintaining function during drought yet slow to rebound), while B exhibits lower resistance but rapid recovery (e.g., drought-deciduous species sacrificing immediate function for efficient post-rain regrowth). The bidirectional arrow signifies that resistance–recovery tradeoffs in real-world systems are dynamically mediated by species traits, soil properties, and historical climate exposure. Though adaptation and transformation metrics are less frequently quantified in empirical studies compared to resistance and recovery, they remain essential for understanding the macro-level evolution of drought resilience [34,35].

4. Plant Drought Resilience Across Spatial Scales

4.1. Molecular and Cellular Mechanisms

4.1.1. Water Relations Regulation

Under drought stress, plants enhance drought resistance through multi-level synergistic mechanisms. Plants optimize water utilization via stomatal behavior and osmotic adjustment: Isohydric species strictly limit stomatal aperture to avoid hydraulic failure, while anisohydric species permit tissue water potential decline to sustain carbon assimilation [36]. Concurrently, plants accumulate osmolytes (e.g., proline, betaine) to lower cellular osmotic potential, ensuring turgor maintenance [37]. Molecular studies reveal that abscisic acid (ABA) signaling mediates rapid stomatal closure by regulating ion channels in guard cells, with transcription factors (e.g., DREB (Dehydration-Responsive Element-Binding protein)) fine-tuning stomatal dynamics for water balance [38].
Plants employ osmotic adjustment to maintain cellular turgor and water homeostasis during drought, a critical physiological adaptation achieved through three synergistic mechanisms: (1) Preservation of plasma and endosomal membrane integrity to prevent turgor loss [39]; (2) Accumulation of compatible solutes that reduce cellular osmotic potential, thereby enhancing soil water extraction efficiency [40]; and (3) Responsiveness to exogenous interventions exemplified by nitric oxide (NO) application, which elevates shoot length (66.60%), root length (29.38%), shoot dry weight (26.69%), and root dry weight (28.52%), while concurrently boosting photosynthesis, antioxidant defenses, and osmotic adjustment to confer comprehensive drought resilience [9].

4.1.2. Carbon–Water Coupling Balance

Plants maintain carbon–water equilibrium through photosynthesis-transpiration decoupling and carbon allocation: C4 (C4 carbon fixation pathway) and CAM (Crassulacean Acid Metabolism) plants enhance water use efficiency (WUE) by spatially or temporally separating CO2 fixation from transpiration. Despite drought-induced photosynthesis suppression leading to carbon starvation, plants prioritize carbon allocation to critical organs (e.g., roots) and leverage non-structural carbohydrate (NSC) reserves as metabolic substrates and osmoregulators [41], sustaining survival under limited carbon supply. Advances in molecular biology and genomics have propelled in-depth research into drought resilience-related genes. Studies reveal that specific genes—such as transcription factors like DREB and NAC—play critical roles in regulating plant drought responses by controlling water management and carbon metabolism pathways. For instance, gene-editing technologies like CRISPR are being deployed to enhance plant drought tolerance [42].

4.1.3. Oxidative Stress Defense

The plant antioxidant defense system primarily protects cells from oxidative damage by balancing the clearance of reactive oxygen species (ROS), involving multiple biochemical processes. This mechanism is particularly crucial under environmental stresses such as drought or light stress. To counter drought-triggered reactive oxygen species (ROS) accumulation, plants activate multi-tiered protection: Antioxidant enzymes (e.g., SOD (Superoxide Dismutase), CAT (Catalase)) scavenge ROS to safeguard membrane integrity [43], while heat shock proteins (HSPs) stabilize denatured proteins to maintain cellular homeostasis [44].

4.2. Leaf-Level Responses

4.2.1. Physiological and Biochemical Responses

Photosynthesis, the cornerstone of plant growth and productivity, exhibits drought-driven dynamics that directly determine resistance capacity and post-stress recovery resilience. Under severe or prolonged drought—and co-occurring waterlogging—photosynthetic impairment manifests through reduced stomatal conductance, disrupted electron transport, and Rubisco inactivation [45,46]. While most photosynthetic parameters recover after rehydration, the extent and velocity of recovery demonstrate significant genotypic variation and scale inversely with stress intensity [45]. Critically, persistent photochemical damage serves as a threshold metric for resilience, where sustained depression of the photosystem II (PSII) maximum quantum yield (Fv/Fm) beyond 15% post-rehydration predicts elevated mortality risk during subsequent droughts in tropical rainforest species [38]. This legacy effect confirms that drought impacts transcend transient physiological disruption, driving cumulative functional deterioration that compromises long-term plant performance.

4.2.2. Photosynthetic Regulation

Physiologically, drought reduces leaf relative water content (RWC)—a critical water status indicator [47]. Stomatal regulation minimizes water loss through closure but concurrently limits photosynthesis and growth [48,49,50]. Under drought conditions, plants adjust carbon metabolism and allocation strategies. Research indicates that while drought suppresses photosynthesis and induces carbon starvation, plants counteract this challenge by preferentially allocating carbon resources to critical organs such as roots [38]. Through reducing carbon investment in non-essential tissues and organs, plants sustain vital functions under constrained carbon supply. Osmotic adjustment counters low water potential via proline, soluble sugars, and mannitol accumulation, preserving cell turgor and function [47]. To mitigate drought-induced reactive oxygen species (ROS), plants elevate antioxidant enzymes (SOD, CAT, POD (Peroxidase)) and non-enzymatic antioxidants (glutathione, vitamin C) [51,52,53]. Photosynthesis declines markedly, evidenced by reduced net photosynthetic rate, transpiration, and stomatal conductance [48,54]. Regulation of carbon metabolism and carbon reserves plays a critical role in plant drought resilience, particularly under prolonged drought stress. Under drought conditions, plants adjust carbon metabolism and allocation strategies. Research indicates that while drought suppresses photosynthesis and induces carbon starvation, plants counteract this challenge by preferentially allocating carbon resources to critical organs such as roots [38]. Through reducing carbon investment in non-essential tissues and organs, plants sustain vital functions under constrained carbon supply.

4.3. Whole-Plant and Species-Level Traits

4.3.1. Morphological Adaptations

Under drought stress, plants undergo significant morphological changes to enhance drought resilience. Root architecture—the key organ for water uptake—adapts through increased root-to-shoot ratio and development of deeper/extensive root systems, improving water absorption efficiency [55,56]. Plant drought resilience is fundamentally governed by interconnected root architectural traits and rhizosphere processes, with root system functionality serving as the primary determinant of water acquisition efficiency under moisture deficit [57]. Adaptive plasticity in root morphology and physiology during drought-rehydration cycles critically drives climate resilience and productivity optimization, though such plasticity can be compromised by concurrent stresses—exemplified by cadmium contamination disrupting root circadian rhythms and suppressing lateral root development [39,57]. Notably, steep lateral root angles (LRA) enabling deeper soil exploration significantly enhance drought resilience through access to subsoil water reserves, as demonstrated by the superior drought survival of narrow-LRA cotton genotypes [58]. Furthermore, drought-adaptive rhizosheath formation—mediated by cultivar-specific biophysical and biochemical traits—orchestrates plant–soil–microbe interactions, where root exudate-driven communications constitute underexplored pathways for drought mitigation [59,60]. Aboveground, plants reduce height, decrease leaf area, or shed leaves to minimize transpirational water loss [49,61], exemplified by eucalyptus seedlings adjusting photosynthesis and growth post-water deficit pretreatment [62].

4.3.2. Microbial and Metabolic Contributions

Plant–microbe interactions and endogenous metabolic processes constitute critical determinants of drought resilience. Rhizospheric microbiomes manifest drought-responsive adaptations, as evidenced by wheat rhizospheres where stress-enriched microbial consortia and metabolites enhance drought tolerance through plant–microbe crosstalk, exhibiting persistent legacy effects [63]. Arbuscular mycorrhizal fungi (AMF) significantly potentiate plant drought resilience via symbiotic water-nutrient exchange [64], while plant growth-promoting rhizobacteria (PGPR) synergistically reinforce resistance to combined drought–salinity stressors [65]. Metabolically, trehalose-6-phosphate (T6P) signaling precursor treatment demonstrates efficacy in boosting spring wheat yield and drought resilience through osmotic regulation [66].

4.3.3. Hydraulic Traits

Hydraulic traits critically govern drought survival. Xylem embolism—gas bubble formation in conduits—impedes water transport and can cause mortality. Plants combat this through anatomical adaptations like modified vessel diameter/density to enhance hydraulic safety [67]. A key strategy is hydraulic vulnerability segmentation, where distal organs (e.g., leaves, fine roots) are sacrificed before core structures, preserving overall hydraulic integrity [68]. Anatomical modifications further optimize drought resistance. Thickened leaf cuticles reduce non-stomatal water loss and improve water-use efficiency [69]. Vascular bundle organization—including xylem vessel size, density, and arrangement—determines hydraulic efficiency and embolism resistance. Drought-tolerant species often exhibit smaller vessel diameters and higher vessel density to enhance hydraulic safety [70].

4.4. Community and Ecosystem Dynamics

4.4.1. Plant Biodiversity and Community Properties

At the ecosystem scale, biodiversity exerts critical influences on drought resilience. Hydraulic diversity—particularly the variability in plant functional traits related to water transport—serves as a key modulator of land–atmosphere feedbacks during drought events, thereby regulating ecosystem resilience [71]. Community composition and species diversity determine ecosystem resistance and recovery capacity, with warming and nitrogen deposition demonstrating drought-contingent effects on grassland resilience through compositional shifts [72,73]. Precipitation anomalies (including drought and pluvial episodes) further reconfigure productivity resilience via altered community dynamics, evidenced by drought-avoidant seed dormancy mechanisms in annual plant communities where wet seasons counteract prior drought impacts [74].

4.4.2. Trait-Based Adaptive Strategies in Community Drought Response

Drought resilience manifests through changes in community composition, functional traits, and species interactions [75]. Ecological theory predicts that community assembly processes may lead to functional traits of coexisting species being more divergent (“divergence”) or more similar (“convergence”) than expected by random chance [76]. Backhaus et al. [77] aimed to identify functional trait patterns across spatial scales and successional stages, discussing their underlying community assembly mechanisms. This implies that under varying drought conditions, communities adaptively adjust their trait composition to maintain stability and functionality. For instance, under prolonged drought stress, communities may selectively favor species with enhanced drought-tolerant traits, resulting in trait convergence phenomena.

4.4.3. Biodiversity Dimensions Mediating Ecosystem Resilience

Species diversity, particularly the diversity of functional traits, is recognized as a critical factor enhancing community drought resilience [78,79,80]. Research indicates that the positive effect of species richness on community drought resistance may be modulated by species evenness, operating through altered complementarity effects [81]. In forest communities, the coexistence of species with divergent drought-resistance strategies can bolster ecosystem-wide stability [13,78]. For instance, studies have shown that increased abundance of drought-tolerant species may compensate for declines in forest productivity by enhancing richness in drought-tolerant traits [82]. However, other research suggests that in multispecies grasslands, drought tolerance is more strongly determined by species identity and functional group diversity than by simple species diversity [83].

4.4.4. Successional Dynamics and Biotic–Abiotic Interactions

Community composition and functional traits undergo changes during succession to adapt to environmental pressures. Under drought conditions, trait convergence may signify that communities enhance overall resilience through selective retention of drought-tolerant species [77]. Interactions among species—including competitive and facilitative effects—can significantly influence community responses to drought [84]. In tropical seedling communities, neighboring diversity modulates seedling drought resistance [85]. Interactions between soil water and plants, along with their legacy effects, also play pivotal roles in drought resistance and resilience within arid ecosystems [33].

5. Quantification and Assessment Methods for Plant Drought Resilience

5.1. Multidimensional Metrics for Plant Drought Resilience Assessment

Current research employs diverse metrics and methodologies to evaluate plant drought resilience from multiple perspectives [19,83,86]. Physiological, biochemical, morphological, and resilience indices are critical for quantifying plant drought resilience (Table 2). Physiological indicators include water status markers, osmoprotectants (proline, soluble sugars), membrane stability (CMS), photosynthetic efficiency (Pn, Fv/Fm), antioxidant enzymes, and stomatal regulation. Morphological traits encompass biomass, root architecture, and leaf structure. Resilience is evaluated via Lloret [24] indices (resistance/recovery/resilience), satellite-derived productivity metrics (NDVI (Normalized Difference Vegetation Index), EVI (Enhanced Vegetation Index), GPP (Gross Primary Productivity)), and integrated statistical methods (PCA (Principal Component Analysis), membership functions). Multi-dimensional assessment enables robust analysis of plant adaptive capacity under drought stress.
Tree-ring analysis serves as the primary technique for quantifying resilience patterns through tree-ring width (TRWi) and density indices. In a representative study, Kang et al. [87] analyzed Betula platyphylla in semi-arid northern China, defining resistance as growth reduction magnitude during drought and resilience as post-disturbance recovery rate. This approach is constrained by its reliance on manual sampling for high-resolution data and limited capacity for capturing short-term dynamic responses. Notably, competition effects (e.g., neighbor interference) significantly modulate local resilience: increased stand density reduces individual tree resistance while enhancing resilience, necessitating quantification via mixed-effects models (as validated by a 166-case meta-analysis) [88]. Integrated remote sensing and ecological indices enable regional resilience mapping through satellite data (e.g., NDVI, GPP) coupled with ground validation. Zhang et al. [27] combined Leaf Area Index (LAI) with the Standardized Precipitation Evapotranspiration Index (SPEI), revealing that drought typology (meteorological vs. agricultural) governs vegetation response patterns, where drought timing (dry vs. wet seasons) critically alters recovery trajectories. Cross-scale evidence confirms biodiversity enhances resilience: Bai and Tang’s [89] analysis of 4072 global sites verified that species richness elevates resistance and resilience through functional redundancy and resource partitioning mechanisms, though climate zone dependencies (humid vs. arid regions) require differential validation.

5.2. Limitations and Emerging Directions in Plant Drought Resilience Assessment

However, these assessment approaches have certain limitations. For instance, physiological indicators are often sensitive to short-term environmental fluctuations, making it difficult to reflect the long-term drought adaptation of plants. Morphological traits, on the other hand, are relatively stable but have a slow response speed, which may lag behind the actual drought stress process. In recent years, with the development of molecular biology techniques, molecular markers have gradually been incorporated into the quantitative evaluation system of plant drought resilience. Genes related to drought resistance, such as those encoding dehydrins and transcription factors involved in stress signaling pathways, their expression levels and polymorphisms can provide new insights into the genetic basis of plant drought resilience [37,65]. Metabolomics analysis, which identifies and quantifies the changes in small molecule metabolites under drought stress, can also complement the traditional physiological and biochemical indicators, revealing the metabolic regulatory networks underlying plant drought adaptation.
Another important aspect is the integration of assessments at different scales. From the cellular level to the whole plant, and further to the community and ecosystem levels, there are complex interactions and feedbacks. For example, the drought resilience of individual plants may affect the structure and function of the entire community, and the microclimate and soil conditions of the community can in turn influence the drought response of individual plants [21,25,45]. Therefore, developing a multi-scale integrated evaluation framework is crucial for a comprehensive understanding of plant drought resilience.
In practical applications, there are also challenges in the quantitative evaluation of plant drought resilience. One of them is the lack of unified standards and criteria for different indicators and methods, which makes it difficult to compare and integrate results from different studies. For example, the threshold values of physiological indicators for different plant species under drought stress may vary greatly, and there is no universal reference standard. In addition, the cost and complexity of some advanced techniques, such as metabolomics and high-throughput sequencing, limit their widespread application in large-scale field studies.

6. Conclusions and Future Perspectives

Plant responses to drought stress represent a critical challenge for their survival and growth, particularly under global climate change where the intensity and frequency of drought events are escalating, posing severe threats to agricultural production and ecosystem stability [35,90,91]. To adapt to drought, plants employ complex multiscale and hierarchical mechanisms spanning morphological, physiological, biochemical, hydraulic, and anatomical traits [48,92,93]. A deeper understanding of these drought resilience mechanisms is essential for developing drought-tolerant crops and advancing ecological restoration. Despite significant advances in understanding plant drought resilience at the individual scale, key limitations persist. Current limitations in plant drought resilience research:
(a)
Lack of Standardized Metrics and Scale Effects: Inconsistent definitions and quantification methods for “resilience” across scales hinder cross-study comparisons. Understanding how micro-scale mechanisms (e.g., leaf physiology, gene expression) translate to macro-scale stability remains unclear.
(b)
Oversimplified Drought Characterization: Treating drought as a single event neglects the independent and combined impacts of key dimensions like severity, duration, and timing (seasonality). Modeling these complex interactions at large scales is challenging.
(c)
Neglect of Long-Term Effects and Recovery Dynamics: Focus remains on short-term resistance and immediate recovery. Understanding cumulative stress effects from repeated droughts, full recovery timelines/mechanisms, and global trends in the resistance–recovery trade-off is insufficient.
(d)
Complexity of Soil–Microbiome Interactions: While soil microbes (e.g., AMF) are known to enhance drought resilience, predicting how soil depth, plant community composition, and highly diverse, dynamic microbial communities collectively influence resilience at ecosystem scales is a major unresolved challenge.
(e)
Quantifying Diversity’s Role: Precisely measuring how different types of plant diversity (species, functional, genetic) contribute to resilience and integrating this into large-scale models is difficult. Simply increasing species number may not suffice; functional complementarity and the ecosystem-level impacts of genetic engineering need deeper investigation.
(f)
Data and Modeling Constraints: Scarcity of high-resolution, long-term ecosystem flux data limits fine-scale drought response analysis. Global hydrological resilience assessment needs better data/methods. Models struggle to capture complex drought dimension interactions and lack validation due to limited field control and long-term data.
Future research should target the identified gaps and limitations, with priorities on the following:
(a)
Integrated Multi-Scale Methodologies: Advancing frameworks that synergize physiological, biochemical, and molecular regulation through multi-omics approaches to identify resilience biomarkers, while integrating ecohydrology with the Soil–Plant–Atmosphere Continuum (SPAC) to quantify hydraulic traits and stomatal dynamics across drought types. Development of novel metrics for resistance (stress-withstanding capacity) and recovery (post-stress restoration) remains essential.
(b)
Long-Term Experimental Validation: Implementing multi-year, multi-scenario drought simulations to assess ecosystem resilience, with emphasis on rainfall pattern variations and Plant–Soil Feedback (PSF) dynamics. Research should elucidate how drought severity and historical precipitation modulate PSF-driven community assembly and ecosystem recovery.
(c)
Harnessing Plant–Microbe Synergies: Prioritizing rhizosphere microbiome engineering for staple crops to quantify trait relationships and legacy effects. Synergistic applications of bioengineered microbes, plant growth regulators (PGRs), and targeted miRNA editing warrant mechanistic exploration to enhance drought tolerance.
In summary, future research on plant drought resilience will constitute a comprehensive research framework that integrates multidisciplinary approaches and combines macro- and micro-level perspectives. By deepening our understanding of physiological, biochemical, and molecular mechanisms, leveraging advanced experimental methods and data analysis techniques, and harnessing the potential of plant–microbe interactions, we will be better equipped to address drought challenges posed by climate change, thereby securing global food security and ecosystem health.

Author Contributions

Conceptualization, W.L. and B.W.; methodology, B.W.; software, W.L.; formal analysis, W.L.; investigation, W.L.; data curation, L.W.; writing—original draft preparation, W.L.; writing—review and editing, B.W.; supervision, Y.G.; project administration, B.W.; funding acquisition, W.L. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by Chinese Academy of Forestry [CAFYBB2024MA043] and National Key R&D Program of China [2023YFF1305303-01].

Data Availability Statement

The data that support the findings of this study are available on request from the corresponding author upon reasonable request.

Conflicts of Interest

The authors declare no conflicts of interest.

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Water 17 02516 g001
Figure 1. Reillustration of scenarios where Lloret’s resilience indices may yield incomplete or misleading assessments of drought response (modified from [19]). Subfigures depict: (a) Lagged growth minimum occurring in year t + 1 following drought at year; (b) Significant pre-drought growth suppression within the reference period; (c) Multi-year drought events leading to prolonged recovery. Calculations (based on simulated detrended data with pre-drought growth set to 1) consider a 3-year pre-drought (blue lines) and post-drought (black lines) reference period. Red lines indicate growth in the drought year.
Figure 1. Reillustration of scenarios where Lloret’s resilience indices may yield incomplete or misleading assessments of drought response (modified from [19]). Subfigures depict: (a) Lagged growth minimum occurring in year t + 1 following drought at year; (b) Significant pre-drought growth suppression within the reference period; (c) Multi-year drought events leading to prolonged recovery. Calculations (based on simulated detrended data with pre-drought growth set to 1) consider a 3-year pre-drought (blue lines) and post-drought (black lines) reference period. Red lines indicate growth in the drought year.
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Figure 2. Dimensions of Drought Resilience. Schematic of vegetation growth dynamics and drought resilience responses under drought events. (a) Conceptual framework of key resilience strategies; (b) Temporal dynamics of resistance and recovery in two distinct systems (A and B). A and B refer to two trees or ecosystems.
Figure 2. Dimensions of Drought Resilience. Schematic of vegetation growth dynamics and drought resilience responses under drought events. (a) Conceptual framework of key resilience strategies; (b) Temporal dynamics of resistance and recovery in two distinct systems (A and B). A and B refer to two trees or ecosystems.
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Table 1. Historical evolution of key conceptual and methodological advances in plant drought resilience research.
Table 1. Historical evolution of key conceptual and methodological advances in plant drought resilience research.
StageTimeframeKey AdvancesSignificance
Conceptual Foundation1970s~90sFocus on drought resistance traits: osmoregulation, deep rooting, stomatal control;
Distinction between avoidance vs. tolerance.		Established physiological basis of drought survival.
Paradigm Shift to ResilienceEarly 2000s~2010sResilience triad: Resistance + Recovery + Elasticity;
Hydraulic failure-carbon starvation model;
Drought memory concept.		Shifted focus to post-stress recovery dynamics; linked physiology to mortality.
Multi-Omics Integration2010s–PresentTranscriptomic networks;
CRISPR (Clustered Regularly Interspaced Short Palindromic Repeats)-based resilience enhancement;
PGPR (Plant Growth-Promoting Rhizobacteria)-mediated rhizosphere resilience;
UAV (Unmanned Aerial Vehicle) phenomics.		Decoded molecular to ecosystem-scale mechanisms; enabled high-throughput screening.
Application and ScalingResilience-prioritized breeding; Microbial bioinoculants;
AI-driven irrigation models;
Mixed-species restoration.		Bridged theory to field practice; optimized resource use.
Table 2. Classification framework for multi-dimensional assessment metrics of plant drought resilience. Integrating physiological, biochemical, morphological, and comprehensive resilience indicators.
Table 2. Classification framework for multi-dimensional assessment metrics of plant drought resilience. Integrating physiological, biochemical, morphological, and comprehensive resilience indicators.
CategoryIndicatorDescription
Physiological and BiochemicalRelative Water Content (RWC)Measures plant tissue water status; declines under drought; correlates with drought resistance.
Osmotic Potential (Ψs)Quantifies osmotic adjustment capacity; critical for maintaining turgor under dehydration.
Proline (Pro) and Soluble Sugar (SS)Osmoprotectants maintain cell turgor and macromolecule integrity; accumulate during osmotic adjustment.
Cell Membrane Stability (CMS)Reflects membrane damage resistance; higher CMS indicates superior drought tolerance.
Photosynthetic Metrics (Pn, Fv/Fm, φPSII)Indicate light-use efficiency and photosynthetic apparatus integrity under drought.
Antioxidant Enzyme Activity Scavenges ROS to mitigate oxidative damage; enhanced under drought.
Stomatal Conductance (gs) Real-time gas exchange regulator; rapid decline triggers drought avoidance.
Stomatal Density (SD)Structural adaptation trait; higher density enhances gas exchange flexibility.
Hydraulic TraitsWater Potential (Ψ)Leaf/cellular water status indicator; negative values intensify with drought severity.
Xylem Embolism Resistance (P50)Critical threshold of water potential causing 50% hydraulic conductivity loss; key survival trait.
Leaf Turgor Loss Point (PTLP)Osmotic potential at cell turgor loss; determines stomatal closure threshold.
Specific Hydraulic Conductivity (Ks)Mass-specific xylem efficiency: conductivity per unit xylem area under pressure gradient.
MorphologicalBiomass and Growth (height, leaf area)Drought suppresses growth, reducing biomass and morphological parameters.
Root Morphology (e.g., root-to-shoot ratio)Deep/extensive roots enhance water uptake; root-to-shoot ratio indicates resource allocation.
Leaf Anatomy (thickness, area)Influences water-use efficiency and transpiration rates.
Resilience IndicesLloret Indices (Resistance/Recovery/Resilience)Quantifies system stability during disturbance (resistance), post-stress recovery, and overall functional maintenance.
Productivity-Based Metrics (NDVI/EVI/GPP)Satellite-derived indices assessing ecosystem-scale resistance and recovery.
Integrated Methods (PCA, membership function)Statistical approaches combining multiple indicators for holistic resilience evaluation.
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Lu, W.; Wu, B.; Wang, L.; Gao, Y. Multi-Scale Drought Resilience in Terrestrial Plants: From Molecular Mechanisms to Ecosystem Sustainability. Water 2025, 17, 2516. https://doi.org/10.3390/w17172516

AMA Style

Lu W, Wu B, Wang L, Gao Y. Multi-Scale Drought Resilience in Terrestrial Plants: From Molecular Mechanisms to Ecosystem Sustainability. Water. 2025; 17(17):2516. https://doi.org/10.3390/w17172516

Chicago/Turabian Style

Lu, Weiwei, Bo Wu, Lili Wang, and Ying Gao. 2025. "Multi-Scale Drought Resilience in Terrestrial Plants: From Molecular Mechanisms to Ecosystem Sustainability" Water 17, no. 17: 2516. https://doi.org/10.3390/w17172516

APA Style

Lu, W., Wu, B., Wang, L., & Gao, Y. (2025). Multi-Scale Drought Resilience in Terrestrial Plants: From Molecular Mechanisms to Ecosystem Sustainability. Water, 17(17), 2516. https://doi.org/10.3390/w17172516

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