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Water-use efficiency
Water-use efficiency
Water-use efficiency
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Water-use efficiency (WUE) refers to the ratio of plant biomass to water lost by transpiration, can be defined either at the leaf, at the whole plant or a population/stand/field level:

  • leaf level : photosynthetic water-use efficiency (also called instantaneous water-use efficiency WUEinst), which is defined as the ratio of the rate of net CO2 carbon assimilation (photosynthesis) to the rate of transpiration or stomatal conductance,[1] then called intrinsic water-use efficiency[2] (iWUE or Wi)
  • plant level : water-use efficiency of productivity (also called integrated water-use efficiency or transpiration efficiency,TE), which is typically defined as the ratio of dry biomass produced to the rate of transpiration.[3]
  • field level : based on measurements of CO2 and water fluxes over a field of a crop or a forest, using the eddy covariance technique[4]

Research to improve the water-use efficiency of crop plants has been ongoing from the early 20th century, however with difficulties to actually achieve crops with increased water-use efficiency.[5]

Intrinsic water-use efficiency Wi usually increases during soil drought, due to stomatal closure and a reduction in transpiration, and is therefore often linked to drought tolerance. Observations from several authors[3][6][7][8] have however suggested that WUE would rather be linked to different drought response strategies, where

  • low WUE plants could either correspond to a drought tolerance strategy, for example by anatomical adaptations reducing vulnerability to xylem cavitation, or to a drought avoidance/water spender strategy through a wide soil exploration by roots or a drought escape strategy due to early flowering
  • whereas high WUE plants could correspond to a drought avoidance/water saving strategy, through drought-sensitive, early closing stomata.

Increases in water-use efficiency are commonly cited as a response mechanism of plants to moderate to severe soil water deficits and have been the focus of many programs that seek to increase crop tolerance to drought.[9] However, there is some question as to the benefit of increased water-use efficiency of plants in agricultural systems, as the processes of increased yield production and decreased water loss due to transpiration (that is, the main driver of increases in water-use efficiency) are fundamentally opposed.[10][11] If there existed a situation where water deficit induced lower transpirational rates without simultaneously decreasing photosynthetic rates and biomass production, then water-use efficiency would be both greatly improved and the desired trait in crop production.

Water-use efficiency is also a much studied trait in Plant ecology, where it has been used already in the early 20th century to study the ecological requirements of Herbaceous plants[12] or forest trees,[13] and is still used today, for example related to a drought-induced limitation of tree growth[14]

References

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Water-use efficiency

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Water-use efficiency (WUE) is defined as the amount of carbon assimilated as or grain yield produced per unit of used by , typically through or . This metric serves as a key indicator of how effectively balances carbon gain and water loss, applicable across scales from individual leaves to entire ecosystems. In , WUE is critical for enhancing productivity under water-limited conditions, where efficient water use can increase yields without proportionally raising demands. For instance, genetic variations in crops like can lead to 19-23% differences in WUE, while management practices such as mulching have been shown to boost it by up to 61%. As global intensifies due to and , improving agricultural WUE is vital for , with elevated atmospheric CO₂ levels potentially enhancing it by up to 30% in C3 plants through improved . In and , ecosystem WUE reflects the coupling of carbon and water cycles, influencing biome-specific fluxes such as those in forests versus croplands. Global trends show terrestrial WUE has increased due to CO₂ fertilization effects on gross primary productivity, though recent saturation is linked to rising vapor pressure deficits from warming, with analyses up to 2018 indicating a continued rate of increase of 0.0016 gC m⁻² mm⁻¹ H₂O per year. Vegetation and meteorological droughts further modulate WUE, with lags in response affecting and ecosystem resilience. Challenges include scaling leaf-level measurements to canopy or ecosystem scales and predicting WUE under projected global temperature increases of 2.0–5.0°C by 2100 under various future emissions scenarios, underscoring the need for integrated genetic, management, and modeling approaches.

Definition and Concepts

Basic Definition

Water-use efficiency (WUE) is a fundamental concept in , defined as the ratio of carbon assimilated—typically measured as production or grain yield—per unit of consumed by or crops. This metric quantifies how effectively convert into productive output, reflecting the inherent trade-off between carbon fixation through and loss primarily via . Common units for WUE include kilograms of dry per cubic meter of (kg/m³) for crop-level assessments or grams of per of transpired (g/kg) for physiological evaluations. The concept of WUE originated in early 20th-century research, where it was first formalized by Lyman J. Briggs and Homer L. Shantz in their 1913 USDA bulletin The Water Requirement of Plants. In this seminal work, they experimentally determined the water requirements of various species grown in controlled conditions, establishing WUE as the produced divided by the water transpired, based on field trials in the . Their findings highlighted species-specific variations, laying the groundwork for understanding in arid environments and influencing subsequent agricultural practices. WUE can be distinguished at different scales: intrinsic WUE refers to the instantaneous leaf-level of net photosynthetic rate (A, in μmol CO₂ m⁻² s⁻¹) to (gₛ, in mol H₂O m⁻² s⁻¹), capturing real-time dynamics under varying environmental conditions. In contrast, integrated WUE encompasses the cumulative carbon assimilation relative to total use over longer growth periods, such as a full cropping cycle, integrating factors like and whole-plant . For instance, in agricultural contexts, integrated WUE is commonly expressed as (e.g., kg/ha) divided by seasonal use (e.g., plus in mm or m³), providing a practical measure for yield optimization.

Types of Water-use Efficiency

Water-use efficiency (WUE) encompasses several distinct types, differentiated primarily by the scale of measurement and the context of application, ranging from physiological processes in individual to broader economic considerations in . Instantaneous WUE refers to the ratio of net to at the leaf level, capturing short-term physiological responses to environmental conditions and proving valuable for studies on dynamics. This type emphasizes momentary carbon assimilation relative to water loss through stomata, often varying with factors like light intensity and deficit. Integrated or seasonal WUE measures accumulation or per unit of total consumed over an entire growing period, serving as a key metric in agricultural evaluations of long-term resource utilization. It integrates inputs from , , and soil storage to assess overall productivity, commonly applied to field crops like and to gauge varietal improvements. Ecosystem-level WUE quantifies net primary productivity or gross primary productivity per unit of across natural or managed landscapes, reflecting the balance of carbon and water cycles at larger scales. In arid regions, this type typically ranges from 0.5 to 2 g C/kg H₂O, influenced by vegetation composition such as shrubs or grasses that adapt to . Economic WUE extends the concept to incorporate financial outcomes, defined as the in a sector—such as agricultural yield value—per unit volume of water used, including irrigation costs to evaluate cost-effectiveness in water-scarce areas. This approach is particularly relevant for irrigated agriculture, where it accounts for from crops excluding rainfed components, aiding policy decisions on water allocation.

Measurement and Calculation

Field-scale Methods

Field-scale methods for measuring water-use efficiency (WUE) focus on integrating water inputs and outputs over entire plots or fields to quantify overall performance, typically expressed as seasonal integrated WUE. These approaches rely on agronomic measurements of water fluxes and yield, providing practical insights for management without delving into plant-level processes. The water balance method estimates (ET) by accounting for all water components in a defined field area, calculated as plus inputs minus the change in storage minus runoff and drainage. This technique allows for direct computation of crop water use, enabling WUE assessment through the ratio of or yield to total ET. It is particularly useful for evaluating efficiency in field trials, where boundary conditions like runoff collection are monitored to close the balance accurately. Yield-based calculations determine WUE as the ratio of harvestable yield to seasonal ET, often employing weighing lysimeters for precise quantification of water fluxes. Weighing lysimeters consist of soil monoliths containing , mounted on scales to measure mass changes attributable to ET, with accuracy typically within 5% for field-scale WUE estimates. Invented in the , these devices have become a standard for developing crop coefficients and validating schedules by capturing daily to seasonal water use dynamics. The general equation for crop WUE at the field scale is: WUE=grain yield (kg/ha)water use (mm)\text{WUE} = \frac{\text{grain yield (kg/ha)}}{\text{water use (mm)}} For , representative values range from 10 to 20 kg/ha/mm, reflecting variations in management and environment under optimal conditions. approaches estimate field-scale WUE by combining satellite-derived vegetation indices, such as the (NDVI), with models like the Surface Energy Balance Algorithm for Land (SEBAL) to derive production and ET. NDVI assesses crop canopy development to proxy , while SEBAL partitions surface energy fluxes for ET estimation, achieving accuracies of 85% on a daily basis and up to 95% seasonally at the field scale. This method facilitates large-area monitoring, supporting scalable WUE assessments in irrigated .

Physiological Methods

Physiological methods for assessing water-use efficiency (WUE) at the and level primarily involve direct measurements of photosynthetic and indirect proxies such as stable isotopes and , enabling insights into cellular and stomatal mechanisms under controlled conditions. These techniques focus on the ratio of carbon assimilation to water loss, providing mechanistic understanding without relying on whole-plant or field-scale integrations. Gas exchange analysis quantifies intrinsic WUE, defined as the ratio of net rate (A, in μmol CO₂ m⁻² s⁻¹) to (g_s, in mmol H₂O m⁻² s⁻¹), yielding units of μmol CO₂ per mmol H₂O. This instantaneous measure, often denoted as A/g_s, reflects the of CO₂ uptake relative to at the level and is commonly assessed using portable systems like the LI-6400 or LI-6800 gas analyzers (IRGAs), which simultaneously monitor CO₂ and H₂O fluxes in a controlled leaf chamber. Handheld porometers, such as the AP4 or SC-1, offer a complementary, rapid assessment of g_s by measuring diffusive resistance to through stomata, facilitating high-throughput -level evaluations. In C₃ under ambient CO₂ concentrations (around 400 μmol mol⁻¹), intrinsic WUE typically ranges from 50 to 100 μmol mol⁻¹, varying with environmental cues like vapor pressure deficit. Stable isotope analysis serves as a non-destructive proxy for long-term intrinsic WUE by examining carbon isotope discrimination (Δ¹³C) in leaf tissue, which inversely correlates with A/g_s. The relationship is approximated by the equation Δ¹³C ≈ a + (b - a) (c_i/c_a), where a (≈4.4‰) is the fractionation during CO₂ diffusion, b (≈27‰) is the net fractionation during carboxylation in C₃ plants, and c_i/c_a is the ratio of intercellular to ambient CO₂ concentration; higher WUE corresponds to lower Δ¹³C and thus less negative δ¹³C values (typically -28‰ to -22‰ in C₃ leaves). Developed in the 1980s, this method enables efficient screening of plant genotypes for inherent WUE traits by analyzing bulk leaf or seed carbon isotopes via mass spectrometry, avoiding the need for repeated gas exchange measurements over time. Chlorophyll fluorescence provides an indirect evaluation of WUE under water stress by monitoring (PSII) efficiency, as drought-induced stomatal closure limits CO₂ availability, reducing photochemical quenching and increasing non-photochemical dissipation. Key parameters include the maximum quantum efficiency of PSII (F_v/F_m, typically 0.75–0.85 in unstressed leaves) and the effective (Φ_PSII), which decline under water deficit, signaling impaired photosynthetic performance and associated WUE trade-offs. This optical technique, using pulse-amplitude modulated (PAM) fluorometers, allows real-time, non-invasive assessment of stress impacts on electron transport without direct gas measurements.

Factors Affecting Water-use Efficiency

Environmental Factors

Environmental factors play a critical role in influencing water-use efficiency (WUE) in , primarily through their effects on and rates. is a key driver, with optimal ranges varying by but generally between 15-30°C promoting balanced and photosynthetic activity, thereby maximizing WUE. Extremes beyond this range, such as high temperatures above 30°C, reduce WUE by elevating the vapor pressure deficit (VPD), which accelerates without a proportional increase in carbon assimilation, leading to greater loss relative to production. Atmospheric CO₂ concentration also significantly impacts WUE, as elevated levels—such as an increase of +200 ppm above ambient—can enhance it by 20-50% through partial closure of stomata, which conserves water while sustaining . This mechanism reduces rates, allowing plants to maintain higher intrinsic WUE under ambient to moderately elevated CO₂ conditions. availability and further modulate WUE; conditions lower it by inducing stomatal closure that limits both water uptake and , resulting in reduced overall per unit water used. decreases WUE in sensitive crops by impairing root water absorption and exacerbating osmotic stress, which induces partial stomatal closure and reduces yield; for example, in tomatoes, levels around 1 dS/m can reduce yield by 20-30%. Light intensity also affects WUE, with higher levels generally increasing carbon assimilation but potentially raising demands under high VPD. Humidity and wind interact to alter evaporative demand, with low humidity increasing transpiration rates and reduces WUE by widening the VPD gradient across leaf surfaces. Wind exacerbates this by enhancing boundary layer conductance, promoting faster water vapor diffusion from leaves and soil, which further strains plant water status without benefiting carbon gain. In regions like Mediterranean climates, seasonal rainfall variability can reduce WUE compared to more stable temperate zones due to erratic water availability that disrupts consistent transpiration control and growth.

Biological Factors

Biological factors significantly influence water-use efficiency (WUE) in through intrinsic physiological, genetic, and symbiotic mechanisms that optimize carbon assimilation relative to loss. These traits enable to maintain under varying internal constraints, primarily by enhancing resource capture and regulating . The photosynthetic pathway is a key determinant of WUE, with C4 such as demonstrating approximately 50% higher WUE than C3 like and . This advantage arises from the CO2 concentrating mechanism in C4 , which minimizes and allows for reduced stomatal opening while sustaining high rates of carbon fixation. Root architecture also plays a crucial role, particularly through the development of deep root systems that enhance access in dry soils. Deep roots can improve WUE by 15-25% by tapping into subsurface reserves, thereby increasing water uptake without proportionally raising . Stomatal regulation, mediated by (ABA), further modulates WUE by promoting stomatal closure in response to water stress signals. This ABA-induced response conserves by limiting , though it simultaneously restricts CO2 uptake and can constrain photosynthetic rates. Microbial interactions, especially with mycorrhizal fungi, bolster WUE by extending the root system's absorptive capacity. Arbuscular mycorrhizal fungi can increase WUE by about 20% through improved uptake and enhanced water absorption via their hyphal networks, which facilitate nutrient and water transport to the host plant. At the genetic level, quantitative trait loci (QTLs) for WUE have been identified in crops like since the early , often linking to genes such as ERECTA that control stomatal density. These QTLs influence transpiration efficiency by altering stomatal characteristics, providing targets for breeding higher WUE varieties.

Importance and Applications

Agricultural Applications

In agricultural systems, water-use efficiency (WUE) is pivotal for optimizing crop yields while conserving limited , particularly in irrigated and rainfed farming contexts where affects productivity. Deficit irrigation, a key scheduling strategy, applies less water than full crop needs during non-critical growth stages, thereby enhancing WUE without severely compromising yields. For instance, in production, deficit irrigation strategies have been shown to enhance WUE compared to full irrigation, as it aligns water supply more closely with physiological demands, reducing losses and promoting deeper systems. Crop rotation combined with residue management further bolsters WUE by improving and water retention. Incorporating cereal crop residues into the elevates content, which enhances infiltration and reduces runoff, leading to WUE improvements of 7.5-25.5% depending on initial organic levels; in soils with 10-15 g/kg , gains average around 19%. This practice is especially beneficial for cereals like and , where residue retention mitigates and sustains moisture availability during dry spells. Benchmarking WUE in major crops provides context for global agricultural performance. For maize, a staple crop, global averages range from 1.2-2.3 kg/m³ of water, reflecting variations across irrigated and rainfed systems; however, in rainfed agriculture of sub-Saharan Africa, WUE often falls below 1 kg/m³ due to erratic rainfall, poor soil fertility, and limited management, exacerbating yield gaps of up to 47% relative to potential. These benchmarks underscore the need for targeted interventions in low-efficiency regions. Economically, enhancing WUE in water-scarce areas like yields substantial benefits by lowering input costs and bolstering . Improved WUE through efficient practices can reduce overall -related expenses by up to 20-30%, including pumping and infrastructure, while supporting higher net farm incomes—potentially increasing them by 30% in groundwater-dependent regions—and enabling sustained production for over 600 million agricultural dependents. FAO analyses from 2020 on precision farming highlight how such approaches can boost yields by 10-15% with 20-40% less , amplifying these economic and outcomes without delving into specific agronomic details.

Ecological Applications

In natural ecosystems, water-use efficiency (WUE) serves as a key indicator of drought resilience, particularly in forests and grasslands where water availability shapes ecosystem dynamics. During drought events, many forest and grassland systems exhibit increased WUE as plants reduce transpiration while maintaining or recovering carbon assimilation, demonstrating adaptive resilience. For instance, in semi-arid savannas, long-term WUE typically ranges from 1 to 3 g C per kg H₂O, reflecting efficient resource use that allows these ecosystems to persist under variable rainfall regimes. This resilience is evident in post-drought recovery, where arid ecosystems show significant WUE elevation, underscoring their capacity to rebound from water stress without long-term productivity losses. Elevated atmospheric CO₂ concentrations further enhance ecosystem WUE by improving and stomatal control, which in turn boosts . Models indicate that this CO₂ fertilization effect can increase carbon storage in forests and grasslands by 10-20% under future scenarios, as reduced water loss per unit of carbon fixed amplifies net productivity. Such improvements are particularly pronounced in semi-arid regions, where higher WUE mitigates water limitations and supports greater accumulation over time. In arid biomes, species exhibiting high intrinsic WUE often dominate community structures, influencing overall and composition. These adaptations, such as enhanced stomatal regulation and deep root systems, allow drought-tolerant like shrubs and perennial grasses to outcompete others, thereby shaping patterns and maintaining stability amid . This selective dominance fosters hotspots in resilient communities, where high-WUE species contribute to and provision. Ecological restoration projects leverage WUE monitoring to assess success in degraded landscapes, as seen in China's , where large-scale since 2000 has significantly elevated WUE through increased vegetation cover and gross primary productivity relative to . in tropical forests diminishes global carbon sinks, as it leads to carbon release and weakens the forests' role in mitigating .

Strategies for Improvement

Agronomic Practices

Agronomic practices play a crucial role in enhancing water-use efficiency (WUE) in agricultural systems by optimizing retention, reducing unproductive water losses, and aligning resource inputs with needs. These field-level techniques, including mulching, reduced , and , promote sustainable water use without relying on genetic modifications or advanced technologies. By minimizing and improving , such practices can lead to measurable gains in WUE, particularly in water-limited environments like production. Mulching and cover cropping are effective strategies for conserving soil water in row crops such as and . These methods create a physical barrier on the surface, significantly reducing losses and thereby increasing the proportion of available for and crop growth. For instance, mulching has been shown to boost WUE by approximately 20% in both and under varying inputs, while mulching can achieve up to 28% improvement in systems at higher inputs (>250 mm). Cover crops further enhance this by improving and infiltration, contributing to long-term retention in row crop fields. Reducing , particularly through no-till systems, conserves soil by preserving residue cover and minimizing soil disturbance. In production, such practices have been associated with enhanced WUE due to better moisture storage and reduced runoff, with yield and efficiency gains in semi-arid regions. Similarly, optimizing application, especially balanced (N) rates, avoids excessive luxury uptake that diverts to non-productive growth, potentially enhancing WUE by 10-15%. This is achieved by synchronizing N supply with demand, as seen in where integrated and N improved by 15%. Crop selection tailored to local conditions also elevates WUE; for example, drought-tolerant varieties like in semi-arid zones exhibit higher WUE than less adapted cereals such as due to efficient systems and reduced under stress. These practices collectively underscore the potential for agronomic management to sustain amid .

Genetic and Technological Approaches

Genetic approaches to enhancing water-use efficiency (WUE) in crops have advanced through marker-assisted breeding and transgenic methods, targeting genes associated with stomatal control to optimize water loss and carbon assimilation. For instance, transgenic manipulation of genes like TaEPF1 has reduced stomatal density in , leading to improved intrinsic WUE without severe yield loss. Breeding programs continue to identify traits for reduced under conditions. Genome editing technologies, particularly , offer precise modifications to genes involved in water transport. Studies have demonstrated improvements in and reduced water loss in crops like through genetic modifications related to aquaporins, enhancing WUE under water-limited conditions. Technological innovations in integrate sensors, IoT devices, and to enable real-time management, particularly in water-intensive crops like grapes. In vineyards, AI algorithms analyze data from sensors, weather forecasts, and plant stress indicators to automate , achieving water savings of 25-40% while sustaining or improving fruit quality and yield. Recent studies report broader of 30-60% in precision systems across crops. Controlled environment agriculture (CEA), including , represents a high-tech for maximizing WUE by recirculating solutions and eliminating soil evaporation losses. In , use can be as low as 10% of that required in open-field production for crops like and tomatoes, translating to 5-10 times higher WUE due to precise delivery and reuse of . This approach is especially impactful in urban or arid settings where is paramount. A notable application of these genetic advancements is the Water Efficient Maize for Africa (WEMA) project, which has developed and disseminated drought-resistant varieties incorporating traits for improved WUE and resistance. As of October 2025, these and related TELA varieties have reached millions of smallholder farmers across , impacting up to 44 million people and resulting in yield improvements of 24-35% under stress compared to traditional , with some cases reporting up to 54% increases.

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