Assessment of lima bean genotypes for high-temperature cultivation based on physiological and agronomic traitsABSTRACT
Rising temperatures resulting from climate change present a significant limitation to lima bean (Phaseolus lunatus L.) production, especially in hot and semiarid regions. This study evaluated lima bean genotypes under elevated-temperature conditions to identify heat-adapted lines based on vegetative, physiological, and yield-related traits. Measurements included vegetative growth parameters, gas exchange, photosystem II quantum efficiency, photosynthetic pigments, agronomic traits, and grain yield across 16 genotypes, encompassing both improved lines and landrace varieties. Significant differences were observed among genotypes for all evaluated characteristics, demonstrating considerable genetic variability. Lines H25-64, H94-31, and H25-54 exhibited superior physiological performance, as indicated by higher photosynthetic efficiency and favorable agronomic traits under elevated temperatures. Notably, line H81-32 displayed low growth and physiological performance but achieved the highest grain yield and hundred-grain weight, suggesting efficient allocation of photoassimilates to reproductive sinks. In contrast, lines H25-56 and H25-67 demonstrated high CO2 assimilation and stomatal conductance, yet this physiological advantage did not translate into increased grain yield. Landrace varieties exhibited traits linked to rusticity and morphological adaptation, such as larger pod dimensions, underscoring their value as sources of adaptive alleles. Multivariate analyses revealed that productivity under elevated temperatures is more closely associated with physiological efficiency than with vegetative biomass accumulation. The identified lines constitute promising parental materials for breeding programs targeting the development of heat-tolerant lima bean cultivars suited to warming environments, thereby supporting the sustainability of production systems under climate change.
Key words
heat tolerance; landraces;
Phaseolus lunatus
; plant breeding
INTRODUCTION
Population growth, together with climate change, poses an increasing challenge to ensuring food security in the 21st century. Projections indicate an average increase of 1.5°C in global temperature over the coming decades, making heat stress a significant threat to agricultural productivity (IPCC 2022, Wang et al. 2024). High temperature scenarios impair plant development, particularly during sensitive stages such as grain filling, thereby threatening the stability of global food production (Chaturvedi et al. 2021).
Heat stress causes damage throughout the entire phenological cycle, with effects varying by developmental stage. At the physiological level, heat stress promotes protein denaturation and disrupts metabolic pathways, leading to the formation of toxic intermediates and reactive oxygen species (ROS) that destabilize cellular membranes, proteins, and nucleic acids (Angon et al. 2024, Zhao et al. 2025).
These physiological damages manifest as morphological alterations. During vegetative growth, heat stress commonly results in reduced plant height, leaf area, and stem diameter, as well as chlorosis and leaf scorching (Zhao et al. 2025). At the reproductive stage, heat stress negatively affects fecundity, ovule viability, stigma receptivity, and pollen formation, and impairs fertilization and post-fertilization processes (Lohani et al. 2020, Ernest and Wisser 2024).
High temperatures during grain filling can decrease yield due to the production of small and malformed grains and inefficient allocation of photoassimilates. Under heat stress, energetic efficiency and photoassimilate allocation are key mechanisms for plant defense, especially during the reproductive stage when plants face a physiological trade-off between maintaining leaf photosynthetic activity and directing resources toward reproductive development (Chaturvedi et al. 2021). Consequently, plants dynamically adjust energy distribution among growth, maintenance, and defense processes.
Lima bean (Phaseolus lunatus L.) is a pulse crop of high nutritional value that is widely cultivated in regions facing environmental constraints (Garcia et al. 2021). Despite its importance, the species remains underrepresented in global statistics and is often grouped within the generic category of “dry beans” (Ernest and Wisser 2024). The crop grows across Africa, Asia, and the Americas, but there is not consistent data on productivity and economic value, which limits a comprehensive understanding of its socio-economic relevance. In Brazil, the estimated cultivated area in 2024 reached 33,928 ha, concentrated predominantly in the Northeast region (99.5%), with an average yield of 328 kg∙ha-1 (IBGE 2025).
Several reports indicate that lima bean is tolerant to drought and excess soil moisture but exhibits marked sensitivity to high temperatures, in contrast to common bean (Chacón-Sánchez and Martínez-Castillo 2017, Jacinto-Júnior et al. 2019, Machado et al. 2022). Optimal development occurs within a temperature range of 20–30°C (Sadok et al. 2020). However, information regarding the genetic variability of lima beans for heat-stress tolerance remains limited, constituting a major constraint for breeding programs.
The development of heat-tolerant cultivars, combined with a comprehensive understanding of plant physiological responses to elevated temperatures, constitutes a promising approach to ensuring food security in the context of global warming (Jin et al. 2024). It is hypothesized that physiological adjustments serve as critical determinants of growth and productive performance in lima bean under high-temperature conditions.
Accordingly, this study evaluated lima bean genotypes, including breeding lines and landraces, under heat stress to identify lines adapted to elevated temperatures based on vegetative, physiological, and yield-related traits. The identification of heat-tolerant genotypes is expected to support breeding programs, improve yield stability under climate warming, and enhance the sustainability of agricultural systems in regions highly vulnerable to climate change.
MATERIAL AND METHODS
Experimental site and climatic conditions
The experiment was conducted in the experimental area of the Department of Crop Science, Center for Agricultural Sciences, Universidade Federal do Piauí (UFPI), Teresina, Piauí state, Brazil (74.4 m above sea level; 05°05’21”S, 42°48’07”W). This location was selected due to its consistently high air temperatures, providing suitable conditions for evaluating crop performance under thermal stress. The regional climate is classified as Aw according to Köppen’s classification, characterized by a rainy summer and a dry winter. Environmental conditions during the experimental period are presented in Fig. 1, showing that daily air temperatures frequently exceeded 30°C, with maximum values reaching 38°C, representing temperature conditions considered stressful for the crop.
Maximum, minimum, and average monthly temperature in Teresina (PI, Brazil) during the period from February to August 2025.
Soil moisture was monitored and maintained at 80% of field capacity throughout the experimental period to ensure that the observed effects were solely attributable to heat stress. Irrigation was scheduled based on daily crop evapotranspiration, with irrigation depths (mm) calculated to maintain the prescribed soil moisture level.
Plant material
Sixteen genotypes of lima bean (Phaseolus lunatus L.) were evaluated—14 F10 advanced elite lines with determinate growth habit, and two landraces displaying indeterminate growth (Table 1; Fig. 2). Determinate genotypes are characterized by limited vegetative growth, compact architecture, reduced branching, and synchronized flowering.
Genotypes (lines and landraces), genealogy, and growth patterns were evaluated for thermal tolerance.
The elite lines originated from biparental crosses and were advanced via the modified bulk method to the F6 generation. Subsequently, lines were fixed at F7 and selected at F8 for agronomic traits such as earliness, short and erect lateral branches, and white seed color. The evaluated lines included H94-30, H94-31, H81-32, H81-33, H81-34, H25-53, H25-54, H25-56, H25-57, H25-58, H25-63, H25-64, H25-66, and H25-67.
Two landraces, UFPI-1356 (“fava-branca”) and VG5 (“boca-de-moça”), were included in the study (Fig. 2). The VG5 landrace, widely cultivated and preferred by smallholder farmers in Piauí state, Brazil, was used as the reference genotype to estimate the relative performance of the evaluated genotypes for each trait (Oliveira et al. 2026). Both landraces exhibit indeterminate growth, continuous vegetative development, increased branch elongation, and prolonged flowering.
Experimental design and crop management
A randomized complete block design with three replications was employed. Each plot contained 20 plants arranged in four rows of five, with 0.5 m between plants, 1 m between rows, 1.5 m between plots, and 2 m between blocks. Sowing involved three seeds per hill. Organic manure and basal mineral fertilization were applied in accordance with recommendations based on chemical analyses of soil samples collected from the 0–20-cm layer (Lopes et al. 2010).
The soil had a pH of 5.6, an organic matter content of 12.4 g∙kg-1, available phosphorus of 62.7 mg∙dm-3, potassium of 0.25 cmolc∙dm-3, calcium of 1.9 cmolc∙dm-3, magnesium of 0.93 cmolc∙dm-3, and base saturation of 86.4%. Gas exchange, photosynthetic pigments, and vegetative growth parameters were evaluated during the grain filling phase, 140 days after germination.
Agronomic assessment
Agronomic performance of all genotypes was evaluated using 13 descriptors for P. lunatus L., as proposed by Bioversity International (2007): days to flowering, defined as the period from emergence to when 50% of plants exhibited at least one open flower; days to maturity, the interval from emergence to when 50% of pods reached full maturity; pod length, pod width, pod thickness, seed length, seed width, and seed thickness, all measured in mm on 10 randomly selected mature pods; number of locules per pod, determined by counting funiculi in 10 pods; number of seeds per pod; 100-seed weight, calculated from the mass of 100 randomly sampled seeds and expressed in grams; pod mass of 10 pods, corresponding to the mass of 10 randomly selected pods; and grain yield, determined from the total grain weight in the usable plot area and expressed in grams, then converted to kg∙ha-1.
Physiological parameters
Gas exchange measurements were conducted on fully expanded, healthy leaves between 8 a.m. and 10:30 a.m. using a LI-6800 portable photosynthesis system (LI-COR Biosciences, Lincoln, NE, United States of America) under a constant photosynthetically active radiation of 1,000 µmol photons m-2∙s-1 and a CO2 concentration of 400 ppm. The following parameters were determined: net photosynthetic rate (µmol CO2 m-2∙s-1), transpiration rate (mmol H2O m-2∙s-1), internal CO2 concentration (µmol∙mol-1), ambient CO2 concentration (µmol∙mol-1), and stomatal conductance (mol H2O m-2∙s-1). Derived indices included Ci/Ca ratio, water-use efficiency (WUE = A/E), intrinsic water-use efficiency (iWUE = A/gs), and instantaneous carboxylation efficiency (A/Ci). The effective quantum yield of photosystem II (ΦPSII) was measured using a LI-600 fluorometer (LI-COR Biosciences, Lincoln, NE, United States of America).
Chlorophyll a, chlorophyll b, total chlorophyll, and carotenoids, quantified as mg∙g-1, were determined according to the protocol described by Wellburn (1994). Pigments were extracted from 1-cm2 leaf discs in dimethyl sulfoxide (DMSO) saturated with calcium carbonate. Following 72 h of incubation in darkness, the extracts were maintained in a water bath at 65°C for 1 h. Absorbance was measured at 665, 649, and 480 nm using a spectrophotometer, and pigment concentrations were calculated according to established methodology.
Vegetative traits
Fresh mass (g), plant dry mass (g), leaf dry mass (g), number of leaves, leaf area (cm2), relative water content, specific leaf area (cm2∙g-1), and leaf area index were quantified. Leaf area and number of leaves were assessed via a destructive sampling approach, wherein one plant per plot was harvested, scanned, and analyzed using the EBImage package in R. Fresh and dry masses were measured with an analytical balance. Specific leaf area was calculated as the ratio of leaf area to leaf dry mass. Relative water content was estimated based on the equation adapted from Mantovani (1999) (Eq. 1):
Leaf area index was calculated as the ratio of total leaf area to the ground area occupied per plant (0.16 m2; 40 × 40 cm).
Statistical analysis
Normality (Shapiro-Wilk’s test), homogeneity of variances (Levene’s test), and residual independence were assessed prior to analysis of variance (ANOVA). When appropriate, data were transformed using the Box-Cox method. Genotypes were grouped using the Scott-Knott’s test at a 5% significance level. Principal component analysis (PCA) and Pearson’s correlation analysis were performed, and the significance of the correlation coefficients was evaluated using Student’s t-test (p ≤ 0.05). Cluster analysis for all traits was performed using hierarchical grouping via the unweighted pair group method with arithmetic mean (UPGMA). Statistical analyses were carried out in R (version 4.5.0, 2025) with the ExpDes.pt, FactoMineR, factoextra, and metan packages, as well as GENES software (Cruz, 2013).
RESULTS
Vegetative performance genotypes
Lima bean genotypes, including both breeding lines and landraces, exhibited differential responses to the environmental conditions evaluated. Cultivation under elevated temperatures and thermal stress induced significant alterations in vegetative, physiological, and agronomic traits (Table 2).
Scott-Knott’s test results for fresh mass, plant dry mass, leaf dry mass, number of leaves, leaf area, relative water content, specific leaf area, and leaf area index in 16 lima bean genotypes.
The data demonstrated that H25-58, H94-30, UFPI-1356, H25-66, H25-67, and H81-33 exhibited superior vegetative performance compared to the VG5 reference genotype, as indicated by higher values of fresh mass, plant dry mass, leaf dry mass, number of leaves, specific leaf area, leaf area, and leaf area index. However, nearly all these lines displayed lower relative water content than VG5.
Agronomic performance
The analysis presented in Table 3 revealed variation in yield-related traits among the experimental lines and the VG5 landrace. For the phenological traits, all experimental lines exhibited lower values for days to flowering and days to maturity than the VG5 landrace, indicating a trend toward accelerated growth cycles. The only exception was the UFPI-1356 landrace, which exhibited higher days to flowering and days to maturity values and was grouped with VG5 for these traits.
Scott-Knott’s test for the traits: days to flowering (DF), days to maturity (DM), pod length (PL), pod width (PW), pod thickness (PT), seed length (SL), seed width (SW), seed thickness (ST), number of locules per pod (NLP), number of seeds per pod (NSP); 100-seed weight (HSW), pod mass of ten pods (PM10) and grain yield (GY).
Regarding seed and pod morphology, the lines exhibited reduced pod length, pod width, seed length, and seed width compared to the reference genotype, as indicated by higher-intensity values for these parameters. In contrast, a divergent trend was observed for grain yield: despite reduced seed size and shorter growth cycle, most experimental lines achieved higher productivity than VG5. Notably, lines H81-32, H25-64, H81-33, and H25-56 displayed the greatest increases in grain yield.
Physiological performance
Physiological responses to thermal stress varied among the genotypes relative to the VG5 reference (Table 4). For the quantum efficiency of photosystem II, most lines were grouped with VG5, whereas H94-30 exhibited the highest value and H81-34 the lowest.
Scott-Knott’s test results for the following traits in 16 lima bean genotypes: net photosynthetic rate, transpiration rate, stomatal conductance, effective quantum yield of photosystem II, ratio of internal to external CO2 concentration, water-use efficiency, intrinsic water-use efficiency, instantaneous carboxylation efficiency, chlorophyll a, chlorophyll b, total chlorophyll, and carotenoids.
For gas-exchange parameters, lines H25-54, H25-56, H25-57, H25-64, and H25-67 exhibited superior photosynthetic performance, with significantly higher net CO2 assimilation, stomatal conductance, and transpiration compared with VG5. Line H25-56 presented the highest values across these metrics.
Although most lines exhibited elevated photosynthetic rates, nearly all had water-use efficiency lower than that of VG5. However, instantaneous carboxylation efficiency was higher in lines such as H25-56 and H25-67. Considerable variability in photosynthetic pigment content was observed, with H25-58 and H81-33 exhibiting the highest chlorophyll a and b indices, whereas UFPI-1356 and H25-63 had the lowest total chlorophyll and carotenoid concentrations.
Correlations and multivariate analyses
Heatmap analysis identified a predominant pattern among genotypes under high-temperature stress: CO2 assimilation rate was negatively correlated with grain- and pod-filling parameters, as well as with the number of loci and seeds per pod, indicating a competitive partitioning of photoassimilates under stress (Fig. 3).
Pearson’s correlation coefficient between 32 traits evaluated in 16 lima bean genotypes under high temperature conditions.
Additionally, physiological traits were not significantly correlated with vegetative growth variables. A negative correlation between vegetative development and total grain yield supports the hypothesis that, under thermal stress, photoassimilate allocation is prioritized for vegetative maintenance over reproductive sinks.
Relative water content exhibited a negative correlation with productivity and seed development traits. These findings suggest that osmolyte production to enhance water regulation may reduce resource allocation for pod and seed development. In contrast, higher relative water content was positively associated with the quantum efficiency of photosystem II, possibly indicating that effective osmoregulation mitigates ROS damage, thereby maintaining photosynthetic efficiency.
PCA integrating vegetative, reproductive, and physiological traits explained 61.5% of the total variance (Fig. 4). Lines H25-56, H25-54, H25-64, and H25-67 clustered with gas-exchange parameters (net photosynthetic rate, stomatal conductance, transpiration rate), whereas landraces VG5 and UFPI-1356 were in opposing quadrants, characterized by extended phenological cycles and larger seed size but lower physiological efficiency and yield. Grain yield was positively associated with photosynthetic pigment content and was specifically linked to H81-32, which exhibited enhanced resilience to thermal stress.
Principal component analysis. Scatter plot of variables measured in lima bean genotypes under high temperature conditions.
Vegetative vigor traits (leaf area, leaf area index, and biomass) formed a distinct cluster associated with lines H25-58 and H25-66, indicating a dissociation between vegetative investment and reproductive success under thermal stress. A clear trade-off was evident between water-use efficiency and carboxylation efficiency, as genotypes with higher photosynthetic rates generally exhibited lower water-use efficiency due to increased transpirational demand for leaf cooling.
Hierarchical clustering (UPGMA) delineated four groups, highlighting genetic dissimilarity between improved lines and landraces with respect to thermal tolerance (cophenetic correlation coefficient, rc = 0.938; Fig. 5). Landraces displayed inferior performance under the imposed conditions, whereas breeding lines developed more effective tolerance mechanisms.
Dendrogram showing the dissimilarity among 16 lima bean genotypes from the core collection of the Germplasm Bank of Phaseolus at the Universidade Federal do Piauí (BGP-UFPI), cultivated under high temperature conditions. The dendrogram was generated using the unweighted pair group method with arithmetic mean (UPGMA) based on Gower’s distance (1971).
Line H25-58 was identified as an outlier, exhibiting extensive investment in vegetative traits to the detriment of yield. The most promising cluster, primarily composed of lines, combined high productivity with moderate CO2 assimilation and biomass production, indicating an optimized source-sink balance.
DISCUSSION
Lima bean cultivation under heat stress imposes physiological and metabolic constraints that vary among genotypes, reflecting distinct adaptive strategies related to biomass allocation, water management, and photoassimilate partitioning. The observed phenotypic variability among genotypes for most traits indicates that heat stress tolerance is not a uniform response. It rather comprises multiple strategies with different implications for yield. Morphological differences among genotypes highlight these diverse adaptive strategies for biomass allocation under elevated temperatures.
The landrace varieties UFPI-1356 and VG5 exhibited higher fresh and dry biomass, larger leaf area, and increased leaf area index, traits historically selected by farmers for robustness and environmental adaptation (Garcia et al. 2021, Lopes et al. 2024). However, this vegetative vigor incurs a high-water cost, as shown by the significant reduction in relative water content, indicating that maintaining large leaf area under heat stress increases transpiration and compromises water balance (Hočevar et al. 2025).
In contrast, lines such as H25-67 demonstrated a resource conservation strategy, characterized by a reduced leaf area and a high relative water content, suggesting more efficient water loss control under heat stress (Sadok et al. 2020). High relative water content is linked to effective osmotic adjustment through proline and compatible sugar synthesis, which also modulates ROS metabolism (Raza et al. 2023).
The phenotypic plasticity of improved lines, which can modulate plant architecture to reduce heat load, contrasts with the lower adaptive flexibility of landraces that, under high temperatures, remain vegetative without translating this investment into yield. Photoassimilate partitioning is a key determinant of productivity under thermal stress, as shown by the inverse relationship between vegetative growth and grain yield. The H81-32 line exemplified this, displaying the lowest biomass but achieving over 600 kg∙ha-1 in productivity, surpassing all other genotypes. This suggests a sink efficiency strategy, in which reduced vegetative investment directs carbon allocation to grain filling (Suzuki et al. 2023).
Similar strategies have been observed in other crops under heat stress, such as common beans, in which photoassimilate allocation and energy efficiency are crucial for plant defense during reproduction. This phase involves a physiological trade-off between sustaining photosynthesis and investing resources in reproductive structures (Lohani et al. 2020, Chaturvedi et al. 2021).
Landraces exhibited a longer vegetative cycle, twice as long as the lines, and higher vegetative investment, resulting in lower yield, a pattern also seen in H25-58. Although some species shorten their vegetative phase under high temperatures, certain Brassica spp. genotypes are insensitive to thermal changes and do not flower earlier, as regulation involves histone protein overexpression linked to flowering genes (Olmo et al. 2019).
Conversely, H81-32 overcame this phenological constraint by resolving the survival–reproduction balance early in favor of reproductive development. With the lowest biomass and shortest cycle, H81-32 demonstrates enhanced carbohydrate utilization, allocating photoassimilates primarily to grain filling (Suárez et al. 2020, Suzuki 2023, Vicentin et al. 2024).
The lack of significant differences in gas exchange among most lines and the reference genotype suggests that higher productivity is not due to increased photosynthetic capacity, but rather to metabolic adjustments that reduce respiratory costs and improve fixed carbon use efficiency (Sadok et al. 2020). This supports the view that the productivity bottleneck is associated with post-fixation carbon metabolism rather than carbon fixation itself (Wang et al. 2024, Palermo et al. 2025).
Lines such as H25-56 and H25-64, which sustained high transpiration rates and stomatal conductance, adopted a leaf cooling strategy, maintaining stomatal opening and CO2 uptake under high temperatures (Shao et al. 2021). Most genotypes preserved photosystem II quantum efficiency under stress, except for H81-34, which showed the lowest ΦPSII, indicating heat sensitivity and reduced capacity for energy dissipation (Shanker et al. 2022).
These results support previous findings that thermal tolerance is a quantitative trait governed by multiple genes (Jin et al. 2024). Therefore, heat stress tolerance in lima bean is multidimensional and cannot be attributed to a single physiological or agronomic trait. Integrated analysis identified two main strategies: water conservation and phenological escape, adopted by the most productive lines, and high vegetative investment, typical of landraces, which resulted in lower productivity under stress.
The analyses indicated that while lima bean genotypes experience limitations under heat stress, these are not directly associated with damage to the photosynthetic apparatus or reduced CO2 assimilation. Instead, the crop displays efficient water management for vegetative cooling and heat tolerance. However, this cooling strategy incurs energetic costs, which may explain reduced productivity and grain vigor, as stress conditions can lead to a 30% loss of produced energy due to increased respiratory demand.
The negative correlation between grain filling and CO2 assimilation rates suggests that, under heat stress, photoassimilates are preferentially directed toward osmolyte synthesis and maintaining vegetative integrity instead of grain reserve accumulation (Zaman et al. 2022). Machado et al. (2022) also reported that local lima bean varieties under heat stress produced many flowers but few seeds, indicating high energy investment in reproductive sinks without success due to reproductive structure sensitivity to temperature.
CONCLUSION
The evaluation of lima bean genotypes under heat-stress conditions revealed variation in physiological, vegetative, and yield-related responses, highlighting differences among the evaluated materials in maintaining performance under elevated temperatures. Among the breeding lines, H25-64, H94-31, and H25-54 showed promising responses associated with physiological and agronomic traits, while H81-32 exhibited favorable performance for grain yield and hundred-grain weight. These genotypes may represent useful sources of variability for further investigation in breeding programs targeting improved performance under high-temperature conditions. The landraces also contributed relevant phenotypic diversity, with traits related to plant architecture and adaptation that may support the exploration of genetic resources for crop improvement. Overall, the findings provide insights into the response of lima bean germplasm to thermal stress and may contribute to future breeding efforts aimed at developing cultivars with improved stability under high-temperature conditions.
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All data generated or analyzed during this study are included in this article.
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FM: fresh mass; PDM: plant dry mass; LDM: leaf dry mass; NL: number of leaves; LA: leaf area; RWC: relative water content; SLA: specific leaf area; LAI: leafarea index; DF: days to flowering; DM: days to maturity; PL: pod length; PW: pod width; PT: pod thickness; SL: seed length; SW: seed width; ST: seed thickness;NLP: number of locules per pod; NSP: number of seeds per pod; HSW: 100-seed weight; PM10: ten-pod mass; GY: grain yield; A: net photosynthetic rate; E:transpiration rate; gs: stomatal conductance; ΦPSII: effective quantum yield of photosystem II; WUE: water-use efficiency; iWUE: intrinsic water-use efficiency;EiC: instantaneous carboxylation efficiency; Chla: chlorophyll a; Chlb: chlorophyll b; Chlt: total chlorophyll; Car: carotenoids.
PCA: principal component analysis; FM: fresh mass; PDM: plant dry mass; LDM: leaf dry mass; NL: number of leaves; LA: leaf area; RWC: relative water content; SLA: specific leaf area; LAI: leaf area index; DF: days to flowering; DM: days to maturity; PL: pod length; PW: pod width; PT: pod thickness; SL: seed length; SW: seed width; ST: seed thickness; NLP: number of locules per pod; NSP: number of seeds per pod; HSW: 100-seed weight; PM10: pod mass of ten pods; GY: grain yield; A: net photosynthetic rate; E: transpiration rate; gs: stomatal conductance; ΦPSII: effective quantum yield of photosystem II; Ci/Ca: ratio of internal to external CO₂ concentration; WUE: water-use efficiency; iWUE: intrinsic water-use efficiency; EiC: instantaneous carboxylation efficiency; Chla: chlorophyll a; Chlb: chlorophyll b; Chlt: total chlorophyll; Car: carotenoids.
