Agricultural Crops Subjected to Drought and Salinity Stress

A special issue of Agriculture (ISSN 2077-0472). This special issue belongs to the section "Crop Production".

Deadline for manuscript submissions: 31 August 2024 | Viewed by 15789

Special Issue Editors


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Guest Editor
Department of Agronomic and Forest Science, Federal Rural University of the Semi-Arid—UFERSA, Mossoró 59625-900, Brazil
Interests: salt stress; drought stress; irrigation; water management; wastewater; alkaline soils; horticulture; plant physiology; plant ecophysiology; plant nutrition
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Guest Editor
Departamento de Biologia, Universidade Estadual da Paraíba, Bairro Universitário, Campina Grande 58429-500, PB, Brazil
Interests: salt stress; drought stress; irrigation; water management; horticulture; plant physiology; plant ecophysiology; Vigna unguiculata; phenotyping
Special Issues, Collections and Topics in MDPI journals

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Guest Editor
Ciências Ambientais e Tecnológicas, Universidade Federal Rural Do Semi-Árido, Mossoró, Belize
Interests: salt stress; drought stress; irrigation; water management; wastewater; Alkaline soils; horticulture; plant nutrition; Cocos nucifera

Special Issue Information

Dear Colleagues,

Agriculture is historically vital to the prosperity of civilizations and has withstood the pressure of the environment and population growth due to genetic improvement and plant management. Most agricultural crops are subject to environmental stresses such as drought and salinity. In many cases, these stresses act together, limiting crop productivity. In this view, innovative management strategies can improve the productivity of agricultural crops subjected to unfavorable environmental conditions, such as drought and salinity.

This Special Issue focuses on developing and evaluating management strategies for crops subjected to drought and salt stress. For this reason, it welcomes studies of an interdisciplinary nature from research fields related to agriculture, including horticulture, genetics, plant ecophysiology, irrigation, soils, and plant nutrition. Research articles will cover various agricultural crops and solutions for growing them under drought and salt stress conditions. Original research articles and reviews will be accepted.

Prof. Dr. Francisco Vanies Da Silva Sá
Prof. Dr. Alberto Soares De Melo
Prof. Dr. Miguel Ferreira Neto
Guest Editors

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Keywords

  • horticulture
  • field crops
  • cereal crops
  • industrial crops
  • plant physiology
  • plant ecophysiology
  • plant nutrition
  • irrigation management
  • plant breeding
  • crosstalk stress

Published Papers (9 papers)

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15 pages, 5392 KiB  
Article
The Identification of Drought Tolerance Candidate Genes in Oryza sativa L. ssp. Japonica Seedlings through Genome-Wide Association Study and Linkage Mapping
by Tao Liu, Shuangshuang Li, Haoqiang Du, Jingnan Cui, Shanbin Xu, Jingguo Wang, Hualong Liu, Detang Zou, Wenhe Lu and Hongliang Zheng
Agriculture 2024, 14(4), 603; https://doi.org/10.3390/agriculture14040603 - 10 Apr 2024
Viewed by 1040
Abstract
Drought stress poses a significant threat to rice production, necessitating the identification of genes associated with drought tolerance. This study employed a combination of genome-wide association study (GWAS) and linkage mapping to pinpoint seedling drought tolerance genes in Japonica rice. Using the leaf [...] Read more.
Drought stress poses a significant threat to rice production, necessitating the identification of genes associated with drought tolerance. This study employed a combination of genome-wide association study (GWAS) and linkage mapping to pinpoint seedling drought tolerance genes in Japonica rice. Using the leaf rolling scale (LRS) as the phenotypic index, we assessed rice drought tolerance under polyethylene glycol-induced drought during the seedling stage. A lead SNP C8_28933410 by GWAS was identified, which was located within qLRS-8-1 identified by linkage mapping on chromosome 8. Combing the LD block analyses and QTL interval, a 138.6 kb overlap interval was considered as the candidate region. Haplotype analysis, qRT-PCR, sequence analysis, and mutant phenotype verification led to the speculation that LOC_Os08g05520 is a candidate gene associated with drought tolerance. Our findings provide a valuable reference for breeders aiming to enhance rice drought tolerance. Full article
(This article belongs to the Special Issue Agricultural Crops Subjected to Drought and Salinity Stress)
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23 pages, 1515 KiB  
Article
Effects of Silicon Alone and Combined with Organic Matter and Trichoderma harzianum on Sorghum Yield, Ions Accumulation and Soil Properties under Saline Irrigation
by José Orlando Nunes da Silva, Luiz Guilherme Medeiros Pessoa, Emanuelle Maria da Silva, Leonardo Raimundo da Silva, Maria Betânia Galvão dos Santos Freire, Eduardo Soares de Souza, Sérgio Luiz Ferreira-Silva, José Geraldo Eugênio de França, Thieres George Freire da Silva and Eurico Lustosa do Nascimento Alencar
Agriculture 2023, 13(11), 2146; https://doi.org/10.3390/agriculture13112146 - 14 Nov 2023
Cited by 1 | Viewed by 1588
Abstract
The action of silicon as a salt stress mitigator has been investigated in isolation, and its combined efficacy with other salt stress mitigators needs to be addressed. This work verified whether silicon, in combination with organic matter and Trichoderma harzianum, enhances the [...] Read more.
The action of silicon as a salt stress mitigator has been investigated in isolation, and its combined efficacy with other salt stress mitigators needs to be addressed. This work verified whether silicon, in combination with organic matter and Trichoderma harzianum, enhances the production of forage sorghum under saline irrigation and its effects on soil properties. The field experiment was conducted in Parnamirim (PE), a semiarid region of Brazil. Forage sorghum (Sorghum sudanense (Piper) Stapf) was irrigated with saline water (3.12 dS m−1) and subjected to the application of non-silicon, silicon alone, and silicon combined with Trichoderma and organic matter over three consecutive cuts (every three months after germination). Silicon applied in combination significantly increased the content of nutrient ions K+, P, Ca2+, and Mg2+ in sorghum leaves, stems, and panicles and increased P content in the soil by 170, 288, and 92% for the first, second, and third cuts, respectively. When silicon was applied in combination, sorghum’s dry and fresh matter (total yield for the three cuts) increased to 62.53 and 182.43 t ha−1, respectively. In summary, applying silicon (Si) combined with Trichoderma and organic matter promotes higher nutrient ion contents in soil and sorghum plants and a higher forage sorghum yield. Full article
(This article belongs to the Special Issue Agricultural Crops Subjected to Drought and Salinity Stress)
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28 pages, 2608 KiB  
Article
Integrating Agro-Morpho-Physiological Traits and SSR Markers for Detecting the Salt Tolerance of Advanced Spring Wheat Lines under Field Conditions
by Muhammad Bilawal Junaid, Salah El-Hendawy, Ibrahim Al-Ashkar, Nasser Al-Suhaibani and Majed Alotaibi
Agriculture 2023, 13(11), 2135; https://doi.org/10.3390/agriculture13112135 - 12 Nov 2023
Cited by 1 | Viewed by 1366
Abstract
To successfully enhance the salt tolerance of genotypes, it is crucial to conduct field-based trials, establish effective screening criteria and analysis tools, evaluate salt tolerance at various growth stages, and integrate phenotypic assessment-based traits with molecular markers. This study aimed to assess the [...] Read more.
To successfully enhance the salt tolerance of genotypes, it is crucial to conduct field-based trials, establish effective screening criteria and analysis tools, evaluate salt tolerance at various growth stages, and integrate phenotypic assessment-based traits with molecular markers. This study aimed to assess the salt tolerance of 16 F8 recombinant inbred lines (RILs) and eight genotypes by analyzing 13 agro-morpho-physiological traits using various analysis tools and SSR markers under both control and high salinity levels (15 dS m−1) in real field conditions. Analysis of variance (ANOVA), comparison of mean values, calculation of reduction percentage, and multivariate analysis were used to compare the assessed traits among genotypes and identify which traits are the most effective ones in describing the salt tolerance of these genotypes. A heatmap cluster analysis (HMCA) was also employed to categorize the salt tolerance of genotypes into different clusters based on the stress tolerance index (STI) for all traits. The ANOVA results revealed significant statistical differences (p ≤ 0.05) between the genotypes and salinity levels for all assessed traits in each season and their combined data. Moreover, the 150 mM NaCl treatment led to decreases in the assessed traits by 10.2% to 36.9% when compared to the control treatments. Furthermore, the mean values of assessed traits for certain genotypes were approximately one to three times greater than those of other genotypes. Principal component analysis has identified plant dry weight, green leaf area, leaf area index, and grain yield per hectare as effective screening criteria for explaining the substantial variation observed among the genotypes. The HMCA successfully grouped genotypes into three distinct clusters and distinguished the salt-tolerant genotypes from the salt-sensitive and intermediate ones. The 24 genotypes/RILs were classified into three main groups according to the allelic data of 40 SSRs associated with salt-tolerant genes. A weak yet significant correlation was observed between the similarity coefficients of agro-morpho-physiological traits and SSR markers, as determined by the Mantel test (r = 0.13, p < 0.03, and alpha = 0.05). In conclusion, this study has successfully identified several traits, particularly those associated with SSR markers, that greatly contribute to our understanding of the phenotypic and genotypic basis influencing the salt tolerance of wheat genotypes in real field conditions. Consequently, assessing these traits for a large number of wheat plant materials in a rapid and cost-effective manner will be greatly importance in breeding programs aimed at improving salt stress tolerance in this vital food crop. This will be the main focus of our forthcoming research. Full article
(This article belongs to the Special Issue Agricultural Crops Subjected to Drought and Salinity Stress)
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14 pages, 1278 KiB  
Article
Effect of Electrical Conductivity Levels and Hydrogen Peroxide Priming on Nutrient Solution Uptake by Chives in a Hydroponic System
by Patrícia Ferreira da Silva, Bárbara Davis Brito dos Santos, José Dantas Neto, Alberto Soares de Melo, Rigoberto Moreira de Matos, Semako Ibrahim Bonou, Tonny José Araújo da Silva, Edna Maria Bonfim-Silva, Ana Paula Candido Gabriel Berilli and Thiago Franco Duarte
Agriculture 2023, 13(7), 1346; https://doi.org/10.3390/agriculture13071346 - 3 Jul 2023
Cited by 1 | Viewed by 2146
Abstract
The use of water of high electrical conductivity has become common in hydroponic systems, especially in regions with water scarcity. However, the use of inferior-quality water can affect crop yields. In this scenario, some studies have tested the use of chemical conditioning agents [...] Read more.
The use of water of high electrical conductivity has become common in hydroponic systems, especially in regions with water scarcity. However, the use of inferior-quality water can affect crop yields. In this scenario, some studies have tested the use of chemical conditioning agents such as hydrogen peroxide to minimize the negative effects of stress on plants. From this perspective, this study aimed to evaluate the action of priming with hydrogen peroxide as a salt stress attenuator on the nutrient solution uptake and productivity of chives in a hydroponic system. The study was conducted in a protected environment with a randomized block design with a split-plot arrangement. The treatments consisted of a main plot consisting of the electrical conductivity of the nutrient solution (1.0, 2.0, 3.0, 4.0, and 5.0 dSm−1) and a subplot with five hydrogen peroxide concentrations (0.0, 0.15, 0.30, 0.45, and 0.60 mM). The increase in the electrical conductivity of the nutrient solution reduced bulb length, the solution volume applied, water uptake, total fresh mass, and the solution use efficiency by plants. Throughout the cultivation cycle in the hydroponic system, the consumption of nutrient solution was 459 mm lost by evapotranspiration. Acclimation with 0.60 mM hydrogen peroxide associated with 1 dSm−1 of electrical conductivity of the nutrient solution favors bulb diameter in chives. The increase in electrical conductivity compromises the productive yield of chives. Full article
(This article belongs to the Special Issue Agricultural Crops Subjected to Drought and Salinity Stress)
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19 pages, 3885 KiB  
Article
Beneficial Microorganisms Affect Soil Microbiological Activity and Corn Yield under Deficit Irrigation
by Josinaldo Lopes Araujo, Jackson de Mesquita Alves, Railene Hérica Carlos Rocha, José Zilton Lopes Santos, Rodolfo dos Santos Barbosa, Francisco Marcelo Nascimento da Costa, Geovani Soares de Lima, Leandro Nunes de Freitas, Adriana Silva Lima, Antonio Elizeneudo Peixoto Nogueira, André Alisson Rodrigues da Silva, Leônidas Canuto dos Santos, Francisco Bezerra Neto and Francisco Vaniés da Silva Sá
Agriculture 2023, 13(6), 1169; https://doi.org/10.3390/agriculture13061169 - 31 May 2023
Cited by 1 | Viewed by 1898
Abstract
Water scarcity is one of the main factors that decrease the growth and productivity of corn, since it negatively affects gas exchange and the general metabolism of the crop. The use of beneficial microorganisms (BM) has been considered a potential attenuator of water [...] Read more.
Water scarcity is one of the main factors that decrease the growth and productivity of corn, since it negatively affects gas exchange and the general metabolism of the crop. The use of beneficial microorganisms (BM) has been considered a potential attenuator of water stress. This study aimed to evaluate the effect of BM and water deficit on growth, gas exchange, grain yield, and soil microbial activity. A field experiment was carried out, in which the treatments were composed of a 2 × 4 factorial scheme, corresponding to two irrigation levels (100% of ETc and 50% of ETc) and to four treatments (T) referring to the soil inoculation with BM (C: control; T1: Bacillus amyloliquefaciens + Azospirillum brasiliense; T2: B. subtilis; and T3: A. brasiliense). The evaluations were carried out in the flowering phase (plant growth, gas exchange, and foliar nitrogen content) and at the end of the plant cycle (grains yield, mineral nitrogen, and microbiological activity). The 50% reduction in irrigation depth severely restricted corn growth and gas exchange and decreased the grain yield by 38%. The water deficit increased the protein content in the grains and the concentration of mineral nitrogen in the soil when the plants were inoculated with BM. Under water stress, inoculation with BM increased corn productivity by 35% and increased soil microbial activity. The inoculation of plants with BM, either in combination (Bacillus amyloliquefaciens + A. brasiliense) or alone (B. subtilis), attenuated the adverse effects of water deficit in maize. Full article
(This article belongs to the Special Issue Agricultural Crops Subjected to Drought and Salinity Stress)
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14 pages, 1094 KiB  
Article
Discontinuous Hydration Cycles with Elicitors Improve Germination, Growth, Osmoprotectant, and Salt Stress Tolerance in Zea mays L.
by Kleane Targino Oliveira Pereira, Salvador Barros Torres, Emanoela Pereira de Paiva, Tatianne Raianne Costa Alves, Maria Lilia de Souza Neta, Jefferson Bittencourt Venâncio, Lauter Silva Souto, Clarisse Pereira Benedito, Tayd Dayvison Custódio Peixoto, Miguel Ferreira Neto, Nildo da Silva Dias and Francisco Vanies da Silva Sá
Agriculture 2023, 13(5), 964; https://doi.org/10.3390/agriculture13050964 - 27 Apr 2023
Cited by 2 | Viewed by 1528
Abstract
Saline stress impairs germination and initial plant growth. However, discontinuous hydration cycles induce osmotic tolerance in seeds and can improve the response of maize seeds to saline stress. The objective of this study was to evaluate the action of discontinuous hydration cycles with [...] Read more.
Saline stress impairs germination and initial plant growth. However, discontinuous hydration cycles induce osmotic tolerance in seeds and can improve the response of maize seeds to saline stress. The objective of this study was to evaluate the action of discontinuous hydration cycles with different salt stress tolerance elicitors on germination, growth, and osmotic adjustment of maize cultivars. Maize seeds of BR 206 and BRS 5037 Cruzeta cultivars were subjected to the following treatments: 0.0 mmol of NaCl (control), 250 mmol of NaCl (salt stress), salt stress + three discontinuous hydration cycles (DHCs) of seeds in water, salt stress + DHCs with gibberellic acid, salt stress + DHCs with hydrogen peroxide, salt stress + DHCs with salicylic acid, and salt stress + DHCs with ascorbic acid. Salt stress reduced the germination, growth, and biomass accumulation in maize seedlings—the BR 206 cultivar outperformed BRS 5037 Cruzeta. Discontinuous hydration cycles with water failed to improve the salt stress tolerance of maize seeds. However, discontinuous hydration cycles with gibberellic acid, hydrogen peroxide, and salicylic acid promoted salt stress tolerance in maize due to increased synthesis of osmoprotectants. Our results revealed salicylic acid is appropriate for discontinuous hydration cycles in maize seeds. Full article
(This article belongs to the Special Issue Agricultural Crops Subjected to Drought and Salinity Stress)
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22 pages, 4451 KiB  
Article
Growth, Solute Accumulation, and Ion Distribution in Sweet Sorghum under Salt and Drought Stresses in a Brazilian Potiguar Semiarid Area
by Gabriela Carvalho Maia de Queiroz, José Francismar de Medeiros, Rodrigo Rafael da Silva, Francimar Maik da Silva Morais, Leonardo Vieira de Sousa, Maria Vanessa Pires de Souza, Elidayane da Nóbrega Santos, Fagner Nogueira Ferreira, Juliana Maria Costa da Silva, Maria Isabela Batista Clemente, Jéssica Christie de Castro Granjeiro, Matheus Nathan de Araújo Sales, Darcio Cesar Constante, Reginaldo Gomes Nobre and Francisco Vanies da Silva Sá
Agriculture 2023, 13(4), 803; https://doi.org/10.3390/agriculture13040803 - 31 Mar 2023
Cited by 6 | Viewed by 1781
Abstract
Agriculture in semiarid regions commonly face problems because of salt and availability of irrigation water. Considering this, studies on cultures resistant to salt and water stresses involving sweet sorghum are required. Therefore, the aim was to evaluate the growth and other mechanisms of [...] Read more.
Agriculture in semiarid regions commonly face problems because of salt and availability of irrigation water. Considering this, studies on cultures resistant to salt and water stresses involving sweet sorghum are required. Therefore, the aim was to evaluate the growth and other mechanisms of tolerance to salinity and water deficit in BRS 506 sweet sorghum. The experimental design was conducted in Upanema-RN, Brazil, in randomized blocks, where the isolated and interactive effect of 3 salinity levels, expressed as the electrical conductivity of irrigation water (1.5, 3.8, and 6.0 dS m−1), and 3 irrigation depths (55, 83, and 110% of crop evapotranspiration) were evaluated. During the cycle, sorghum adapted to the salinity and deficit irrigation depth, since stem height reduced only −5.5% with increasing salinity and −11.95% with decreasing irrigation depth, and aerial dry mass was affected by interaction only at the end of the cycle. Proline, total amino acids, and total soluble sugars were not differenced by stresses. Additionally, around 68.71% of total Na+ was at roots at the end of the cycle. In summary, sorghum BRS 506 was more tolerant to salt than water stress and used Na+ compartmentalization in root cells as the main tolerance mechanism. Full article
(This article belongs to the Special Issue Agricultural Crops Subjected to Drought and Salinity Stress)
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16 pages, 4418 KiB  
Article
Irrigation Depth and Potassium Doses Affect Fruit Yield and Quality of Figs (Ficus carica L.)
by Elias Ariel Moura, Vander Mendonça, Vladimir Batista Figueirêdo, Luana Mendes Oliveira, Marlenildo Ferreira Melo, Toni Halan Silva Irineu, Alex Danilo Monte Andrade, Edvan Alves Chagas, Pollyana Cardoso Chagas, Enoch Souza Ferreira, Luciana Freitas Medeiros Mendonça and Francisco Romário Andrade Figueiredo
Agriculture 2023, 13(3), 640; https://doi.org/10.3390/agriculture13030640 - 8 Mar 2023
Cited by 3 | Viewed by 1816
Abstract
The need to diversify agricultural production has fostered the cultivation of several crops under environmental conditions atypical to their origin, justifying the extreme importance of studies on the agricultural management of crops in semiarid regions. In this context, this study aimed to evaluate [...] Read more.
The need to diversify agricultural production has fostered the cultivation of several crops under environmental conditions atypical to their origin, justifying the extreme importance of studies on the agricultural management of crops in semiarid regions. In this context, this study aimed to evaluate the effects of irrigation depth and potassium doses on fig quality under semiarid conditions. The experiment was conducted in a 4 × 4 split-split-plot design, in randomized block design, with three replicates. The plots corresponded to four irrigation levels (50%, 75%, 100%, and 125% ETc), the subplots consisted of four potassium doses (0, 60, 120, and 240 g K2O plant−1), and the sub-subplot corresponded to the crop years (2018/19 and 2019/20). Results showed that water deficit reduced fig productivity, and the irrigation levels equal to or greater than 100% ETc performed cumulatively throughout the growing cycles. Therefore, irrigation depths from 85.19% to 95.16% ETc are recommended for greater water-use efficiency and fruit quality. Furthermore, potassium fertilization mitigated water stress in fig plants, allowing for reduced irrigation levels, especially in the second year, without compromising fruit traits. Full article
(This article belongs to the Special Issue Agricultural Crops Subjected to Drought and Salinity Stress)
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10 pages, 955 KiB  
Brief Report
Variability in Stomatal Adaptation to Drought among Grapevine Cultivars: Genotype-Dependent Responses
by Luca Nerva, Walter Chitarra, Gianni Fila, Lorenzo Lovat and Federica Gaiotti
Agriculture 2023, 13(12), 2186; https://doi.org/10.3390/agriculture13122186 - 22 Nov 2023
Cited by 2 | Viewed by 1169
Abstract
Leaf stomata are the primary determinants of the plant water relations. Physiological adaptations of stomata in response to water stress have been extensively reported for grapevine. On the contrary, little is known about how the plasticity in stomatal anatomical features may affect their [...] Read more.
Leaf stomata are the primary determinants of the plant water relations. Physiological adaptations of stomata in response to water stress have been extensively reported for grapevine. On the contrary, little is known about how the plasticity in stomatal anatomical features may affect their adaptability to drought conditions. In this study, we investigated, at the molecular and anatomical level, the effect of water stress on the stomatal anatomical features of four grapevine varieties extensively cultivated in the north of Italy. Potted plants of Garganega, Glera, Moscato giallo, and Merlot varieties were subjected to a 12–13 day period of water restriction during two consecutive seasons. Stomatal density and size were investigated in newly developed young leaves, 7 days after tip separation, following the occurrence of a water stress event. Furthermore, the gene expression of three key stomagenesis genes (VvEPFL9, VvEPF1, and VvEPF2) was analysed. The response of stomatal anatomical features to drought varied among the studied varieties. Moscato and Glera showed an increase in stomatal density and a decrease in stomatal size. On the contrary, Merlot displayed a reduction in stomatal number, while Garganega remained unchanged in terms of these values. Transcript levels of VvEPFL9 were overall in agreement with stomatal densities measured in the four varieties, showing an up-regulation when drought induced an increase in stomatal density or a down-regulation when the stomatal number decreased. The wide variability in stomatal response observed in the four varieties under study suggests that anatomical changes in stomatal characteristics are genotype dependent. These variations contribute to the intra-specific variability in grapevine’s response to water stress. Full article
(This article belongs to the Special Issue Agricultural Crops Subjected to Drought and Salinity Stress)
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Planned Papers

The below list represents only planned manuscripts. Some of these manuscripts have not been received by the Editorial Office yet. Papers submitted to MDPI journals are subject to peer-review.

Title: Increasing tolerance to salt stress in lettuce cultivars by applying elicitors
Authors: Miguel Ferreira Neto
Affiliation: Universidade Federal Rural do Semi-Árido
Abstract: Lettuce productivity can be limited by the concentration of salts in both water and soil. We evaluated the influence of salt stress tolerance elicitors on photosynthesis, yield, and ion homeostasis of lettuce cultivars. The research was conducted in a greenhouse with pots containing 12 dm3 of soil classified as Oxisol. For irrigation, a mixture of reject brine with an electrical conductivity of 9.0 dS m-1 and supply water was used, obtaining an electrical conductivity of 4.0 dS m-1. The experiment was conducted in a randomized block design, in a 3 x 5 factorial scheme, with five replications. The first factor corresponded to three cultivars of curly lettuce: SVR 2005, Simpson, and Grand Rapids. In the second factor, the lettuce cultivars were submitted to five combinations of treatments, including the foliar application of elicitors: E1 - irrigation with low salinity water (0.53 dS m-1, control); E2 - irrigation with high salinity water (4.0 dS m-1, saline stress); E3 - saline stress + exogenous application of ascorbic acid (50 µM L-1); E4 - saline stress + exogenous application of gibberellic acid (50 µM L-1); and E5 - saline stress + exogenous application of salicylic acid (50 µM L-1). The irrigation with saline water of 4.0 dS m-1 decreased the assimilation rate of CO2, stomatal conductance, growth, and biomass production of the salinity-sensitive cultivars SRV 2005 and Simpson. Salicylic acid attenuated salt stress in cultivar SRV 2005, improving growth, stomatal conductance, photosynthesis, and maximum efficiency of photosystem II.

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