Abiotic stress in quinoa: A comprehensive review on the impact of salinity and mitigation strategies

Mohamed ABDELAZEEM MOUSA; Szilvia VERES; Oqba BASAL

doi:10.15835/nbha53414860

Authors

Mohamed ABDELAZEEM MOUSA 1) University of Debrecen, Faculty of Agricultural and Food Sciences and Environmental Management, Institute of Applied Plant Biology, Böszörményi road 138, 4032, Debrecen; 2) Qena University, Faculty of Science, Department of Botany and Microbiology, Qena, 83523, Egypt (HU)
Szilvia VERES University of Debrecen, Faculty of Agricultural and Food Sciences and Environmental Management, Institute of Applied Plant Biology, Böszörményi road 138, 4032, Debrecen (HU)
Oqba BASAL University of Debrecen, Faculty of Agricultural and Food Sciences and Environmental Management, Institute of Applied Plant Biology, Böszörményi road 138, 4032, Debrecen (HU)

DOI:

https://doi.org/10.15835/nbha53414860

Keywords:

Na⁺ sequestration, osmotic adjustment, plant growth regulators, quinoa, reactive oxygen species, tolerance

Abstract

Quinoa (Chenopodium quinoa wild) is a gluten-free pseudocereal with an exceptionally nutritious, balanced profile of lipids, carbohydrates, proteins, minerals, vitamins, and beneficial secondary metabolites. These nutritional qualities, combined with Its capacity to grow successfully under drought, cold, salinity, and heavy metal stress, have earned special attention worldwide and positioned quinoa as a promising future crop for addressing global food security challenges amid climate change. This review highlights the physiological, morphological, and metabolic characteristics of quinoa that enable quinoa to tolerate a variety of abiotic stresses, with a particular emphasis on salinity. Quinoa exhibits various mechanisms under salt stress, including efficient Na⁺ sequestration in leaf vacuoles, controlled xylem Na⁺ loading, accumulation of organic and inorganic osmolytes, enhanced ROS resistance, improved K⁺ retention, and precise stomatal regulation. Quinoa’s tolerance to salinity can be significantly enhanced through seed priming, foliar applications of plant growth regulators, organic amendments, and microbial inoculants.

References

Abbas G, Abrar MM, Naeem MA, Siddiqui MH, Ali HM, Li Y, Ahmed K, Sun N, Xu M (2022). Biochar increases salt tolerance and grain yield of quinoa on saline-sodic soil: multivariate comparison of physiological and oxidative stress attributes. Journal of Soils and Sediments 22(5):1446-1459. https://doi.org/10.1007/s11368-022-03159-2

Abbas G, Areej F, Asad SA, Saqib M, Anwar-ul-Haq M, Afzal S, … Siddique HM (2023). Differential effect of heat stress on drought and salt tolerance potential of quinoa genotypes: A physiological and biochemical investigation. Plants 12(4):774. https://doi.org/10.3390/plants12040774

Abbas G, Rehman S, Siddiqui MH, Ali HM, Farooq MA, Chen Y (2022). Potassium and humic acid synergistically increase salt tolerance and nutrient uptake in contrasting wheat genotypes through ionic homeostasis and activation of antioxidant enzymes. Plants 11(3):263. https://doi.org/10.3390/plants11030263

Abdallah MM-S, El Sebai TN, Ramadan AAE-M, El-Bassiouny HMS (2020). Physiological and biochemical role of proline, trehalose, and compost on enhancing salinity tolerance of quinoa plant. Bulletin of the National Research Centre 44(1):96. https://doi.org/10.1186/s42269-020-00354-4

Abdrabou MR, Gomah H, Darweesh A, Eissa MA, Selmy SAH (2022). Response of saline irrigated quinoa (Chenopodium quinoa Wild) grown on coarse texture soils to organic manure. Egyptian Journal of Soil Science 62(2):169-178. https://doi.org/10.21608/ejss.2022.146571.1511

Abdulmajeed AM (2023). Salinity stress amelioration and morpho-physiological growth stimulation by silicon priming and biochar supplementation in Chenopodium quinoa. Notulae Botanicae Horti Agrobotanici Cluj-Napoca 51(2):13043. https://doi.org/10.15835/nbha51213043

Abo-Kassem EEM (2007). Effects of salinity: Calcium interaction on growth and nucleic acid metabolism in five species of Chenopodiaceae. Turkish Journal of Botany 31(2):125-134.

Adolf VI, Jacobsen S-E, Shabala S (2013). Salt tolerance mechanisms in quinoa (Chenopodium quinoa Willd.). Environmental and Experimental Botany 92:43-54. https://doi.org/10.1016/j.envexpbot.2012.07.004

Adolf VI, Shabala S, Andersen MN, Razzaghi F, Jacobsen S-E (2012). Varietal differences of quinoa’s tolerance to saline conditions. Plant and Soil 357(1):117-129. https://doi.org/10.1007/s11104-012-1133-7

Ahmed M, Qadeer U, Ahmed ZI, Hassan F (2016). Improvement of wheat (Triticum aestivum) drought tolerance by seed priming with silicon. Archives of Agronomy and Soil Science 62(3):299-315. https://doi.org/10.1080/03650340.2015.1048235

Ain QT, Siddique K, Bawazeer S, Ali I, Mazhar M, Rasool R, … Jafar TH (2023). Adaptive mechanisms in quinoa for coping in stressful environments: an update. PeerJ 11: e14832. https://doi.org/10.7717/peerj.14832

Alam A, Ullah H, Cha-um S, Tisarum R, Datta A (2021). Effect of seed priming with potassium nitrate on growth, fruit yield, quality and water productivity of cantaloupe under water-deficit stress. Scientia Horticulturae 288:110354. https://doi.org/10.1016/j.scienta.2021.110354

Alandia G, Rodriguez JP, Jacobsen S-E, Bazile D, Condori B (2020). Global expansion of quinoa and challenges for the Andean region. Global Food Security 26:100429. https://doi.org/10.1016/j.gfs.2020.100429

Alcívar M, Zurita-Silva A, Sandoval M, Muñoz C, Schoebitz M (2018). Reclamation of saline-sodic soils with combined amendments: Impact on quinoa performance and biological soil quality. Sustainability (Switzerland) 10(9):3083. https://doi.org/10.3390/su10093083

Alghamdi SA, Alharby HF, Abbas G, Al-Solami HM, Younas A, Aldehri M, … Chen Y (2023). Salicylic acid-and potassium-enhanced resilience of quinoa (Chenopodium quinoa Willd.) against salinity and cadmium stress through mitigating ionic and oxidative stress. Plants 12(19):3450. https://doi.org/10.3390/plants12193450

Almodares A, Hadi MR, Dosti B (2007). Effects of salt stress on germination percentage and seedling growth in sweet sorghum cultivars. Journal of Biological Sciences 7(8):1492-1495.

Aloisi I, Parrotta L, Ruiz KB, Landi C, Bini L, Cai G, Biondi S, Del Duca S (2016). New insight into quinoa seed quality under salinity: Changes in proteomic and amino acid profiles, phenolic content, and antioxidant activity of protein extracts. Frontiers in Plant Science 7:656. https://doi.org/10.3389/fpls.2016.00656

Andrews D (2017). Race, status, and biodiversity: the social climbing of quinoa. Culture, Agriculture, Food and Environment 39(1):15-24. https://doi.org/10.1111/cuag.12084

Arif Y, Singh P, Siddiqui H, Bajguz A, Hayat S (2020). Salinity induced physiological and biochemical changes in plants: An omic approach towards salt stress tolerance. Plant Physiology and Biochemistry 156:64-77. https://doi.org/10.1016/j.plaphy.2020.08.042

Awadalla A, Morsy ASM, Sherif MM (2020). Impact of foliar spray by proline on the production and quality of quinoa under saline soil conditions at Toshka region. Journal of Plant Production 11(5):391-397. https://doi.org/10.21608/jpp.2020.102750

Azooz MM, Youssef AM, Ahmad P (2011). Evaluation of salicylic acid (SA) application on growth, osmotic solutes and antioxidant enzyme activities on broad bean seedlings grown under diluted seawater. International Journal of Plant Physiology and Biochemistry3(14):253-264. https://doi.org/10.5897/IJPPB11.052

Barkla BJ, Vera‐Estrella R, Pantoja O (2012). Protein profiling of epidermal bladder cells from the halophyte Mesembryanthemum crystallinum. Proteomics 12(18):2862-2865. https://doi.org/10.1002/pmic.201200152

Barragán V, Leidi EO, Andrés Z, Rubio L, De Luca A, Fernandez JA, … Pardo JM (2012). Ion exchangers NHX1 and NHX2 mediate active potassium uptake into vacuoles to regulate cell turgor and stomatal function in Arabidopsis. The Plant Cell 24(3):1127-1142. https://doi.org/10.1105/tpc.111.095273

Bazile D, Jacobsen S-E, Verniau A (2016). The global expansion of quinoa: trends and limits. Frontiers in Plant Science 7:622. https://doi.org/doi.org/10.3389/fpls.2016.00622

Bazile D, Martínez EA, Fuentes F (2014). Diversity of quinoa in a biogeographical Island: A review of constraints and potential from arid to temperate regions of Chile. Notulae Botanicae Horti Agrobotanici Cluj-Napoca 42(2):289-298. https://doi.org/10.15835/nbha4229733

Becker VI, Goessling JW, Duarte B, Caçador I, Liu F, Rosenqvist E, Jacobsen S-E (2017). Combined effects of soil salinity and high temperature on photosynthesis and growth of quinoa plants (Chenopodium quinoa). Functional Plant Biology 44(7):665-678. https://doi.org/10.1071/FP16370

Biondi S, Antognoni F, Marincich L, Lianza M, Tejos R, Ruiz KB (2022). The polyamine “multiverse” and stress mitigation in crops: A case study with seed priming in quinoa. Scientia Horticulturae 304:111292. https://doi.org/10.1016/j.scienta.2022.111292

Bonales-Alatorre E, Shabala S, Chen Z-H, Pottosin I (2013). Reduced tonoplast fast-activating and slow-activating channel activity is essential for conferring salinity tolerance in a facultative halophyte, quinoa. Plant Physiology 162(2):940-952. https://doi.org/10.1104/pp.113.216572

Bouras H, Choukr-Allah R, Amouaouch Y, Bouaziz A, Devkota KP, El Mouttaqi A, … Hirich A (2022). How does quinoa (Chenopodium quinoa Willd.) respond to phosphorus fertilization and irrigation water salinity? Plants 11(2):216. https://doi.org/10.3390/plants11020216

Bourhim MR, Cheto S, Qaddoury A, Hirich A, Ghoulam C (2022). Chemical seed priming with zinc sulfate improves quinoa tolerance to salinity at germination stage. Environmental Sciences Proceedings 16(1):23. https://doi.org/10.3390/environsciproc2022016023

Bourhim MR, El Mouttaqi A, Oukhey F, Farissi M, Khadraji A, Qaddoury A, … Ghoulam C (2025). Priming enhances germination and reserve mobilisation in quinoa seeds under salt stress. Seed Science and Technology 53(2):311-326. https://doi.org/10.15258/sst.2025.53.2.13

Brini F, Hanin M, Lumbreras V, Amara I, Khoudi H, Hassairi A, … Masmoudi K (2007). Overexpression of wheat dehydrin DHN-5 enhances tolerance to salt and osmotic stress in Arabidopsis thaliana. Plant Cell Reports 26(11):2017-2026. https://doi.org/10.1007/s00299-007-0412-x

Bruin A de (1964). Investigation of the food value of quinua and cañihua seed. Journal of Food Science 29(6):872-876. https://doi.org/10.1111/j.1365-2621.1964.tb00464.x

Bueno M, Cordovilla MP (2021). Ecophysiology and uses of halophytes in diverse habitats. In: Grigore MN (Eds). Handbook of halophytes: from molecules to ecosystems towards biosaline agriculture. Springer, Cham pp 1613-1636. https://doi.org/10.1007/978-3-030-57635-6_57

Burrieza HP, Koyro H-W, Tosar LM, Kobayashi K, Maldonado S (2012). High salinity induces dehydrin accumulation in Chenopodium quinoa Willd. cv. Hualhuas embryos. Plant and Soil 354(1):69-79. https://doi.org/10.1007/s11104-011-1045-y

Cai Z-Q, Gao Q (2020). Comparative physiological and biochemical mechanisms of salt tolerance in five contrasting highland quinoa cultivars. BMC Plant Biology 20(1):70. https://doi.org/10.1186/s12870-020-2279-8

Causin HF, Bordón DAE, Burrieza H (2020). Salinity tolerance mechanisms during germination and early seedling growth in Chenopodium quinoa Wild. genotypes with different sensitivity to saline stress. Environmental and Experimental Botany 172:103995. https://doi.org/10.1016/j.envexpbot.2020.103995

Chauhan A, AbuAmarah BA, Kumar A, Verma JS, Ghramh HA, Khan KA, Ansari MJ (2019). Influence of gibberellic acid and different salt concentrations on germination percentage and physiological parameters of oat cultivars. Saudi Journal of Biological Sciences 26(6):1298-1304. https://doi.org/10.1016/j.sjbs.2019.04.014

Chen T-W, Gomez Pineda IM, Brand AM, Stützel H (2020). Determining ion toxicity in cucumber under salinity stress. Agronomy 10(5):677. https://doi.org/10.3390/agronomy10050677

Choudhary S, Wani KI, Naeem M, Khan MMA, Aftab T (2023). Cellular responses, osmotic adjustments, and role of osmolytes in providing salt stress resilience in higher plants: Polyamines and nitric oxide crosstalk. Journal of Plant Growth Regulation 42(2):539-553. https://doi.org/10.1007/s00344-022-10584-7

Cifuentes L, González M, Pinto-Irish K, Álvarez R, Coba de la Peña T, Ostria-Gallardo E, … Castro PA (2023). Metabolic imprint induced by seed halo-priming promotes a differential physiological performance in two contrasting quinoa ecotypes. Frontiers in Plant Science 13:1034788 https://doi.org/10.3389/fpls.2022.1034788

Cocozza C, Pulvento C, Lavini A, Riccardi M, D’Andria R, Tognetti R (2013). Effects of increasing salinity stress and decreasing water availability on ecophysiological traits of quinoa (Chenopodium quinoa Willd.) grown in a mediterranean‐type agroecosystem. Journal of Agronomy and Crop Science 199(4):229-240. https://doi.org/10.1111/jac.12012

Craine EB, Murphy KM (2020). Seed composition and amino acid profiles for quinoa grown in Washington State. Frontiers in Nutrition 7:126. https://doi.org/10.3389/fnut.2020.00126

Cuin TA, Tian Y, Betts SA, Chalmandrier R, Shabala S (2009). Ionic relations and osmotic adjustment in durum and bread wheat under saline conditions. Functional Plant Biology. Functional Plant Biology 36(12):1110-1119. https://doi.org/10.1071/FP09051

Daur I (2018). Effects of hydro and hormonal priming on quinoa (Chenopodium quinoa Willd.) seed germination under salt and drought stress. Pakistan Journal of Botany 50(5):1669-1673.

De Pascale S, Ruggiero C, Barbieri G, Maggio A (2003). Physiological responses of pepper to salinity and drought. Journal-American Society for Horticultural Science 128(1):48-54.

Dehghanian Z, Ahmadabadi M, Asgari Lajayer B, Gougerdchi V, Hamedpour-Darabi M, … Dell B (2024). Quinoa: A promising crop for resolving the bottleneck of cultivation in soils affected by multiple environmental abiotic stresses. Plants 13(15):2117. https://doi.org/10.3390/plants13152117

Delatorre-Herrera J, Pinto M (2009). Importancia de los componentes iónico y osmótico del estrés salino sobre la germinación de cuatro selecciones de quinua (Chenopodium quinoa Willd.). Chilean journal of Agricultural Research 69(4):477-485. http://dx.doi.org/10.4067/S0718-58392009000400001

Derbali I, Derbali W, Gharred J, Manaa A, Slama I, Koyro H-W (2024). Mitigating salinity stress in quinoa (Chenopodium quinoa Willd.) with biochar and superabsorber polymer amendments. Plants 13(1):92. https://doi.org/10.3390/plants13010092

Derbali W, Manaa A, Goussi R, Derbali I, Abdelly C, Koyro H-W (2021). Post-stress restorative response of two quinoa genotypes differing in their salt resistance after salinity release. Plant Physiology and Biochemistry 164:222-236. https://doi.org/10.1016/j.plaphy.2021.04.024

Devika OS, Singh S, Sarkar D, Barnwal P, Suman J, Rakshit A (2021). Seed priming: a potential supplement in integrated resource management under fragile intensive ecosystems. Frontiers in Sustainable Food Systems 5:654001. https://doi.org/10.3389/fsufs.2021.654001

Ding Z, Kheir AMS, Ali MGM, Ali OAM, Abdelaal AIN, Lin X, … He Z (2020). The integrated effect of salinity, organic amendments, phosphorus fertilizers, and deficit irrigation on soil properties, phosphorus fractionation and wheat productivity. Scientific Reports 10(1):2736. https://doi.org/10.1038/s41598-020-59650-8

Dini A, Rastrelli L, Saturnino P, Schettino O (1992). A compositional study of Chenopodium quinoa seeds. Nahrung, 36(4):400-404. https://doi.org/10.1002/food.19920360412

Dinneny JR (2015). Traversing organizational scales in plant salt-stress responses. Current Opinion in Plant Biology 23:70-75. https://doi.org/10.1016/j.pbi.2014.10.009

Ehtaiwesh AF (2022). The effect of salinity on nutrient availability and uptake in crop plants. Scientific Journal of Applied Sciences of Sabratha University 9:55-73.

Eisa S, Eid MA, Abd El-Samad EH, Hussin SA, Abdel-Ati AA, El-Bordeny NE, … Masoud AM (2017). “Chenopodium quinoa” Willd. A new cash crop halophyte for saline regions of Egypt. Australian Journal of Crop Science 11(3):343-351.

Eisa S, Hussin S, Geissler N, Koyro HW (2012). Effect of NaCl salinity on water relations, photosynthesis and chemical composition of Quinoa (Chenopodium quinoa Willd.) as a potential cash crop halophyte. Australian Journal of Crop Science 6(2):357-368. https://doi.org/ 10.3316/informit.054900975531667

El Mouttaqi A, Sabraoui T, Belcaid M, Ibourki M, Mnaouer I, Lazaar K, ... Rafik S (2023). Agro-morphological and biochemical responses of quinoa (Chenopodium quinoa Willd. var: ICBA-Q5) to organic amendments under various salinity conditions. Frontiers in Plant Science 14:1143170. https://doi.org/10.3389/fpls.2023.1143170

El Sebai TN, Abdallah MM-S, El-Bassiouny HMS, Ibrahim FM (2016). Amelioration of the adverse effects of salinity stress by using compost, Nigella sativa extract or ascorbic acid in quinoa plants. International Journal of PharmTech Research 9(6):127-144.

FAO (2024). Global map of salt-affected soils. FAO. Rome, Italy. https://www.fao.org/soils-portal/data-hub/soil-maps-and-databases/global-map-of-salt-affected-soils/en/

Farsiani A, Ghobadi ME (2009). Effects of PEG and NaCl stress on two cultivars of corn (Zea mays L.) at germination and early seedling stages. International Journal of Biological, Life and Agricultural Sciences 57:382-385.

Fischer S, Wilckens R, Jara J, Aranda M, Valdivia W, Bustamante L, … Obal I (2017). Protein and antioxidant composition of quinoa (Chenopodium quinoa Willd.) sprout from seeds submitted to water stress, salinity and light conditions. Industrial Crops and Products 107:558-564. https://doi.org/10.1016/j.indcrop.2017.04.035

Flowers TJ (2004). Improving crop salt tolerance. Journal of Experimental Botany 55(396):307-319. https://doi.org/10.1093/jxb/erh003

Flowers TJ, Colmer TD (2008). Salinity tolerance in halophytes. New Phytologist 179(4):945-963. https://www.jstor.org/stable/25150520

Flowers TJ, Munns R, Colmer TD (2015). Sodium chloride toxicity and the cellular basis of s alt tolerance in halophytes. Annals of Botany 115(3):419-431. https://doi.org/10.1093/aob/mcu217

Freitas H, Breckle S-W (1992). Importance of bladder hairs for salt tolerance of field-grown Atriplex species from a Portuguese salt marsh. Flora 187:283-297. https://doi.org/10.1016/S0367-2530(17)32233-8

Geissler N, Hussin S, El-Far MMM, Koyro H-W (2015). Elevated atmospheric CO2 concentration leads to different salt resistance mechanisms in a C3 (Chenopodium quinoa) and a C4 (Atriplex nummularia) halophyte. Environmental and Experimental Botany 118:67-77. https://doi.org/10.1016/j.envexpbot.2015.06.003

Gill SS, Tuteja N (2010). Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant Physiology and Biochemistry 48(12):909-930. https://doi.org/10.1016/j.plaphy.2010.08.016

Gómez-Caravaca AM, Iafelice G, Lavini A, Pulvento C, Caboni MF, Marconi E (2012). Phenolic compounds and saponins in quinoa samples (Chenopodium quinoa Willd.) grown under different saline and nonsaline irrigation regimens. Journal of Agricultural and Food Chemistry 60(18):4620-4627. https://doi.org/10.1021/jf3002125

Gómez‐Pando LR, Álvarez‐Castro R, Eguiluz‐De La Barra A (2010). Effect of salt stress on Peruvian germplasm of Chenopodium quinoa Willd.: a promising crop. Journal of Agronomy and Crop Science 196(5):391-396. https://doi.org/10.1111/j.1439-037X.2010.00429.x

González-Teuber M, Vilo C, Bascuñán-Godoy L (2017). Molecular characterization of endophytic fungi associated with the roots of Chenopodium quinoa inhabiting the Atacama Desert, Chile. Genomics Data 11:109-112. https://doi.org/10.1016/j.gdata.2016.12.015

Gul Z, Arif M, Hayat M, Quyen L (2026). Salinity resilience in quinoa (Chenopodium quinoa); investigating adaptive mechanisms across different varieties to combat salt stress for sustainable agriculture. Pakistan Journal of Botany 58(2):1-16. https://doi.org/10.30848/PJB2026-2(11)

Hao Y, Hong Y, Guo H, Qin P, Huang A, Yang X, Ren G (2022). Transcriptomic and metabolomic landscape of quinoa during seed germination. BMC Plant Biology 22(1):237. https://doi.org/10.1186/s12870-022-03621-w

Hariadi Y, Marandon K, Tian Y, Jacobsen S-E, Shabala S (2011). Ionic and osmotic relations in quinoa (Chenopodium quinoa Willd.) plants grown at various salinity levels. Journal of Experimental Botany 62(1):185-193. https://doi.org/10.1093/jxb/erq257

Hasanuzzaman M, Nahar K, Alam MM, Roychowdhury R, Fujita M (2013). Physiological, biochemical, and molecular mechanisms of heat stress tolerance in plants. International Journal of Molecular Sciences 14(5):9643-9684. https://doi.org/10.3390/ijms14059643

Hassani A, Azapagic A, Shokri N (2021). Global predictions of primary soil salinization under changing climate in the 21st century. Nature Communications 12:6663. https://doi.org/10.1038/s41467-021-26907-3

Hinojosa L, González JA, Barrios-Masias FH, Fuentes F, Murphy KM (2018). Quinoa abiotic stress responses: A review. Plants 7(4):106. https://doi.org/10.3390/plants7040106

Hussain MI, Al-Dakheel AJ, Reigosa MJ (2018). Genotypic differences in agro-physiological, biochemical and isotopic responses to salinity stress in quinoa (Chenopodium quinoa Willd.) plants: Prospects for salinity tolerance and yield stability. Plant Physiology and Biochemistry 129:411-420. https://doi.org/10.1016/j.plaphy.2018.06.023

Hussain MI, Muscolo A, Ahmed M, Asghar MA, Al-Dakheel AJ (2020). Agro-morphological, yield and quality traits and interrelationship with yield stability in quinoa (Chenopodium quinoa Willd.) genotypes under saline marginal environment. Plants 9(12):1763. https://doi.org/10.3390/plants9121763

Hussein SO, Kovács F, Tobak Z (2017). Spatiotemporal assessment of vegetation indices and land cover for Erbil city and its surrounding using MODIS imageries. Journal of Environmental Geography 10(1-2):31-39. https://doi.org/10.1515/jengeo-2017-0004

Hussin SA, Ali SH, Lotfy ME, El-Samad EHA, Eid MA, Abd-Elkader AM, Eisa SS (2023). Morpho-physiological mechanisms of two different quinoa ecotypes to resist salt stress. BMC Plant Biology 23(1):374. https://doi.org/10.1186/s12870-023-04342-4

Imamura T, Yasui Y, Koga H, Takagi H, Abe A, Nishizawa K, … Mori M (2020). A novel WD40-repeat protein involved in formation of epidermal bladder cells in the halophyte quinoa. Communications Biology 3(1):513. https://doi.org/10.1038/s42003-020-01249-w

Jacobsen S (2017). The scope for adaptation of quinoa in Northern latitudes of Europe. Journal of Agronomy and Crop Science 203(6):603-613. https://doi.org/10.1111/jac.12228

Jacobsen S-E, Liu F, Jensen CR (2009). Does root-sourced ABA play a role for regulation of stomata under drought in quinoa (Chenopodium quinoa Willd.). Scientia Horticulturae 122(2):281-287. https://doi.org/10.1016/j.scienta.2009.05.019

Jacobsen S-E, Mujica A (2002). Genetic resources and breeding of the Andean grain crop quinoa (Chenopodium quinoa Willd.). Plant Genetic Resources Newsletter 130:54-61.

Jacobsen S-E, Mujica A, Jensen CR (2003). The resistance of quinoa (Chenopodium quinoa Willd.) to adverse abiotic factors. Food Reviews International 19(1-2):99-109. https://doi.org/10.1081/FRI-120018872

Jafari T, Iranbakhsh A, Aliabad KK, Daneshmand F, Seifati SE (2024). Nitric oxide reduced saponin metabolite in Chenopodium quinoa seedlings cultivated under salinity. Russian Journal of Plant Physiology 71(3):66. https://doi.org/10.1134/S1021443723603518

Jiang X-W, Zhang C-R, Wang W-H, Xu G-H, Zhang H-Y (2020). Seed priming improves seed germination and seedling growth of Isatis indigotica Fort. under salt stress. HortScience 55(5):647-650. https://doi.org/10.21273/HORTSCI14854-20

Karimi G, Pourakbar L, Siavash Moghaddam S, Rezaee Danesh Y, Popovi´c-Djordjevi´c J (2022). Effectiveness of fungal bacterial biofertilizers on agrobiochemical attributes of quinoa (Chenopodium quinoa willd.) under salinity stress. International Journal of Environmental Science and Technology 19(12):11989-12002. https://doi.org/10.1007/s13762-022-04427-x

Karyotis T, Iliadis C, Noulas C, Mitsibonas TH (2003). Preliminary research on seed production and nutrient content for certain quinoa varieties in a saline-sodic soil. Journal of Agronomy and Crop Science 189(6):402-408. https://doi.org/10.1046/j.0931-2250.2003.00063.x

Khalofah A, Migdadi H, El-Harty E (2021). Antioxidant enzymatic activities and growth response of quinoa (Chenopodium quinoa Willd) to exogenous selenium application. Plants 10(4):719. https://doi.org/10.3390/plants10040719

Kiani‐Pouya A, Roessner U, Jayasinghe NS, Lutz A, Rupasinghe T, Bazihizina N, … Shabala S (2017). Epidermal bladder cells confer salinity stress tolerance in the halophyte quinoa and Atriplex species. Plant, Cell & Environment 40(9):1900-1915. https://doi.org/10.1111/pce.12995

Killi D, Haworth M (2017). Diffusive and metabolic constraints to photosynthesis in quinoa during drought and salt stress. Plants 6(4):49. https://doi.org/10.3390/plants6040049

Koca YO (2017). Effects of different salt concentrations on Quinoa seedling quality. International Journal of Secondary Metabolite 4(3):20-26. https://doi.org/10.21448/ijsm.356248

Koyro H-W, Eisa SS (2008). Effect of salinity on composition, viability and germination of seeds of Chenopodium quinoa Willd. Plant and Soil 302(1):79-90. https://doi.org/10.1007/s11104-007-9457-4

Koyro H-W, Lieth H, Eisa SS (2008). Salt tolerance of Chenopodium quinoa Willd., grains of the Andes: Influence of salinity on biomass production, yield, composition of reserves in the seeds, water and solute relations. In: Lieth H, Sucre MG, Herzog B (Eds). Mangroves and halophytes: Restoration and utilisation. Springer, Dordrecht pp 133-145. https://doi.org/10.1007/978-1-4020-6720-4_13

Kozioł MJ (1992). Chemical composition and nutritional evaluation of quinoa (Chenopodium quinoa Willd.). Journal of Food Composition and Analysis 5(1):35-68. https://doi.org/10.1016/0889-1575(92)90006-6

Lallouche B, Kouider BH (2024). Effects of hydropriming, halopriming, and hormopriming seed treatments on the subsequent salt stress tolerance of quinoa (Chenopodium quinoa Willd.) in Algeria. Acta Scientiarum Polonorum Hortorum Cultus 23(5):59-70. https://doi.org/10.24326/asphc.2024.5417

Leng BY, Yuan F, Dong XX, Wang J, Wang BS (2018). Distribution pattern and salt excretion rate of salt glands in two recretohalophyte species of Limonium (Plumbaginaceae). South African Journal of Botany 115:74-80. https://doi.org/10.1016/j.sajb.2018.01.002

Leogrande R, Vitti C (2019). Use of organic amendments to reclaim saline and sodic soils: a review. Arid Land Research and Management 33(1):1-21. https://doi.org/10.1080/15324982.2018.1498038

Li M, Fu Q, Singh VP, Liu D, Li T (2019). Stochastic multi-objective modeling for optimization of water-food-energy nexus of irrigated agriculture. Advances in water resources 127:209-224. https://doi.org/10.1016/j.advwatres.2019.03.015

LoPresti EF (2014). Chenopod salt bladders deter insect herbivores. Oecologia 174(3):921-930. https://doi.org/10.1007/s00442-013-2827-0

Maas E V, Grattan SR (1999). Crop yields as affected by salinity. Agricultural Drainage 38:55-108. https://doi.org/10.2134/agronmonogr38.c3

Maksimovic I, Ilin Z (2012). Effects of salinity on vegetable growth and nutrients uptake. In: Irrigation systems and practices in challenging environments. InTech Open Croatia 9:169-190.

Maleki P, Saadat S, Bahrami HA, Rezaei H, Esmaeelnejad L (2019). Accumulation of ions in shoot and seed of quinoa (Chenopodium‎ quinoa Willd.) under salinity stress. Communications in Soil Science and Plant Analysis 50(6):782-793. https://doi.org/10.1080/00103624.2019.1589486

Mamedi A, Sharifzadeh F (2023). Hydrotime analysis of Chenopodium quinoa seed germination responses to combined effects of nacl-induced osmotic stress (Ψ), chilling temperature (T), and Ca2+-priming treatment. Journal of Soil Science and Plant Nutrition 23(2):2299-2315. https://doi.org/10.1007/s42729-023-01180-z

Mamedi A, Sharifzadeh F, Maali-Amiri R, Divargar F, Rasoulnia A (2022). Seed osmopriming with Ca2+ and K+ improves salt tolerance in quinoa seeds and seedlings by amplifying antioxidant defense and ameliorating the osmotic adjustment process. Physiology and Molecular Biology of Plants 28(1):251-274. https://doi.org/10.1007/s12298-022-01125-3

Manaa A, Goussi R, Derbali W, Cantamessa S, Abdelly C, Barbato R (2019). Salinity tolerance of quinoa (Chenopodium quinoa Willd) as assessed by chloroplast ultrastructure and photosynthetic performance. Environmental and Experimental Botany 162:103-114. https://doi.org/10.1016/j.envexpbot.2019.02.012

Mastebroek HD, Van Loo EN, Dolstra O (2002). Combining ability for seed yield traits of Chenopodium quinoa breeding lines. Euphytica 125(3):427-432. https://doi.org/10.1023/A:1016030129541

Meddich A, Jaiti F, Bourzik W, El Asli A, Hafidi M (2015). Use of mycorrhizal fungi as a strategy for improving the drought tolerance in date palm (Phoenix dactylifera). Scientia Horticulturae 192:468-474. https://doi.org/10.1016/j.scienta.2015.06.024

Mhada M, Metougui ML, El Hazzam K, El Kacimi K, Yasri A (2020). Variations of saponins, minerals and total phenolic compounds due to processing and cooking of quinoa (Chenopodium quinoa Willd.) seeds. Foods 9(5):660. https://doi.org/10.3390/foods9050660

Mittler R (2006). Abiotic stress, the field environment and stress combination. Trends in Plant Science 11(1):15-19. https://doi.org/10.1016/j.tplants.2005.11.002

Mizuno N, Toyoshima M, Fujita M, Fukuda S, Kobayashi Y, Ueno M, … Fujita Y (2020). The genotype-dependent phenotypic landscape of quinoa in salt tolerance and key growth traits. DNA Research 27(4):dsaa022. https://doi.org/10.1093/dnares/dsaa022

Mohamed Ahmed IA, Al Juhaimi F, Özcan MM (2021). Insights into the nutritional value and bioactive properties of quinoa (Chenopodium quinoa): past, present and future prospective. International Journal of Food Science and Technology 56(8):3726–3741. https://doi.org/10.1111/ijfs.15011

Mohammadi H, Rahimpour B, Pirasteh-Anosheh H, Race M (2022). Salicylic acid manipulates ion accumulation and distribution in favor of salinity tolerance in Chenopodium quinoa. International Journal of Environmental Research and Public Health 19:1576. https://doi.org/10.3390/ijerph19031576

Moog MW, Trinh MDL, Nørrevang AF, Bendtsen AK, Wang C, Østerberg JT, … Palmgren M (2022). The epidermal bladder cell‐free mutant of the salt‐tolerant quinoa challenges our understanding of halophyte crop salinity tolerance. New phytologist 236(4):1409-1421. https://doi.org/10.1111/nph.18420

Moog MW, Yang X, Bendtsen AK, Dong L, Crocoll C, Imamura T, … Palmgren M (2023). Epidermal bladder cells as a herbivore defense mechanism. Current Biology 33(21):4662-4673. https://doi.org/10.1016/j.cub.2023.09.063

Moreno C, Seal CE, Papenbrock J (2018). Seed priming improves germination in saline conditions for Chenopodium quinoa and Amaranthus caudatus. Journal of Agronomy and Crop Science 204:40-48. https://doi.org/10.1111/jac.12242

Munns R, James RA, Läuchli A (2006). Approaches to increasing the salt tolerance of wheat and other cereals. Journal of Experimental Botany 57(5):1025-1043. https://doi.org/10.1093/jxb/erj100

Nachtergaele F, Van Velthuizen H, Verelst L, Batjes N, Dijkshoorn K, van Engelen V, ... Petri M (2010). Harmonized world soil database. In: Proceedings Of The 19th World Congress Of Soil Science, Soil Solutions For a Changing World, Brisbane, Australia. International Union of Soil Sciences pp 34-37.

Nanduri KR, Hirich A, Salehi M, Saadat S, Jacobsen SE (2019). Quinoa: A new crop for harsh environments. In: Gul B, Böer B, Khan M, Clüsener-Godt M, Hameed A (Eds). Sabkha Ecosystems: Tasks for Vegetation Science. Springer pp 301-333. https://doi.org/10.1007/978-3-030-04417-6_19

Naveed M, Ramzan N, Mustafa A, Samad A, Niamat B, Yaseen M, … Xu M (2020). Alleviation of salinity induced oxidative stress in Chenopodium quinoa by Fe biofortification and biochar-endophyte interaction. Agronomy 10(2):168. https://doi.org/10.3390/agronomy10020168

Nowak V, Du J, Charrondière UR (2016). Assessment of the nutritional composition of quinoa (Chenopodium quinoa Willd.). Food chemistry 193:47-54. https://doi.org/10.1016/j.foodchem.2015.02.111

Okon OG (2019). Effect of salinity on physiological processes in plants. In: B Giri B, Varma A(Eds). Microorganisms in saline environments: strategies and functions. Soil biology. Springer, Cham 56:237-262. https://doi.org/10.1007/978-3-030-18975-4_10

Olmos E, Jimenez-Perez B, Roman-Garcia I, Fernandez-Garcia N (2024). Salt-tolerance mechanisms in quinoa: Is Glycine betaine the missing piece of the puzzle? Plant Physiology and Biochemistry 206:108276. https://doi.org/10.1016/j.plaphy.2023.108276

Orsini F, Accorsi M, Gianquinto G, Dinelli G, Antognoni F, Carrasco KBR, ... Bosi S (2011). Beyond the ionic and osmotic response to salinity in Chenopodium quinoa: functional elements of successful halophytism. Functional Plant Biology 38(10):818-831. https://doi.org/10.1071/FP11088

Panuccio MR, Jacobsen SE, Akhtar SS, Muscolo A (2014). Effect of saline water on seed germination and early seedling growth of the halophyte quinoa. AoB Plants 6:plu047. https://doi.org/10.1093/aobpla/plu047

Paul A, Mondal S, Chakraborty K, Biswas AK (2024). Moving forward to understand the alteration of physiological mechanism by seed priming with different halo-agents under salt stress. Plant Molecular Biology 114(2):24. https://doi.org/10.1007/s11103-024-01425-0

Pellegrini M, Lucas-Gonzales R, Ricci A, Fontecha J, Fernández-López J, Pérez-Álvarez JA, Viuda-Martos M (2018). Chemical, fatty acid, polyphenolic profile, techno-functional and antioxidant properties of flours obtained from quinoa (Chenopodium quinoa Willd) seeds. Industrial Crops and Products 111:38-46. https://doi.org/10.1016/j.indcrop.2017.10.006

Pereira A (2016). Plant abiotic stress challenges from the changing environment. Frontiers in Plant Science 7:1123. https://doi.org/10.3389/fpls.2016.01123

Peterson A, Murphy K (2015). Tolerance of lowland quinoa cultivars to sodium chloride and sodium sulfate salinity. Crop Science 55(1): 331-338. https://doi.org/10.2135/cropsci2014.04.0271

Pirasteh-Anosheh H, Emam Y (2020). Role of chlormequat chloride and salicylic acid in improving cereal crops production under saline. In: Improving cereal productivity through climate smart practices. Woodhead Publishing pp 145. https://doi.org/10.1016/b978-0-12-821316-2.00009-1

Pirasteh-Anosheh H, Emam Y, Pessarakli M (2019). Grain filling pattern of Hordeum vulgare as affected by salicylic acid and salt stress. Journal of Plant Nutrition 42(3):278-286. https://doi.org/10.1080/01904167.2018.1554680

Pooja, Munjal R (2019). Oxidative stress and antioxidant defense in plants under high temperature, Reactive oxygen, nitrogen and sulfur species in plants: Production, metabolism, signaling and defense mechanisms. Wiley Online Library pp 337-352. https://doi.org/10.1002/9781119468677.ch14

Prado FE, Fernández‐Turiel JL, Tsarouchi M, Psaras GK, González JA (2014). Variation of seed mineral concentrations in seven quinoa cultivars grown in two agroecological sites. Cereal Chemistry 91(5):453-459. https://doi.org/10.1094/CCHEM-08-13-0157-R

Präger A, Munz S, Nkebiwe PM, Mast B, Graeff-Hönninger S (2018). Yield and quality characteristics of different quinoa (Chenopodium quinoa Willd.) cultivars grown under field conditions in Southwestern Germany. Agronomy 8(10):197. https://doi.org/10.3390/agronomy8100197

Pulvento C, Riccardi M, Lavini A, Iafelice G, Marconi E, D’Andria R (2012). Yield and quality characteristics of quinoa grown in open field under different saline and non‐saline irrigation regimes. Journal of Agronomy and Crop Science 198(4):254-263. https://doi.org/10.1111/j.1439-037X.2012.00509.x

Qadir M, Quillérou E, Nangia V, Murtaza G, Singh M, Thomas RJ, Drechsel P, Noble AD (2014). Economics of salt‐induced land degradation and restoration. In: Natural resources forum. Wiley Online Library pp 282-295. https://doi.org/10.1111/1477-8947.12054

Qureshi AS, Daba AW (2020). Evaluating growth and yield parameters of five quinoa (Chenopodium quinoa w.) genotypes under different salt stress conditions. Journal of Agricultural Science 12(128):10-5539. https://doi.org/10.5539/jas.v12n3p128

Razzaghi F, Ahmadi SH, Adolf VI, Jensen CR, Jacobsen S-E, Andersen MN (2011). Water relations and transpiration of quinoa (Chenopodium quinoa Willd.) under salinity and soil drying. Journal of Agronomy and Crop Science 197(5):348-360. https://doi.org/10.1111/j.1439-037X.2011.00473.x

Razzaghi F, Ahmadi SH, Jacobsen S-E, Jensen CR, Andersen MN (2012). Effects of salinity and soil-drying on radiation use efficiency, water productivity, seed set and final yield of field-grown quinoa (Chenopodium quinoa Willd.). Journal of Agronomy and Crop Science 198(3):173-184. https://doi.org/10.1111/j.1439-037X.2011.00496.x

Reisizadeh A, Amerian M, Gholami A (2024). Effect of spermidine and salicylic acid application on the morphological and physiological characteristics of quinoa (Chenopodium quinoa) under salt stress conditions. Agricultural Research 13(3):450-464. https://doi.org/10.1007/s40003-024-00710-0

Rekaby SA, Awad M, Majrashi A, Ali EF, Eissa MA (2021). Corn cob-derived biochar improves the growth of saline-irrigated quinoa in different orders of Egyptian soils. Horticulturae 7(8):221. https://doi.org/10.3390/horticulturae7080221

Repo-Carrasco R, Espinoza C, Jacobsen S-E (2003). Nutritional value and use of the Andean crops quinoa (Chenopodium quinoa) and kañiwa (Chenopodium pallidicaule). Food Reviews International 19(1-2):179-89. https://doi.org/10.1081/FRI-120018884

Rezzouk FZ, Shahid MA, Elouafi IA, Zhou B, Araus JL, Serret MD (2020). Agronomic performance of irrigated quinoa in desert areas: Comparing different approaches for early assessment of salinity stress. Agricultural Water Management 240:106205. https://doi.org/10.1016/j.agwat.2020.106205

Rhoades JD (1996) 5 Methods of soil analysis: Part 3 Chemical methods salinity: Electrical conductivity and total dissolved solids. Wiley Online Library pp 417-435. https://doi.org/10.2136/sssabookser5.3.c14

Riaz F, Abbas G, Saqib M, Amjad M, Farooq A, Ahmad S, ... Ahmed K (2020). Comparative effect of salinity on growth, ionic and physiological attributes of two quinoa genotypes. Pakistan Journal of Agricultural Sciences 57(1). https://doi.org/10.21162/PAKJAS/20.9018

Riccardi M, Mele G, Pulvento C, Lavini A, D’Andria R, Jacobsen S-E (2014). Non-destructive evaluation of chlorophyll content in quinoa and amaranth leaves by simple and multiple regression analysis of RGB image components. Photosynthesis Research 120(3):263-272. https://doi.org/10.1007/s11120-014-9970-2

Roman VJ, den Toom LA, Gamiz CC, van der Pijl N, Visser RGF, van Loo EN, van der Linden CG (2020). Differential responses to salt stress in ion dynamics, growth and seed yield of European quinoa varieties. Environmental and Experimental Botany 177:104146. https://doi.org/10.1016/j.envexpbot.2020.104146

Romano N, Ureta MM, Guerrero-Sanchez M, Gómez-Zavaglia A (2020). Nutritional and technological properties of a quinoa (Chenopodium quinoa Willd.) spray-dried powdered extract. Food Research International 129:108884. https://doi.org/10.1016/j.foodres.2019.108884

Rorat T (2006). Plant dehydrins-tissue location, structure and function. Cellular & Molecular Biology Letters 11(4):536-556. https://doi.org/10.2478/s11658-006-0044-0

Ruffino AMC, Rosa M, Hilal M, González JA, Prado FE (2010). The role of cotyledon metabolism in the establishment of quinoa (Chenopodium quinoa) seedlings growing under salinity. Plant and Soil 326(1):213-224. https://doi.org/10.1007/s11104-009-9999-8

Ruiz KB, Biondi S, Martínez EA, Orsini F, Antognoni F, Jacobsen S-E (2016). Quinoa-a model crop for understanding salt-tolerance mechanisms in halophytes. Plant Biosystems-An International Journal Dealing with all Aspects of Plant Biology 150(2):357-371. https://doi.org/10.1080/11263504.2015.1027317

Ruiz KB, Stefania B, Rómulo O, Ian A-R, Fabiana A, Enrique, ... Molina-Montenegro MA (2013). Quinoa biodiversity and sustainability for food security under climate change. A review. Agronomy for Sustainable Development 34:349-359. https://doi.org/10.1007/s13593-013-0195-0

Ruiz-Carrasco K, Antognoni F, Coulibaly AK, Lizardi S, Covarrubias A, Martínez EA, ... Zurita-Silva A (2011). Variation in salinity tolerance of four lowland genotypes of quinoa (Chenopodium quinoa Willd.) as assessed by growth, physiological traits, and sodium transporter gene expression. Plant Physiology and Biochemistry 49(11):1333-1341. https://doi.org/10.1016/j.plaphy.2011.08.005

Saavedra L, Svensson J, Carballo V, Izmendi D, Welin B, Vidal S (2006). A dehydrin gene in Physcomitrella patens is required for salt and osmotic stress tolerance. The Plant Journal 45(2):237-249. https://doi.org/10.1111/j.1365-313X.2005.02603.x

Salama AM, Seleem E, Abd El Salam R, Ghoniem A (2021). Response of quinoa plant grown under drought stress to foliar application with salicylic acid, paclobutrazol and algae extract. Scientific Journal of Agricultural Sciences 3(2):87-104. https://doi.org/10.21608/sjas.2021.81529.1118

Salma T, Mohamed A, Abderrahim B, Raja B-L, Wissal B, Hela BA, Meddich A (2023). Combined use of mycorrhizae and green compost for reducing the deleterious effects of salt stress in two genotypes of quinoa (Chenopodium quinoa). Journal of Soil Science and Plant Nutrition 23(1):1254-1271. https://doi.org/10.1007/s42729-022-01118-x

Santos J, Al-Azzawi M, Aronson J, Flowers TJ (2016). eHALOPH a database of salt-tolerant plants: helping put halophytes to work. Plant and Cell Physiology 57(1):10. https://doi.org/10.1093/pcp/pcv155

Schubert S, Qadir M (2024). Important parameters for the characterization of salt-affected soils. In: Schubert S, Qadir M (Eds). Soil salinity and salt resistance of crop plants. Springer International Publishing, Cham pp 15-32. https://doi.org/10.1007/978-3-031-73250-8_2

Scotti R, Pane C, Spaccini R, Palese AM, Piccolo A, Celano G, Zaccardelli M (2016). On-farm compost: a useful tool to improve soil quality under intensive farming systems. Applied Soil Ecology 107:13-23. https://doi.org/10.1016/j.apsoil.2016.05.004

Sellami MH, Pulvento C, Lavini A (2020). Agronomic practices and performances of quinoa under field conditions: A systematic review. Plants 10(1):72. https://doi.org/10.3390/plants10010072

Sen A, Johnson R, Puthur JT (2021). Seed priming: A cost-effective strategy to impart abiotic stress tolerance In: Husen A (Eds). Plant performance under environmental stress: Hormones, biostimulants and sustainable plant growth management. Springer International Publishing, Cham pp 459-480. https://doi.org/10.1007/978-3-030-78521-5_18

Serrat X, Quello A, Manikan B, Lino G, Nogués S (2024). Comparative salt-stress responses in salt-tolerant (Vikinga) and salt-sensitive (Regalona) quinoa varieties. Physiological, Anatomical and Biochemical Perspectives. Agronomy 14(12):3003. https://doi.org/10.3390/agronomy14123003

Shabala L, Cuin TA, Newman IA, Shabala S (2005). Salinity-induced ion flux patterns from the excised roots of Arabidopsis sos mutants. Planta 222(6):1041-1050. https://doi.org/10.1007/s00425-005-0074-2

Shabala L, Mackay A, Tian Y, Jacobsen S, Zhou D, Shabala S (2012). Oxidative stress protection and stomatal patterning as components of salinity tolerance mechanism in quinoa (Chenopodium quinoa). Physiologia Plantarum 146(1):26-38. https://doi.org/10.1111/j.1399-3054.2012.01599.x

Shabala S, Demidchik V, Shabala L, Cuin TA, Smith SJ, Miller AJ, Davies JM, Newman IA (2006). Extracellular Ca2+ ameliorates NaCl-induced K+ loss from Arabidopsis root and leaf cells by controlling plasma membrane K+ permeable channels. Plant Physiology 141(4):1653-1665. https://doi.org/10.1104/pp.106.082388

Shabala S, Hariadi Y, Jacobsen S-E (2013). Genotypic difference in salinity tolerance in quinoa is determined by differential control of xylem Na+ loading and stomatal density. Journal of Plant Physiology 170(10):906-914. https://doi.org/10.1016/j.jplph.2013.01.014

Shabala S, Munns R (2017). Salinity stress: physiological constraints and adaptive mechanisms. In: Plant stress physiology. Cabi Wallingford pp 24-63. https://doi.org/10.1079/9781780647296.0024

Shabala S, Shabala L (2011). Ion transport and osmotic adjustment in plants and bacteria. Biomolecular concepts 2(5):151-187. https://doi.org/10.1515/BMC.2011.032

Shahid SA, Zaman M, Heng L (2018). Soil salinity: Historical perspectives and a world overview of the problem. In: Guideline for salinity assessment, mitigation and adaptation using nuclear and related techniques. SpringeR, Cham pp 43-53. https://doi.org/10.1007/978-3-319-96190-3_2

Shani U, Ben-Gal A (2005) Long-term response of grapevines to salinity: osmotic effects and ion toxicity. American Journal of Enology and Viticulture 56(2):148-154. https://doi.org/10.5344/ajev.2005.56.2.148

Shannon MC, Grieve CM (1998). Tolerance of vegetable crops to salinity. Scientia Horticulturae 78(1-4):5-38. https://doi.org/10.1016/S0304-4238(98)00189-7

Sharma P, Jha AB, Dubey RS, Pessarakli M (2012). Reactive oxygen species, oxidative damage, and antioxidative defense mechanism in plants under stressful conditions. Journal of botany 2012(1):217037. https://doi.org/10.1155/2012/217037

Shuyskaya E, Rakhmankulova Z, Prokofieva M, Kazantseva V, Lunkova N (2023). Impact of salinity, elevated temperature, and their interaction with the photosynthetic efficiency of halophyte crop Chenopodium quinoa Willd. Agriculture 13(6):1198. https://doi.org/10.3390/agriculture13061198

Slatni T, Ben Slimene I, Harzalli Z, Taamalli W, Smaoui A, Abdelly C, Elkahoui S (2024). Enhancing quinoa (Chenopodium quinoa) growth in saline environments through salt‐tolerant rhizobacteria from halophyte biotope. Physiologia Plantarum 176(4):14466. https://doi.org/10.1111/ppl.14466

Slimani N, Arraouadi S, Hajlaoui H, Borgi MA, Boughattas NE, De Feo V, Snoussi M (2023). the impact of greenhouse and field growth conditions on Chenopodium quinoa willd accessions’ response to salt stress: A comparative approach. Agronomy 13(9):2303. https://doi.org/10.3390/agronomy13092303

Sofy MR, Sharaf AEM, Fouda HM (2016). Stimulatory effect of hormones, vitamin C on growth, yield and some metabolic activities of Chenopodium quinoa plants in Egypt. Journal of Plant Biochemistry & Physiology 4(161):4-10. https://doi.org/10.4172/2329-9029.1000161

Stavi I, Thevs N, Priori S (2021). Soil salinity and sodicity in drylands: a review of causes, effects, monitoring, and restoration measures. Frontiers in Environmental Science 9:712831. https://doi.org/10.3389/fenvs.2021.712831

Stoleru V, Slabu C, Vitanescu M, Peres C, Cojocaru A, Covasa M, Mihalache G (2019). Tolerance of three quinoa cultivars (Chenopodium quinoa Willd.) to salinity and alkalinity stress during germination stage. Agronomy 9(6):287. https://doi.org/10.3390/agronomy9060287

Sun W, Yao M, Wang Z, Chen Y, Zhan J, Yan J, ... Wu Q (2022). Involvement of auxin-mediated CqEXPA50 contributes to salt tolerance in quinoa (Chenopodium quinoa) by interaction with auxin pathway genes. International Journal of Molecular Sciences 23(15):8480. https://doi.org/10.3390/ijms23158480

Sun Y, Lindberg S, Shabala L, Morgan S, Shabala S, Jacobsen S-E (2017). A comparative analysis of cytosolic Na+ changes under salinity between halophyte quinoa (Chenopodium quinoa) and glycophyte pea (Pisum sativum). Environmental and Experimental Botany 141:154-160. https://doi.org/10.1016/j.envexpbot.2017.07.003

Sun Y, Liu F, Bendevis M, Shabala S, Jacobsen S (2014). Sensitivity of two quinoa (Chenopodium quinoa Willd.) varieties to progressive drought stress. Journal of Agronomy and Crop Science 200(1):12-23. https://doi.org/10.1111/jac.12042

Surekha CH, Kumari KN, Aruna L V, Suneetha G, Arundhati A, Kavi Kishor PB (2014). Expression of the Vigna aconitifolia P5CSF129A gene in transgenic pigeonpea enhances proline accumulation and salt tolerance. Plant Cell, Tissue and Organ Culture (PCTOC) 116(1):27-36. https://doi.org/10.1007/s11240-013-0378-z

Surekha CH, Neelapu NRR, Kamala G, Prasad BS, Ganesh PS (2013). Efficacy of Trichoderma viride to induce disease resistance and antioxidant responses in legume Vigna mungo infested by Fusarium oxysporum and Alternaria alternata. International Journal of Agricultural Science and Research 3(2):285-294.

Suzuki N, Rivero RM, Shulaev V, Blumwald E, Mittler R (2014). Abiotic and biotic stress combinations. New Phytologist 203(1):32-43. https://doi.org/10.1111/nph.12797

Talebnejad R, Sepaskhah AR (2016). Physiological characteristics, gas exchange, and plant ion relations of quinoa to different saline groundwater depths and water salinity. Archives of Agronomy and Soil Science 62(10):1347-1367. https://doi.org/10.1080/03650340.2016.1144925

Tanaka Y, Sasaki N, Ohmiya A (2008). Biosynthesis of plant pigments: anthocyanins, betalains and carotenoids. The Plant Journal 54:733-749. https://doi.org/10.1111/j.1365-313X.2008.03447.x

Tang Y, Li X, Zhang B, Chen PX, Liu R, Tsao R (2015). Characterisation of phenolics, betanins and antioxidant activities in seeds of three Chenopodium quinoa Willd. genotypes. Food Chemistry 166:380-388. https://doi.org/10.1016/j.foodchem.2014.06.018

Tarolli P, Luo J, Park E, Barcaccia G, Masin R (2024). Soil salinization in agriculture: Mitigation and adaptation strategies combining nature-based solutions and bioengineering. Iscience 27(2):108830. https://doi.org/10.1016/j.isci.2024.108830

Teakle NL, Tyerman SD (2010). Mechanisms of Cl‐transport contributing to salt tolerance. Plant, Cell & Environment 33(4):566-589. https://doi.org/10.1111/j.1365-3040.2009.02060.x

Toderich KN, Mamadrahimov AA, Khaitov BB, Karimov AA, Soliev AA, Nanduri KR, Shuyskaya E V (2020). Differential impact of salinity stress on seeds minerals, storage proteins, fatty acids, and squalene composition of new quinoa genotype, grown in hyper-arid desert environments. Frontiers in Plant Science 11:607102. https://doi.org/10.3389/fpls.2020.607102

Toubali S, Meddich A (2023). Role of combined use of mycorrhizae fungi and plant growth promoting rhizobacteria in the tolerance of quinoa plants under salt stress. Gesunde Pflanzen 75(5):1855-1869. https://doi.org/10.1007/s10343-023-00847-y

Turcios AE, Papenbrock J, Tränkner M (2021). Potassium, an important element to improve water use efficiency and growth parameters in quinoa (Chenopodium quinoa) under saline conditions. Journal of Agronomy and Crop Science 207(4):618-630. https://doi.org/10.1111/jac.12477

Ullah A, Bano A, Khan N (2021). Climate change and salinity effects on crops and chemical communication between plants and plant growth-promoting microorganisms under stress. Frontiers in Sustainable Food Systems 5:618092. https://doi.org/10.3389/fsufs.2021.618092

Vali A, Zahedi H, Alipour A, Sharghi Y, Naeini MR (2024). Differential effects of foliar and seed priming glycine betaine application on quinoa physiology under varying salinity level. South African Journal of Botany 175:253-267. https://doi.org/10.1016/j.sajb.2024.10.009

Wakeda M, Mohamed WH, Gad HI (2023). Evaluating the role of some soaking materials in enhancing the growth characteristics and nutrient contents of quinoa crop grown under saline soil condition. Alexandria Journal of Soil and Water Sciences 7(1):14-27. https://doi.org/10.21608/ajsws.2023.183147.1007

Wang X, Bai J, Wang W, Zhang G, Yin S, Wang D (2021). A comparative metabolomics analysis of the halophyte Suaeda salsa and Salicornia europaea. Environmental Geochemistry and Health 43(3):1109-1122. https://doi.org/10.1007/s10653-020-00569-4

Wicke B, Smeets E, Dornburg V, Vashev B, Gaiser T, Turkenburg W, Faaij A (2011). The global technical and economic potential of bioenergy from salt-affected soils. Energy & Environmental Science 4(8):2669-2681. https://doi.org/10.3390/environsciproc2022016023

Wilson C, Read JJ, Abo-Kassem E (2002). Effect of mixed-salt salinity on growth and ion relations of a quinoa and a wheat variety. Journal of Plant Nutrition 25(12):2689-2704. https://doi.org/10.1081/PLN-120015532

Wright KH, Pike OA, Fairbanks DJ, Huber CS (2002). Composition of Atriplex hortensis, sweet and bitter Chenopodium quinoa seeds. Journal of Food Science 67(4):1383-1385. https://doi.org/10.1111/j.1365-2621.2002.tb10294.x

Wu G, Peterson AJ, Morris CF, Murphy KM (2016). Quinoa seed quality response to sodium chloride and sodium sulfate salinity. Frontiers in Plant Science 7:790. https://doi.org/10.3389/fpls.2016.00790

Yañez-Yazlle MF, Romano-Armada N, Acreche MM, Rajal VB, Irazusta VP (2021). Halotolerant bacteria isolated from extreme environments induce seed germination and growth of chia (Salvia hispanica L.) and quinoa (Chenopodium quinoa Willd.) under saline stress. Ecotoxicology and Environmental Safety 218:112273. https://doi.org/10.1016/j.ecoenv.2021.112273

Yang A, Akhtar SS, Iqbal S, Amjad M, Naveed M, Zahir ZA, Jacobsen S-E (2016). Enhancing salt tolerance in quinoa by halotolerant bacterial inoculation. Functional Plant Biology 43(7):632-642. https://doi.org/10.1071/FP15265

Yang A, Akhtar SS, Iqbal S, Qi Z, Alandia G, Saddiq MS, Jacobsen S (2018). Saponin seed priming improves salt tolerance in quinoa. Journal of Agronomy and Crop Science 204(1):31-39. https://doi.org/10.1111/jac.12229

Yang A, Akhtar SS, Li L, Fu Q, Li Q, Naeem MA, ... Jacobsen S-E (2020). Biochar mitigates combined effects of drought and salinity stress in quinoa. Agronomy 10(6):912. https://doi.org/10.3390/agronomy10060912

Zhang J, Jia W, Yang J, Ismail AM (2006). Role of ABA in integrating plant responses to drought and salt stresses. Field Crops Research 97(1):111-119. https://doi.org/10.1016/j.fcr.2005.08.018