Free Standard AU & NZ Shipping For All Book Orders Over $80!
Register      Login
Crop and Pasture Science Crop and Pasture Science Society
Plant sciences, sustainable farming systems and food quality
REVIEW

Effects, tolerance mechanisms and management of salt stress in lucerne (Medicago sativa)

Safaa Mohammed Al-Farsi A B , Ahmad Nawaz C , Anees-ur-Rehman C , Saleem K. Nadaf D , Abdullah M. Al-Sadi A , Kadambot H. M. Siddique E and Muhammad Farooq https://orcid.org/0000-0003-4368-9357 A E F
+ Author Affiliations
- Author Affiliations

A Department of Plant Sciences, College of Agricultural and Marine Sciences, Sultan Qaboos University, Al-Khoud 123, Oman.

B Directorate General of Agriculture and Livestock Research, Ministry of Agriculture and Fisheries, Al-Seeb 121, Oman.

C College of Agriculture, Bahauddin Zakariya University, Bahadur Sub-Campus, Layyah 31200, Pakistan.

D Oman Animal and Plant Genetic Resources Center, The Research Council, PO Box 92, Muscat 123, Oman.

E The UWA Institute of Agriculture, The University of Western Australia, Stirling Highway, Crawley, WA 6009, Australia.

F Corresponding author. Email: farooqcp@squ.edu.om

Crop and Pasture Science 71(5) 411-428 https://doi.org/10.1071/CP20033
Submitted: 10 February 2020  Accepted: 19 April 2020   Published: 15 May 2020

Abstract

Lucerne (alfalfa, Medicago sativa L.) is a forage legume that is widely cultivated in arid and semi-arid regions of the world. The main aim of this review was to highlight the effects of salt stress on the performance of lucerne and to suggest different tolerance mechanisms and management strategies for improving its yield under salt stress. Salt stress significantly affects seed germination, carbon fixation, light harvesting, biological N2 fixation, mineral uptake and assimilation and dry-matter accumulation in lucerne. Accumulation of osmolytes or compatible solutes such as proline, polyamines, trehalose and soluble sugars confers salt tolerance in lucerne. Maintenance of low Na+ : K+ ratios, antioxidant enzyme activation, and hormonal regulation also help lucerne to withstand salt stress. The screening of diverse genotypes on the basis of germination indices, gas exchange, biomass production, lipid peroxidation and antioxidant enzymes might be useful for breeding salt-tolerant lucerne genotypes. Novel biotechnological tools and functional genomics used to identify salt-conferring genes and quantitative trait loci will help to improve salt tolerance. Use of rhizobial and non-rhizobial plant growth-promoting bacteria, arbuscular mycorrhizal fungi, exogenous application of osmoprotectants, and seed priming with brassinolide, gibberellic acid and salicylic acid may help to improve lucerne performance in saline environments.

Additional keywords: antioxidants, arbuscular mycorrhizal fungi, osmoprotectants, seed priming, ion toxicity.


References

Agarwal S, Singh B, Kumar P (2010) Salinity tolerance of some cultivars of berseem (Trifolium alexandrinum L.) during germination and early seedling growth. Vegetos—An International Journal of Plant Research 23, 63–82.

Ai‐Ke B, Zheng‐Gang G, Hong‐Fei Z, Suo‐Min W (2009) A procedure for assessing the salt tolerance of lucerne (Medicago sativa L.) cultivar seedlings by combining agronomic and physiological indicators. New Zealand Journal of Agricultural Research 52, 435–442.
A procedure for assessing the salt tolerance of lucerne (Medicago sativa L.) cultivar seedlings by combining agronomic and physiological indicators.Crossref | GoogleScholarGoogle Scholar |

Al Khanjari S, Al Khathiri A, Esechie HA (2002) Variation in chlorophyll meter readings, nodulation and dry matter yield of alfalfa (Medicago sativa L.) cultivars differing in salt tolerance. Crop Research 24, 350–356.

Al-Khateeb SA (2006) Effect of calcium/sodium ratio on growth and ion relations of alfalfa (Medicago sativa L.) seedling grown under saline condition. Journal of Agronomy 5, 175–181.
Effect of calcium/sodium ratio on growth and ion relations of alfalfa (Medicago sativa L.) seedling grown under saline condition.Crossref | GoogleScholarGoogle Scholar |

Al-Khatib MM (1991) An assessment of the potential for improving salt tolerance in alfalfa (Medicago sativa L.). PhD Thesis, University of Liverpool, Liverpool, UK.

Al-Khatib M, McNeilly T, Collins JC (1992) The potential of selection and breeding for improved salt tolerance in lucerne (Medicago sativa L.). Euphytica 65, 43–51.
The potential of selection and breeding for improved salt tolerance in lucerne (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar |

Al-Khatib MM, McNeilly T, Collins JC (1994) Between and within cultivar variability in salt tolerance in lucerne (Medicago sativa L.). Genetic Resources and Crop Evolution 41, 159–164.
Between and within cultivar variability in salt tolerance in lucerne (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar |

Al-Niemi TS, Campbell WF, Rumbaugh MD (1992) Response of alfalfa cultivars to salinity during germination and post-germination growth. Crop Science 32, 976–980.
Response of alfalfa cultivars to salinity during germination and post-germination growth.Crossref | GoogleScholarGoogle Scholar |

Allen SG, Dobrenz AK, Scharnhorst M, Stoner JEA (1985) Heritability of NaCl tolerance in germinating alfalfa seeds. Agronomy Journal 77, 99–101.
Heritability of NaCl tolerance in germinating alfalfa seeds.Crossref | GoogleScholarGoogle Scholar |

Amooaghaie R (2011) The effect of hydro and osmopriming on alfalfa seed germination and antioxidant defenses under salt stress. African Journal of Biotechnology 10, 6269–6275.

Amooaghaie R, Tabatabaie F (2017) Osmopriming-induced salt tolerance during seed germination of alfalfa most likely mediates through H2O2 signaling and upregulation of heme oxygenase. Protoplasma 254, 1791–1803.
Osmopriming-induced salt tolerance during seed germination of alfalfa most likely mediates through H2O2 signaling and upregulation of heme oxygenase.Crossref | GoogleScholarGoogle Scholar | 28093607PubMed |

Anand A, Baig MJ, Mandal PK (2000) Response of alfalfa genotypes to saline water irrigation. Biologia Plantarum 43, 455–457.
Response of alfalfa genotypes to saline water irrigation.Crossref | GoogleScholarGoogle Scholar |

Arraouadi S, Chardon F, Huguet T, Aouani ME, Badri M (2011) QTLs mapping of morphological traits related to salt tolerance in Medicago truncatula. Acta Physiologiae Plantarum 33, 917–926.
QTLs mapping of morphological traits related to salt tolerance in Medicago truncatula.Crossref | GoogleScholarGoogle Scholar |

Arraouadi S, Badri M, Abdelly C, Huguet T, Aouani ME (2012) QTL mapping of physiological traits associated with salt tolerance in Medicago truncatula recombinant inbred lines. Genomics 99, 118–125.
QTL mapping of physiological traits associated with salt tolerance in Medicago truncatula recombinant inbred lines.Crossref | GoogleScholarGoogle Scholar | 22178264PubMed |

Ashraf M, McNeilly T, Bradshaw AD (1987) Selection and heritability of tolerance to sodium chloride in four forage species. Crop Science 27, 232–234.
Selection and heritability of tolerance to sodium chloride in four forage species.Crossref | GoogleScholarGoogle Scholar |

Ashrafi E, Zahedi M, Razmjoo J (2014a) Co-inoculations of arbuscular mycorrhizal fungi and rhizobia under salinity in alfalfa. Soil Science and Plant Nutrition 60, 619–629.
Co-inoculations of arbuscular mycorrhizal fungi and rhizobia under salinity in alfalfa.Crossref | GoogleScholarGoogle Scholar |

Ashrafi E, Razmjoo J, Zahedi M, Pessarakli M (2014b) Selecting alfalfa cultivars for salt tolerance based on some physiochemical traits. Agronomy Journal 106, 1758–1764.
Selecting alfalfa cultivars for salt tolerance based on some physiochemical traits.Crossref | GoogleScholarGoogle Scholar |

Ashrafi E, Razmjoo J, Zahedi M, Pessarakli M (2015) Screening alfalfa for salt tolerance based on lipid peroxidation and antioxidant enzymes. Agronomy Journal 107, 167–173.
Screening alfalfa for salt tolerance based on lipid peroxidation and antioxidant enzymes.Crossref | GoogleScholarGoogle Scholar |

Ashrafi E, Razmjoo J, Zahedi M (2018) Effect of salt stress on growth and ion accumulation of alfalfa (Medicago sativa L.) cultivars. Journal of Plant Nutrition 41, 818–831.
Effect of salt stress on growth and ion accumulation of alfalfa (Medicago sativa L.) cultivars.Crossref | GoogleScholarGoogle Scholar |

Aydi S, Drevon JJ, Abdelly C (2004) Effect of salinity on root-nodule conductance to the oxygen diffusion in the Medicago truncatulaSinorhizobium meliloti symbiosis. Plant Physiology and Biochemistry 42, 833–840.
Effect of salinity on root-nodule conductance to the oxygen diffusion in the Medicago truncatulaSinorhizobium meliloti symbiosis.Crossref | GoogleScholarGoogle Scholar | 15596103PubMed |

Azcón R, El-Atrash F (1997) Influence of arbuscular mycorrhizae and phosphorus fertilization on growth, nodulation and N2 fixation (15N) in Medicago sativa at four salinity levels. Biology and Fertility of Soils 24, 81–86.
Influence of arbuscular mycorrhizae and phosphorus fertilization on growth, nodulation and N2 fixation (15N) in Medicago sativa at four salinity levels.Crossref | GoogleScholarGoogle Scholar |

Badran AE, ElSherebeny EA, Salama YA (2015) Performance of some alfalfa cultivars under salinity stress conditions. The Journal of Agricultural Science 7, 281–290.

Baha N, Bekki A (2015) An approach of improving plant salt tolerance of lucerne (Medicago sativa) grown under salt stress: use of bio-inoculants. Journal of Plant Growth Regulation 34, 169–182.
An approach of improving plant salt tolerance of lucerne (Medicago sativa) grown under salt stress: use of bio-inoculants.Crossref | GoogleScholarGoogle Scholar |

Bansal S (2016) Effect of salinity on biological nitrogen fixation in alfalfa (Medicago sativa) and its response to applied mineral nitrogen. PhD Thesis, Jordon College of Agricultural Sciences and Technology, California State University, Fresno, CA, USA.

Basalah MO, Mohammad S (1999) Effect of salinity and plant growth regulators on seed germination of Medicago sativa L. Pakistan Journal of Biological Sciences 2, 651–653.
Effect of salinity and plant growth regulators on seed germination of Medicago sativa L.Crossref | GoogleScholarGoogle Scholar |

Ben Salah IB, Albacete A, Andújar CM, Haouala R, Labidi N, Zribi F, Martinez V, Pérez-Alfocea F, Abdelly C (2009) Response of nitrogen fixation in relation to nodule carbohydrate metabolism in Medicago ciliaris lines subjected to salt stress. Journal of Plant Physiology 166, 477–488.
Response of nitrogen fixation in relation to nodule carbohydrate metabolism in Medicago ciliaris lines subjected to salt stress.Crossref | GoogleScholarGoogle Scholar |

Bernstein L, Ogata G (1966) Effects of salinity on nodulation, nitrogen fixation, and growth of soybeans and alfalfa 1. Agronomy Journal 58, 201–203.
Effects of salinity on nodulation, nitrogen fixation, and growth of soybeans and alfalfa 1.Crossref | GoogleScholarGoogle Scholar |

Bertrand A, Dhont C, Bipfubusa M, Chalifour FP, Drouin P, Beauchamp CJ (2015) Improving salt stress responses of the symbiosis in alfalfa using salt-tolerant cultivar and rhizobial strain. Applied Soil Ecology 87, 108–117.
Improving salt stress responses of the symbiosis in alfalfa using salt-tolerant cultivar and rhizobial strain.Crossref | GoogleScholarGoogle Scholar |

Bertrand A, Bipfubusa M, Dhont C, Chalifour FP, Drouin P, Beauchamp CJ (2016) Rhizobial strains exert a major effect on the amino acid composition of alfalfa nodules under NaCl stress. Plant Physiology and Biochemistry 108, 344–352.
Rhizobial strains exert a major effect on the amino acid composition of alfalfa nodules under NaCl stress.Crossref | GoogleScholarGoogle Scholar | 27508354PubMed |

Bhardwaj S, Sharma NK, Srivastava PK, Gaurav S (2010) Salt tolerance assessment in alfalfa (Medicago sativa L.) ecotypes. Botany Research Journal 3, 1–6.
Salt tolerance assessment in alfalfa (Medicago sativa L.) ecotypes.Crossref | GoogleScholarGoogle Scholar |

Bose J, Rodrigo-Moreno A, Shabala S (2014) ROS homeostasis in halophytes in the context of salinity stress tolerance. Journal of Experimental Botany 65, 1241–1257.
ROS homeostasis in halophytes in the context of salinity stress tolerance.Crossref | GoogleScholarGoogle Scholar | 24368505PubMed |

Boughanmi N, Michonneau P, Daghfous D, Fleurat‐Lessard P (2005) Adaptation of Medicago sativa cv. Gabès to long‐term NaCl stress. Journal of Plant Nutrition and Soil Science 168, 262–268.
Adaptation of Medicago sativa cv. Gabès to long‐term NaCl stress.Crossref | GoogleScholarGoogle Scholar |

Campanelli A, Ruta C, Morone-Fortunato I, De Mastro G (2013a) Alfalfa (Medicago sativa L.) clones tolerant to salt stress: in vitro selection. Central European Journal of Biology 8, 765–776.

Campanelli A, Ruta C, De Mastro G, Morone-Fortunato I (2013b) The role of arbuscular mycorrhizal fungi in alleviating salt stress in Medicago sativa L. var. icon. Symbiosis 59, 65–76.
The role of arbuscular mycorrhizal fungi in alleviating salt stress in Medicago sativa L. var. icon.Crossref | GoogleScholarGoogle Scholar |

Cornacchione MV, Suarez DL (2015) Emergence, forage production, and ion relations of alfalfa in response to saline waters. Crop Science 55, 444–457.
Emergence, forage production, and ion relations of alfalfa in response to saline waters.Crossref | GoogleScholarGoogle Scholar |

Cornacchione MV, Suarez DL (2017) Evaluation of alfalfa (Medicago sativa L.) populations’ response to salinity stress. Crop Science 57, 137–150.
Evaluation of alfalfa (Medicago sativa L.) populations’ response to salinity stress.Crossref | GoogleScholarGoogle Scholar |

Deinlein U, Stephan AB, Horie T, Luo W, Xu G, Schroeder J (2014) Plant salt tolerance mechanisms. Trends in Plant Science 19, 371–379.
Plant salt tolerance mechanisms.Crossref | GoogleScholarGoogle Scholar | 24630845PubMed |

Delgado MJ, Ligero F, Lluch C (1994) Effects of salt stress on growth and nitrogen fixation by pea, faba-bean, common bean, and soybean plants. Soil Biology & Biochemistry 26, 371–376.
Effects of salt stress on growth and nitrogen fixation by pea, faba-bean, common bean, and soybean plants.Crossref | GoogleScholarGoogle Scholar |

Díaz FJ, Grattan SR, Reyes JA, de la Roza-Delgado B, Benes SE, Jiménez C, Dorta M, Tejedor M (2018) Using saline soil and marginal quality water to produce alfalfa in arid climates. Agricultural Water Management 199, 11–21.
Using saline soil and marginal quality water to produce alfalfa in arid climates.Crossref | GoogleScholarGoogle Scholar |

Dissing-Nielsen J (1990) The effect of VAM on growth and uptake of nutrients in lucerne. Agriculture, Ecosystems & Environment 29, 99–102.
The effect of VAM on growth and uptake of nutrients in lucerne.Crossref | GoogleScholarGoogle Scholar |

Djilianov D, Prinsen E, Oden S, Van Onckelen H, Müller J (2003) Nodulation under salt stress of alfalfa lines obtained after in vitro selection for osmotic tolerance. Plant Science 165, 887–894.
Nodulation under salt stress of alfalfa lines obtained after in vitro selection for osmotic tolerance.Crossref | GoogleScholarGoogle Scholar |

Duc G, Agrama H, Bao S, Berger J, Bourion V, De Ron AM, Gowda CLL, Mikic A, Millot D, Singh KB, Tullu A, Vandenberg A, Vaz Patto MC, Warkentin TD, Zong X (2015) Breeding annual grain legumes for sustainable agriculture: new methods to approach complex traits and target new cultivar ideotypes. Critical Reviews in Plant Sciences 34, 381–411.
Breeding annual grain legumes for sustainable agriculture: new methods to approach complex traits and target new cultivar ideotypes.Crossref | GoogleScholarGoogle Scholar |

El-Nakhlawy FS, Shaheen MA, Al-Shareef AR (2012) Response of forage yield, protein and proline contents of alfalfa genotypes to irrigation water salinity and phosphorus fertilizer. Journal of Food, Agriculture and Environment 10, 551–557.

Endo T, Kubo‐Nakano Y, Lopez RA, Serrano RR, Larrinaga JA, Yamamoto S, Honna T (2014) Growth characteristics of kochia (Kochia scoparia L.) and alfalfa (Medicago sativa L.) in saline environments. Grassland Science 60, 225–232.
Growth characteristics of kochia (Kochia scoparia L.) and alfalfa (Medicago sativa L.) in saline environments.Crossref | GoogleScholarGoogle Scholar |

Esechie HA, Rodriguez V (1999) Does salinity inhibit alfalfa leaf growth by reducing tissue concentration of essential mineral nutrients? Journal of Agronomy & Crop Science 182, 273–278.
Does salinity inhibit alfalfa leaf growth by reducing tissue concentration of essential mineral nutrients?Crossref | GoogleScholarGoogle Scholar |

Evelin H, Kapoor R, Giri B (2009) Arbuscular mycorrhizal fungi in alleviation of salt stress: a review. Annals of Botany 104, 1263–1280.
Arbuscular mycorrhizal fungi in alleviation of salt stress: a review.Crossref | GoogleScholarGoogle Scholar | 19815570PubMed |

Fakhari F, Sadeghi H (2016) Investigating the effects of pod elimination on salinity tolerance in annual medic (Medicago scutellata L.). Journal of Rangeland Science 6, 232–241.

Fan X, Chang W, Feng F, Song F (2018) Responses of photosynthesis-related parameters and chloroplast ultrastructure to atrazine in alfalfa (Medicago sativa L.) inoculated with arbuscular mycorrhizal fungi. Ecotoxicology and Environmental Safety 166, 102–108.
Responses of photosynthesis-related parameters and chloroplast ultrastructure to atrazine in alfalfa (Medicago sativa L.) inoculated with arbuscular mycorrhizal fungi.Crossref | GoogleScholarGoogle Scholar | 30253284PubMed |

Farissi M, Bouizgaren A, Faghire M, Bargaz A, Ghoulam C (2011) Agro-physiological responses of Moroccan alfalfa (Medicago sativa L.) populations to salt stress during germination and early seedling stages. Seed Science and Technology 39, 389–401.
Agro-physiological responses of Moroccan alfalfa (Medicago sativa L.) populations to salt stress during germination and early seedling stages.Crossref | GoogleScholarGoogle Scholar |

Farissi M, Ghoulam C, Bouizgaren A (2013) Changes in water deficit saturation and photosynthetic pigments of alfalfa populations under salinity and assessment of proline role in salt tolerance. Agricultural Science Research Journal 3, 29–35.

Farooq M, Hussain M, Wakeel A, Siddique KHM (2015) Salt stress in maize effects resistance mechanisms and management: a review. Agronomy for Sustainable Development 35, 461–481.
Salt stress in maize effects resistance mechanisms and management: a review.Crossref | GoogleScholarGoogle Scholar |

Farooq M, Gogoi N, Hussain M, Barthakur S, Paul S, Bharadwaj N, Migdadi HM, Alghamdi SS, Siddique KHM (2017) Effects, tolerance mechanisms and management of salt stress in grain legumes. Plant Physiology and Biochemistry 118, 199–217.
Effects, tolerance mechanisms and management of salt stress in grain legumes.Crossref | GoogleScholarGoogle Scholar | 28648997PubMed |

Fernandez-Cornejo J, Wechsler SJ, Milkove DL (2016) The adoption of genetically engineered alfalfa, canola and sugarbeets in the United States. United States Department of Agriculture, Economic Research Service, Washington, DC, USA.

Ferreira J, Cornacchione M, Liu X, Suarez D (2015) Nutrient composition, forage parameters, and antioxidant capacity of alfalfa (Medicago sativa L.) in response to saline irrigation water. Agriculture 5, 577–597.
Nutrient composition, forage parameters, and antioxidant capacity of alfalfa (Medicago sativa L.) in response to saline irrigation water.Crossref | GoogleScholarGoogle Scholar |

Flexas J, Bota J, Loreto F, Cornic G, Sharkey TD (2004) Diffusive and metabolic limitations to photosynthesis under drought and salinity in C3 plants. Plant Biology 6, 269–279.
Diffusive and metabolic limitations to photosynthesis under drought and salinity in C3 plants.Crossref | GoogleScholarGoogle Scholar | 15143435PubMed |

Fougère F, Le Rudulier D, Streeter JG (1991) Effects of salt stress on amino acid, organic acid, and carbohydrate composition of roots, bacteroids, and cytosol of alfalfa (Medicago sativa L.). Plant Physiology 96, 1228–1236.
Effects of salt stress on amino acid, organic acid, and carbohydrate composition of roots, bacteroids, and cytosol of alfalfa (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar | 16668324PubMed |

Garg N, Chandel S (2011) Effect of mycorrhizal inoculation on growth, nitrogen fixation and nutrient uptake in Cicer arietinum (L.) under salt stress. Turkish Journal of Agriculture and Forestry 35, 205–214.

Garg N, Manchanda G (2008) Effect of arbuscular mycorrhizal inoculation of salt induced nodule senescence in Cajanus cajan L. (pigeon pea). Journal of Plant Growth Regulation 27, 115–124.
Effect of arbuscular mycorrhizal inoculation of salt induced nodule senescence in Cajanus cajan L. (pigeon pea).Crossref | GoogleScholarGoogle Scholar |

Garg N, Noor Z (2009) Genotypic differences in plant growth, osmotic and antioxidative defence of Cajanus cajan (L.) Millsp. modulated by salt stress. Archives of Agronomy and Soil Science 55, 3–33.
Genotypic differences in plant growth, osmotic and antioxidative defence of Cajanus cajan (L.) Millsp. modulated by salt stress.Crossref | GoogleScholarGoogle Scholar |

Geilfus CM, Mühling KH (2013) Ratiometric monitoring of transient apoplastic alkalinizations in the leaf apoplast of living Vicia faba plants: chloride primes and PM–H+‐ATPase shapes NaCl‐induced systemic alkalinizations. New Phytologist 197, 1117–1129.
Ratiometric monitoring of transient apoplastic alkalinizations in the leaf apoplast of living Vicia faba plants: chloride primes and PM–H+‐ATPase shapes NaCl‐induced systemic alkalinizations.Crossref | GoogleScholarGoogle Scholar | 23176077PubMed |

Ghasem F, Poustini K, Besharati H, Mohammadi VA, Abooei Mehrizi F, Goettfert M (2012) Pre-incubation of Sinorhizobium meliloti with luteolin, methyl jasmonate and genistein affecting alfalfa (Medicago sativa L.) growth, nodulation and nitrogen fixation under salt stress conditions. Journal of Agricultural Science and Technology 14, 1255–1264.

Goicoechea N, Antolín MC, Sánchez-Díaz M (1997) Influence of arbuscular mycorrhizae and Rhizobium on nutrient content and water relations in drought stressed alfalfa. Plant and Soil 192, 261–268.
Influence of arbuscular mycorrhizae and Rhizobium on nutrient content and water relations in drought stressed alfalfa.Crossref | GoogleScholarGoogle Scholar |

Goicoechea N, Szalai G, Antolin MC, Sanchez-Diaz M, Paldi E (1998) Influence of arbuscular mycorrhizae and Rhizobium on free polyamines and proline levels in water-stressed alfalfa. Journal of Plant Physiology 153, 706–711.
Influence of arbuscular mycorrhizae and Rhizobium on free polyamines and proline levels in water-stressed alfalfa.Crossref | GoogleScholarGoogle Scholar |

Gordon AJ, Minchin FR, Skot L, James CL (1997) Stress induced declines in soybean N2 fixation are related to nodule sucrose synthase activity. Plant Physiology 114, 937–946.
Stress induced declines in soybean N2 fixation are related to nodule sucrose synthase activity.Crossref | GoogleScholarGoogle Scholar | 12223754PubMed |

Grattan SR, Grieve CM, Poss JA, Robinson PH, Suarez DL, Benes SE (2004) Evaluation of salt-tolerant forages for sequential water reuse systems: III. Potential implications for ruminant mineral nutrition. Agricultural Water Management 70, 137–150.

Guan B, Zhou D, Zhang H, Tian Y, Japhet W, Wang P (2009) Germination responses of Medicago ruthenica seeds to salinity, alkalinity, and temperature. Journal of Arid Environments 73, 135–138.
Germination responses of Medicago ruthenica seeds to salinity, alkalinity, and temperature.Crossref | GoogleScholarGoogle Scholar |

Hager A (2003) Role of the plasma membrane H+-ATPase in auxin induced elongation growth: historical and new aspects. Journal of Plant Research 116, 483–505.
Role of the plasma membrane H+-ATPase in auxin induced elongation growth: historical and new aspects.Crossref | GoogleScholarGoogle Scholar | 12937999PubMed |

Hasegawa PM, Bressan RA, Zhu J, Bohnert BJ (2000) Plant cellular and molecular response to high salinity. Annual Review of Plant Biology 51, 463–499.
Plant cellular and molecular response to high salinity.Crossref | GoogleScholarGoogle Scholar |

Isla R, Aragüés R (2009) Response of alfalfa (Medicago sativa L.) to diurnal and nocturnal saline sprinkler irrigations. I: total dry matter and hay quality. Irrigation Science 27, 497–505.
Response of alfalfa (Medicago sativa L.) to diurnal and nocturnal saline sprinkler irrigations. I: total dry matter and hay quality.Crossref | GoogleScholarGoogle Scholar |

Jame YW, Biederbeck VO, Nicholaichuk W, Korven HC (1984) Salinity and alfalfa yield under effluent irrigation in southwestern Saskatchewan. Canadian Journal of Soil Science 64, 323–332.
Salinity and alfalfa yield under effluent irrigation in southwestern Saskatchewan.Crossref | GoogleScholarGoogle Scholar |

Jin T, Chang Q, Li W, Lin D, Li Z, Liu B, Liu L (2010a) Stress-inducible expression of GmDREB1 conferred salt tolerance in transgenic alfalfa. Plant Cell, Tissue and Organ Culture 100, 219–227.
Stress-inducible expression of GmDREB1 conferred salt tolerance in transgenic alfalfa.Crossref | GoogleScholarGoogle Scholar |

Jin H, Sun Y, Yang Q, Chao Y, Kang J, Jin H, Li Y, Margaret G (2010b) Screening of genes induced by salt stress from alfalfa. Molecular Biology Reports 37, 745–753.
Screening of genes induced by salt stress from alfalfa.Crossref | GoogleScholarGoogle Scholar | 19572213PubMed |

Jorjandi M, Sirchi GRS (2012) The effect of priming on germination and seedling growth of alfalfa (Medicago sativa L.) under salinity stress. Journal of Stress Physiology & Biochemistry 8, 234–239.

Julier B, Huyghe C, Ecalle C (2000) Within- and among cultivar genetic variation in alfalfa: forage quality, morphology, and yield. Crop Science 40, 365–369.
Within- and among cultivar genetic variation in alfalfa: forage quality, morphology, and yield.Crossref | GoogleScholarGoogle Scholar |

Kazemeini SA, Pirasteh-Anosheh HADI, Basirat A, Akram NA (2018) Salinity tolerance threshold of berseem clover (Trifolium alexandrinum) at different growth stages. Pakistan Journal of Botany 50, 1675–1680.

Khajeh-Hosseini M, Powell AA, Bingham IJ (2003) The interaction between salinity stress and seed vigor during germination of soybean seeds. Seed Science and Technology 31, 715–725.
The interaction between salinity stress and seed vigor during germination of soybean seeds.Crossref | GoogleScholarGoogle Scholar |

Khalifa AY, Almalki MA (2015) Isolation and characterization of an endophytic bacterium, Bacillus megaterium BMN1, associated with root-nodules of Medicago sativa L. growing in Al-Ahsaa region, Saudi Arabia. Annals of Microbiology 65, 1017–1026.
Isolation and characterization of an endophytic bacterium, Bacillus megaterium BMN1, associated with root-nodules of Medicago sativa L. growing in Al-Ahsaa region, Saudi Arabia.Crossref | GoogleScholarGoogle Scholar |

Khan MG, Silverbusch M, Lips SH (1994) Physiological studies on salinity and nitrogen interaction in alfalfa. I. Biomass production and root development. Journal of Plant Nutrition 17, 657–668.
Physiological studies on salinity and nitrogen interaction in alfalfa. I. Biomass production and root development.Crossref | GoogleScholarGoogle Scholar |

Khan MG, Silberbush M, Lips SH (1997) Responses of alfalfa to potassium, calcium, and nitrogen under stress induced by sodium chloride. Biologia Plantarum 40, 251–259.
Responses of alfalfa to potassium, calcium, and nitrogen under stress induced by sodium chloride.Crossref | GoogleScholarGoogle Scholar |

Khan MJ, Jan MT, Khan AU, Arif M, Shafi M (2010) Management of saline sodic soils through cultural practices and gypsum. Pakistan Journal of Botany 42, 4143–4155.

Khorshidi MB, Yarnia M, Hassanpanah D (2009) Salinity effect on nutrients accumulation in alfalfa shoots in hydroponic condition. Journal of Food, Agriculture and Environment 7, 787–790.

Larose G, Chênevert R, Moutoglis P, Gagné S, Piché Y, Vierheilig H (2002) Flavonoid levels in roots of Medicago sativa are modulated by the developmental stage of the symbiosis and the root colonizing arbuscular mycorrhizal fungus. Journal of Plant Physiology 159, 1329–1339.
Flavonoid levels in roots of Medicago sativa are modulated by the developmental stage of the symbiosis and the root colonizing arbuscular mycorrhizal fungus.Crossref | GoogleScholarGoogle Scholar |

Latrach L, Farissi M, Mouradi M, Makoudi B, Bouizgaren A, Ghoulam C (2014) Growth and nodulation of alfalfa–rhizobia symbiosis under salinity: electrolyte leakage, stomatal conductance, and chlorophyll fluorescence. Turkish Journal of Agriculture and Forestry 38, 320–326.
Growth and nodulation of alfalfa–rhizobia symbiosis under salinity: electrolyte leakage, stomatal conductance, and chlorophyll fluorescence.Crossref | GoogleScholarGoogle Scholar |

Lei Y, Xu Y, Hettenhausen C, Lu C, Shen G, Zhang C, Li J, Song J, Lin H, Wu J (2018) Comparative analysis of alfalfa (Medicago sativa L.) leaf transcriptomes reveals genotype-specific salt tolerance mechanisms. BMC Plant Biology 18, 35
Comparative analysis of alfalfa (Medicago sativa L.) leaf transcriptomes reveals genotype-specific salt tolerance mechanisms.Crossref | GoogleScholarGoogle Scholar | 29448940PubMed |

Li WYF, Wong FL, Tsai SN, Phang TH, Shao G, Lam HM (2006) Tonoplast located GmCLC1 and GmNHX1 from soybean enhance NaCl tolerance in transgenic bright yellow (BY)-2 cells. Plant, Cell & Environment 29, 1122–1137.
Tonoplast located GmCLC1 and GmNHX1 from soybean enhance NaCl tolerance in transgenic bright yellow (BY)-2 cells.Crossref | GoogleScholarGoogle Scholar |

Li R, Shi F, Fukuda K, Yang Y (2010a) Effects of salt and alkali stresses on germination, growth, photosynthesis and ion accumulation in alfalfa (Medicago sativa L.). Soil Science and Plant Nutrition 56, 725–733.
Effects of salt and alkali stresses on germination, growth, photosynthesis and ion accumulation in alfalfa (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar |

Li Y, Liu G, Gao H, Sun G, Zhao H, Xie N (2010b) A comprehensive evaluation of salt-tolerance and the physiological response of Medicago sativa at the seedling stage. Acta Prataculturae Sinica 19, 79–86.

Li H, Wang Z, Ke Q, Ji CY, Jeong JC, Lee HS, Lim YP, Xu B, Deng XP, Kwak SS (2014) Overexpression of codA gene confers enhanced tolerance to abiotic stresses in alfalfa. Plant Physiology and Biochemistry 85, 31–40.
Overexpression of codA gene confers enhanced tolerance to abiotic stresses in alfalfa.Crossref | GoogleScholarGoogle Scholar | 25394798PubMed |

Liu XP, Yu LX (2017) Genome-wide association mapping of loci associated with plant growth and forage production under salt stress in alfalfa (Medicago sativa L.). Frontiers in Plant Science 8, 853
Genome-wide association mapping of loci associated with plant growth and forage production under salt stress in alfalfa (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar | 28596776PubMed |

Liu ZH, Zhang HM, Li GL, Guo XL, Chen SY, Liu GB, Zhang YM (2011) Enhancement of salt tolerance in alfalfa transformed with the gene encoding for betaine aldehyde dehydrogenase. Euphytica 178, 363–372.
Enhancement of salt tolerance in alfalfa transformed with the gene encoding for betaine aldehyde dehydrogenase.Crossref | GoogleScholarGoogle Scholar |

Liu M, Wang T-Z, Zhang W-H (2015) Sodium extrusion associated with enhanced expression of SOS1 underlies different salt tolerance between Medicago falcata and Medicago truncatula seedlings. Environmental and Experimental Botany 110, 46–55.
Sodium extrusion associated with enhanced expression of SOS1 underlies different salt tolerance between Medicago falcata and Medicago truncatula seedlings.Crossref | GoogleScholarGoogle Scholar |

López M, Herrera-Cervera JA, Iribarne C, Tejera NA, Lluch C (2008a) Growth and nitrogen fixation in Lotus japonicus and Medicago truncatula under NaCl stress: nodule carbon metabolism. Journal of Plant Physiology 165, 641–650.
Growth and nitrogen fixation in Lotus japonicus and Medicago truncatula under NaCl stress: nodule carbon metabolism.Crossref | GoogleScholarGoogle Scholar | 17728011PubMed |

López M, Tejera NA, Iribarne C, Lluch C, Herrera‐Cervera JA (2008b) Trehalose and trehalase in root nodules of Medicago truncatula and Phaseolus vulgaris in response to salt stress. Physiologia Plantarum 134, 575–582.
Trehalose and trehalase in root nodules of Medicago truncatula and Phaseolus vulgaris in response to salt stress.Crossref | GoogleScholarGoogle Scholar | 18823327PubMed |

López M, Tejera NA, Lluch C (2009) Validamycin A improves the response of Medicago truncatula plants to salt stress by inducing trehalose accumulation in the root nodules. Journal of Plant Physiology 166, 1218–1222.
Validamycin A improves the response of Medicago truncatula plants to salt stress by inducing trehalose accumulation in the root nodules.Crossref | GoogleScholarGoogle Scholar | 19232773PubMed |

López-Gómez M, Tejera NA, Iribarne C, Herrera-Cervera JA, Lluch C (2012) Different strategies for salt tolerance in determined and indeterminate nodules of Lotus japonicus and Medicago truncatula. Archives of Agronomy and Soil Science 58, 1061–1073.
Different strategies for salt tolerance in determined and indeterminate nodules of Lotus japonicus and Medicago truncatula.Crossref | GoogleScholarGoogle Scholar |

López-Gómez M, Hidalgo-Castellanos J, Iribarne C, Lluch C (2014) Proline accumulation has prevalence over polyamines in nodules of Medicago sativa in symbiosis with Sinorhizobium meliloti during the initial response to salinity. Plant and Soil 374, 149–159.
Proline accumulation has prevalence over polyamines in nodules of Medicago sativa in symbiosis with Sinorhizobium meliloti during the initial response to salinity.Crossref | GoogleScholarGoogle Scholar |

Lou Y, Guan R, Sun M, Han F, He W, Wang H, Song F, Cui X, Zhuge Y (2018) Spermidine application alleviates salinity damage to antioxidant enzyme activity and gene expression in alfalfa. Ecotoxicology 27, 1323–1330.
Spermidine application alleviates salinity damage to antioxidant enzyme activity and gene expression in alfalfa.Crossref | GoogleScholarGoogle Scholar | 30244325PubMed |

Lu S, Guo H, Wang SM, Zhang X (2011) Effects of AM fungi on growth and physiological characters of Medicago sativa L. under NaCl stress. Journal of Soil and Water Conservation 10, 1–15.
Effects of AM fungi on growth and physiological characters of Medicago sativa L. under NaCl stress.Crossref | GoogleScholarGoogle Scholar |

Mahmood A, Athar M, Qadri R, Mahmood N (2008) Effect of NaCl salinity on growth, nodulation and total nitrogen content in Sesbania sesban. ACS. Agriculturae Conspectus Scientificus 73, 137–141.

McKimmie T, Dobrenz AK (1991) Ionic concentrations and water relations of alfalfa seedlings differing in salt tolerance. Agronomy Journal 83, 363–367.
Ionic concentrations and water relations of alfalfa seedlings differing in salt tolerance.Crossref | GoogleScholarGoogle Scholar |

Mezni M, Bizid E, Hamza M (1999) Effects of salinity of the irrigation waters on the survival and the growth of 3 cultivars of Prenne alfalfa (Medicago sativa L.). In ‘Proceedings 6th National Days on Results of the Agronomic Research’. Nabeult, Tunisia. pp. 558–565. (L'Institution de la Recherche et de l'Enseignement Supérieur Agricoles (IRESA): Tunis)

Mezni M, Albouchi A, Bizid E, Hamza M (2010) Minerals uptake, organic osmotica contents and water balance in alfalfa under salt stress. Journal of Phytology 2, 1–12.

Mohammadi H, Poustini K, Ahmadi A (2008) Root nitrogen remobilization and ion status of two alfalfa (Medicago sativa L.) cultivars in response to salinity stress. Journal of Agronomy & Crop Science 194, 126–134.
Root nitrogen remobilization and ion status of two alfalfa (Medicago sativa L.) cultivars in response to salinity stress.Crossref | GoogleScholarGoogle Scholar |

Monirifar H, Barghi M (2009) Identification and selection for salt tolerance in alfalfa (Medicago sativa L.) ecotypes via physiological traits. Notulae Scientia Biologicae 1, 63–66.
Identification and selection for salt tolerance in alfalfa (Medicago sativa L.) ecotypes via physiological traits.Crossref | GoogleScholarGoogle Scholar |

Moradi A (2016) Effect of mycorrhizal inoculation on growth, nitrogen fixation and nutrient uptake in alfalfa (Medicago sativa) under salt stress. Cercetari Agronomice În Moldova 49, 67–80.
Effect of mycorrhizal inoculation on growth, nitrogen fixation and nutrient uptake in alfalfa (Medicago sativa) under salt stress.Crossref | GoogleScholarGoogle Scholar |

Morgan SH, Maity PJ, Geilfus CM, Karl SL, Mühling H (2014) Leaf ion homeostasis and plasma membrane H+-ATPase activity in Vicia faba change after extra calcium and potassium supply under salinity. Plant Physiology and Biochemistry 82, 244–253.
Leaf ion homeostasis and plasma membrane H+-ATPase activity in Vicia faba change after extra calcium and potassium supply under salinity.Crossref | GoogleScholarGoogle Scholar | 25010036PubMed |

Mouradi M, Latrach L, Farissi M, Bouizgarne A, Ghoulam C (2018) Impact of the salt stress on the agronomic potential of the Moroccan populations of alfalfa (Medicago sativa L.) under the field conditions of Marrakesh. Applied Journal of Environmental Engineering Science 4, 4–13.

Munns R, Tester M (2008) Mechanisms of salinity tolerance. Annual Review of Plant Biology 59, 651–681.
Mechanisms of salinity tolerance.Crossref | GoogleScholarGoogle Scholar | 18444910PubMed |

Nabizadeh E, Jalilnejad N, Armakani M (2011) Effect of salinity on growth and nitrogen fixation of alfalfa (Medicago sativa). World Applied Sciences Journal 13, 1895–1900.

Noble CL, Halloran MC, West DW (1984) Identification and selection for salt tolerance in lucerne (Medicago sativa L.). Australian Journal of Agricultural Research 35, 239–252.
Identification and selection for salt tolerance in lucerne (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar |

Noori F, Etesami H, Zarini HN, Khoshkholgh-Sima NA, Salekdeh GH, Alishahi F (2018) Mining alfalfa (Medicago sativa L.) nodules for salinity tolerant non-rhizobial bacteria to improve growth of alfalfa under salinity stress. Ecotoxicology and Environmental Safety 162, 129–138.
Mining alfalfa (Medicago sativa L.) nodules for salinity tolerant non-rhizobial bacteria to improve growth of alfalfa under salinity stress.Crossref | GoogleScholarGoogle Scholar | 29990724PubMed |

Okcu G, Kaya MD, Atak M (2005) Effects of salt and drought stresses on germination and seedling growth of pea (Pisum sativum L.). Turkish Journal of Agriculture and Forestry 29, 237–242.

Palma F, López-Gómez M, Tejera NA, Lluch C (2013) Salicylic acid improves the salinity tolerance of Medicago sativa in symbiosis with Sinorhizobium meliloti by preventing nitrogen fixation inhibition. Plant Science 208, 75–82.
Salicylic acid improves the salinity tolerance of Medicago sativa in symbiosis with Sinorhizobium meliloti by preventing nitrogen fixation inhibition.Crossref | GoogleScholarGoogle Scholar | 23683932PubMed |

Palma F, López-Gómez M, Tejera NA, Lluch C (2014) Involvement of abscisic acid in the response of Medicago sativa plants in symbiosis with Sinorhizobium meliloti to salinity. Plant Science 223, 16–24.
Involvement of abscisic acid in the response of Medicago sativa plants in symbiosis with Sinorhizobium meliloti to salinity.Crossref | GoogleScholarGoogle Scholar | 24767111PubMed |

Peel M, Waldron B, Jensen K, Chatterton N, Horton H, Dudley L (2004) Screening for salinity tolerance in alfalfa: a repeatable method. Crop Science 44, 2049
Screening for salinity tolerance in alfalfa: a repeatable method.Crossref | GoogleScholarGoogle Scholar |

Peng YL, Gao ZW, Gao Y, Liu GF, Sheng LX, Wang DL (2008) Eco‐physiological characteristics of alfalfa seedlings in response to various mixed salt‐alkaline stresses. Journal of Integrative Plant Biology 50, 29–39.
Eco‐physiological characteristics of alfalfa seedlings in response to various mixed salt‐alkaline stresses.Crossref | GoogleScholarGoogle Scholar | 18666949PubMed |

Petcu E, Schitea M, Badea D (2007) The behavior of some Romanian alfalfa genotypes to salt and water stress. Romanian Agricultural Research 24, 51–54.

Petrusa LM, Winicov I (1997) Proline status in salt tolerant and salt sensitive alfalfa cell lines and plants in response to NaCl. Plant Physiology and Biochemistry 35, 303–310.

Pitann B, Kranz T, Zorb C, Walter A, Schurr U, Mühling KH (2011) Apoplastic pH and growth in expanding leaves of Vicia faba under salinity. Environmental and Experimental Botany 74, 31–36.
Apoplastic pH and growth in expanding leaves of Vicia faba under salinity.Crossref | GoogleScholarGoogle Scholar |

Postnikova OA, Shao J, Nemchinov LG (2013) Analysis of the alfalfa root transcriptome in response to salinity stress. Plant & Cell Physiology 54, 1041–1055.
Analysis of the alfalfa root transcriptome in response to salinity stress.Crossref | GoogleScholarGoogle Scholar |

Pottosin I, Velarde-Buendía AM, Bose J, Zepeda-Jazo I, Shabala S, Dobrovinskaya O (2014) Cross-talk between reactive oxygen species and polyamines in regulation of ion transport across the plasma membrane: implications for plant adaptive responses. Journal of Experimental Botany 65, 1271–1283.
Cross-talk between reactive oxygen species and polyamines in regulation of ion transport across the plasma membrane: implications for plant adaptive responses.Crossref | GoogleScholarGoogle Scholar | 24465010PubMed |

Provorov NA, Onishchuk OP, Kurchak ON (2016) Impacts of inoculation with Sinorhizobium meliloti strains differing in salt tolerance on the productivity and habitus of alfalfa (Medicago sativa L.). Agricultural Biology 51, 343–350.
Impacts of inoculation with Sinorhizobium meliloti strains differing in salt tolerance on the productivity and habitus of alfalfa (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar |

Putnam DH, Benes S, Galdi G, Hutmacher B, Grattan S (2017) Alfalfa (Medicago sativa L.) is tolerant to higher levels of salinity than previous guidelines indicated: implications of field and greenhouse studies. EGU General Assembly Conference Abstracts 19, 18266

Quan W, Liu X, Wang H, Chan Z (2016) Physiological and transcriptional responses of contrasting alfalfa (Medicago sativa L.) varieties to salt stress. Plant Cell, Tissue and Organ Culture 126, 105–115.
Physiological and transcriptional responses of contrasting alfalfa (Medicago sativa L.) varieties to salt stress.Crossref | GoogleScholarGoogle Scholar |

Rahman MA, Alam I, Kim YG, Ahn NY, Heo SH, Lee DG, Liu G, Lee BH (2015) Screening for salt-responsive proteins in two contrasting alfalfa cultivars using a comparative proteome approach. Plant Physiology and Biochemistry 89, 112–122.
Screening for salt-responsive proteins in two contrasting alfalfa cultivars using a comparative proteome approach.Crossref | GoogleScholarGoogle Scholar | 25743099PubMed |

Redondo FJ, de la Pena TC, Lucas MM, Pueyo JJ (2012) Alfalfa nodules elicited by a flavodoxin-overexpressing Ensifer meliloti strain display nitrogen-fixing activity with enhanced tolerance to salinity stress. Planta 236, 1687–1700.
Alfalfa nodules elicited by a flavodoxin-overexpressing Ensifer meliloti strain display nitrogen-fixing activity with enhanced tolerance to salinity stress.Crossref | GoogleScholarGoogle Scholar | 22864594PubMed |

Robinson PH, Grattan SR, Getachew G, Grieve CM, Poss JA, Suarez DL, Benes SE (2004) Biomass accumulation and potential nutritive value of some forages irrigated with saline-sodic drainage water. Animal Feed Science and Technology 111, 175–189.
Biomass accumulation and potential nutritive value of some forages irrigated with saline-sodic drainage water.Crossref | GoogleScholarGoogle Scholar |

Roger ME (1998) Salinity effects on irrigated lucerne. In ‘Proceeding 9th Australian Agronomy Conference’. Wagga Wagga, NSW, Australia. (Australian Society of Agronomy)

Rogers ME (2001) The effect of saline irrigation on lucerne production: shoot and root growth, ion relations and flowering incidence in six cultivars grown in northern Victoria, Australia. Irrigation Science 20, 55–64.
The effect of saline irrigation on lucerne production: shoot and root growth, ion relations and flowering incidence in six cultivars grown in northern Victoria, Australia.Crossref | GoogleScholarGoogle Scholar |

Rogers ME, Grieve CM, Shannon MC (2003) Plant growth and ion relations in lucerne (Medicago sativa L.) in response to the combined effects of NaCl and P. Plant and Soil 253, 187–194.
Plant growth and ion relations in lucerne (Medicago sativa L.) in response to the combined effects of NaCl and P.Crossref | GoogleScholarGoogle Scholar |

Rokebul Anower MR, Mott IW, Peel MD, Wu Y (2013) Characterization of physiological responses of two alfalfa half-sib families with improved salt tolerance. Plant Physiology and Biochemistry 71, 103–111.
Characterization of physiological responses of two alfalfa half-sib families with improved salt tolerance.Crossref | GoogleScholarGoogle Scholar |

Rokebul Anower MR, Peel MD, Mott IW, Wu Y (2017) Physiological processes associated with salinity tolerance in an alfalfa half‐sib family. Journal of Agronomy & Crop Science 203, 506–518.
Physiological processes associated with salinity tolerance in an alfalfa half‐sib family.Crossref | GoogleScholarGoogle Scholar |

Roy SJ, Negrao S, Tester M (2014) Salt resistant crop plants. Current Opinion in Biotechnology 26, 115–124.
Salt resistant crop plants.Crossref | GoogleScholarGoogle Scholar | 24679267PubMed |

Sadeghi H, Khaef N (2011) Priming-induced metabolic changes in three annual medics species improve germination and early growth under drought and salt stress conditions. Genetics and Plant Physiology 1, 186–198.

Sah SK, Reddy KR, Li J (2016) Abscisic acid and abiotic stress tolerance in crop plants. Frontiers in Plant Science 7, 571
Abscisic acid and abiotic stress tolerance in crop plants.Crossref | GoogleScholarGoogle Scholar | 27200044PubMed |

Sandhu D, Cornacchione MV, Ferreira JF, Suarez DL (2017) Variable salinity responses of 12 alfalfa genotypes and comparative expression analyses of salt-response genes. Scientific Reports 7, 42958
Variable salinity responses of 12 alfalfa genotypes and comparative expression analyses of salt-response genes.Crossref | GoogleScholarGoogle Scholar | 28225027PubMed |

Sandhu D, Pudussery MV, Kaundal R, Suarez DL, Kaundal A, Sekhon RS (2018) Molecular characterization and expression analysis of the Na+/H+ exchanger gene family in Medicago truncatula. Functional & Integrative Genomics 18, 141–153.
Molecular characterization and expression analysis of the Na+/H+ exchanger gene family in Medicago truncatula.Crossref | GoogleScholarGoogle Scholar |

Sepehri A, Najari S, Rouhi HR (2015) Seed priming to overcome salinity stress in Persian cultivars of alfalfa (Medicago sativa L.). Notulae Scientia Biologicae 7, 96–101.
Seed priming to overcome salinity stress in Persian cultivars of alfalfa (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar |

Sharifi M, Ghorbanli M, Ebrahimzadeh H (2007) Improved growth of salinity stressed soybean after inoculation with salt pretreated mycorrhizal fungi. Journal of Plant Physiology 164, 1144–1151.
Improved growth of salinity stressed soybean after inoculation with salt pretreated mycorrhizal fungi.Crossref | GoogleScholarGoogle Scholar | 16919369PubMed |

Sharma S, Upadhyaya H, Varshney R, Gowda C (2013) Pre-breeding for diversification of primary gene pool and genetic enhancement of grain legumes. Frontiers in Plant Science 4, 309
Pre-breeding for diversification of primary gene pool and genetic enhancement of grain legumes.Crossref | GoogleScholarGoogle Scholar | 23970889PubMed |

Sharp RG, Davies WJ (2009) Variability among species in the apoplastic pH signalling response to drying soils. Journal of Experimental Botany 60, 4363–4370.
Variability among species in the apoplastic pH signalling response to drying soils.Crossref | GoogleScholarGoogle Scholar | 19726633PubMed |

Singh NK, LaRosa PC, Handa AK, Hasegawa PM, Bressan RA (1987) Hormonal regulation of protein synthesis associated with salt tolerance in plant cells. Proceedings of the National Academy of Sciences of the United States of America 84, 739–743.
Hormonal regulation of protein synthesis associated with salt tolerance in plant cells.Crossref | GoogleScholarGoogle Scholar | 16593808PubMed |

Smýkal P, Coyne CJ, Ambrose MJ, Maxted N, Schaefer H, Blair MW, Berger J, Greene SL, Nelson MN, Besharat N, Vymyslický T, Toker C, Saxena RK, Roorkiwal M, Pandey MK, Hu J, Li YH, Wang LX, Guo Y, Qiu LJ, Redden RJ, Varshney RK (2015) Legume crops phylogeny and genetic diversity for science and breeding. Critical Reviews in Plant Sciences 34, 43–104.
Legume crops phylogeny and genetic diversity for science and breeding.Crossref | GoogleScholarGoogle Scholar |

Soltani A, Khodarahmpour Z, Jafari AA, Nakhjavan S (2012) Selection of alfalfa (Medicago sativa L.) cultivars for salt stress tolerance using germination indices. African Journal of Biotechnology 11, 7899–7905.

Soussi M, Ocana A, Lluch C (1998) Effects of salt stress on growth, photosynthesis and nitrogen fixation in chick-pea (Cicer arietinum L.). Journal of Experimental Botany 49, 1329–1337.
Effects of salt stress on growth, photosynthesis and nitrogen fixation in chick-pea (Cicer arietinum L.).Crossref | GoogleScholarGoogle Scholar |

Srivastava D, Mukerji KG (1995) Field response of mycorrhizal and nonmycorrhizal Medicago sativa var. local in the F1 generation. Mycorrhiza 5, 219–221.
Field response of mycorrhizal and nonmycorrhizal Medicago sativa var. local in the F1 generation.Crossref | GoogleScholarGoogle Scholar |

Steppuhn H, Acharya SN, Iwaasa AD, Gruber M, Miller DR (2012) Inherent responses to root-zone salinity in nine alfalfa populations. Canadian Journal of Plant Science 92, 235–248.
Inherent responses to root-zone salinity in nine alfalfa populations.Crossref | GoogleScholarGoogle Scholar |

Strogonov BP (1974) ‘Structure and function of plant cells in saline habitats.’ (Halstead Press: New York)

Suárez R, Calderón C, Iturriaga G (2009) Enhanced tolerance to multiple abiotic stresses in transgenic alfalfa accumulating trehalose. Crop Science 49, 1791–1799.
Enhanced tolerance to multiple abiotic stresses in transgenic alfalfa accumulating trehalose.Crossref | GoogleScholarGoogle Scholar |

Suyama H, Benes SE, Robinson PH, Grattan SR, Grieve CM, Getachew G (2007) Forage yield and quality under irrigation with saline-sodic drainage water: greenhouse evaluation. Agricultural Water Management 88, 159–172.
Forage yield and quality under irrigation with saline-sodic drainage water: greenhouse evaluation.Crossref | GoogleScholarGoogle Scholar |

Swaraj K, Bishnoi NR (1999) Effect of salt stress on nodulation and nitrogen fixation in legumes. Indian Journal of Experimental Biology 37, 843–848.

Tang L, Cai H, Ji W, Luo X, Wang Z, Wu J, Wang X, Cui L, Wang Y, Zhu Y, Bai X (2013) Overexpression of GsZFP1 enhances salt and drought tolerance in transgenic alfalfa (Medicago sativa L.). Plant Physiology and Biochemistry 71, 22–30.
Overexpression of GsZFP1 enhances salt and drought tolerance in transgenic alfalfa (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar | 23867600PubMed |

Tang L, Cai H, Zhai H, Luo X, Wang ZY, Cui L, Bai X (2014) Overexpression of Glycine soja WRKY20 enhances both drought and salt tolerance in transgenic alfalfa (Medicago sativa L.). Plant Cell, Tissue and Organ Culture 118, 77–86.
Overexpression of Glycine soja WRKY20 enhances both drought and salt tolerance in transgenic alfalfa (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar |

Tavakoli M, Poustini K, Besharati H, Ali S (2019) Variable salinity responses of 25 alfalfa genotypes and comparative salt-response ion distribution. Russian Journal of Plant Physiology 66, 231–239.
Variable salinity responses of 25 alfalfa genotypes and comparative salt-response ion distribution.Crossref | GoogleScholarGoogle Scholar |

Teakle NL, Real D, Colmer TD (2006) Growth and ion relations in response to combined salinity and waterlogging in the perennial forage legumes Lotus corniculatus and Lotus tenuis. Plant and Soil 289, 369
Growth and ion relations in response to combined salinity and waterlogging in the perennial forage legumes Lotus corniculatus and Lotus tenuis.Crossref | GoogleScholarGoogle Scholar |

Tian Y, Wu Y, Zheng W, Zhang WX, Zhou DW (2009) Effect of NaCl on germination and early seedling growth of seven accessions of Medicago falcata L. Chinese Journal of Grassland 6, 25–29.

Tilaki GAD, Behtari B, Behtari B (2009) Effect of salt and water stress on the germination of alfalfa (Medicago sativa L.) seed. Volga Ecological Magazine (ПOBOлжcкий экOлOrичecкий жypHaл) 2, 158–164.

Torabi M, Halim MRA (2010) Variation of root and shoot growth and free proline accumulation in Iranian alfalfa ecotypes under salt stress. Journal of Food, Agriculture and Environment 8, 323–327.

Torabi M, Halim RA, Sinniah UR, Choukan R (2011) Influence of salinity on the germination of Iranian alfalfa ecotypes. African Journal of Agricultural Research 6, 4624–4630.

Trinchant JC, Boscari A, Spennato G, Van de Sype G, Le Rudulier D (2004) Proline betaine accumulation and metabolism in alfalfa plants under sodium chloride stress. Exploring its compartmentalization in nodules. Plant Physiology 135, 1583–1594.
Proline betaine accumulation and metabolism in alfalfa plants under sodium chloride stress. Exploring its compartmentalization in nodules.Crossref | GoogleScholarGoogle Scholar | 15235114PubMed |

Vaughan LV, MacAdam JW, Smith SE, Dudley LM (2002) Root growth and yield of differing alfalfa rooting populations under increasing salinity and zero leaching. Crop Science 42, 2064–2071.
Root growth and yield of differing alfalfa rooting populations under increasing salinity and zero leaching.Crossref | GoogleScholarGoogle Scholar |

Wang XS, Han JG (2009) Changes of proline content, activity, and active isoforms of antioxidative enzymes in two alfalfa cultivars under salt stress. Agricultural Sciences in China 8, 431–440.
Changes of proline content, activity, and active isoforms of antioxidative enzymes in two alfalfa cultivars under salt stress.Crossref | GoogleScholarGoogle Scholar |

Wang YX, Zhang B, Wang T (2009a) Effect of salt stress on the contents of chlorophyll and betaine and its membrane permeability of Medicago sativa [J]. Pratacultural Science 3, S541.9 http://en.cnki.com.cn/Article_en/CJFDTotal-CYKX200903012.htm

Wang WB, Kim YH, Lee HS, Kim KY, Deng XP, Kwak SS (2009b) Analysis of antioxidant enzyme activity during germination of alfalfa under salt and drought stresses. Plant Physiology and Biochemistry 47, 570–577.
Analysis of antioxidant enzyme activity during germination of alfalfa under salt and drought stresses.Crossref | GoogleScholarGoogle Scholar | 19318268PubMed |

Wang X, Yang X, Chen L, Feng G, Zhang J, Jin L (2011) Genetic diversity among alfalfa (Medicago sativa L.) cultivars in Northwest China. Acta Agriculturæ Scandinavica. Section B, Soil and Plant Science 61, 60–66.
Genetic diversity among alfalfa (Medicago sativa L.) cultivars in Northwest China.Crossref | GoogleScholarGoogle Scholar |

Wang Z, Li H, Ke Q, Jeong JC, Lee HS, Xu B, Deng XP, Lim YP, Kwak SS (2014) Transgenic alfalfa plants expressing AtNDPK2 exhibit increased growth and tolerance to abiotic stresses. Plant Physiology and Biochemistry 84, 67–77.
Transgenic alfalfa plants expressing AtNDPK2 exhibit increased growth and tolerance to abiotic stresses.Crossref | GoogleScholarGoogle Scholar | 25240265PubMed |

Wang Y, Zhang Z, Zhang P, Cao Y, Hu T, Yang P (2016) Rhizobium symbiosis contribution to short-term salt stress tolerance in alfalfa (Medicago sativa L.). Plant and Soil 402, 247–261.
Rhizobium symbiosis contribution to short-term salt stress tolerance in alfalfa (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar |

Wang XS, Ren HL, Wei ZW, Wang YW, Ren WB (2017) Effects of neutral salt and alkali on ion distributions in the roots, shoots, and leaves of two alfalfa cultivars with differing degrees of salt tolerance. Journal of Integrative Agriculture 16, 1800–1807.
Effects of neutral salt and alkali on ion distributions in the roots, shoots, and leaves of two alfalfa cultivars with differing degrees of salt tolerance.Crossref | GoogleScholarGoogle Scholar |

Winicov I, Bastola DR (1999) Transgenic overexpression of the transcription factor Alfin1 enhances expression of the endogenous MsPRP2 gene in alfalfa and improves salinity tolerance of the plants. Plant Physiology 120, 473–480.
Transgenic overexpression of the transcription factor Alfin1 enhances expression of the endogenous MsPRP2 gene in alfalfa and improves salinity tolerance of the plants.Crossref | GoogleScholarGoogle Scholar | 10364398PubMed |

Winicov I, Seemann JR (1990) Expression of genes for photosynthesis and the relationship to salt tolerance of alfalfa (Medicago sativa) cells. Plant & Cell Physiology 31, 1155–1161.

Wolf O, Jeschke WD, Hartung W (1990) Long distance transport of abscisic acid in NaCl-treated intact plants of Lupinus albus. Journal of Experimental Botany 41, 593–600.
Long distance transport of abscisic acid in NaCl-treated intact plants of Lupinus albus.Crossref | GoogleScholarGoogle Scholar |

Yacoubi R, Job C, Belghazi M, Chaibi W, Job D (2011) Toward characterizing seed vigor in alfalfa through proteomic analysis of germination and priming. Journal of Proteome Research 10, 3891–3903.
Toward characterizing seed vigor in alfalfa through proteomic analysis of germination and priming.Crossref | GoogleScholarGoogle Scholar | 21755932PubMed |

Yacoubi R, Job C, Belghazi M, Chaibi W, Job D (2013) Proteomic analysis of the enhancement of seed vigour in osmoprimed alfalfa seeds germinated under salinity stress. Seed Science Research 23, 99–110.
Proteomic analysis of the enhancement of seed vigour in osmoprimed alfalfa seeds germinated under salinity stress.Crossref | GoogleScholarGoogle Scholar |

Yarnia M, Heidari Sharif Abad H, Hashemi Dezfuli SA, Rahimzade Khoei F, Ghalavand A (2001) Evaluation of alfalfa (Medicago sativa L.) lines to salinity tolerance. Majallah’-i ‘Ulum-i Zira’i-i Iran 3, 12–26.

Younesi O, Moradi A (2014) Effect of priming of seeds of Medicago sativa ‘bami’ with gibberellic acid on germination, seedlings growth and antioxidant enzymes activity under salinity stress. Journal of Horticultural Research 22, 167–174.
Effect of priming of seeds of Medicago sativa ‘bami’ with gibberellic acid on germination, seedlings growth and antioxidant enzymes activity under salinity stress.Crossref | GoogleScholarGoogle Scholar |

Younesi O, Baghbani A, Namdari A (2013) The effects of Pseudomonas fluorescence and Rhizobium meliloti co-inoculation on nodulation and mineral nutrient contents in alfalfa (Medicago sativa) under salinity stress. International Journal of Agriculture and Crop Sciences 5, 1500

Yu LX, Liu X, Boge W, Liu XP (2016) Genome-wide association study identifies loci for salt tolerance during germination in autotetraploid alfalfa (Medicago sativa L.) using genotyping-by-sequencing. Frontiers in Plant Sciences 7, 956
Genome-wide association study identifies loci for salt tolerance during germination in autotetraploid alfalfa (Medicago sativa L.) using genotyping-by-sequencing.Crossref | GoogleScholarGoogle Scholar |

Yurtseven S (2011) The nutrient and energy contents of Medicago varieties growth in salt-affected soils of the Harran plain. Hayvansal Üretim 52, 39–45.

Zahaf O, Blanchet S, De Zélicourt A, Alunni B, Plet J, Laffont C, De Lorenzo L, Imbeaud S, Ichanté JL, Diet A, Badri M (2012) Comparative transcriptomic analysis of salt adaptation in roots of contrasting Medicago truncatula genotypes. Molecular Plant 5, 1068–1081.
Comparative transcriptomic analysis of salt adaptation in roots of contrasting Medicago truncatula genotypes.Crossref | GoogleScholarGoogle Scholar | 22419822PubMed |

Zhang WJ, Wang T (2015) Enhanced salt tolerance of alfalfa (Medicago sativa) by rstB gene transformation. Plant Science 234, 110–118.
Enhanced salt tolerance of alfalfa (Medicago sativa) by rstB gene transformation.Crossref | GoogleScholarGoogle Scholar | 25804814PubMed |

Zhang S, Hu J, Zhang Y, Xie XJ, Knapp A (2007) Seed priming with brassinolide improves lucerne (Medicago sativa L.) seed germination and seedling growth in relation to physiological changes under salinity stress. Australian Journal of Agricultural Research 58, 811–815.
Seed priming with brassinolide improves lucerne (Medicago sativa L.) seed germination and seedling growth in relation to physiological changes under salinity stress.Crossref | GoogleScholarGoogle Scholar |

Zhang L, Zhang Q, Ye BX (2010) Effect of arbuscular mycorrhizal fungi (AMF) on growth of Medicago sativa L. under salt stress. Shandong Agricultural Sciences 3, 10

Zhang Q, Xu L, Tang J, Bai M, Chen X (2011) Arbuscular mycorrhizal mediation of biomass–density relationship of Medicago sativa L. under two water conditions in a field experiment. Mycorrhiza 21, 269–277.
Arbuscular mycorrhizal mediation of biomass–density relationship of Medicago sativa L. under two water conditions in a field experiment.Crossref | GoogleScholarGoogle Scholar | 20652365PubMed |

Zhang L, Niu Y, Huridu H, Hao J, Qi Z, Hasi A (2014) Salicornia europaea L. Na+/H+ antiporter gene improves salt tolerance in transgenic alfalfa (Medicago sativa L.). Genetics and Molecular Research 13, 5350–5360.
Salicornia europaea L. Na+/H+ antiporter gene improves salt tolerance in transgenic alfalfa (Medicago sativa L.).Crossref | GoogleScholarGoogle Scholar | 25078591PubMed |

Zhanwu G, Hui Z, Jicai G, Chunwu Y, Chunsheng M, Deli W (2011) Germination responses of alfalfa (Medicago sativa L.) seeds to various salt alkaline mixed stress. African Journal of Agricultural Research 6, 3793–3803.

Zhu JK (2002) Salt and drought stress signal transduction in plants. Annual Review of Plant Biology 53, 247–273.
Salt and drought stress signal transduction in plants.Crossref | GoogleScholarGoogle Scholar | 12221975PubMed |

Zhu JK (2016) Abiotic stress signaling and responses in plants. Cell 167, 313–324.
Abiotic stress signaling and responses in plants.Crossref | GoogleScholarGoogle Scholar | 27716505PubMed |