摘要
Iron(Fe) bioavailability to plants is reduced in saline soils; however, the exact mechanisms underlying this effect are not yet completely understood. Siderophore-expressing rhizobacteria may represent a promising alternative to chemical fertilizers by simultaneously tackling salt-stress effects and Fe limitation in saline soils. In addition to draught, plants growing in arid soils face two other major challenges: high salinity and Fe deficiency. Salinity attenuates growth, affects plant physiology, and causes nutrient imbalance,which is, in fact, one of the major consequences of saline stress. Iron is a micronutrient essential for plant development, and it is required by several metalloenzymes involved in photosynthesis and respiration. Iron deficiency is associated with chlorosis and low crop productivity. The role of microbial siderophores in Fe supply to plants and the effect of plant growth-promoting rhizobacteria(PGPR) on the mitigation of saline stress in crop culture are well documented. However, the dual effect of siderophore-producing PGPR, both on salt stress and Fe limitation, is still poorly explored. This review provides a critical overview of the combined effects of Fe limitation and soil salinization as challenges to modern agriculture and intends to summarize some indirect evidence that argues in favour of siderophore-producing PGPR as biofertilization agents in salinized soils. Recent developments and future perspectives on the use of PGPR are discussed as clues to sustainable agricultural practices in the context of present and future climate change scenarios.
Iron(Fe) bioavailability to plants is reduced in saline soils; however, the exact mechanisms underlying this effect are not yet completely understood. Siderophore-expressing rhizobacteria may represent a promising alternative to chemical fertilizers by simultaneously tackling salt-stress effects and Fe limitation in saline soils. In addition to draught, plants growing in arid soils face two other major challenges: high salinity and Fe deficiency. Salinity attenuates growth, affects plant physiology, and causes nutrient imbalance,which is, in fact, one of the major consequences of saline stress. Iron is a micronutrient essential for plant development, and it is required by several metalloenzymes involved in photosynthesis and respiration. Iron deficiency is associated with chlorosis and low crop productivity. The role of microbial siderophores in Fe supply to plants and the effect of plant growth-promoting rhizobacteria(PGPR) on the mitigation of saline stress in crop culture are well documented. However, the dual effect of siderophore-producing PGPR, both on salt stress and Fe limitation, is still poorly explored. This review provides a critical overview of the combined effects of Fe limitation and soil salinization as challenges to modern agriculture and intends to summarize some indirect evidence that argues in favour of siderophore-producing PGPR as biofertilization agents in salinized soils. Recent developments and future perspectives on the use of PGPR are discussed as clues to sustainable agricultural practices in the context of present and future climate change scenarios.
引文
Abadía J,López-Millán A F,Rombolà A,Abadía A.2002.Organic acids and Fe deficiency:A review.Plant Soil.241:75-86.
Abbas G,Saqib M,Akhtar J.2015.Interactive effects of salinity and iron deficiency on different rice genotypes.J Plant Nutr Soil Sci.178:306-311.
Abbasi H,Jamil M,Haq A,Ali S,Ahmad R,Malik Z,Parveen.2016.Salt stress manifestation on plants,mechanism of salt tolerance and potassium role in alleviating it:A review.Zemdirbyste Agric.103:229-238.
Acosta-Motos J R,Ortu?o M F,Bernal-Vicente A,Diaz-Vivancos P,Sanchez-Blanco M J,Hernandez J A.2017.Plant responses to salt stress:Adaptive mechanisms.Agronomy.7:18.
Aguado-Santacruz G A,Moreno-Gómez B,Jiménez-Francisco B,García-Moya E,Preciado-Ortiz R E.2012.Impact of the microbial siderophores and phytosiderophores on the iron assimilation by plants:A synthesis.Rev Fitotec Mex.35:9-21.
Ali S,Charles T C,Glick B R.2014.Amelioration of high salinity stress damage by plant growth-promoting bacterial endophytes that contain ACC deaminase.Plant Physiol Biochem.80:160-167.
Ali S S,Vidhale N N.2013.Bacterial siderophore and their application:A review.Int J Curr Microbiol Appl Sci.2:303-312.
Ameixa O M C C,Marques B,Fernandes V S,Soares A M VM,Calado R,Lilleb?A I.2016.Dimorphic seeds of Salicornia ramosissima display contrasting germination responses under different salinities.Ecol Eng.87:120-123.
Aragüés R,Medina E T,Martínez-Cob A,Faci J.2014.Effects of deficit irrigation strategies on soil salinization and sodification in a semiarid drip-irrigated peach orchard.Agric Water Manage.142:1-9.
Ashraf M,Hasnain S,Berge O,Mahmood T.2004.Inoculating wheat seedlings with exopolysaccharide-producing bacteria restricts sodium uptake and stimulates plant growth under salt stress.Biol Fert Soils.40:157-162.
Baetz U,Martinoia E.2014.Root exudates:The hidden part of plant defense.Trends Plant Sci.19:90-98.
Bais H P,Prithiviraj B,Jha A K,Ausubel F M,Vivanco J M.2005.Mediation of pathogen resistance by exudation of antimicrobials from roots.Nature.434:217-221.
Becker V I,Goessling J W,Duarte B,Ca?cador I,Liu F L,Rosenqvist E,Jacobsen S E.2017.Combined effects of soil salinity and high temperature on photosynthesis and growth of quinoa plants(Chenopodium quinoa).Funct Plant Biol.44:665-678.
Bertrand S.2011.Siderophore Base-the web data base of microbial siderophores.Available online at http://bertrandsamuel.free.fr/siderophore base/siderophores.php(verified on August 24,2018).
Bogdan A R,Miyazawa M,Hashimoto K,Tsuji Y.2016.Regulators of iron homeostasis:New players in metabolism,cell death,and disease.Trends Biochem Sci.41:274-286.
Briat J F,Dubos C,Gaymard F.2015.Iron nutrition,biomass production,and plant product quality.Trends Plant Sci.20:33-40.
Cassat J E,Skaar E P.2013.Iron in infection and immunity.Cell Host Microbe.13:509-519.
Chakraborty K,Sairam R K,Bhaduri D.2016.Effects of different levels of soil salinity on yield attributes,accumulation of nitrogen,and micronutrients in Brassica spp.J Plant Nutr.39:1026-1037.
Chen J H,Lin Y H.2010.Sodium chloride causes variation in organic acids and proteins in tomato root.Afr J Biotechnol.9:8161-8167.
Chen W W,Yang J L,Qin C,Jin C W,Mo J H,Ye T,Zheng SJ.2010.Nitric oxide acts downstream of auxin to trigger root ferric-chelate reductase activity in response to iron deficiency in Arabidopsis.Plant Physiol.154:810-819.
Chen Y,Barak P.1982.Iron nutrition of plants in calcareous soils.Adv Agron.35:217-240.
Chen Y,Hadar Y.1991.Iron Nutrition and Interactions in Plants.Springer,Dordrecht.
Cheng Z Y,Park E,Glick B R.2007.1-aminocyclopropane-1-carboxylate deaminase from Pseudomonas putida UW4 facilitates the growth of canola in the presence of salt.Can JMicrobiol.53:912-918.
Ciancio A,Pieterse C M J,Mercado-Blanco J.2016.Editorial:Harnessing useful rhizosphere microorganisms for pathogen and pest biocontrol.Front Microbiol.7:1620.
Cohen G N.2014.Microbial Biochemistry.3rd Edn.Springer,Dordrecht.
Coleman D C,Callaham M A,Crossley Jr D A.2017.Fundamentals of Soil Ecology.3rd Edn.Academic Press,San Diego.
Colombo C,Palumbo G,He J Z,Pinton R,Cesco S.2014.Review on iron availability in soil:Interaction of Fe minerals,plants,and microbes.J Soil Sedimet.14:538-548.
Comensoli L,Bindschedler S,Junier P,Joseph E.2017.Chapter two-iron and fungal physiology:A review of biotechnological opportunities.Adv Appl Microbiol.98:31-60.
Cotsaftis O,Plett D,Johnson A A T,Walia H,Wilson C,Ismail A M,Close T J,Tester M,Baumann U.2011.Root-specific transcript profiling of contrasting rice genotypes in response to salinity stress.Mol Plant.4:25-41.
Curie C,Mari S.2017.New routes for plant iron mining.New Phytol.214:521-525.
Daliakopoulos I N,Tsanis I K,Koutroulis A,Kourgialas N N,Varouchakis A E,Karatzas G P,Ritsema C J.2016.The threat of soil salinity:A European scale review.Sci Total Environ.573:727-739.
de Santiago A,García-López A M,Quintero J M,Avilés M,Delgado A.2013.Effect of Trichoderma asperellum strain T34and glucose addition on iron nutrition in cucumber grown on calcareous soils.Soil Biol Biochem.57:598-605.
de Souza R,Ambrosini A,Passaglia L M P.2015.Plant growthpromoting bacteria as inoculants in agricultural soils.Genet Mol Biol.38:401-419.
El Fouly M M,Mobarak Z M,Salama Z A.2010.Improving tolerance of faba bean during early growth stages to salinity through micronutrients foliar spray.Not Sci Biol.2:98-102.
Enthaler S,Junge K,Beller M.2008.Sustainable metal catalysis with iron:From rust to a rising star?Angew Chem Int Edit.47:3317-3321.
Estévez J,Dardanelli M S,Megías M,Rodríguez-Navarro D.2009.Symbiotic performance of common bean and soybean co-inoculated with rhizobia and Chryseobacterium balustinum Aur9 under moderate saline conditions.Symbiosis.49:29-36.
Etesami H,Maheshwari D K.2018.Use of plant growth promoting rhizobacteria(PGPRs)with multiple plant growth promoting traits in stress agriculture:Action mechanisms and future prospects.Ecotoxicol Environ Saf.156:225-246.
Falkowski P G,Lin H Z,Gorbunov M Y.2017.What limits photosynthetic energy conversion efficiency in nature?Lessons from the oceans.Philos Trans Roy Soc B Biol Sci.372:20160376.
Faraldo-Gómez J D,Sansom M S P.2003.Acquisition of siderophores in Gram-negative bacteria.Nat Rev Mol Cell Biol.4:105-116.
Ferchichi S,El Khouni A,Zorrig W,Atia A,Rabhi M,Gharsalli M,Abdelly C.2016.Nutrient uptake and use efficiencies in Medicago ciliaris under salinity.J Plant Nutr.39:932-941.
Freitas M A,Medeiros F H V,Carvalho S P,Guilherme L RG,Teixeira W D,Zhang H M,Paré P W.2015.Augmenting iron accumulation in cassava by the beneficial soil bacterium Bacillus subtilis(GBO3).Front in Plant Sci.6:596.
Fukushima T,Allred B E,Raymond K N.2014.Direct evidence of iron uptake by the Gram-positive siderophoreshuttle mechanism without iron reduction.ACS Chem Biol.9:2092-2100.
García M J,Suárez V,Romera F J,Alcántara E,Pérez-Vicente R.2011.A new model involving ethylene,nitric oxide and Fe to explain the regulation of Fe-acquisition genes in Strategy I plants.Plant Physiol Biochem.49:537-544.
Gargallo-Garriga A,Preece C,Sardans J,Oravec M,Urban O,Pe?uelas J.2018.Root exudate metabolomes change under drought and show limited capacity for recovery.Sci Rep.8:12696.
Grattan S R,Grieve C M.1998.Salinity-mineral nutrient relations in horticultural crops.Sci Hortic.78:127-157.
Grobelak A,Hiller J.2017.Bacterial siderophores promote plant growth:Screening of catechol and hydroxamate siderophores.Int J Phytorem.19:825-833.
Gunes A,Karagoz K,Turan M,Kotan R,Yildirim E,Cakmakci R,Sahin F.2015.Fertilizer efficiency of some plant growth promoting rhizobacteria for plant growth.Res J Soil Biol.7:28-45.
Gutiérrez Ma?ero F J,Probanza A,Ramos B,Colón Flores JJ,Lucas García J A.2003.Effects of culture filtrates of rhizobacteria isolated from wild lupine on germination,growth,and biological nitrogen fixation of lupine seedlings.J Plant Nutr.26:1101-1115.
Hahm M S,Son J S,Hwang Y J,Kwon D K,Ghim S Y.2017.Alleviation of salt stress in pepper(Capsicum annum L.)plants by plant growth-promoting rhizobacteria.J Microbiol Biotechnol.27:1790-1797.
Hallenbeck P C(ed.).2017.Modern Topics in the Phototrophic Prokaryotes:Metabolism,Bioenergetics,and Omics.Springer,Cham.
Hassani M A,Durán P,Hacquard S.2018.Microbial interactions within the plant holobiont.Microbiome.6:58.
Heidari M,Sarani S.2012.Growth,biochemical components and ion content of chamomile(Matricaria chamomilla L.)under salinity stress and iron deficiency.J Saudi Soc Agric Sci.11:37-42.
Hindt M N,Guerinot M L.2012.Getting a sense for signals:Regulation of the plant iron deficiency response.Biochim Biophys Acta.1823:1521-1530.
Hu X Y,Page M T,Sumida A,Tanaka A,Terry M J,Tanaka R.2017.The iron-sulfur cluster biosynthesis protein sufb is required for chlorophyll synthesis,but not phytochrome signaling.Plant J.89:1184-1194.
Jin C W,Li G X,Yu X H,Zheng S J.2010.Plant Fe status affects the composition of siderophore-secreting microbes in the rhizosphere.Ann Bot.105:835-841.
Jin C W,Ye Y Q,Zheng S J.2014.An underground tale:Contribution of microbial activity to plant iron acquisition via ecological processes.Ann Bot.113:7-18.
Johnstone T C,Nolan E M.2015.Beyond iron:Non-classical biological functions of bacterial siderophores.Dalton Trans.44:6320-6339.
Kavamura V N,Santos S N,da Silva J L,Parma M M,ávila LA,Visconti A,Zucchi T D,Taketani R G,Andreote F D,de Melo I S.2013.Screening of Brazilian cacti rhizobacteria for plant growth promotion under drought.Microbiol Res.168:183-191.
Khalid M,Bilal M,Hassani D,Iqbal H M N,Wang H,Huang D F.2017.Mitigation of salt stress in white clover(Trifolium repens)by Azospirillum brasilense and its inoculation effect.Bot Stud.58:5.
Kim H J,Fonseca J M,Choi J H,Kubota C,Kwon D Y.2008.Salt in irrigation water affects the nutritional and visual properties of romaine lettuce(Lactuca sativa L.).J Agric Food Chem.56:3772-3776.
Kim S A,Guerinot M L.2007.Mining iron:Iron uptake and transport in plants.FEBS Lett.581:2273-2280.
Klebba P E.2016.ROSET model of tonB action in Gramnegative bacterial iron acquisition.J Bacteriol.198:1013-1021.
Kloepper J W,Leong J,Teintze M,Schroth M N.1980.Enhanced plant growth by siderophores produced by plant growth-promoting rhizobacteria.Nature.286:885-886.
Kobayashi T,Nishizawa N K.2012.Iron uptake,translocation,and regulation in higher plants.Annu Rev Plant Biol.63:131-152.
Konu?skan?,G?zübenli H,Ati?s˙I,Atak M.2017.Effects of salinity stress on emergence and seedling growth parameters of some maize genotypes(Zea mays L.).Turk J Agric-Food Sci Technol.5:1668-1672.
Lewicka-Rataj K,′Swi?atecki A,Górniak D.2018.The effect of Lobelia dortmanna L.on the structure and bacterial activity of the rhizosphere.Aquat Bot.145:10-20.
Li H Q,Jiang X W.2017.Inoculation with plant growthpromoting bacteria(PGPB)improves salt tolerance of maize seedling.Russ J Plant Physiol.64:235-241.
Li Q,Yang A,Zhang W H.2016.Efficient acquisition of iron confers greater tolerance to saline-alkaline stress in rice(Oryza sativa L.).J Exp Bot.67:6431-6444.
Liu K,McInroy J A,Hu C H,Kloepper J W.2018.Mixtures of plant-growth-promoting rhizobacteria enhance biological control of multiple plant diseases and plant-growth promotion in the presence of pathogens.Plant Dis.102:67-72.
Liu W,Li R J,Han T T,Cai W,Fu Z W,Lu Y T.2015.Salt stress reduces root meristem size by nitric oxide-mediated modulation of auxin accumulation and signaling in Arabidopsis.Plant Physiol.168:343-356.
Loper J E,Buyer J S.1991.Siderophores in microbial interactions on plant surfaces.Mol Plant Microbe Interact.4:5-13.
López-Berenguer C,Martínez-Ballesta M C,Moreno D A,Carvajal M,García-Viguera C.2009.Growing hardier crops for better health:Salinity tolerance and the nutritional value of broccoli.J Agric Food Chem.57:572-578.
Ma J,Lv C F,Hao P F,Yuan Z,Wang Y W,Shen W J,Xu C,Lv C G,Chen G X,Gao Z P.2017.Sub-cellular distribution of nutrient elements and photosynthesis performance in Oryza sativa L.seedlings under salt stress.Pak J Bot.49:405-411.
Maheshwari D K,Saraf M.2015.Halophiles:Biodiversity and Sustainable Exploitation.Springer,Cham.
Mahmood S,Daur I,Al-Solaimani S G,Ahmad S,Madkour MH,Yasir M,Hirt H,Ali S,Ali Z.2016.Plant growth promoting rhizobacteria and silicon synergistically enhance salinity tolerance of mung bean.Front Plant Sci.7:Article 876.
Malhi S S,Nyborg M,Harapiak J T.1998.Effects of long-term Nfertilizer-induced acidification and liming on micronutrients in soil and in bromegrass hay.Soil Till Res.48:91-101.
Martínez R,Espejo A,Sierra M,Ortiz-Bernad I,Correa D,Bedmar E,López-Jurado M,Porres J M.2015.Co-inoculation of Halomonas maura and Ensifer meliloti to improve alfalfa yield in saline soils.Appl Soil Ecol.87:81-86.
Masalha J,Kosegarten H,Elmaci?,Mengel K.2000.The central role of microbial activity for iron acquisition in maize and sunflower.Biol Fert Soils.30:433-439.
Mavi M S,Marschner P,Chittleborough D J,Cox J W,Sanderman J.2012a.Salinity and sodicity affect soil respiration and dissolved organic matter dynamics differentially in soils varying in texture.Soil Biol Biochem.45:8-13.
Mavi M S,Sanderman J,Chittleborough D J,Cox J W,Marschner P.2012b.Sorption of dissolved organic matter in salt-affected soils:Effect of salinity,sodicity and texture.Sci Total Environ.435-436:337-344.
Mc Near Jr D H.2013.The rhizosphere-roots,soil and everything in between.Nat Educ Knowl.4:1.
Meng R,Saade S,Kurtek S,Berger B,Brien C,Pillen K,Tester M,Sun Y.2017.Growth curve registration for evaluating salinity tolerance in barley.Plant Methods.13:18.
Mimmo T,Del Buono D,Terzano R,Tomasi N,Vigani G,Crecchio C,Pinton R,Zocchi G,Cesco S.2014.Rhizospheric organic compounds in the soil-microorganism-plant system:Their role in iron availability.Eur J Soil Sci.65:629-642.
Miransari M.2014.Plant growth promoting rhizobacteria.JPlant Nutr.37:2227-2235.
Mozafari A A,Ghadakchi Asl A,Ghaderi N.2018.Grape response to salinity stress and role of iron nanoparticle and potassium silicate to mitigate salt induced damage under in vitro conditions.Physiol Mol Biol Plants.24:25-35.
Munns R,Schachtman D P,Condon A G.1995.The significance of a two-phase growth response to salinity in wheat and barley.Aust J Plant Physiol.22:561-569.
Naz I,Bano A,Ul-Hassan T.2009.Isolation of phytohormones producing plant growth promoting rhizobacteria from weeds growing in Khewra salt range,Pakistan and their implication in providing salt tolerance to Glycine max L.Afr J Biotechnol.8:5762-5766.
Negr?o S,Schm?ckel S M,Tester M.2017.Evaluating physiological responses of plants to salinity stress.Ann Bot.119:1-11.
Neilands J B.1974.Microbial Iron Metabolism:A Comprehensive Treatise.Academic Press,New York.
Nenova V.2008.Growth and mineral concentrations of pea plants under different salinity levels and iron supply.Gen Appl Plant Physiol.34:189-202.
Nikiforova E M,Kasimov N S,Kosheleva N E.2014.Long-term dynamics of the anthropogenic salinization of soils in Moscow(by the example of the eastern district).Eurasian Soil Sci.47:203-215.
Nóbrega F M,Santos I S,Cunha M D,Carvalho A O,Gomes V M.2005.Antimicrobial proteins from cowpea root exudates:Inhibitory activity against Fusarium oxysporum and purification of a chitinase-like protein.Plant Soil.272:223-232.
Nozoye T,Nagasaka S,Kobayashi T,Takahashi M,Sato Y,Sato Y,Uozumi N,Nakanishi H,Nishizawa N K.2011.Phytosiderophore efflux transporters are crucial for iron acquisition in graminaceous plants.J Biol Chem.286:5446-5454.
Oliveira V,Gomes N C M,Almeida A,Silva A M S,Silva H,Cunha?.2015.Microbe-assisted phytoremediation of hydrocarbons in estuarine environments.Microb Ecol.69:1-12.
Oliveira V,Santos A L,Aguiar C,Santos L,Salvador? C,Gomes N C M,Silva H,Rocha S M,Almeida A,Cunha?.2012.Prokaryotes in salt marsh sediments of Ria de Aveiro:Effects of halophyte vegetation on abundance and diversity.Estuar Coast Shelf Sci.110:61-68.
Orhan F.2016.Alleviation of salt stress by halotolerant and halophilic plant growth-promoting bacteria in wheat(Triticum aestivum).Braz J Microbiol.47:621-627.
Paul D,Lade H.2014.Plant-growth-promoting rhizobacteria to improve crop growth in saline soils:A review.Agron Sustain Dev.34:737-752.
Pausch J,Kuzyakov Y.2018.Carbon input by roots into the soil:Quantification of rhizodeposition from root to ecosystem scale.Glob Change Biol.24:1-12.
Pessarakli M(ed.).1999.Handbook of Plant and Crop Stress.2nd Edn.Marcel Dekker,New York.
Pessarakli M.2014.Handbook of Plant and Crop Physiology.3rd Edn.CRC Press,Boca Raton.
Petropoulos S A,Levizou E,Ntatsi G,Fernandes?,Petrotos K,Akoumianakis K,Barros L,Ferreira I C F R.2017.Salinity effect on nutritional value,chemical composition and bioactive compounds content of Cichorium spinosum L.Food Chem.214:129-136.
Pii Y,Marastoni L,Springeth C,Fontanella M C,Beone G M,Cesco S,Mimmo T.2016.Modulation of Fe acquisition process by Azospirillum brasilense in cucumber plants.Environ Exp Bot.130:216-225.
Pii Y,Mimmo T,Tomasi N,Terzano R,Cesco S,Crecchio C.2015a.Microbial interactions in the rhizosphere:Beneficial influences of plant growth-promoting rhizobacteria on nutrient acquisition process.A review.Biol Fert Soils.51:403-415.
Pii Y,Penn A,Terzano R,Crecchio C,Mimmo T,Cesco S.2015b.Plant-microorganism-soil interactions influence the Fe availability in the rhizosphere of cucumber plants.Plant Physiol Biochem.87:45-52.
Posada R H,Heredia-Abarca G,Sieverding E,Sánchez de Prager M.2013.Solubilization of iron and calcium phosphates by soil fungi isolated from coffee plantations.Arch Agron Soil Sci.59:185-196.
Puig S,Ramos-Alonso L,Romero A M,Martínez-Pastor M T.2017.The elemental role of iron in DNA synthesis and repair.Metallomics.9:1483-1500.
Purohit D,Sankararamasubramanian H M,Kumar Pal A,Kumar Parida A.2016.Identification and characterization of a novel iron deficiency and salt stress responsive transcription factor IDEF1 in Porteresia coarctata.Biol Plant.60:469-481.
Qadir M,Noble A D,Oster J D,Schubert S,Ghafoor A.2005.Driving forces for sodium removal during phytoremediation of calcareous sodic and saline-sodic soils:A review.Soil Use Manage.21:173-180.
Qi W Z,Zhao L.2013.Study of the siderophore-producing Trichoderma asperellum Q1 on cucumber growth promotion under salt stress.J Basic Microbiol.53:355-364.
Rabhi M,Barhoumi Z,Ksouri R,Abdelly C,Gharsalli M.2007.Interactive effects of salinity and iron deficiency in Medicago ciliaris.CR Biol.330:779-788.
Ramadoss D,Lakkineni V K,Bose P,Ali S,Annapurna K.2013.Mitigation of salt stress in wheat seedlings by halotolerant bacteria isolated from saline habitats.SpringerPlus.2:6.
Reich M,Aghajanzadeh T,Helm J,Parmar S,Hawkesford M J,De Kok L J.2017.Chloride and sulfate salinity differently affect biomass,mineral nutrient composition and expression of sulfate transport and assimilation genes in Brassica rapa.Plant Soil.411:319-332.
Rodríguez-Celma J,Lin W D,Fu G M,Abadía J,López-Míllán A F,Schmidt W.2013.Mutually exclusive alterations in secondary metabolism are critical for the uptake of insoluble iron compounds by Arabidopsis and Medicago truncatula.Plant Physiol.162:1473-1485.
Rueda-Puente E O,García-Hernández J L,Preciado-Rangel P,Murillo-Amador B,Tarazón-Herrera M A,Flores-Hernández A,Holguin-Pe?a J,Aybar A N,Barrón Hoyos J M,Weimers D,Mwandemele O,Kaaya G,Mayoral J L,Troyo-Diéguez E.2007.Germination of Salicornia bigelovii ecotypes under stressing conditions of temperature and salinity and ameliorative effects of plant growth-promoting bacteria.J Agron Crop Sci.193:167-176.
Rungin S,Indananda C,Suttiviriya P,Kruasuwan W,Jaemsaeng R,Thamchaipenet A.2012.Plant growth enhancing effects by a siderophore-producing endophytic streptomycete isolated from a Thai jasmine rice plant(Oryza sativa L.cv.KDML105).Anton Leeuw J Microbiol.102:463-472.
Saha M,Sarkar S,Sarkar B,Sharma B K,Bhattacharjee S,Tribedi P.2016.Microbial siderophores and their potential applications:A review.Environ Sci Pollut Res.23:3984-3999.
Sandmann G.1985.Consequences of iron deficiency on photosynthetic and respiratory electron transport in blue-green algae.Photosynth Res.6:261-271.
Saravanakumar D,Samiyappan R.2007.ACC deaminase from Pseudomonas fluorescens mediated saline resistance in groundnut(Arachis hypogea)plants.J Appl Microbiol.102:1283-1292.
Sarkar A,Ghosh P K,Pramanik K,Mitra S,Soren T,Pandey S,Mondal M H,Maiti T K.2018.A halotolerant Enterobacter sp.displaying ACC deaminase activity promotes rice seedling growth under salt stress.Res Microbiol.169:20-32.
Scagliola M,Pii Y,Mimmo T,Cesco S,Ricciuti P,Crecchio C.2016.Characterization of plant growth promoting traits of bacterial isolates from the rhizosphere of barley(Hordeum vulgare L.)and tomato(Solanum lycopersicon L.)grown under Fe sufficiency and deficiency.Plant Physiol Biochem.107:187-196.
Schalk I J,Mislin G L A.2017.Bacterial iron uptake pathways:Gates for the import of bactericide compounds.J Med Chem.60:4573-4576.
Séguéla M,Briat J F,Vert G,Curie C.2008.Cytokinins negatively regulate the root iron uptake machinery in Arabidopsis through a growth-dependent pathway.Plant J.55:289-300.
Shameer S,Prasad T N V K V.2018.Plant growth promoting rhizobacteria for sustainable agricultural practices with special reference to biotic and abiotic stresses.Plant Growth Regul.84:603-615.
Shannon M C,Grieve C M.1998.Tolerance of vegetable crops to salinity.Sci Hortic.78:5-38.
Shirley M,Avoscan L,Bernaud E,Vansuyt G,Lemanceau P.2011.Comparison of iron acquisition from Fe-pyoverdine by strategy I and strategy II plants.Botany.89:731-735.
Singh R P,Jha P,Jha P N.2015.The plant-growth-promoting bacterium Klebsiella sp.SBP-8 confers induced systemic tolerance in wheat(Triticum aestivum)under salt stress.JPlant Physiol.184:57-67.
Sisó-Terraza P,Rios J J,Abadía J,Abadía A,álvarez-Fernández A.2016.Flavins secreted by roots of iron-deficient Beta vulgaris enable mining of ferric oxide via reductive mechanisms.New Phytol.209:733-745.
T??bi K,T??bi F,Ait Abderrahim L,Ennajah A,Belkhodja M,Mulet J M.2016.Effect of salt stress on growth,chlorophyll content,lipid peroxidation and antioxidant defence systems in Phaseolus vulgaris L.South Afr J Bot.105:306-312.
Tank N,Saraf M.2010.Salinity-resistant plant growth promoting rhizobacteria ameliorates sodium chloride stress on tomato plants.J Plant Interact.5:51-58.
Tiwari S,Singh P,Tiwari R,Meena K K,Yandigeri M,Singh D P,Arora D K.2011.Salt-tolerant rhizobacteria-mediated induced tolerance in wheat(Triticum aestivum)and chemical diversity in rhizosphere enhance plant growth.Biol Fert Soils.47:907-916.
Torabian S,Zahedi M,Khoshgoftar A H.2017.Effects of foliar spray of nano-particles of FeSO4 on the growth and ion content of sunflower under saline condition.J Plant Nutr.40:615-623.
Trapet P,Avoscan L,Klinguer A,Pateyron S,Citerne S,Chervin C,Mazurier S,Lemanceau P,Wendehenne D,Besson-Bard A.2016.The Pseudomonas fluorescens siderophore pyoverdine weakens Arabidopsis thaliana defense in favor of growth in iron-deficient conditions.Plant Physiol.171:675-693.
Tsai H H,Schmidt W.2017.Mobilization of iron by plant-borne coumarins.Trends Plant Sci.22:538-548.
Turan M A,Elkarim A H A,Taban N,Taban S.2010.Effect of salt stress on growth and ion distribution and accumulation in shoot and root of maize plant.Afr J Agric Res.5:584-588.
Vivas A,Marulanda A,Ruiz-Lozano J M,Barea J M,Azcón R.2003.Influence of a Bacillus sp.on physiological activities of two arbuscular mycorrhizal fungi and on plant responses to peg-induced drought stress.Mycorrhiza.13:249-256.
Wallender W W,Tanji K K(eds.).2011.Agricultural Salinity Assessment and Management.2nd Edn.American Society of Civil Engineers,Reston.
Wang X P,Yang J S,Liu G M,Yao R J,Yu S P.2015.Impact of irrigation volume and water salinity on winter wheat productivity and soil salinity distribution.Agric Water Manage.149:44-54.
Winkelmann G.2007.Ecology of siderophores with special reference to the fungi.BioMetals.20:379-392.
Witzel K,Weidner A,Surabhi G K,B?rner A,Mock H P.2009.Salt stress-induced alterations in the root proteome of barley genotypes with contrasting response towards salinity.J Exp Bot.60:3545-3557.
Yang A Z,Akhtar S S,Iqbal S,Amjad M,Naveed M,Zahir ZA,Jacobsen S E.2016.Enhancing salt tolerance in quinoa by halotolerant bacterial inoculation.Funct Plant Biol.43:632-642.
Younesi O,Moradi A.2014.Effects of plant growth-promoting rhizobacterium(PGPR)and arbuscular mycorrhizal fungus(AMF)on antioxidant enzyme activities in salt-stressed bean(Phaseolus vulgaris L.).Agriculture.60:10-21.
Yousfi S,Wissal M,Mahmoudi H,Abdelly C,Gharsalli M.2007.Effect of salt on physiological responses of barley to iron deficiency.Plant Physiol Biochem.45:309-314.
Zhang H M,Sun Y,Xie X T,Kim M S,Dowd S E,Paré P W.2009.A soil bacterium regulates plant acquisition of iron via deficiency-inducible mechanisms.Plant J.58:568-577.
