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Plant sciences, sustainable farming systems and food quality

Growth, ion content, gas exchange, and water relations of wheat genotypes differing in salt tolerances

Salah E. El-Hendawy A B , Yuncai Hu A C and Urs Schmidhalter A

A Department of Plant Science, Technical University of Munich, Am Hochanger 2, D-85350 Freising-Weihenstephan, Germany.

B Suez Canal University, Faculty of Agriculture, Agronomy Department, Ismalia, Egypt.

C Corresponding author. Email:

Australian Journal of Agricultural Research 56(2) 123-134
Submitted: 27 January 2004  Accepted: 13 January 2005   Published: 28 February 2005


Although the mechanisms of salt tolerance in plants have received much attention for many years, genotypic differences influencing salt tolerance still remain uncertain. To investigate the key physiological factors associated with genotypic differences in salt tolerance of wheat and their relationship to salt stress, 13 wheat genotypes from Egypt, Australia, India, and Germany, that differ in their salt tolerances, were grown in a greenhouse in soils of 4 different salinity levels (control, 50, 100, and 150 mm NaCl). Relative growth rate (RGR), net assimilation rate (NAR), leaf area ratio (LAR), photosynthesis, chlorophyll content (SPAD value), and leaf water relations were measured at Days 45 and 60 after sowing. Mineral nutrient content in leaves and stems was determined at Day 45 and final harvest. Salinity reduced RGR, NAR, photosynthetic rate, stomatal conductance, water and osmotic potentials, and K+ and Ca2+ content in stems and leaves at all times, whereas it increased leaf respiration, and Na+ and Cl content in leaves and stems. LAR was not affected by salinity and the effect of salinity on SPAD value was genotype-dependent. Growth of salt-tolerant genotypes (Sakha 8, Sakha 93, and Kharchia) was affected by salinity primarily due to a decline in photosynthetic capacity rather than a reduction in leaf area, whereas NAR was the more important factor in determining RGR of moderately tolerant and salt-sensitive genotypes. We conclude that Na+ and Cl exclusion did not always reflect the salt tolerance, whereas K+ in the leaves and Ca2+ in the leaves and stems were closely associated with genotypic differences in salt tolerance among the 13 genotypes even at Day 45. Calcium content showed a greater difference in salt tolerance among the genotypes than did K+ content. The genotypic variation in salt tolerance was also observed for the parameters involved in photosynthesis, and water and osmotic potentials, but not for turgor pressure.

Additional keywords: mineral elements, photosynthesis, plant growth, salinity, salt tolerance.


Ashraf M 1999 Interactive effect of salt (NaCl) and nitrogen form on growth, water relations and photosynthetic capacity of sunflower (Helianthus annuus L.). Annals of Applied Biology 135 509 513

Ashraf M 2001 Relationships between growth and gas exchange characteristics in some salt-tolerant amphidiploid Brassica species in relation to their diploid parents. Environmental and Experimental Botany 45 155 163

Ashraf M 2004 Some important physiological selection criteria for salt tolerance in plants. Flora 199 361 376

Ashraf M Mcneilly T Bradshaw AD 1986 The response to NaCl and ionic content of selected salt-tolerant and normal lines of 3 legume forage species in sand culture. New Phytologist 104 463 471

Ashraf M Waheed A 1993 Responses of some local exotic accessions of lentil (Lens culinaris Medic) to salt stress. Journal of Agronomy and Crop Science – Zeitschrift für Acker Und Pflanzenbau 170 103 112

Cheeseman JM 1988 Mechanisms of salinity tolerance in plants. Plant Physiology 87 547 550

Chow WS Ball MC Anderson JM 1990 Growth and photosynthetic responses of spinach to salinity—implications of K+ nutrition for salt tolerance. Australian Journal of Plant Physiology 17 563 578

Cramer GR 2002 Sodium–calcium interactions under salinity stress. ‘Salinity: environment–plants–molecules’. Läuchli A Lüttge U 205 227 Kluwer Dordrecht, The Netherlands

Cramer GR Alberico GJ Schmidt C 1994 Salt tolerance is not associated with the sodium accumulation of 2 maize hybrids. Australian Journal of Plant Physiology 21 675 692

Curtis PS Läuchli A 1986 The role of leaf-area development and photosynthetic capacity in determining growth of kenaf under moderate salt stress. Australian Journal of Plant Physiology 13 553 565

Delfine S Alvino A Villani MC Loreto F 1999 Restrictions to carbon dioxide conductance and photosynthesis in spinach leaves recovering from salt stress. Plant Physiology 119 1101 1106

Demidchik V Tester M 2002 Sodium fluxes through nonselective cation channels in the plasma membrane of protoplasts from Arabidopsis roots. Plant Physiology 128 379 387 DOI

El-Hendawy SE Hu Y Yakout GM Awad AM Hafiz SE Schmidhalter U 2004 Evaluating salt tolerance of wheat genotypes using multiple parameters. European Journal of Agronomy (in press)

Essah PA Davenport R Tester M 2003 Sodium influx and accumulation in Arabidopsis. Plant Physiology 133 307 318

Ghassemi F Jakeman AJ Nix HA 1995 ‘Salinization of land and water resources.’ University of New South Wales Press Ltd Canberra, ACT

Gorham J 1992 Salt tolerance of plants. Science in Progress 76 273 285

Gorham J Hardy C Wyn Jones RG Joppa LR Law CN 1987 Chromosomal location of a K/Na discrimination character in the D genome of wheat. Theoretical and Applied Genetics 74 584 588

Gorham J Wyn Jones RG Bristol A 1990 Partial characterization of the trait for enhanced K+–Na+ discrimination in the D-genome of wheat. Planta 180 590 597

Greenway H Munns R 1980 Mechanisms of salt tolerance in non-halophytes. Annual Review of Plant Physiology and Plant Molecular Biology 31 149 190

Hasegawa PM Bressan RA Zhu JK Bohnert HJ 2000 Plant cellular and molecular responses to high salinity. Annual Review of Plant Physiology and Plant Molecular Biology 51 463 499

Hawkins HJ Lewis OAM 1993 Combination effect of NaCl salinity, nitrogen form and calcium-concentration on the growth, ionic content and gaseous exchange properties of Triticum aestivum L. cv. gamtoos. New Phytologist 124 161 170

He T Cramer GR 1993 a Salt tolerance of rapid-cycling Brassica species in relation to potassium–sodium ratio and selectivity at the whole plant and callus levels. Journal of Plant Nutrition 16 1263 1277

He T Cramer GR 1993 b Cellular-responses of 2 rapid-cycling Brassica species, Brassica napus and B. carinata, to seawater salinity. Physiologia Plantarum 87 54 60

Heuer B Plaut Z 1989 Photosynthesis and osmotic adjustment of 2 sugarbeet cultivars grown under saline conditions. Journal of Experimental Botany 40 437 440

Hunt R 1990 ‘Basic growth analysis: plant growth analysis for beginners.’ Academic Press London, UK

Ishikawa S Oikawa T Furukawa A 1991 Responses of photosynthesis, leaf conductance and growth to different salinities in 3 coastal dune plants. Ecological Research 6 217 226

James RA Rivelli AR Munns R Von Caemmerer S 2002 Factors affecting CO2 assimilation, leaf injury and growth in salt-stressed durum wheat. Functional Plant Biology 29 1393 1403

Knight H 2000 Calcium signaling during abiotic stress in plants. International Review of Cytology, A Survey of Cell Biology 195 269 324

LaHaye PA Epstein E 1971 Calcium and salt toleration by bean plants. Physiologia Plantarum 25 213 218

Leidi EO Saiz JF 1997 Is salinity tolerance related to Na accumulation in upland cotton (Gossypium hirsutum) seedlings? Plant and Soil 190 67 75

Maathuis FJM Amtmann A 1999 K+ Nutrition and Na+ toxicity: the basis of cellular K+/Na+ ratios. Annals of Botany 84 123 133 DOI

Marschner H 1995 ‘Mineral nutrition of higher plants.’ Academic Press London, UK

Morales MA Sanchez-Blanco MJ Olmos E Torrecillas A Alarcon JJ 1998 Changes in the growth, leaf water relations and cell ultrastructure in Argyranthemum coronopifolium plants under saline conditions. Journal of Plant Physiology 153 174 180

Munns R 1993 Physiological processes limiting plant-growth in saline soils—some dogmas and hypotheses. Plant, Cell and Environment 16 15 24

Munns R Husain S Rivelli AR James RA Condon AG Lindsay MP Lagudah ES Schachtman DP Hare RA 2002 Avenues for increasing salt tolerance of crops, and the role of physiologically based selection traits. Plant and Soil 247 93 105

Munns R James RA 2003 Screening methods for salinity tolerance: a case study with tetraploid wheat. Plant and Soil 253 201 218 DOI

Murillo-Amador B Troyo-Dieguez E Lopezaguilar R Lopez-Cortes A Tinoco-Ojanguren CL Jones HG Kaya C 2002 Matching physiological traits and ion concentrations associated with salt stress in cowpea genotypes. Australian Journal of Agricultural Research 53 1243 1255 DOI

Rengel Z 1992 The role of calcium in salt toxicity. Plant, Cell and Environment 15 625 632

Rivelli AR James RA Munns R Condon AG 2002 Effect of salinity on water relations and growth of wheat genotypes with contrasting sodium uptake. Functional Plant Biology 29 1065 1074

Robinson SP Downton WJS Millhouse JA 1983 Photosynthesis and ion content of leaves and isolated-chloroplasts of salt-stressed spinach. Plant Physiology 73 238 242

Rogers ME Noble CL 1992 Variation in growth and ion accumulation between 2 selected populations of Trifolium repens L. differing in salt tolerance. Plant and Soil 146 131 136

SAS Institute 2000 ‘SAS User’s guide (Version 4.0.2).’ SAS Inst. Cary, NC

Schachtman DP Munns R 1992 Sodium accumulation in leaves of Triticum species that differ in salt tolerance. Australian Journal of Plant Physiology 19 331 340

Scholander PF Hamme HT Bradstreet ED Hemmingsen EA 1965 Sap pressure in vascular plants. Science 148 339 346

Schwarz M Gale J 1981 Maintenance respiration and carbon balance of plants at low-levels of sodium-chloride salinity. Journal of Experimental Botany 32 933 941

Seemann JR Critchley C 1985 Effects of salt stress on the growth, ion content, stomatal behavior and photosynthetic capacity of a salt-sensitive species, Phaseolus vulgaris L. Planta 164 151 162

Semikhatova OA Ivanova TI Yudina OS 1993 Respiratory cost of plant-growth under conditions of salinity. Russian Plant Physiology 40 490 497

Serraj R Sinclair TR 2002 Osmolyte accumulation: Can it really help increase crop yield under drought conditions? Plant, Cell and Environment 25 333 341

Shabala S 2000 Ionic and osmotic components of salt stress specifically modulate net ion fluxes from bean leaf mesophyll. Plant, Cell and Environment 23 825 837 DOI

Shabala S Shabala L Van Volkenburgh E 2003 Effect of calcium on root development and root ion fluxes in salinised barley seedlings. Functional Plant Biology 30 507 514 DOI

Soussi M Ocana A Liuch C 2001 Growth, nitrogen fixation and ion accumulation in two chickpea cultivars under salt stress. Agricoltura Mediterranea 131 1 8

Tattini M Gucci R Coradeschi MA Ponzio C Everard JD 1995 Growth, gas-exchange and ion content in Olea europaea plants during salinity stress and subsequent relief. Physiologia Plantarum 95 203 210

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

Yeo AR Flowers TJ 1983 Varietal differences in the toxicity of sodium-ions in rice leaves. Physiologia Plantarum 59 189 195

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