Free Standard AU & NZ Shipping For All Book Orders Over $80!
Register      Login
Functional Plant Biology Functional Plant Biology Society
Plant function and evolutionary biology
RESEARCH ARTICLE

Root traits and δ13C and δ18O of durum wheat under different water regimes

Abdelhalim Elazab A , Gemma Molero A B , Maria Dolores Serret A and José Luis Araus A B C
+ Author Affiliations
- Author Affiliations

A Unitat de Fisiologia Vegetal, Facultat de Biologia, Universitat de Barcelona, Av. Diagonal, 647, 08028 Barcelona, Spain.

B International Maize and Wheat Improvement Center (CIMMYT), El Batán, Texcoco, CP 56130, Mexico.

C Corresponding author. Email: jaraus@ub.edu

Functional Plant Biology 39(5) 379-393 https://doi.org/10.1071/FP11237
Submitted: 22 October 2011  Accepted: 13 March 2012   Published: 27 April 2012

Abstract

Plant growth, root characteristics and the stable carbon (δ13C) and oxygen (δ18O) composition were studied in durum wheat. Four recombinant inbred lines with good agronomic adaptation were grown under well watered (WW) and water stress (WS) conditions until mid-grain filling in lysimeters. Gas exchange was measured in the flag leaf just before harvest and then the aerial dry matter (Aerial DM), root weight density (RWD) and root length density (RLD) and the specific root length (SRL) were evaluated and the δ13C and δ18O of the roots, the flag leaf blade and the spike were analysed. Water stress decreased stomatal conductance, plant accumulated transpiration and Aerial DM, whereas δ13C and δ18O increased. Genotypic differences were found for all gas-exchange and root traits and isotope signatures. Aerial DM was positively correlated with RLD, regardless of the water regime, whereas it was negatively correlated with δ13C and δ18O, but only under WW conditions. Moreover, RWD and RLD were negatively related to both δ13C and δ18O under the WW regime, but no clear pattern existed under WS. Our study supports the use of δ13C and δ18O as proxies for selecting root traits associated with higher growth in the absence of water stress.

Additional keywords: leaf gas exchange, lysimeter, Triticum turgidum ssp. durum, root traits, δ13C, δ18O.


References

Araki H, Iijima M (2001) Deep rooting in winter wheat: rooting of nodes of deep roots in two cultivars with deep and shallow root systems. Plant Production Science 4, 215–219.
Deep rooting in winter wheat: rooting of nodes of deep roots in two cultivars with deep and shallow root systems.Crossref | GoogleScholarGoogle Scholar |

Araus JL (2004) The problem of sustainable water use in the Mediterranean and research requirements for agriculture. Annals of Applied Biology 144, 229–272.

Araus JL, Brown HR, Febrero A, Bort J, Serret MD (1993) Ear photosynthesis, carbon isotope discrimination and the respiratory CO2 to differences in grain mass in durum wheat. Plant, Cell & Environment 16, 383–392.
Ear photosynthesis, carbon isotope discrimination and the respiratory CO2 to differences in grain mass in durum wheat.Crossref | GoogleScholarGoogle Scholar |

Araus JL, Amaro T, Casadesus J, Asbati A, Nachit MM (1998) Relationships between ash content, carbon isotope discrimination and yield in durum wheat. Australian Journal of Plant Physiology 25, 835–842.
Relationships between ash content, carbon isotope discrimination and yield in durum wheat.Crossref | GoogleScholarGoogle Scholar |

Araus JL, Slafer GA, Reynolds MP, Royo C (2002) Plant breeding and drought in C3 cereals: what should we breed for? Annals of Botany 89, 925–940.
Plant breeding and drought in C3 cereals: what should we breed for?Crossref | GoogleScholarGoogle Scholar |

Araus JL, Villegas D, Aparicio N, del Moral LFG, El Hani S, Rharrabti Y, Ferrio JP, Royo C (2003) Environmental factors determining carbon isotope discrimination and yield in durum wheat under Mediterranean conditions. Crop Science 43, 170–180.
Environmental factors determining carbon isotope discrimination and yield in durum wheat under Mediterranean conditions.Crossref | GoogleScholarGoogle Scholar |

Araus JL, Slafer GA, Royo C, Serret MD (2008) Breeding for yield potential and stress adaptation in cereals. Critical Reviews in Plant Sciences 27, 377–412.
Breeding for yield potential and stress adaptation in cereals.Crossref | GoogleScholarGoogle Scholar |

Baburai Nagesh AK (2006) The physiological and genetic bases of water-use efficiency in winter wheat. PhD thesis, The University of Nottingham, UK.

Badeck F-W, Tcherkez G, Nogués S, Piel C, Ghashgaie J (2005) Post-photosynthetic fractionation of carbon stable isotopes between plant organs – a widespread phenomenon. Rapid Communications in Mass Spectrometry 19, 1381–1391.
Post-photosynthetic fractionation of carbon stable isotopes between plant organs – a widespread phenomenon.Crossref | GoogleScholarGoogle Scholar |

Barbour MM, Farquhar GD (2000) Relative humidity- and ABA-induced variation in carbon and oxygen isotope ratios of cotton leaves. Plant, Cell & Environment 23, 473–485.
Relative humidity- and ABA-induced variation in carbon and oxygen isotope ratios of cotton leaves.Crossref | GoogleScholarGoogle Scholar |

Barbour MM, Fischer RA, Sayre KD, Farquhar GD (2000) Oxygen isotope ratio of leaf and grain material correlates with stomatal conductance and grain yield in irrigated wheat. Australian Journal of Plant Physiology 27, 625–637.

Barraclough PB (1989) Root-growth, macro-nutrient uptake dynamics and soil fertility requirements of a high-yielding winter oilseed rape crop. Plant and Soil 119, 59–70.
Root-growth, macro-nutrient uptake dynamics and soil fertility requirements of a high-yielding winter oilseed rape crop.Crossref | GoogleScholarGoogle Scholar |

Blum A (2005) Drought resistance, water-use efficiency and yield potential – are they compatible, dissonant, or mutually exclusive? Australian Journal of Agricultural Research 56, 1159–1168.
Drought resistance, water-use efficiency and yield potential – are they compatible, dissonant, or mutually exclusive?Crossref | GoogleScholarGoogle Scholar |

Blum A (2009) Effective use of water (EUW) and not water-use efficiency (WUE) is the target of crop yield improvement under drought stress. Field Crops Research 112, 119–123.
Effective use of water (EUW) and not water-use efficiency (WUE) is the target of crop yield improvement under drought stress.Crossref | GoogleScholarGoogle Scholar |

Cabrera-Bosquet L, Molero G, Bort J, Nogués S, Araus JL (2009a) The combined effect of constant water deficit and nitrogen supply on WUE, NUE and Δ13C in durum wheat potted plants. Journal of Experimental Botany 60, 1633–1644.
The combined effect of constant water deficit and nitrogen supply on WUE, NUE and Δ13C in durum wheat potted plants.Crossref | GoogleScholarGoogle Scholar |

Cabrera-Bosquet L, Sanchez C, Araus JL (2009b) Oxygen isotope enrichment (Δ18O) reflects yield potential and drought resistance in maize. Plant, Cell & Environment 32, 1487–1499.
Oxygen isotope enrichment (Δ18O) reflects yield potential and drought resistance in maize.Crossref | GoogleScholarGoogle Scholar |

Cabrera-Bosquet L, Albrizio R, Nogués S, Araus JL (2011) Dual Δ13C/δ18O response to water and nitrogen availability and its relationship with yield in field-grown durum wheat. Plant, Cell & Environment 34, 418–433.
Dual Δ13C/δ18O response to water and nitrogen availability and its relationship with yield in field-grown durum wheat.Crossref | GoogleScholarGoogle Scholar |

Carvalho PM (2009) Optimising root growth to improve uptake and utilization of water and nitrogen in wheat and barley. PhD thesis, University of Nottingham, UK.

Cernusak LA, Winter K, Aranda J, Turner BL, Marshall JD (2007) Transpiration efficiency of a tropical pioneer tree (Ficus insipida) in relation to soil fertility. Journal of Experimental Botany 58, 3549–3566.
Transpiration efficiency of a tropical pioneer tree (Ficus insipida) in relation to soil fertility.Crossref | GoogleScholarGoogle Scholar |

Cernusak LA, Tcherkez G, Keitel C, Cornwell WK, Santiago LS, Knohl A, Barbour MM, Williams DG, Reich PB, Ellsworth DS, Dawson TE, Griffiths HG, Farquhar GD, Wright IJ (2009a) Why are non-photosynthetic tissues generally 13C enriched compared with leaves in C3 plants? Review and synthesis of current hypotheses. Functional Plant Biology 36, 199–213.
Why are non-photosynthetic tissues generally 13C enriched compared with leaves in C3 plants? Review and synthesis of current hypotheses.Crossref | GoogleScholarGoogle Scholar |

Cernusak LA, Winter K, Turner BL (2009b) Physiological and isotopic (δ13C and δ18O) responses of three tropical tree species to water and nutrient availability. Plant, Cell & Environment 32, 1441–1455.
Physiological and isotopic (δ13C and δ18O) responses of three tropical tree species to water and nutrient availability.Crossref | GoogleScholarGoogle Scholar |

Christopher JT, Manschadi AM, Hammer GL, Borrell AK (2008) Development and physiological traits associated with high yield and stay-green phenotype in wheat. Australian Journal of Agricultural Research 59, 354–364.
Development and physiological traits associated with high yield and stay-green phenotype in wheat.Crossref | GoogleScholarGoogle Scholar |

Condon AG, Richards RA (1992) Broad sense heritability and genotypes-environment interaction for carbon isotope discrimination in field-grown wheat. Australian Journal of Agricultural Research 43, 921–934.
Broad sense heritability and genotypes-environment interaction for carbon isotope discrimination in field-grown wheat.Crossref | GoogleScholarGoogle Scholar |

Condon AG, Richards RA, Rebetzke GJ, Farquhar GD (2002) Improving intrinsic water-use efficiency and crop yield. Crop Science 42, 122–132.
Improving intrinsic water-use efficiency and crop yield.Crossref | GoogleScholarGoogle Scholar |

Coplen TB (2008) ‘Explanatory glossary of terms used in expression of relative isotope ratios and gas ratios. IUPAC Recommendations 2008’. (International Union of Pure and Applied Chemistry Inorganic Chemistry Division, Commission on Isotopic Abundances and Atomic Weights: Research Triangle Park, NC, USA)

Cregg B, Zhang J (2000) Carbon isotope discrimination as a tool to screen for improved drought tolerance. In ‘Proceedings of the eleventh conference of the Metropolitan Tree Improvement Alliance’. (Gresham. OR, USA) Available at http://www.ces.ncsu.edu/fletcher/programs/nursery/metria/metria11/cregg/index.html [Verified May 2011]

Davies WJ, Bacon MA (2003) Adaptation of roots to drought. In ‘Root ecology’. (Eds H Kroon, EJW Visser) pp. 173–192. (Springer: Berlin)

Dwyer LM, Stewart DW, Balchin D (1988) Rooting characteristics of corn, soybeans and barley as a function of available water and soil physical characteristics. Canadian Journal of Soil Science 68, 121–132.
Rooting characteristics of corn, soybeans and barley as a function of available water and soil physical characteristics.Crossref | GoogleScholarGoogle Scholar |

Ehdaie B, Hall AE, Farquhar GD, Nguyen HT, Waines JG (1991) Water-use efficiency and carbon isotope discrimination in wheat. Crop Science 31, 1282–1288.
Water-use efficiency and carbon isotope discrimination in wheat.Crossref | GoogleScholarGoogle Scholar |

Eissenstat DM (1992) Costs and benefits of constructing roots of small diameter. Journal of Plant Nutrition 15, 763–782.
Costs and benefits of constructing roots of small diameter.Crossref | GoogleScholarGoogle Scholar |

Farquhar GD, Richards RA (1984) Isotopic composition of plant carbon correlates with water-use-efficiency of wheat genotypes. Australian Journal of Plant Physiology 11, 539–552.
Isotopic composition of plant carbon correlates with water-use-efficiency of wheat genotypes.Crossref | GoogleScholarGoogle Scholar |

Farquhar GD, Ehleringer JR, Hubick KT (1989) Carbon isotope discrimination and photosynthesis. Annual Review of Plant Physiology 40, 503–537.
Carbon isotope discrimination and photosynthesis.Crossref | GoogleScholarGoogle Scholar |

Farquhar GD, Cernusak LA, Barnes B (2007) Heavy water fractionation during transpiration. Plant Physiology 143, 11–18.
Heavy water fractionation during transpiration.Crossref | GoogleScholarGoogle Scholar |

Ferrio JP, Mateo MA, Bort J, Abdalla O, Voltas J, Araus JL (2007) Relationships of grain δ13C and δ18O with wheat phenology and yield under water-limited conditions. The Annals of Applied Biology 150, 207–215.
Relationships of grain δ13C and δ18O with wheat phenology and yield under water-limited conditions.Crossref | GoogleScholarGoogle Scholar |

Food and Agriculture Organization of the United Nations (FAO) (2008) ‘FAO Statistical Yearbook 2007–2008.’ Available at http://www.fao.org/economic/ess/ess-publications/ess-yearbook/fao-statistical-yearbook-2007-2008/b-agricultural-production/jp/ [Verified May 2011].

Galmés J, Pou A, Alsina MM, Tomàs M, Medrano H, Flexas J (2007) Aquaporin expression in response to different water stress intensities and recovery in Richter-110 (Vitis sp.): relationship with ecophysiological status. Planta 226, 671–681.
Aquaporin expression in response to different water stress intensities and recovery in Richter-110 (Vitis sp.): relationship with ecophysiological status.Crossref | GoogleScholarGoogle Scholar |

Gan KS, Wong SC, Yong JWH, Farquhar GD (2003) Evaluation of models of leaf water 18O enrichment using measurements of spatial patterns of vein xylem water, leaf water and dry matter in maize leaves. Plant, Cell & Environment 26, 1479–1495.
Evaluation of models of leaf water 18O enrichment using measurements of spatial patterns of vein xylem water, leaf water and dry matter in maize leaves.Crossref | GoogleScholarGoogle Scholar |

Głąb T (2007) Effect of soil compaction on root system development and yields of tall fescue. International Agrophysics 21, 233–239.

Gregory PJ (1994) Resource capture by root networks. In ‘Resource capture by crops’. (Eds JL Monteith, RK Scott, MH Unsworth) pp. 77–97. (Nottingham University Press: Nottingham)

Gregory PJ (2006) ‘Plant roots: their growth, activity and interactions with soil.’ (Blackwell Scientific Publications: Oxford, UK)

Gregory PJ, McGowan M, Biscoe PV (1978) Water relations of winter wheat. 2 Soil water relations. The Journal of Agricultural Science 91, 103–116.
Water relations of winter wheat. 2 Soil water relations.Crossref | GoogleScholarGoogle Scholar |

Gregory PJ, Tennant D, Belford RK (1992) Root and shoot growth and water and light-use efficiency of barley and wheat crops grown on a shallow duplex soil in a Mediterranean-type environment. Australian Journal of Agricultural Research 43, 555–573.
Root and shoot growth and water and light-use efficiency of barley and wheat crops grown on a shallow duplex soil in a Mediterranean-type environment.Crossref | GoogleScholarGoogle Scholar |

Gregory PJ, Palta JA, Batts GR (1997) Root systems and root : mass ratio – carbon allocation under current and projected atmospheric conditions in arable crops. Plant and Soil 187, 221–228.
Root systems and root : mass ratio – carbon allocation under current and projected atmospheric conditions in arable crops.Crossref | GoogleScholarGoogle Scholar |

Gregory PJ, Bengough AG, Grinev D, Schmidt S, Thomas WTB, Wojciechowski T, Young IM (2009) Root phenomics of crops: opportunities and challenges. Functional Plant Biology 36, 922–929.
Root phenomics of crops: opportunities and challenges.Crossref | GoogleScholarGoogle Scholar |

Ho MD, Rosas JC, Brown KM, Lynch JP (2005) Root architectural tradeoffs for water and phosphorus acquisition. Functional Plant Biology 32, 737–748.
Root architectural tradeoffs for water and phosphorus acquisition.Crossref | GoogleScholarGoogle Scholar |

Hoagland DR, Arnon DI (1938) The water-culture method for growing plants without soil. California Agricultural Experiment Station Circular 347, 1–39.

Hobbie EA, Werner RA (2004) Intramolecular, compound-specific and bulk carbon isotope patterns in C3 and C4 plants: a review and synthesis. New Phytologist 161, 371–385.
Intramolecular, compound-specific and bulk carbon isotope patterns in C3 and C4 plants: a review and synthesis.Crossref | GoogleScholarGoogle Scholar |

Kage H, Kochler M, Stützel H (2004) Root growth and dry matter partitioning of cauliflower under drought stress conditions: measurement and simulation. European Journal of Agronomy 20, 379–394.
Root growth and dry matter partitioning of cauliflower under drought stress conditions: measurement and simulation.Crossref | GoogleScholarGoogle Scholar |

Kang SY, Morita S, Yamazaki K (1994) Root growth and distribution in some Japonica-Indica hybrid and Japonica type rice cultivars under field conditions. Nihon Sakumotsu Gakkai Kiji 63, 118–124.
Root growth and distribution in some Japonica-Indica hybrid and Japonica type rice cultivars under field conditions.Crossref | GoogleScholarGoogle Scholar |

King J, Gay A, Sylvester-Bradley R, Bingham I, Foulkes J, Gregory PJ, Robinson D (2003) Modelling cereal root systems for water and nitrogen capture: towards an economic optimum. Annals of Botany 91, 383–390.
Modelling cereal root systems for water and nitrogen capture: towards an economic optimum.Crossref | GoogleScholarGoogle Scholar |

Kramer PJ (1983) ‘Water relations of plants.’ (Academic Press: Orlando, FL)

Lafitte R, Blum A, Atlin G (2003) Using secondary traits to help identify drought-tolerant genotypes. In ‘Breeding rice for drought-prone environments’. (Eds KS Fischer, R Lafitte, S Fukai, G Atlin, B Hardy) pp. 37–48. (International Rice Research Institute: Manila, Philippines)

Lopes MS, Reynolds MP (2010) Partitioning of assimilates to deeper roots is associated with cooler canopies and increased yield under drought in wheat. Functional Plant Biology 37, 147–156.
Partitioning of assimilates to deeper roots is associated with cooler canopies and increased yield under drought in wheat.Crossref | GoogleScholarGoogle Scholar |

Lovisolo C, Tramontini S, Flexas J, Schubert A (2008) Mercurial inhibition of root hydraulic conductance in Vitis spp. rootstocks under water stress. Environmental and Experimental Botany 63, 178–182.
Mercurial inhibition of root hydraulic conductance in Vitis spp. rootstocks under water stress.Crossref | GoogleScholarGoogle Scholar |

Ma Z, Wood CW, Bransby D (2000) Impacts of soil management on root characteristics of switchgrass. Biomass and Bioenergy 18, 105–112.
Impacts of soil management on root characteristics of switchgrass.Crossref | GoogleScholarGoogle Scholar |

Maccaferri M, Sanguineti MC, Corneti S, Araus JL, Ben Salern M, Bort J, DeAmbrogio E, del Moral LG, Demontis A, El-Ahmed A, Maalouf F, Machlab H, Martos V, Moragues M, Motawaj J, Nachit MM, Nserallah N, Ouabbou H, Royo C, Slama A, Tuberosa R (2008) Quantitative trait loci for grain yield and adaptation of durum wheat (Triticum durum Desf.) across a wide range of water availability. Genetics 178, 489–511.
Quantitative trait loci for grain yield and adaptation of durum wheat (Triticum durum Desf.) across a wide range of water availability.Crossref | GoogleScholarGoogle Scholar |

Manschadi AM, Hammer GL, Christopher JT, deVoil P (2008) Genotypic variation in seedling root architectural traits and implications for drought adaptation in wheat (Triticum aestivum L.). Plant and Soil 303, 115–129.
Genotypic variation in seedling root architectural traits and implications for drought adaptation in wheat (Triticum aestivum L.).Crossref | GoogleScholarGoogle Scholar |

Manske GGB, Vlek PLG (2002) Root architecture – wheat as a model plant. In ‘Plant roots: the hidden half’. (Eds Y Waisel, A Eshel, U Kafkafi) pp. 249–259. (Marcel Dekker Inc.: New York)

Merah O, Deléens E, Teulat B, Monneveux P (2002) Association between yield and carbon isotope discrimination value in different organs of durum wheat under drought. Journal Agronomy & Crop Science 188, 426–434.
Association between yield and carbon isotope discrimination value in different organs of durum wheat under drought.Crossref | GoogleScholarGoogle Scholar |

Monneveux P, Reynolds MP, Trethowan R, González-Santoyo H, Peña RJ, Zapata F (2005) Relationship between grain yield and carbon isotope discrimination in bread wheat under four water regimes. European Journal of Agronomy 22, 231–242.
Relationship between grain yield and carbon isotope discrimination in bread wheat under four water regimes.Crossref | GoogleScholarGoogle Scholar |

Monti A, Brugnoli E, Scartazza A, Amaducci MT (2006) The effect of transient and continuous drought on yield, photosynthesis and carbon isotope discrimination in sugar beet (Beta vulgaris L). Journal of Experimental Botany 57, 1253–1262.
The effect of transient and continuous drought on yield, photosynthesis and carbon isotope discrimination in sugar beet (Beta vulgaris L).Crossref | GoogleScholarGoogle Scholar |

Nachit MM, Elouafi I, Pagnotta MA, Salen AE, Iacono E, Labhilili M, Asbati A, Azrak M, Hazzam H, Bensher D, Khairallah M, Ribaut JM, Tanzarella OA, Porceddu E, Sorrells ME (2001) Molecular linkage map for an intraspecific recombinant inbred population of durum wheat (Triticum turgidum L. var durum). Theoretical and Applied Genetics 102, 177–186.
Molecular linkage map for an intraspecific recombinant inbred population of durum wheat (Triticum turgidum L. var durum).Crossref | GoogleScholarGoogle Scholar |

Olivares-Villegas JJ, Reynolds MP, McDonald GK (2007) Drought-adaptive attributes in the Seri/Babax hexaploid wheat population. Functional Plant Biology 34, 189–203.
Drought-adaptive attributes in the Seri/Babax hexaploid wheat population.Crossref | GoogleScholarGoogle Scholar |

Ostonen I, Püttsepp Ü, Biel C, Alberton O, Bakker MR, Lõhmus K, Majdi H, Metcalfe D, Olsthoorn AFM, Pronk A, Vanguelova E, Weih M, Brunner I (2007) Specific root length as an indicator of environmental change. Plant Biosystems 141, 426–442.
Specific root length as an indicator of environmental change.Crossref | GoogleScholarGoogle Scholar |

Passioura JB (1983) Root and drought resistance. Agricultural Water Management 7, 265–280.
Root and drought resistance.Crossref | GoogleScholarGoogle Scholar |

Passioura JB (2006) Increasing crop productivity when water is scarce – from breeding to field management. Agricultural Water Management 80, 176–196.
Increasing crop productivity when water is scarce – from breeding to field management.Crossref | GoogleScholarGoogle Scholar |

Razmjoo K, Heydarizadeh P, Sabzalian MR (2008) Effect of salinity and drought stresses on growth parameters and essential oil content of Matricaria chamomile. International Journal of Agriculture and Biology 10, 451–454.

Rebetzke GJ, Richards RA, Condon AG, Farquhar GD (2006) Inheritance of reduced carbon isotope discrimination in bread wheat (Triticum aestivum L). Euphytica 150, 97–106.
Inheritance of reduced carbon isotope discrimination in bread wheat (Triticum aestivum L).Crossref | GoogleScholarGoogle Scholar |

Richards RA, Watt M, Rebetzke GJ (2007) Physiological traits and cereal germplasm for sustainable agricultural systems. Euphytica 154, 409–425.
Physiological traits and cereal germplasm for sustainable agricultural systems.Crossref | GoogleScholarGoogle Scholar |

Richards RA, Rebetzke GJ, Watt M, Condon AG, Spielmeyer W, Dolferus R (2010) Breeding for improved water productivity in temperate cereals: phenotyping, quantitative trait loci, markers and the selection environment. Functional Plant Biology 37, 85–97.
Breeding for improved water productivity in temperate cereals: phenotyping, quantitative trait loci, markers and the selection environment.Crossref | GoogleScholarGoogle Scholar |

Robinson D (2004) Scaling the depths: below-ground allocation in plants, forests and biomes. Functional Ecology 18, 290–295.
Scaling the depths: below-ground allocation in plants, forests and biomes.Crossref | GoogleScholarGoogle Scholar |

Ryser P (1998) Intra- and interspecific variation in root length, root turnover and the underlying parameters In ‘Inherent variation in plant growth, physiological mechanisms and ecological consequences’. (Eds H Lambers, H Poorter, MMI van Vuuren) pp. 441–465. (Backhuys Publishers: Leiden)

Ryser P (2006) The mysterious root length. Plant and Soil 286, 1–6.
The mysterious root length.Crossref | GoogleScholarGoogle Scholar |

Sacks MM, Silk WK, Burman P (1997) Effect of water stress on cortical cell division rates within the apical meristem of primary roots of maize. Plant Physiology 114, 519–527.

Schonfeld MA, Johnson RC, Carver BF, Mornhinweg DW (1988) Water relation in winter wheat as drought resistance indicators. Crop Science 28, 526–531.
Water relation in winter wheat as drought resistance indicators.Crossref | GoogleScholarGoogle Scholar |

Schraut D, Ullrich CI, Hartung W (2004) Lateral ABA transport in maize roots (Zea mays): visualization by immunolocalization. Journal of Experimental Botany 55, 1635–1641.
Lateral ABA transport in maize roots (Zea mays): visualization by immunolocalization.Crossref | GoogleScholarGoogle Scholar |

Sharp RE, Davies WJ (1979) Solute regulation and growth by roots and shoots of water-stressed maize plants. Planta 147, 43–49.
Solute regulation and growth by roots and shoots of water-stressed maize plants.Crossref | GoogleScholarGoogle Scholar |

Sharp RE, LeNoble ME (2002) ABA, ethylene and the control of shoot and root growth under water stress. Journal of Experimental Botany 53, 33–37.
ABA, ethylene and the control of shoot and root growth under water stress.Crossref | GoogleScholarGoogle Scholar |

Sheshshayee MS, Bindumadhava H, Ramesh R, Prasad TG, Lakshminarayana MR, Udayakumar M (2005) Oxygen isotope enrichment (Delta O-18) as a measure of time-averaged transpiration rate. Journal of Experimental Botany 56, 3033–3039.
Oxygen isotope enrichment (Delta O-18) as a measure of time-averaged transpiration rate.Crossref | GoogleScholarGoogle Scholar |

Slafer GA, Araus JL (2007) Physiological traits for improving wheat yield under a wide range of conditions. In ‘Proceedings of the Frontis Workshop on scale and complexity in plant systems research: gene–plant–crop relations’. (Eds JHJ Spiertz, PC Struik, HH van Laar) pp. 147–156. (Springer: Dordecht, The Netherlands)

Song L, Zhang D-W, Li F-M, Fan X-W, Ma Q, Turner NC (2010) Soil water availability alters the inter- and intra-cultivar competition of three spring wheat cultivars bred in different eras. Journal Agronomy & Crop Science 196, 323–335.
Soil water availability alters the inter- and intra-cultivar competition of three spring wheat cultivars bred in different eras.Crossref | GoogleScholarGoogle Scholar |

Tambussi EA, Bort J, Araus JL (2007) Water use efficiency in C3 cereals under Mediterranean conditions: a review of physiological aspects. Annals of Applied Biology 150, 307–321.
Water use efficiency in C3 cereals under Mediterranean conditions: a review of physiological aspects.Crossref | GoogleScholarGoogle Scholar |

Tcherkez G, Farquhar GD (2008) On the effect of heavy water (D2O) on carbon isotope fractionation in photosynthesis. Functional Plant Biology 35, 201–212.
On the effect of heavy water (D2O) on carbon isotope fractionation in photosynthesis.Crossref | GoogleScholarGoogle Scholar |

Wang XF, Yakir D (1995) Temporal and spatial variations in the oxygen-18 content of leaf water in different plant species. Plant, Cell & Environment 18, 1377–1385.
Temporal and spatial variations in the oxygen-18 content of leaf water in different plant species.Crossref | GoogleScholarGoogle Scholar |

Watt M, Kirkegaard JA, Rebetzke GJ (2005) A wheat genotype developed for rapid leaf growth copes well with the physical and biological constraints of unploughed soil. Functional Plant Biology 32, 695–706.
A wheat genotype developed for rapid leaf growth copes well with the physical and biological constraints of unploughed soil.Crossref | GoogleScholarGoogle Scholar |

White JW, Castillo JA, Ehleringer JR (1990) Associations between productivity, root growth and carbon isotope discrimination in Phaseolus vulgaris under water deficit. Australian Journal of Plant Physiology 17, 189–198.
Associations between productivity, root growth and carbon isotope discrimination in Phaseolus vulgaris under water deficit.Crossref | GoogleScholarGoogle Scholar |

Yousfi S, Serret MD, Araus JL (2009) Shoot δ15N gives a better indication than ion concentration or Δ13C of genotypic differences in the response of durum wheat to salinity. Functional Plant Biology 36, 144–155.
Shoot δ15N gives a better indication than ion concentration or Δ13C of genotypic differences in the response of durum wheat to salinity.Crossref | GoogleScholarGoogle Scholar |

Yousfi S, Serret MD, Márquez AJ, Voltas J, Araus JL (2012) Combined use of δ13C, δ18O and δ15N tracks nitrogen metabolism and genotypic adaptation of durum wheat to salinity and water deficit. New Phytologist 194, 230–244.
Combined use of δ13C, δ18O and δ15N tracks nitrogen metabolism and genotypic adaptation of durum wheat to salinity and water deficit.Crossref | GoogleScholarGoogle Scholar |

Zaman-Allah M, Jenkinson DM, Vadez V (2011) A conservative pattern of water use, rather than deep or profuse rooting, is critical for the terminal drought tolerance of chickpea. Journal of Experimental Botany 62, 4239–4252.
A conservative pattern of water use, rather than deep or profuse rooting, is critical for the terminal drought tolerance of chickpea.Crossref | GoogleScholarGoogle Scholar |

Zhang DY (1995) Analysis of growth redundancy of crop root system in semi-arid area. Acta Botanica Boreali-Occidentalia Sinaca 15, 110–114.

Zhang X, Chen S, Sun H, Wang Y, Shao L (2009) Root size, distribution and soil water depletion as affected by cultivars and environmental factors. Field Crops Research 114, 75–83.
Root size, distribution and soil water depletion as affected by cultivars and environmental factors.Crossref | GoogleScholarGoogle Scholar |