Crop and Pasture Science Crop and Pasture Science Society
Plant sciences, sustainable farming systems and food quality

Quantitative trait loci for water-soluble carbohydrates and associations with agronomic traits in wheat

G. J. Rebetzke A C E , A. F. van Herwaarden B , C. Jenkins A C , M. Weiss A , D. Lewis C , S. Ruuska A C , L. Tabe A C , N. A. Fettell D and R. A. Richards A C

A CSIRO Plant Industry, GPO Box 1600, Canberra, ACT 2601, Australia.

B CSIRO Plant Industry, 306 Carmody Rd, St Lucia, Qld 4067, Australia.

C GrainGene, PO Box 1600, Canberra, ACT 2601, Australia.

D NSW Department of Primary Industries, Condobolin, NSW 2877, Australia.

E Corresponding author. Email:

Australian Journal of Agricultural Research 59(10) 891-905
Submitted: 20 February 2008  Accepted: 23 June 2008   Published: 18 September 2008


Several environmental factors including drought and disease can reduce leaf area and photosynthesis during grain-filling to decrease grain yield and kernel weight of cereal crops. Water-soluble carbohydrates (WSC) accumulated around anthesis can be mobilised to assist in filling of developing grains when post-anthesis assimilation is low. Cultivar differences support opportunities to select for high WSC but little is known of the extent or nature of genetic control for this trait in wheat. Three wheat mapping populations (Cranbrook/Halberd, Sunco/Tasman, and CD87/Katepwa) were phenotyped for WSC and other agronomic traits across multiple environments. The range for WSC concentration (WSC-C) was large among progeny contributing to moderate-to-high narrow-sense heritabilities within environments (h2 = 0.51–0.77). Modest genotype × environment interaction reduced the correlation of genotype means across environments (rp = 0.37–0.78, P < 0.01) to reduce heritability on a line-mean (h2 = 0.55–0.87) basis. Transgressive segregation was large and genetic control complex, with 7–16 QTLs being identified for WSC-C in each population. Heritability was smaller (h2 = 0.32–0.54) for WSC mass per unit area (WSC-A), reflecting large genotype × environment interaction and residual variance with estimating anthesis biomass. Fewer significant QTLs (4–8) were identified for this trait in each population, while sizes of individual genetic effects varied between populations but were repeatable across environments. Several genomic regions were common across populations including those associated with plant height (e.g. Rht-B1) and/or anthesis date (e.g. Ppd1). Genotypes with high WSC-C were commonly shorter, flowered earlier, and produced significantly (P < 0.01) fewer tillers than those of low WSC-C. This resulted in similar yields, lower final biomass, and fewer grains per m2, but greater dry weight partitioning to grain, kernel weight, and less grain screenings in high compared with low WSC-C genotypes. By contrast, lines high for WSC-A produced more fertile tillers associated with similar or greater anthesis and maturity biomass, grain number, and yield, yet similar kernel weight or size compared with genotypes with low WSC-A. The data support an important role for WSC-A in assuring stable yield and grain size. However, the small effects of many independent WSC QTLs may limit their direct use for marker-aided selection in breeding programs. We suggest using molecular markers to enrich populations for favourable height and anthesis date alleles before the more costly phenotypic selection among partially inbred families for greater WSC-A.

Additional keywords: screenings, QTL, kernel size, fructan, non-structural and stem carbohydrates, nitrogen, breeding.


Asseng S van Herwaarden AF 2003 Analysis of the benefits to wheat yield from assimilates stored prior to grain filling in a range of environments. Plant and Soil 256 217 229 doi:10.1023/A:1026231904221

Austin RB Morgan CL Ford MA Blackwell RD 1980 Contributions to grain yield from pre-anthesis assimilation in tall and dwarf barley phenotypes in two contrasting seasons. Annals of Botany 45 309 319

Batten GD Blakeney AB McGrath VB Ciavarella S 1993 Non-structural carbohydrate: analysis by near infrared reflectance spectroscopy and its importance as an indicator of plant growth. Plant and Soil 155-156 243 246

Bell CJ Incoll LD 1990 The redistribution of assimilate in field-grown winter wheat. Journal of Experimental Botany 41 949 960 doi:10.1093/jxb/41.8.949

Bidinger F Musgrave RB Fischer RA 1977 Contribution of stored pre-anthesis assimilation to grain yield in wheat and barley. Nature 270 431 433 doi:10.1038/270431a0

Blum A Sinmena B Mayer J Golan G Shpiler L 1994 Stem reserve mobilisation supports wheat-grain filling under heat stress. Australian Journal of Plant Physiology 21 771 781

Borrell AK Incoll LD Dalling MJ 1993 The influence of Rht 1 and Rht 2 alleles on the deposition and use or stem reserves in wheat. Annals of Botany 71 317 326

Chalmers KJ Campbell AW Kretschmer J Karakousis A Henschke PH Pierens S Harker N Pallotta M Cornish GB Shariflou MR Rampling LR McLauchlan A Daggard G Sharp PJ Holton TA Sutherland MW Appels R Langridge P 2001 Construction of three linkage maps in bread wheat (Triticum aestivum L.). Australian Journal of Agricultural Research 52 1089 1120 doi:10.1071/AR01081

Churchill GA Doerge RW 1994 Empirical threshold values for quantitative trait mapping. Genetics 138 963 971

Diekmann F Fischbeck G 2005 Differences in wheat cultivar response to nitrogen supply. II: Differences in N-metabolism-related traits. Journal of Agronomy & Crop Science 191 362 376

Ehdaie B Alloush GA Madore MA Waines JG 2006 Genotypic variation for stem reserves and mobilization in wheat: II. Postanthesis changes in internode water-soluble carbohydrates. Crop Science 46 2093 2103 doi:10.2135/cropsci2006.01.0013

Ehdaie B Alloush GA Waines JG 2008 Genotypic variation in linear rate of grain growth and contribution of stem reserves to yield in wheat. Field Crops Research 106 34 43 doi:10.1016/j.fcr.2007.10.012

Evans LT , Wardlaw IF , Fischer RA (1975) Wheat. In ‘Crop physiology: some case studies’. (Ed. LT Evans) pp. 101–150. (Cambridge University Press: Cambridge, UK)

Flood RG Martin PJ Gardner WK 1995 Dry matter accumulation and partitioning and its relationship to grain yield in wheat. Australian Journal of Experimental Agriculture 35 495 502 doi:10.1071/EA9950495

Foulkes MJ Scott RK Sylvester-Bradley R 2002 The ability of wheat cultivars to withstand drought in UK conditions: formation of grain yield. The Journal of Agricultural Science 138 153 169 doi:10.1017/S0021859601001836

Gebbing T Schnyder H Kuhbauch W 1999 The utilization of pre-anthesis reserves in grain filling of wheat. Assessment by steady-state 13CO2/12CO2 labelling. Plant, Cell & Environment 22 851 858 doi:10.1046/j.1365-3040.1999.00436.x

Housley TL (2000) Role of fructans redistributed from vegetative tissues in grain filling of wheat and barley. In ‘Carbohydrate reserves in plants: synthesis and regulation’. Developments in Crop Science 26. (Eds AK Gupta, N Kaur) pp. 207–221. (Elsevier: Dordrecht, The Netherlands)

Hunt LA (1979) Stem weight changes during grain filling in wheat from diverse sources. In ‘Proceedings of the 5th International Wheat Genetics Symposium’. (Ed. S Ramanujam) pp. 923–927. (Indian Society of Genetics and Plant Breeding: New Delhi)

Kammholz SJ Campbell AW Sutherland MW Hollamby G Martin P Eastwood R Barclay I Wilson R Brennan P Sheppard J 2001 Establishment and characterisation of wheat genetic mapping populations. Australian Journal of Agricultural Research 52 1079 1088 doi:10.1071/AR01043

Kiniry JR 1993 Nonstructural carbohydrate utilization by wheat shaded during grain growth. Agronomy Journal 85 844 849

Korol A (2005) ‘MultiQTL: QTL mapping software guide.’ (

Kühbauch W Thorne U 1989 Nonstructural carbohydrates of wheat stems as influenced by sink-source manipulations. Journal of Plant Physiology 134 243 250

Laperche A Brancourt-Hulmel M Heumez E Gardet O Hanocq E Devienne-Barret F Le Gouis J 2007 Using genotype × nitrogen interaction variables to evaluate the QTL involved in wheat tolerance to nitrogen constraints. Theoretical and Applied Genetics 115 399 415

Lehmensiek A Eckermann PJ Verbyla AP Appels R Sutherland MW Daggard GE 2005 Curation of wheat maps to improve map accuracy and QTL detection. Australian Journal of Agricultural Research 56 1347 1354

Littell RC , Milliken GA , Stroup WW , Wolfinger RD (1996) ‘SAS System for mixed models.’ (SAS Institute Inc.: Cary, NC)

López-Castañeda C Richards RA 1994 Variation in temperate cereals in rainfed environments II. Phasic development and growth. Field Crops Research 37 63 75

Morgan JA LeCain DR Wells R 1990 Semidwarfing genes concentrate photosynthetic machinery and affect leaf gas exchange of wheat. Crop Science 30 602 608

Nagata K Shimizu H Terao T 2002 Quantitative trait loci for non-structural carbohydrate accumulation in leaf sheaths and culms of rice (Oryza sativa L.) and their effects on grain filling. Breeding Science 52 275 283

Nicolas ME Turner NC 1993 Use of chemical desiccants and senescing agents to select wheat lines maintaining stable grain size during post-anthesis drought. Field Crops Research 31 155 171 doi:10.1016/0378-4290(93)90058-U

Paynter B , Young K (2001) Morphological traits associated with grain plumpness of barley. In ‘Proceedings of the 10th Australian Barley Technical Symposium’. Canberra, ACT. (

Rawson HM Evans LT 1971 The contribution of stem reserves to grain development in a range of wheat cultivars of different height. Australian Journal of Agricultural Research 22 851 863 doi:10.1071/AR9710851

Ruuska S Rebetzke GJ van Herwaarden A Richards RA Fettell NA Tabe L Jenkins C 2006 Genotypic variation for water soluble carbohydrate accumulation in wheat. Functional Plant Biology 33 799 809 doi:10.1071/FP06062

Salem KFM Röder MS Börner A 2007 Identification and mapping quantitative trait loci for stem reserve mobilisation in wheat (Triticum aestivum L.). Cereal Research Communications 35 1367 1374 doi:10.1556/CRC.35.2007.3.1

Schnyder H 1993 The role of carbohydrate storage and redistribution in the source-sink relations of wheat and barley during grain filling—a review. New Phytologist 123 233 245 doi:10.1111/j.1469-8137.1993.tb03731.x

Setter TL , Anderson WK , Asseng S , Barclay I (1998) Review of the impact of high shoot carbohydrate concentrations on maintenance of high yields in cereals exposed to environmental stress during grain filling. In ‘Wheat research needs beyond 2000 AD’. (Eds S Nagarajan, G Singh, B S Tyagi) pp. 237–255. (Narosa Publishing House: New Delhi)

Shearman VJ Sylvester-Bradley R Scott RK Foulkes MJ 2005 Physiological processes associated with wheat yield progress in UK. Crop Science 45 175 185

Snape JW Foulkes MJ Simmonds J Leverington M Fish LJ Wang Y Ciavarrella M 2007 Dissecting gene × environmental effects on wheat yields via QTL and physiological analysis. Euphytica 154 401 408

Takahashi T Chevalier PM Rupp RI 2001 Storage and remobilization of soluble carbohydrates after heading in different plant parts of a winter wheat cultivar. Plant Production Science 4 160 165

Takai T Fukata Y Shiraiwa T Horie T 2005 Time-related mapping of quantitative trait loci controlling grain-filling of rice (Orzya sativa L.). Journal of Experimental Botany 56 2107 2118

Turner LB Cairns AJ Armstead IP Ashton J Skøt K Whittaker D Humphreys MO 2006 Dissecting the regulation of fructan metabolism in perennial ryegrass (Lolium perenne) with quantitative trait locus mapping. New Phytologist 169 45 58 doi:10.1111/j.1469-8137.2005.01575.x

van Herwaarden AF (1996) Physiological traits for high protein dryland wheat production. In ‘Proceedings of the 8th Assembly Wheat Breeding Society of Australia’. Canberra, ACT. (Eds RA Richards, CW Wrigley, HM Rawson, GJ Rebetzke, JL Davidson, RIS Brettell) pp. 168–172. (Australian Wheat Breeding Society: Canberra)

van Herwaarden AF Angus JF Richards RA Farquhar GD 1998 ‘Haying-off’, the negative grain yield response of dryland wheat to nitrogen fertiliser II. Carbohydrate and protein dynamics. Australian Journal of Agricultural Research 49 1083 1093 doi:10.1071/A97040

van Herwaarden AF , Richards RA (2002) Water soluble carbohydrate accumulation in stems is related to breeding progress in Australia wheats. In ‘Proceedings of the 12th Australasian Plant Breeding Conference’. Perth, W. Aust. pp. 878–882. (Australasian Plant Breeding Society: Perth)

Voorrips RE 2002 MapChart: Software for the graphical presentation of linkage maps and QTLs. The Journal of Heredity 93 77 78 doi:10.1093/jhered/93.1.77

Wilkins PW Humphreys MO 2003 Progress in breeding perennial forage grasses for temperate agriculture. Journal of Agricultural Science (Cambridge) 140 129 150 doi:10.1017/S0021859603003058

Yang D Jing R Chang X Li W 2007 Identification of quantitative trait loci and environmental interactions for accumulation and remobilization of water-soluble carbohydrates in wheat (Triticum aestivum L.) stems. Genetics 176 571 584 doi:10.1534/genetics.106.068361

Yang J Zhang J Wang Z Zhu Q Liu L 2001 Water deficit-induced senescence and its relationship to the remobilization of pre-stored carbon in wheat during grain filling. Agronomy Journal 93 196 206

Youseffian S Kirby EJM Gale MD 1992 Pleiotropic effects of the GA-insensitive Rht dwarfing genes in wheat. II. Effects on leaf, stem, ear and floret growth. Field Crops Research 28 191 210

Zadoks JC Chang TT Konzak CF 1974 A decimal code for the growth stages of cereals. Weed Research 14 415 421

Export Citation