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Engineering >> 2018, Volume 4, Issue 4 doi: 10.1016/j.eng.2018.06.005

Developing Wheat for Improved Yield and Adaptation Under a Changing Climate: Optimization of a Few Key Genes

a School of Veterinary and Life Sciences, Murdoch University, Perth, WA 6150, Australia

b Edstar Genetics Pty. Ltd., Perth, WA 6150, Australia

Received: 2017-04-26 Revised: 2017-08-25 Accepted: 2017-11-20 Available online: 2018-07-04

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Abstract

Wheat grown under rain-fed conditions is often affected by drought worldwide. Future projections from a climate simulation model predict that the combined effects of increasing temperature and changing rainfall patterns will aggravate this drought scenario and may significantly reduce wheat yields unless appropriate varieties are adopted. Wheat is adapted to a wide range of environments due to the diversity in its phenology genes. Wheat phenology offers the opportunity to fight against drought by modifying crop developmental phases according to water availability in target environments. This review summarizes recent advances in wheat phenology research, including vernalization (Vrn), photoperiod (Ppd), and also dwarfing (Rht) genes. The alleles, haplotypes, and copy number variation identified for Vrn and Ppd genes respond differently in different climatic conditions, and thus could alter not only the development phases but also the yield. Compared with the model plant Arabidopsis, more phenology genes have not yet been identified in wheat; quantifying their effects in target environments would benefit the breeding of wheat for improved drought tolerance. Hence, there is scope to maximize yields in water-limited environments by deploying appropriate phenology gene combinations along with Rht genes and other important physiological traits that are associated with drought resistance.

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References

[ 1 ] Braun HJ, Atlin G, Payne T. Multi-location testing as a tool to identify plant response to global climate change. In: Reynolds MP, editor. Climate change and crop production. Wallingford: Center for Agriculture and Biosciences International; 2010. p. 115–38.

[ 2 ] FAOSTAT database collections [Internet]. Rome: Food and Agriculture Organization of the United Nations; c2017 [cited 2015 Oct 23]. link1

[ 3 ] Boyer JS. Plant productivity and environment. Science 1982;218 (4571):443–8. link1

[ 4 ] AQUASTAT main database (2015) [Internet]. Rome: Food and Agriculture Organization of the United Nations; c2017 [cited 2015 Oct 22]. link1

[ 5 ] Zaynali Nezhad K, Weber WE, Röder MS, Sharma S, Lohwasser U, Meyer RC, et al. QTL analysis for thousand-grain weight under terminal drought stress in bread wheat (Triticum aestivum L.). Euphytica 2012;186(1):127–38. link1

[ 6 ] Byerlee D, Morris M. Research for marginal environments: are we underinvested? Food Policy 1993;18(5):381–93. link1

[ 7 ] Morris ML, Belaid A, Byerlee D. Part 1: wheat and barley production in rainfed marginal environments of the developing world. In: 1990–91 CIMMYT world wheat facts and trends: wheat and barley production in rainfed marginal environments of the developing world. Mexico: International Maize and Wheat Improvement Center; 1991. p. 1–28. link1

[ 8 ] Lobell DB, Gourdji SM. The influence of climate change on global crop productivity. Plant Physiol 2012;160(4):1686–97. link1

[ 9 ] Intergovernmental Panel on Climate Change. Summary for policymakers. In: Field CB, Barros VR, Dokken DJ, Mach KJ, Mastrandrea MD, Bilir TE, et al. editors. Climate change 2014: impacts, adaptation, and vulnerability. Part A: global and sectoral aspects. Contribution of Working Group II to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge: Cambridge University Press; 2014. p. 1–32.

[10] Nelson GC, Rosegrant MW, Koo J, Robertson R, Sulser T, Zhu T, et al. Climate change: impact on agriculture and costs of adaptation. Washington, DC: International Food Policy Research Institute; 2009.

[11] Reynolds MP, editor. Climate change and crop production. Wallingford: Center for Agriculture and Biosciences International; 2010.

[12] Turner NC. Sustainable production of crops and pastures under drought in a Mediterranean environment. Ann Appl Biol 2004;144(2):139–47. link1

[13] Nachit MM. Durum breeding research to improve dryland productivity in the Mediterranean region. In: Nachit MM, Baum M, Porceddu E, Monneveux P, Picard E, editors. SEWANA (South Europe, West Asia and North Africa) durum research network: Proceedings of the SEWANA Durum Network Workshop; 1995 Mar 20–23; Aleppo, Syria. Aleppo: International Center for Agricultural Research in the Dry Areas; 1998. p. 1–15. link1

[14] Lopez CG, Banowetz GM, Peterson CJ, Kronstad WE. Dehydrin expression and drought tolerance in seven wheat cultivars. Crop Sci 2003;43 (2):577–82. link1

[15] Serraj R, Hash CT, Rizvi SMH, Sharma A, Yadav RS, Bidinger FR. Recent advances in marker-assisted selection for drought tolerance in pearl millet. Plant Prod Sci 2005;8(3):334–7. link1

[16] Slafer GA, Abeledo LG, Miralles DJ, Gonzalez FG, Whitechurch EM. Photoperiod sensitivity during stem elongation as an avenue to raise potential yield in wheat. Euphytica 2001;119(1–2):191–7. link1

[17] Whitechurch EM, Slafer GA. Contrasting Ppd alleles in wheat: effects on sensitivity to photoperiod in different phases. Field Crops Res 2002;73(2– 3):95–105. link1

[18] Saini HS, Westgate ME. Reproductive development in grain crops during drought. Adv Agron 1999;68:59–96. link1

[19] Richards RA. Crop improvement for temperate Australia: future opportunities. Field Crops Res 1991;26(2):141–69. link1

[20] Worland AJ. The influence of flowering time genes on environmental adaptability in European wheats. Euphytica 1996;89(1):49–57. link1

[21] Debaeke P. Scenario analysis for cereal management in water-limited conditions by the means of a crop simulation model (STICS). Agronomie 2004;24(6–7):315–26. link1

[22] Cockram J, Jones H, Leigh FJ, O’Sullivan D, Powell W, Laurie DA, et al. Control of flowering time in temperate cereals: genes, domestication, and sustainable productivity. J Exp Bot 2007;58(6):1231–44. link1

[23] Barrett B, Bayram M, Kidwell K, Weber WE. Identifying AFLP and microsatellite markers for vernalization response gene Vrn-B1 in hexaploid wheat using reciprocal mapping populations. Plant Breed 2002;121 (5):400–6. link1

[24] Acevedo E. Assessing crop and plant attributes for cereal improvement in water-limited Mediterranean environments. In: Srivastava JP, Porceddu E, Acevedo E, Varma S, editors. Drought tolerance in winter cereals. Chichester: John Wiley and Sons; 1987. p. 303–20. link1

[25] Reynolds M, Foulkes MJ, Slafer GA, Berry P, Parry MA, Snape JW, et al. Raising yield potential in wheat. J Exp Bot 2009;60(7):1899–918. link1

[26] Curtis BC, Rajaram S, Gómez Macpherson H, editors. Bread wheat: improvement and production. Rome: Food and Agriculture Organization of the United Nations; 2002. link1

[27] Kato K, Yamagata H. Method for evaluation of chilling requirement and narrow-sense earliness of wheat cultivars. Jpn J Breed 1988;38(2):172–86. Japanese. link1

[28] Trevaskis B. The central role of the VERNALIZATION1 gene in the vernalization response of cereals. Funct Plant Biol 2010;37(6):479–87. link1

[29] Le Gouis J, Bordes J, Ravel C, Heumez E, Faure S, Praud S, et al. Genome-wide association analysis to identify chromosomal regions determining components of earliness in wheat. Theor Appl Genet 2012;124(3):597–611. link1

[30] Pugsley AT. A genetic analysis of the spring-winter habit of growth in wheat. Aust J Agric Res 1971;22(1):21–31. link1

[31] Dubcovsky J, Lijavetzky D, Appendino L, Tranquilli G. Comparative RFLP mapping of Triticum monococcum genes controlling vernalization requirement. Theor Appl Genet 1998;97(5–6):968–75. link1

[32] Yan L, Fu D, Li C, Blechl A, Tranquilli G, Bonafede M, et al. The wheat and barley vernalization gene VRN3 is an orthologue of FT. Proc Natl Acad Sci USA 2006;103(51):19581–6. link1

[33] Law CN, Worland AJ, Giorgi B. The genetic control of ear-emergence time by chromosomes 5A and 5D of wheat. Heredity 1976;36(1):49–58 link1

[34] Galiba G, Quarrie SA, Sutka J, Morgounov A, Snape JW. RFLP mapping of the vernalization (Vrn1) and frost resistance (Fr1) genes on chromosome 5A of wheat. Theor Appl Genet 1995;90(7–8):1174–9. link1

[35] Law CN, Wolfe MS. Location of genetic factors for mildew resistance and ear emergence time on chromosome 7B of wheat. Can J Genet Cytol 1966;8 (3):462–70. link1

[36] Yan L, Loukoianov A, Tranquilli G, Helguera M, Fahima T, Dubcovsky J. Positional cloning of the wheat vernalization gene VRN1. Proc Natl Acad Sci USA 2003;100(10):6263–8. link1

[37] Yan L, Loukoianov A, Blechl A, Tranquilli G, Ramakrishna W, SanMiguel P, et al. The wheat VRN2 gene is a flowering repressor down-regulated by vernalization. Science 2004;303(5664):1640–4. link1

[38] Stelmakh AF. Genetic effects of the Vrn1-3 loci and specific action of the dominant Vrn3 allele in common bread wheat. Cytol Genet 1987;21 (4):278–86. Russian.

[39] Tranquilli G, Dubcovsky J. Epistatic interaction between vernalization genes Vrn-Am1 and Vrn-Am2 in diploid wheat. J Hered 2000;91(4):304–6. link1

[40] Trevaskis B, Hemming MN, Dennis ES, Peacock WJ. The molecular basis of vernalization-induced flowering in cereals. Trends Plant Sci 2007;12 (8):352–7. link1

[41] Tsunewaki K, Jenkins BC. Monosomic and conventional gene analyses in common wheat. II. Growth habit and awnedness. J Genet 1961;36(11– 12):428–43. Japanese. link1

[42] Roberts DMA, MacDonald MD. Evidence for the multiplicity of alleles at Vrn1, the winter–spring habit locus in common wheat. Can J Genet Cytol 1984;26 (2):191–3. link1

[43] Koval SF, Goncharov NP. Multiple allelism at the VRN1 locus of common wheat. Acta Agron Hung 1998;46(2):113–9. link1

[44] Yan L, Helguera M, Kato K, Fukuyama S, Sherman J, Dubcovsky J. Allelic variation at the VRN-1 promoter region in polyploid wheat. Theor Appl Genet 2004;109(8):1677–86. link1

[45] Fu D, Szu}cs P, Yan L, Helguera M, Skinner JS, von Zitzewitz J, et al. Large deletions within the first intron in VRN-1 are associated with spring growth habit in barley and wheat. Mol Genet Genomics 2005;273(1):54–65. link1

[46] Santra DK, Santra M, Allan RE, Campbell KG, Kidwell KK. Genetic and molecular characterization of vernalization genes Vrn-A1, Vrn-B1, and Vrn-D1 in spring wheat germplasm from the Pacific Northwest region of the USA. Plant Breed 2009;128(6):576–84. link1

[47] Milec Z, Tomková L, Sumíková T, Pánková K. A new multiplex PCR test for the determination of Vrn-B1 alleles in bread wheat (Triticum aestivum L.). Mol Breed 2012;30(1):317–23. link1

[48] Zhang J, Wang Y, Wu S, Yang J, Liu H, Zhou Y. A single nucleotide polymorphism at the Vrn-D1 promoter region in common wheat is associated with vernalization response. Theor Appl Genet 2012;125 (8):1697–704. link1

[49] Nishida H, Yoshida T, Kawakami K, Fujita M, Long B, Akashi Y, et al. Structural variation in the 50 upstream region of photoperiod-insensitive alleles Ppd-A1a and Ppd-B1a identified in hexaploid wheat (Triticum aestivum L.), and their effect on heading time. Mol Breed 2013;31(1):27–37. link1

[50] Wilhelm EP, Turner AS, Laurie DA. Photoperiod insensitive Ppd-A1a mutations in tetraploid wheat (Triticum durum Desf.). Theor Appl Genet 2009;118 (2):285–94. link1

[51] Muterko A, Kalendar R, Cockram J, Balashova I. Discovery, evaluation and distribution of haplotypes and new alleles of the Photoperiod-A1 gene in wheat. Plant Mol Biol 2015;88(1–2):149–64. link1

[52] Díaz A, Zikhali M, Turner AS, Isaac P, Laurie DA. Copy number variation affecting the Photoperiod-B1 and Vernalization-A1 genes is associated with altered flowering time in wheat (Triticum aestivum). PLoS One 2012;7(3). link1

[53] Cane K, Eagles HA, Laurie DA, Trevaskis B, Vallance N, Eastwood RF, et al. Ppd- B1 and Ppd-D1 and their effects in southern Australian wheat. Crop Pasture Sci 2013;64(2):100–14. link1

[54] Beales J, Turner A, Griffiths S, Snape JW, Laurie DA. A pseudo-response regulator is misexpressed in the photoperiod insensitive Ppd-D1a mutant of wheat (Triticum aestivum L.). Theor Appl Genet 2007;115(5):721–33. link1

[55] Guo Z, Song Y, Zhou R, Ren Z, Jia J. Discovery, evaluation and distribution of haplotypes of the wheat Ppd-D1 gene. New Phytol 2010;185(3):841–51. link1

[56] Gotoh T. Genetic studies on growth habit of some important spring wheat cultivars in Japan, with special reference to the identification of the spring genes involved Japanese. Jpn J Breed 1979;29(2):133–45. link1

[57] Stelmakh A. Geographic distribution of Vrn-genes in landraces and improved varieties of spring bread wheat. Euphytica 1990;45(2):113–8. link1

[58] Goncharov NP. Genetic resources of wheat related species: The Vrn genes controlling growth habit (spring vs. winter). Euphytica 1998;100(1– 3):371–6. link1

[59] Stelmakh AF. Genetic systems regulating flowering response in wheat. Euphytica 1998;100(1–3):359–69. link1

[60] Iwaki K, Nakagawa K, Kuno H, Kato K. Ecogeographical differentiation in East Asian wheat, revealed from the geographical variation of growth habit and Vrn genotype. Euphytica 2000;111(2):137–43. link1

[61] Iwaki K, Haruna S, Niwa T, Kato K. Adaptation and ecological differentiation in wheat with special reference to geographical variation of growth habit and Vrn genotype. Plant Breed 2001;120(2):107–14. link1

[62] Trevaskis B, Bagnall DJ, Ellis MH, Peacock WJ, Dennis ES. MADS-box genes control vernalization-induced flowering in cereals. Proc Natl Acad Sci USA 2003;100(22):13099–104. link1

[63] Danyluk J, Kane NA, Breton G, Limin AE, Fowler DB, Sarhan F. TaVRT-1, a putative transcription factor associated with vegetative to reproductive transition in cereals. Plant Physiol 2003;132(4):1849–60. link1

[64] Szu}cs P, Skinner JS, Karsai I, Cuesta-Marcos A, Haggard KG, Corey AE, et al. Validation of the VRN-H2/VRN-H1 epistatic model in barley reveals that intron length variation in VRN-H1 may account for a continuum of vernalization sensitivity. Mol Genet Genomics 2007;277(3):249–61. link1

[65] Hemming MN, Fieg S, James Peacock W, Dennis ES, Trevaskis B. Regions associated with repression of the barley (Hordeum vulgare) VERNALIZATION1 gene are not required for cold induction. Mol Genet Genomics 2009;282 (2):107–17. link1

[66] McIntosh RA, Hart GE, Devos KM, Gale MD, Rogers WJ. Catalogue of gene symbols for wheat. In: Slinkard AE, editor. Proceedings of the 9th International Wheat Genetics Symposium; 1998 Aug 2–7. Saskatoon, SK, Canada: Saskatoon: University of Saskatchewan; 1998. p. 123–72. link1

[67] Goncharov NP. Genetics of growth habit (spring vs winter) in common wheat: confirmation of the existence of dominant gene Vrn4. Theor Appl Genet 2003;107(4):768–72. link1

[68] Foulkes MJ, Sylvester-Bradley R, Worland AJ, Snape JW. Effects of a photoperiod-response gene Ppd-D1 on yield potential and drought resistance in UK winter wheat. Euphytica 2004;135(1):63–73. link1

[69] Kumar S, Sharma V, Chaudhary S, Tyagi A, Mishra P, Priyadarshini A, et al. Genetics of flowering time in bread wheat Triticum aestivum: complementary interaction between vernalization-insensitive and photoperiod-insensitive mutations imparts very early flowering habit to spring wheat. J Genet 2012;91(1):33–47. link1

[70] Welsh JR, Keim DL, Piratesh B, Richards RD. Genetic control of photoperiod response in wheat. In: Sears ER, Sears LMS, editors. Proceedings of the fourth international wheat genetics symposium; 1973 Aug 6–11; Columbia, MO, USA. Columbia: Agricultural Experiment Station, College of Agriculture, University of Missouri; 1973. p. 879–84. link1

[71] Law CN, Sutka J, Worland AJ. A genetic study of day-length response in wheat. Heredity 1978;41(2):185–91. link1

[72] Börner A, Worland AJ, Plaschke J, Schumann E, Law CN. Pleiotropic effects of genes for reduced height (Rht) and day-length insensitivity (Ppd) on yield and its components for wheat grown in middle Europe. Plant Breed 1993;111 (3):204–16. link1

[73] Worland AJ, Börner A, Korzun V, Li WM, Petrovíc S, Sayers EJ. The influence of photoperiod genes on the adaptability of European winter wheats. Euphytica 1998;100(1–3):385–94. link1

[74] Snape JW, Butterworth K, Whitechurch E, Worland AJ. Waiting for fine times: genetics of flowering time in wheat. Euphytica 2001;119(1–2): 185–90. link1

[75] Takenaka S, Kawahara T. Evolution and dispersal of emmer wheat (Triticum sp.) from novel haplotypes of Ppd-1 (photoperiod response) genes and their surrounding DNA sequences. Theor Appl Genet 2012;125 (5):999–1014. link1

[76] Chen Y, Carver BF, Wang S, Cao S, Yan L. Genetic regulation of developmental phases in winter wheat. Mol Breed 2010;26(4):573–82. link1

[77] Law CN, Suarez E, Miller TE, Worland AJ. The influence of the group 1 chromosomes of wheat on ear-emergence times and their involvement with vernalization and day length. Heredity 1998;80(1):83–91. link1

[78] Kulwal PL, Roy JK, Balyan HS, Gupta PK. QTL mapping for growth and leaf characters in bread wheat. Plant Sci 2003;164(2):267–77. link1

[79] Tóth B, Galiba G, Fehér E, Sutka J, Snape JW. Mapping genes affecting flowering time and frost resistance on chromosome 5B of wheat. Theor Appl Genet 2003;107(3):509–14. link1

[80] Hanocq E, Laperche A, Jaminon O, Lainé AL, Le Gouis J. Most significant genome regions involved in the control of earliness traits in bread wheat, as revealed by QTL meta-analysis. Theor Appl Genet 2007;114(3):569–84. link1

[81] Griffiths S, Simmonds J, Leverington M, Wang Y, Fish L, Sayers L, et al. Meta- QTL analysis of the genetic control of ear emergence in elite European winter wheat germplasm. Theor Appl Genet 2009;119(3):383–95. link1

[82] Hoogendoorn J. A reciprocal F1 monosomic analysis of the genetic control of time of ear emergence, number of leaves and number of spikelets in wheat (Triticum aestivum L.). Euphytica 1985;34(2):545–58. link1

[83] Chen F, Gao M, Zhang J, Zuo A, Shang X, Cui D. Molecular characterization of vernalization and response genes in bread wheat from the Yellow and Huai Valley of China. BMC Plant Biol 2013;13:199. link1

[84] Eagles HA, Cane K, Trevaskis B. Veery wheats carry an allele of Vrn-A1 that has implications for freezing tolerance in winter wheats. Plant Breed 2011;130 (4):413–8. link1

[85] Eagles HA, Cane K, Vallance N. The flow of alleles of important photoperiod and vernalisation genes through Australian wheat. Crop Pasture Sci 2009;60 (7):646–57. link1

[86] Iqbal M, Navabi A, Yang RC, Salmon DF, Spaner D. Molecular characterization of vernalization response genes in Canadian spring wheat. Genome 2007;50 (5):511–6. link1

[87] Dubcovsky J, Loukoianov A, Fu D, Valarik M, Sanchez A, Yan L. Effect of photoperiod on the regulation of wheat vernalization genes VRN1 and VRN2. Plant Mol Biol 2006;60(4):469–80. link1

[88] Distelfeld A, Dubcovsky J. Characterization of the maintained vegetative phase deletions from diploid wheat and their effect on VRN2 and FT transcript levels. Mol Genet Genomics 2010;283(3):223–32. link1

[89] Distelfeld A, Li C, Dubcovsky J. Regulation of flowering in temperate cereals. Curr Opin Plant Biol 2009;12(2):178–84. link1

[90] Shimada S, Ogawa T, Kitagawa S, Suzuki T, Ikari C, Shitsukawa N, et al. A genetic network of flowering-time genes in wheat leaves, in which an APETALA1/FRUITFULL-like gene, VRN1, is upstream of FLOWERING LOCUS T. Plant J 2009;58(4):668–81. link1

[91] Rebetzke GJ, Bonnett DG, Ellis MH. Combining gibberellic acid-sensitive and insensitive dwarfing genes in breeding of higher-yielding, sesqui-dwarf wheats. Field Crops Res 2012;127:17–25. link1

[92] Waddington SR, Ransom JK, Osmanzai M, Saunders DA. Improvement in the yield potential of bread wheat adapted to northwest Mexico. Crop Sci 1986;26(4):698–703. link1

[93] Chapman SC, Mathews KL, Trethowan RM, Singh RP. Relationships between height and yield in near-isogenic spring wheats that contrast for major reduced height genes. Euphytica 2007;157(3):391–7. link1

[94] Keyes GJ, Paolillo DJ, Sorrells ME. The effects of dwarfing genes Rht1 and Rht2 on cellular dimensions and rate of leaf elongation in wheat. Ann Bot 1989;64 (6):683–90. link1

[95] Donald CM, Puckridge DW. The ecology of the wheat crop. In: Lazenby A, Matheson EM, editors. Australian field crops. Volume 1. Wheat and other temperate cereals. Sydney: Angus and Robertson; 1975. p. 288–303. link1

[96] Allan RE. Agronomic comparisons between Rht1 and Rht2 semidwarf genes in winter wheat. Crop Sci 1989;29(5):1103–8. link1

[97] Richards RA. The effect of dwarfing genes in spring wheat in dry environments. I. Agronomic characteristics. Aust J Agric Res 1992;43 (3):517–27. link1

[98] Rebetzke GJ, Botwright TL, Moore CS, Richards RA, Condon AG. Genotypic variation in specific leaf area for genetic improvement of early vigour in wheat. Field Crops Res 2004;88(2–3):179–89. link1

[99] Botwright TL, Rebetzke GJ, Condon AG, Richards RA. Influence of the gibberellin-sensitive Rht8 dwarfing gene on leaf epidermal cell dimensions and early vigour in wheat (Triticum aestivum L.). Ann Bot 2005;95(4):631–9. link1

[100] Peng J, Richards DE, Hartley NM, Murphy GP, Devos KM, Flintham JE, et al. ‘‘Green revolution” genes encode mutant gibberellin response modulators. Nature 1999;400(6741):256–61. link1

[101] Leuning R, Condon AG, Dunin FX, Zegelin S, Denmead OT. Rainfall interception and evaporation from soil below a wheat canopy. Agric For Meteorol 1994;67(3–4):221–38. link1

[102] Siddique KHM, Tennant D, Perry MW, Belford RK. Water use and water use efficiency of old and modern wheat cultivars in a Mediterranean-type environment. Aust J Agric Res 1990;41(3):431–47. link1

[103] Regan KL, Siddique KHM, Turner NC, Whan BR. Potential for increasing early vigour and total biomass in spring wheat. II. Characteristics associated with early vigour. Aust J Agric Res 1992;43(3):541–53. link1

[104] López-Castañeda C, Richards RA. Variation in temperate cereals in rainfed environments III. Water use and water-use efficiency. Field Crops Res 1994;39(2–3):85–98. link1

[105] Doyle AD, Marcellos H. Time of sowing and wheat yield in northern New South Wales. Aust J Exp Agric Anim Husb 1974;14(66):93–102. link1

[106] Shackley BJ, Anderson WK. Responses of wheat cultivars to time of sowing in the southern wheatbelt of Western Australia. Aust J Exp Agric 1995;35 (5):579–87. link1

[107] Hadjichristodoulou A, Della A, Photiades J. Effect of sowing depth on plant establishment, tillering capacity and other agronomic characters of cereals. J Agric Sci 1977;89(1):161–7. link1

[108] Gan Y, Stobbe EH, Moes J. Relative date of wheat seedling emergence and its impact on grain yield. Crop Sci 1992;32(5):1275–81. link1

[109] Whan BR. The emergence of semidwarf and standard wheats, and its association with coleoptile length. Aust J Exp Agric Anim Husb 1976;16 (80):411–6. link1

[110] Schillinger WF, Donaldson E, Allan RE, Jones SS. Winter wheat seedling emergence from deep sowing depths. Agron J 1998;90(5):582–6. link1

[111] Richards RA, Rebetzke GJ, Condon AG, van Herwaarden AF. Breeding opportunities for increasing the efficiency of water use and crop yield in temperate cereals. Crop Sci 2002;42(1):111–21. link1

[112] Rebetzke GJ, Richards RA. Genetic improvement of early vigour in wheat. Aust J Agric Res 1999;50(3):291–302. link1

[113] Ellis MH, Rebetzke GJ, Chandler P, Bonnett D, Spielmeyer W, Richards RA. The effect of different height reducing genes on the early growth of wheat. Funct Plant Biol 2004;31(6):583–9. link1

[114] Rebetzke GJ, Richards RA, Fettell NA, Long M, Condon AG, Forrester RI, et al. Genotypic increases in coleoptile length improves stand establishment, vigour and grain yield of deep-sown wheat. Field Crops Res 2007;100 (1):10–23. link1

[115] Borrell AK, Incoll LD, Simpson RJ, Dalling MJ. Partitioning of dry matter and the deposition and use of stem reserves in a semi-dwarf wheat crop. Ann Bot 1989;63(5):527–39. link1

[116] Flintham JE, Börner A, Worland AJ, Gale MD. Optimizing wheat grain yield: effects of Rht (gibberellin-insensitive) dwarfing genes. J Agric Sci 1997;128 (1):11–25. link1

[117] Rebetzke GJ, Richards RA, Fischer VM, Mickelson BJ. Breeding long coleoptile, reduced height wheats. Euphytica 1999;106(2):159–68. link1

[118] Korzun V, Röder MS, Ganal MW, Worland AJ, Law CN. Genetic analysis of the dwarfing gene (Rht8) in wheat. Part I. Molecular mapping of Rht8 on the short arm of chromosome 2D of bread wheat (Triticum aestivum L.). Theor Appl Genet 1998;96(8):1104–9. link1

[119] Ellis MH, Bonnett DG. Rebetzke GJ. A 192bp allele at the Xgwm261 locus is not always associated with the Rht8 dwarfing gene in wheat (Triticum aestivum L.). Euphytica 2007;157(1–2):209–14. link1

[120] Slafer GA, Kantolic AG, Appendino ML, Miralles DJ, Savin R. Crop development: genetic control, environmental modulation and relevance for genetic improvement of crop yield. In: Sadras VO, Calderini DF, editors. Crop physiology: applications for genetic improvement and agronomy. Amsterdam: Elsevier Inc.; 2009. p. 277–308.

[121] Fischer RA. Number of kernels in wheat crops and the influence of solar radiation and temperature. J Agric Sci 1985;105(2):447–61. link1

[122] Passioura JB. Grain yield, harvest index, and water use of wheat. J Aust Inst Agric Sci 1977;43:117–20. link1

[123] Jamieson PD, Francis GS, Wilson DR, Martin RJ. Effects of water deficits on evapotranspiration from barley. Agric For Meteorol 1995;76(1):41–58. link1

[124] Araus JL, Slafer GA, Reynolds MP, Royo C. Plant breeding and drought in C3 cereals: what should we breed for? Ann Bot 2002;89(7):925–40. link1

[125] Roberts DWA. Identification of loci on chromosome 5A of wheat involved in control of cold hardiness, vernalization, leaf length, rosette growth habit, and height of hardened plants. Genome 1990;33(2):247–59. link1

[126] Borràs-Gelonch G, Rebetzke GJ, Richards RA. Romagosa I. Genetic control of duration of pre-anthesis phases in wheat (Triticum aestivum L.) and relationships to leaf appearance, tillering, and dry matter accumulation. J Exp Bot 2012;63(1):69–89. link1

[127] Eagles HA, Cane K, Kuchel H, Hollamby GJ, Vallance N, Eastwood RF, et al. Photoperiod and vernalization gene effects in southern Australian wheat. Crop Pasture Sci 2010;61(9):721–30. link1

[128] Eagles HA, Cane K, Trevaskis B, Vallance N, Eastwood RF, Gororo NN, et al. Ppd1, Vrn1, ALMT1 and Rht genes and their effects on grain yield in lower rainfall environments in southern Australia. Crop Pasture Sci 2014;65 (2):159–70. link1

[129] Rebetzke GJ, Rattey AR, Farquhar GD, Richards RA, Condon ATG. Genomic regions for canopy temperature and their genetic association with stomatal conductance and grain yield in wheat. Funct Plant Biol 2013;40(1):14–33. link1

[130] Asana RD. Physiological analysis of yield of wheat in relation to water-stress and temperature. J Post-Grad Sch Indian Agric Res Inst 1966;4:17–31.

[131] Brooks A, Jenner CF, Aspinall D. Effects of water deficit on endosperm starch granules and on grain physiology of wheat and barley. Aust J Plant Physiol 1982;9(4):423–36. link1

[132] Aggarwal PK, Sinha SK. Effect of water stress on grain growth and assimilate partitioning in two cultivars of wheat contrasting in their yield stability in a drought-environment. Ann Bot 1984;53(3):329–40. link1

[133] Blum A. Effective use of water (EUW) and not water-use efficiency (WUE) is the target of crop yield improvement under drought stress. Field Crops Res 2009;112(2–3):119–23. link1

[134] Blum A, Shpiler L, Golan G, Mayer J, Sinmena B. Mass selection of wheat for grain filling without transient photosynthesis. Euphytica 1991;54(1):111–6. link1

[135] Kiniry JR. Nonstructural carbohydrate utilization by wheat shaded during grain growth. Agron J 1993;85(4):844–9. link1

[136] Schnyder H. The role of carbohydrate storage and redistribution in the source–sink relations of wheat and barley during grain filling—a review. New Phytol 1993;123(2):233–45. link1

[137] Borrell AK, Incoll LD, Dalling MJ. The influence of the Rht1 and Rht2 alleles on the deposition and use of stem reserves in wheat 317 26. Ann Bot 1993;71(4). link1

[138] Izanloo A, Condon AG, Langridge P, Tester M, Schnurbusch T. Different mechanisms of adaptation to cyclic water stress in two South Australian bread wheat cultivars. J Exp Bot 2008;59(12):3327–46. link1

[139] Mir RR, Zaman-Allah M, Sreenivasulu N, Trethowan R, Varshney RK. Integrated genomics, physiology and breeding approaches for improving drought tolerance in crops. Theor Appl Genet 2012;125(4):625–45. link1

[140] Reynolds M, Foulkes J, Furbank R, Griffiths S, King J, Murchie E, et al. Achieving yield gains in wheat. Plant Cell Environ 2012;35(10):1799–823. link1

[141] Law CN, Worland AJ. Genetic analysis of some flowering time and adaptive traits in wheat. New Phytol 1997;137(1):19–28. link1

[142] Servin B, Martin OC, Mézard M, Hospital F. Toward a theory of marker- assisted gene pyramiding. Genetics 2004;168(1):513–23. link1

[143] Mouradov A, Cremer F, Coupland G. Control of flowering time: Interacting pathways as a basis for diversity. Plant Cell 2002;14(Suppl 1):S111–30. link1

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