Abstract
Key message
The review outlines past failures, present status and future prospects of hybrid wheat, and includes information on CMS/CHA/transgenic approaches for male sterility, heterotic groups and cost-effective hybrid seed production.
Abstract
Hybrid varieties give increased yield and improved grain quality in both cross- and self-pollinated crops. However, hybrid varieties in self-pollinated crops (particularly cereals) have not been very successful, except for hybrid rice in China. In case of hybrid wheat, despite the earlier failures, renewed efforts in recent years have been made and hybrid varieties with desirable attributes have been produced and marketed in some European countries. This review builds upon previous reviews, with a new outlook and improved knowledge base, not covered in earlier reviews. New technologies have been described, which include the Hordeum chilense-based CMS–fertility restorer system, chromosomal XYZ-4E-ms system and the following transgenic technologies: (1) conditional male sterility involving use of tapetum-specific expression of a gene that converts a pro-toxin into a phytotoxin causing male sterility; (2) barnase-barstar SeedLink system of Bayer CropScience; (3) split-barnase system that obviates the need of a barstar-based male restorer line; and (4) seed production technology of DuPont-Pioneer that makes use of transgenes in production of male-sterile lines, but gives hybrid seed with no transgenes. This review also includes a brief account of studies for discovery of heterotic QTL, genomic prediction of hybrid vigour and the development of heterotic groups/patterns and their importance in hybrid wheat production. The problem of high cost of hybrid seed due to required high seed rate in wheat relative to hybrid rice has also been addressed. The review concludes with a brief account of the current efforts and future possibilities in making hybrid wheat a commercial success.
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Acknowledgements
This review had a long incubation period, during which the authors received necessary facilities from Head, Department of Genetics and Plant Breeding, CCS University (Meerut, India); HSB continued to hold the position of INSA Senior Scientist, starting in 2015; PKG was initially awarded INSA Senior Scientist position during 2015–2017 and later continued to hold the position of Hony Senior Scientist, starting in June 2017; VG was awarded the position of INSPIRE Faculty by DST [DST/INSPIRE/04/2017/000413] and SG worked as SRF in an ICAR-NASF Project.
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Supplementary Figure 1
XYZ 4E-ms system for production of hybrid wheat. ms1 recessive male-sterile gene located on chromosome 4BS of wheat; fertility can be restored to the homozygous male-sterile mutant (Z line) through the action of Ms1 dominant fertility restorer gene on alien chromosome 4E of Agropyron elongatum ssp. ruthenicum Beldie (Reproduced with permission from Whitford et al. 2013) (TIFF 111 kb)
Supplementary Figure 2a
Pollination control systems based on conditional male sterility. (a) The induction of male sterility through conversion of a non-phytotoxic substance, i.e. protoxin (N-acetyl-L-phosphinothricin) into a phytotoxin (L-phosphinothricin) in the anthers catalysed by the action of the transgene argE from Escherichia coli (encodes N-acetyl-L-ornithine deacetylase) having exclusive expression in male reproductive tissue (T, tapetum-specific promoter) (JPEG 50 kb)
Supplementary Figure 2b
Pollination control systems based on conditional male sterility. (b) Conditional male sterility through herbicide treatment. A transgene-impairing herbicide tolerance (HerbR) is constitutively expressed (C, constitutive promoter). Its inactivation in male tissue results in the selective destruction of cells necessary for pollen development after herbicide treatment. Transgene inactivation is achieved by male-specific expression (MS, male-specific promoter) of a protein that binds to a 50 UTL operator region (OP) of the HerbR-mRNA. Alternatively, a micro-RNA (miRNA) that binds specifically to an exogenous recognition site (RS) of the HerbR-mRNA may lead to suppression of the protein in the male reproductive tissue (modified after Kempe and Gils 2011) (JPEG 64 kb)
Supplementary Figure 3
Dual-component barnase-barstar system (SeedLink hybridization system) for production of hybrid wheat. Tapetal cell-specific expression (Ta) of barnase (bar) induces male sterility. Herbicide resistance gene (HR) allows selection of male-sterile individuals by herbicide spray. Barstar inhibitor (barstar) inactivates barnase resulting in restored fertility (Reproduced with permission from Whitford et al. 2013) (TIFF 92 kb)
Supplementary Figure 4a
(a) Vector constructs used in split-barnase technology (based on T-DNA; only T-DNA part of the vector is shown here); A is the barnase expression vector called provector; B (pICH13130) is the phiC31-integrase expression vector; (C) in F1 plants obtained by crossing two parents (one with the provector and the other with phiC31); (D) assembly of active barnase cytotoxin via the intein-mediated trans-splicing of two translated precursor protein molecules, Bar-N and Bar-C (with HPTII, hygromycin phosphotransferase; IntN and IntC). For details, see text. LB and RB, T-DNA left and right borders; ocs, octopine synthase terminator; NLS, SV40 T antigen nuclear localization signal (amino acids PKKKRKV); nos, nopaline synthase terminator; ocs, octopine synthase terminator; phiC31, phage phiC31 recombinase coding sequence; PSK, intron PSK7-i3 from Petunia hybrida; Ptap, tapetum-specific promoter from rice; Pubi, maize ubiquitin 1 promoter; S, (GGGGS)3 flexible peptide linker; UBQ, intron UBQ10-i1 from Arabidopsis thaliana (TIFF 69 kb)
Supplementary Figure 4b
(b) Experimental design for the production of hybrid seed using split-barnase approach. Genotypes that have an expected male-sterile phenotype are shown in green. (A) shows development of male-sterile line; (B) represents the strategy for maintaining the male-sterile line, and (C) shows hybrid seed production (reproduced from Kempe et al. 2014) (TIFF 95 kb)
Supplementary Figure 5
Pioneer-DuPont Seed Production Technology (SPT) for production of hybrid wheat. ms45, male sterility inducing gene; Ms45, a dominant fertility restorer gene; SC, seed colour marker; SC allows the visual separation of transgenic maintainer seed from non-transgenic male-sterile seed; PGI is the pollen germination inhibitor. F1 hybrid is non-transgenic (Reproduced with permission from Whitford et al. 2013) (TIFF 104 kb)
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Gupta, P.K., Balyan, H.S., Gahlaut, V. et al. Hybrid wheat: past, present and future. Theor Appl Genet 132, 2463–2483 (2019). https://doi.org/10.1007/s00122-019-03397-y
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DOI: https://doi.org/10.1007/s00122-019-03397-y