1. Please check and comment entries here.
Table of Contents

    Topic review

    Genetic Protection of Soft Wheat

    Subjects: Pathology
    View times: 11
    Submitted by: Elena Gultyaeva

    Definition

    The southern Ural is consistently among the 10 best regions in Russia for agricultural production, including wheat. Breeding in the Chelyabinsk Research Institute of Agriculture aims to develop wheat cultivars genetically protected from the main diseases (leaf and stem rust, septoria leaf blotch and tan spot). The genes for resistance to leaf rust, Lr1, Lr9, Lr10 and Lr26/Sr31, alone or in combination, are widespread in cultivars grown in the southern Ural.

    1. Introduction

    Despite the contrasting climatic conditions, in the 2010s, the Chelyabinsk was consistently among the 10 best regions in Russia for agricultural production. The total area sown to various agricultural crops represents 2–3% of the total cropping area of Russia. In the region, soft and durum wheat, barley, oats, rye, triticale and millet are grown [1]. The largest proportion of cropping in the region is soft wheat (mostly spring wheat), comprising 40–50% of the area sown. In terms of area sown to soft wheat, the Chelyabinsk region is in the top 10 regions of Russia [2].

    Until recently, leaf rust was the main contributor to a damaging pathogenic complex in the region. In some conducive years, the yield losses in wheat were more than 35% [3]. From mid-2010, the severity of stem rust and leaf blotches (septoria leaf blotch and tan spot) outbreaks began to increase.

    Breeding of soft spring wheat at ChRIA commenced in 1937 and has two methodological periods. The first period (1937–1960) was characterized by the use of mainly local wheat cultivars and their hybridization. During this period, the cv. Iskra, Vesna and others were developed. These local cultivars of soft spring wheat had drought tolerance, but lacked the economically important traits of resistance to lodging and diseases due to their long stems.

    Breeding for resistance to leaf rust has a long history in the southern Ural, and it has been based on the strategy of producing cultivars with vertical resistance. The use of nonspecific disease resistance has been limited as it has a negative association with drought tolerance. In the mid-2010s, ChRIA, in conjunction with standard breeding for resistance to leaf rust, began using molecular markers and marker assistance selection (MAS).

    2. Cultivars and Promising Lines of Soft Spring Wheat Produced in ChRIA

    Since 1937, ChRIA breeders have produced and introduced to the region 40 cultivars, of which 17 are included in the State Register of Breeding Achievements (SRBA) as approved for use ( Table 1 ). The widely sown cultivars in southern Ural and Western Siberia at different times were cv. Iskra, Vesna, Uralskaya 52, Rossiyanka, Erythrospermum 59, Niva 2, Duet and Chelyaba 75. The ecological optimums for Chelyabinsk cultivars have now shifted towards more efficient use of the moisture resource due to increased resistance to lodging, leaf rust, EMSD and other stresses.

    Table 1. Cultivars of the Chelyabinsk Research Institute of Agriculture included in the State Register of Breeding Achievements (SRBA) and promising cultivars progressed to State Cultivar Testing (SCT).
    Cultivar Pedigree Year of Involvement in the SRBA/
    Transfer to SCT
    Ripening Time Grain Quality Resistance Genes
    Iskra Milturum 321 × Kitchener 1949 late-ripe weak  
    Vesna - 1961 mid-ripe weak  
    Ural’skaya 52 Cesium 111 × Lutescens 324 1974 mid-ripe strong  
    Rossiyanka Saunders × Svenno 1981 mid-ripe strong  
    Uralochka Svenno × (Lee × Kenya Farmer) 1987 mid-ripe strong  
    Eritrospermum 59 Chayka × Irtyshanka 10 1994 mid-ripe strong Lr10
    Izumrudnaya Waldron × Ural’skaya 52 1996 mid-ripe filler Lr26/Sr31/Pm8/Yr9
    Niva 2 Ps 360/76 × Irtyshanka 10 1997 mid-ripe strong  
    Duet Eritrospermum 59 ×
    (Tselinnaya 20 × ANK-02)
    2003 mid-ripe valuable Lr9 Lr10
    Chelyaba 2 {(Tezpishar × Irtyshanka 10) ×
    Irtushanka 10} ×
    Tselinnaya 20 × ANK-102)
    2005 mid-early valuable Lr9 Lr10
    Pamyati Ryuba Tertsiya ×
    Eritrospermum 19542
    2006 mid-ripe valuable Lr9 Lr10
    Chelyaba yubileynaya Eritrospermum 59 × Tertsiya 2010 mid-late filler Lr9 Lr10
    Chelyaba stepnaya Eritrospermum 59 × Tertsiya 2011 mid-early valuable Lr9 Lr10
    Chelyaba 75 Chernyava 13 ×
    Eritrospermum 21338
    2012 mid-ripe valuable Lr1 Lr10 LrSp
    Ural’skaya kukushka Lutescens 4 × Tulunskaya10 ×
    Lutescens 22178
    2016 mid-early filler  
    Chelyaba rannyaya Chelyaba 2 × ANK-104 2016 early-ripe filler Lr9 Lr10
    Silach Lutescens 210/99-10 ×
    Eritrospermum 23090
    2020 mid-late filler Lr10 Lr9 Lr26/Sr31/Pm8/Yr9
    Chelyaba 80 Cuckoo line 210 × Rossiyanka ×
    Novosibirskaya 15
    SCT 2017 mid-late valuable LrSp
    Il’menskaya 2 Chelyaba 75 ×
    (Chelyaba 2 × Fori 7)
    SCT 2018 mid-early valuable LrSp
    Odintsovskaya Chelyaba 75 × ANK-17B SCT 2018 early-ripe valuable LrSp
    Chelyabinka Vatan × Duet SCT 2021 mid-ripe valuable Lr9 Lr26

    Most of the cultivars grown in the southern Ural before the 2000s were characterized by a high level of susceptibility to leaf rust, and many of them are no longer grown commercially, with the exception of cv. Erythrospermum 59, given that it is equivalent to many modern cultivars in grain quality and resistance to hydrological and hydrothermal stresses. Cultivar Erythrospermum 59 is simultaneously resistant to drought and excessive moisture, has high resistant to lodging and is not subject to EMSD. It is also tolerant to leaf rust, but susceptible to stem rust, septoria leaf blotch and other leaf spots.

    In the 2000s, to expand genetic diversity of resistance to leaf rust, alien donors began to be widely used by ChRIA. One of them was a cuckoo-type line with some Aegilops speltoides genetic background. These lines were developed at the N.I. Vavilov All-Russian Institute of Plant Genetic Resources from crosses and backcrosses with soft wheat of the complex-resistant amphidiploid Triticum dicoccum × Ae. speltoides , and they carried a block of effective Lr and Sr genes linked to the gametocidal gene ( Gc ). The Gc -gene expression leads to the elimination of gametes carrying the recessive gc allele in the heterozygous sporophyte tissues. Recessive alleles linked to it ( lr and sr ) are also eliminated, and as a result, F1 plants are semi-sterile, and in F2 they are homozygous for resistance [4]. The Gc -gene was named cuckoo. Interest in Lr and Sr gene identification in cuckoo-type lines considered for selection is associated with their linked inheritance of resistance to two damaging pathogens and with a genetic mechanism of selection for resistance ( Gc -gene). The presence of the Gc -gene in the block excludes the use of established hybridological analysis; therefore, molecular screening is an alternative method for identifying these genes. Ibragimova et al. [5] identified the gametocidal gene of the cuckoo-type lines as Gc1 and determined its localization on chromosome 2B.

    In the 2000s, ChRIA began to use new donors and lines in breeding with alien genes Lr22a Lr24 , Lr21 , Lr25 , Lr38 , Lr47 (Pavon), Lr48 (CSP44), Lr49 (VL404) and others. A new breeding material of soft spring wheat has been created, combining disease resistance with adaptability to adverse environmental factors, high yield and grain quality [6][7]. Molecular analysis showed a high genetic diversity of all lines in terms of resistance to leaf and stem rusts. Many of the breeding lines have a complexed resistance to the foliar diseases (leaf and stem rust, powdery mildew and blotches) and carry resistance genes previously unused in Russian wheat cultivars (e.g., Lr21 and Lr24 ). More important examples of them are presented in Table 2 .

    Table 2. Reaction of breeding lines and cultivars produced at the Chelyabinsk Research Institute of Agriculture (ChRIA) to foliar diseases in seedling and adult plant stages (2019–2020).
    Wheat Line/
    Cultivar
    Idenified Resistance Genes Reaction Type to Foliar Wheat Pathogens at the Seedling Stage Disease Severity in the Field (%)
    Puccinia triticina Puccinia graminis Parastagonospora nodorum Parastagonospora avenae f. sp. tritici Pyrenophora tritici-repentis Stem Rust Leaf Rust Septoria Leaf Blotch Tan Spot
    PtK1 PtK2 PtK3 Pg1 Pn Pa Ptr
    ToxA+
    Ptr
    ToxA−
    Ptr
    ToxB+
     
    Lut. 26534   3 0–1 3 3–4 5 3–4 2–3/2–3 2–3/2–3 1–2/1–2 1–5 MS 0 5 1
    Er. 26596 Lr10 LrSp/SrSp
    Lr34/Yr18/Sr57/Pm38
    0 0 0 1–2 2–3 2–3 2–3/2–3 2–3/2–3 2–3/2–3 01 MR 0 5 1
    Lut. 26708 Lr3 Lr10 Lr6Agi2 0–1 0 0–1 0–1 3–4 3 3/3–4 2–3/2–3 2–3/2–3 0 0 0 0
    Lut. 26720 Lr1 Lr3 Lr10 Tsn1 3 3 3 0–1 5   2–3/2–3 2/2 1/1 0 0 5 0
    Er. 26725 Lr24/Sr24 0 0 0 0 5 4 1–2/1–2 3/3 1–2/1 0 0 5 10
    Ferr. 26727 Lr10 Lr24/Sr24 0 0 0 0–1 5 4 1–2/1–2 1–2/1–2 1–2/1–2 0 0 15 1
    Er. 26762 Lr26/Sr31/Pm8/Yr9
    Sr35
    0–1 0 0 0–1 5 3 1–2/1–2 1–2/1–2 1–2/1–2 0 0 1 1
    Er. 26775 Lr10 1–2 3 3 0 3–4 3 2–3/2–3 2–3/2–3 2–3/2–3 0 0 5 0
    M. 26690 Lr1 Lr3 Lr10 0 0–1 0–1 0 3–4 1–2 1/1 1/1 1/1 0 0 20 1
    Ferr. 26757 Lr1 Lr3 Lr10
    LrSp/SrSp
    0 0 0 1–2 5 3 2–3/2–3 2–3/2–3 2–3/2–3 0 0 10 10
    Ferr. 26774 Lr10 Lr21
    Lr34/Yr18/Sr57/Pm38
    0–1 0 0 0 2–3 1–2 2–3/2–3 2–3/2–3 1–2/2 0 0 5 0
    Chelyaba 75 Lr1 Lr10 LrSp/SrSp 0 0 0 1–2 3–4 2–3 2–3/2–3 2–3/2–3 2–3/2–3 1 MR 0 10 1
    Chelyaba yubileynaya Lr9 Lr10 3 0 0 3 4 3 3/3 3/3 2–3/2–3 20 S 10–20 S 5 5
    Er. 59   3–4 3–4 3–4 3–4 5 3 3/3 3/3 3/3 30 S 70 S 20 10
    Ferr., Ferrugineum; Lut., Lutescens; Er., Eritrospermum; M, Miltrum. MS, moderately susceptible; MR, moderately resistant; S, susceptible. Reaction types for rust were 0–2 for resistance and 3–4 for susceptibility, with MR type 2, MS types 2–3 and S type 4. Avirulence of test-isolates: P. triticina PtK1 for Lr 19, 23, 24, 26, 28, 29 and 44, and vir-ulence for Lr 1, 2a, 2b, 2c, 3a, 3bg, 3ka, 9, 10, 11, 14a, 14b, 15, 16, 17, 18, 20 and 30. PtK2 for Lr 9, 11, 16, 23, 24, 26, 28 and 29, and Lr 1, 2a, 2b, 2c, 3a, 3bg, 3ka, 10, 14a, 14b, 15, 17, 18, 19, 20, 30 and 44. PtK3 or Lr 9, 2a, 15, 16, 19, 20, 23, 24, 26, 28 and 29, and Lr 1, 2b, 2c, 3a, 3bg, 3ka, 10, 11,14a, 14b, 17, 18, 30 and 44. P. graminis Pg for Sr 24, 24 + 31, 24 + 36, 31, 36, and Sr: 5, 6, 7b, 8a, 9a, 9b, 9g, 9e, 10, 11, 17, 21, 30, 9d and Tmp. P. tritici-repentis: PtrToxA+/−, producing/not producing toxin ToxA; PtrToxB+, pro-ducing toxin ToxB. P. nodorum: Pn, isolates from northwestern Russia; P. avenae f. sp. tritici: Pa2 isolate from Chelyabinsk. −/−, reaction necrosis/chlorosis. P. tritici-repentis and Para-stagonospora sp. genotypes with reaction types 0–2 are resistant, with 3 are moderately susceptible and with 4–5 are susceptible. The GS11 stages of screening for seeding based on Zadoks growth scale and GS 61–85 for adult stages.

     

     

    3. Variability of Pathogens’ Population Structure in Response to Genetically Protected Cultivar Production

    Widespread adoption of genetically protected wheat cultivars can increase variability in pathogen populations. Resistance of cultivars to one pathogen produces a favorable niche for the development of another pathogen (in the absence of complex resistance). This can lead to a significant shift in the composition of pathogen populations. For successful disease genetic management, it is necessary to control the variability in pathogen populations and the effectiveness of resistance genes.

    Tan spot: The recent increase in the importance of tan spot is due partly to reduced or zero-tillage practices that retain stubble on the soil surface, shorter crop rotations or continuous wheat cultivation. The tan spot causal agent, Pyrenophora tritici-repentis , is a necrotrophic fungus that produces host-specific toxins. The pathogen has the ToxA gene, which determines the production of necrosis inducing toxin PtrToxA in wheat cultivars having the avirulence gene Tsn1 . In addition to necrosis on wheat leaves, chlorosis caused by toxins PtrToxB (race 5) and PtrToxC (race 3) develops. However, the main critical virulence factor is PtrToxA [8]. It has been shown that races producing exotoxin ToxA are potentially damaging (race 1, ToxA + ToxC ; race 2, ToxA ; race 7, ToxA + ToxB ; race 8, ToxA + ToxB + ToxC ) [9]. However, in our studies, wheat genotypes at the seedling stage mostly had similar reactions to Ptr isolates producing different toxins ( Table 2 ).

    differential wheat lines/cultivars (lines 6B365 and line 6B662, and cv. Glenlea): 6B365 has Tsc1 , the gene controlling sensitivity to PtrToxC (races 1, 3, 6 and 8), 6B662 has Tsc2 , the gene controlling sensitivity to Ptr ToxB (races 5, 6, 7 and 8), and cv. Glenlea has Tsn1 , the gene controlling sensitivity to PtrToxA (races 1, 2, 7 and 8) [10].

    In 2017, wheat leaves with spots were collected from samples studied in Chelyabinsk under the Kazakhstan-Siberia Network for the Spring Wheat Improvement program [11]. Nineteen monoconidial isolates of P. tritici-repentis were assessed and five races were determined ( Table 3 ). The highest proportion was in races producing Ptr ToxA toxin (86%).

    Table 3. Frequency of Pyrenophora tritici-repentis races in Chelyabinsk region on spring soft wheat (%).
    Race Ptr Toxins 2017 2019 2020
    1 PtrToxA, ToxC 26 2 32
    2 ToxA 53 8 11
    3 ToxC 0 3 8
    4 No toxins 5 29 27
    5 PtrToxB 0 14 0
    6 ToxB + Tox 0 12 8
    7 ToxA + ToxB 5 16 0
    8 ToxA + ToxB + ToxC 7 16 11
    Number of isolates   19 86 37

    4. Conclusions

    This article presented an analysis of wheat genetic variability in the southern Ural, Russia, in terms of population compositions of their obligate and hemibiotrophic pathogen complexes. This helps in understanding plant reactions when they are exposed to multiple pathogens with differing mechanisms of infection and pathogenicity under conducive conditions.

    The analysis of the genetic diversity of wheat cultivars grown in the southern Ural, and the pathogenic complex present, indicate that pathogens continuously evolve into novel virulent races. Since leaf rust is the most damaging pathogen in the southern Ural, breeders need to develop cultivars resistant to this disease. The importance of stem rust and leaf blotches increased in the 2010s, when cultivars with the Lr9 gene began to be widely grown in the region. These cultivars were resistant to leaf rust but susceptible to stem rust and leaf blotches. Thus, cultivar resistance to leaf rust provides an opportunity for colonization of leaves by other pathogens. After 10 years of commercial wheat farming, the Lr9 gene has lost effectiveness. However, in the 2010s, the ranged of commercial wheat cultivars grown in the region was expanded to include cultivars having other race-specific resistance genes. The common production of moderately susceptible or only moderately resistant wheat cultivars in farming in the southern Ural, along with changes in agricultural practices, has led to pathogen build-up and providing abundant inoculum to initiate disease cycles, and provided favorable conditions for the leaf blotches to increase in their importance. In addition to these factors, the influence of global climate change favoring disease development in the region cannot be excluded. Knowledge of genetic variation in a pathogen population underpins breeding for disease resistance. Long-term monitoring of the leaf rust pathogen populations in the southern Ural has not revealed significant changes in the pathogen virulence since the 2010s. In 2019–2020, there was a moderate decrease in the frequency of pathogen virulence to a number of Lr genes. This indicates some stability in the genetic management of wheat in the region for this pathogen. However, significant variation in susceptibility to P. tritici-repentis and Parastagonospora races was found; in particular, potentially damaging races of P. tritici-repentis producing the exotoxin ToxA were found.

    Microevolutionary changes in populations of the tan spot pathogen increase genetic diversity, allowing expansion into new areas and increasing virulence in comparison to previous populations [12]. The southern Ural is an area where P. tritici-repentis has appeared recently. Pathogen populations adjust to their environment over time, but the dynamics of emerging pathogens can be difficult to predict. According to Mikhailova et al. [9], the virulence structure is an indicator of population age. Older populations have lower virulence, as natural selection operates against excess virulence. The Tsn1 gene is not present in most wheat cultivars grown in the southern Ural, but some ChRIA wheat genotypes were susceptible to race PtrToxA.

    Under co-infection conditions, pathogens are thought to exploit host-limited resources more efficiently, with natural selection favoring the coexistence of pathogens that are less damaging to their hosts [13][14]. Early during co-infection, the more virulent pathogen may quickly dominate. However, similarly virulent pathogens can coexist and share their hosts, mainly due to conditions favoring the occurrence of multiple pathogens [15]. An unresolved question is how changes in natural (e.g., climate) and man-made (e.g., new cultivars with polygenic or major gene resistances) conditions alter coexistence in the long term. Changes to conditions may favor one pathogen over another, potentially leading to epidemics of more aggressive pathogens [16]. Therefore, to make significant progress in wheat disease management, research efforts need to include field evaluations encompassing the range of environmental conditions and multiple-pathogen infections.

    The entry is from 10.3390/agriculture11080703

    References

    1. Expert and Analytical Center for Agribusiness. Available online: https://ab-centre.ru/page/selskoe-hozyaystvo-chelyabinskoy-oblasti (accessed on 10 May 2021).
    2. Agrovestnik. Available online: https://agrovesti.net/lib/industries/cereals/posevnye-ploshchadi-valovye-sbory-i-urozhajnost-pshenitsy-v-rossii-itogi-2018-goda.html (accessed on 10 May 2021).
    3. Tyunin, V.A.; Shreyder, E.R. Features of the Technology of Selection of Soft Spring Wheat for Resistance to Carbohydrate-Protein Depletion of Seeds and Other Stresses in the Conditions of the Southern Urals; Russian Academy of Agricultural Sciences, State Scientific Institution Chelyabinsk Scientific Research, Institute of Agriculture: Chelyabinsk, Russia, 2010; 119p. (In Russian)
    4. Odintsova, I.G.; Agafonova, N.A.; Boguslavsky, R.L. Introgressive lines of common wheat with resistantance to leaf rust transferred from Aegilops speltoides, in Source material and problems of wheat and triticale breeding. Proc. Appl. Botany. Genet. Breed. 1991, 142, 106–110. (In Russian)
    5. Ibraimova, Z.K.I.; Tankimanova, M.K.; Bersimbaev, R.I. Chromosomal localization of the gamete-killing gene in the wheat line. Tsitol Genet. 2001, 35, 15–19. (In Russian)
    6. Tyunin, V.A.; Shreyder, E.R.; Gultyaeva, E.I.; Shaydayuk, E.L. Characteristics of virulence of Puccinia triticina populations and the potential of the Lr24, Lr25, LrSp genes for spring common wheat breeding in the Southern Ural. Vavilov J. Genet. Breed. 2017, 21, 523–529.
    7. Gultyaeva, E.I.; Shreyder, E.R.; Kushnirenko, I.Y.; Shaydayuk, E.L.; Kovalenko, N.M. Evaluation of advanced bread spring wheat lines for field and seedling resistance to foliar pathogens. In BIO Web of Conferences; EDP Sciences: Les Ulis, France, 2020.
    8. Lamari, L.; Gilbert, J.; Tekauz, A. Race differentiation in Perynophora tritici-repentis and survey of physiologic variation in western Canada. Can. J. Plant Pathol. 1998, 20, 396–400.
    9. Mikhailova, L.A.; Kokorina, N.M.; Kopahnke, D. Genetics of tan spot (Pyrenophora tritici-repentis) resistance in spring wheat lines 181-5, Vicam“S”70, 292. Russ. Agric. Sci. 2003, 1, 20–22. (In Russian)
    10. Friesen, T.L.; Ali, S.; Klein, K.K.; Rasmussen, J.B. Population genetic analysis of a global collection of Pyrenophora tritici-repentis, causal agent of tan spot of wheat. Phytopathology 2005, 95, 1144–1150.
    11. Gultyaeva, E.I.; Kovalenko, N.M.; Shamanin, V.P.; Tyunin, V.A.; Shreyder, E.R.; Shaydayuk, E.L.; Morgunov, A.I. Population structure of leaf pathogens of common spring wheat in the west Asian regions of Russia and north Kazakhstan in 2017. Vavilov J. Genet. Breed. 2018, 22, 363–369.
    12. Friesen, T.L.; Stukenbrock, E.H.; Liu, Z.; Meinhardt, S.; Ling, H.; Faris, J.; Rasmussen, J.B.; Solomon, P.; McDonald, B.; Oliver, R.P. Emergence of a new disease as a result of interspecific virulence gene transfer. Nat. Genet. 2006, 38, 953–956.
    13. Andrie, R.M.; Pandelova, I.; Ciuffetti, L.M. A combination of phenotypic and genotypic characterization strengthens Pyrenophora tritici-repentis race identification. Phytopathology 2007, 97, 694–701.
    14. Brown, S.P.; Hochberg, M.E.; Grenfell, B.T. Does multiple infection select for raised virulence? Trends Microbiol. 2002, 10, 401–405.
    15. Van Baalen, M.; Sabelis, M.W. The scope for virulence management: A comment on Ewald’s view on the evolution of virulence. Trends Microbiol. 1995, 3, 414–416.
    16. Abdullah, A.S.; Moffat, C.S.; Lopez-Ruiz, F.J.; Gibberd, M.R.; Hamblin, J.; Zerihun, A. Host-multi-pathogen warfare: Pathogen interactions in co-infected plants. Front. Plant. Sci. 2017, 8, 1806.
    More