Stem rust
Stem rust | |
---|---|
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Basidiomycota |
Class: | Pucciniomycetes |
Order: | Pucciniales |
Family: | Pucciniaceae |
Genus: | Puccinia |
Species: | P. graminis
|
Binomial name | |
Puccinia graminis Pers., (1794)
| |
Synonyms | |
See § Synonyms |
Stem rust, also known as cereal rust, black rust,
In 1999 a new virulent race of stem rust was identified against which most current wheat strains show no resistance. The race was named TTKSK (e.g. isolate Ug99). An epidemic of stem rust on wheat caused by race TTKSK spread across Africa, Asia and the Middle East, causing major concern due to the large numbers of people dependent on wheat for sustenance, thus threatening global food security.[7]
An outbreak of another virulent race of stem rust, TTTTF, took place in
History
The fungal ancestors of stem rust have infected grasses for millions of years and wheat crops for as long as they have been grown.[7] According to Jim Peterson, professor of wheat breeding and genetics at Oregon State University, "Stem rust destroyed more than 20% of U.S. wheat crops several times between 1917 and 1935, and losses reached 9% twice in the 1950s," with the last U.S. outbreak in 1962 destroying 5.2% of the crop.[7]
Stem rust has been an ongoing problem dating back to
The parasitic nature of stem rust was discovered in the 1700s. Two Italian scientists,
Due to the useful nature of both barberry and wheat plants, they were eventually brought to North America by European colonists. Barberry was used for many things like making wine and jams from the berries to tool handles from the wood. Ultimately, as they did in Europe, the colonists began to notice a relationship between barberry and stem rust epidemics in wheat. Laws were enacted in many New England colonies, but as the farmers moved west, the problem with stem rust moved with them and began to spread to many areas, creating a devastating epidemic in 1916. It was not until two years later in 1918 that the United States created a program to remove barberry. The program was one that was supported by state and federal entities and was partly prompted by the threat it posed to food supplies during the war. The "war against barberries" was waged and called upon the help of citizens through radio and newspaper advertisements, pamphlets, and fair booths asking for help from all in the attempt to rid the barberry bushes of their existence. Later, in 1975–1980, the program was reestablished under state jurisdiction. Once this happened, a federal quarantine was established against the sale of stem rust susceptible barberry in those states that were part of the program. A barberry testing program was created to ensure that only those species and varieties of barberry that are immune to stem rust will be grown in the quarantine area.[2]
In 1969 two races not detected before in Australia were found[11] and for decades one hypothesis was an African origin,[11][12] and in 2018 DNA analysis confirmed that,[12] specifically South African.[11]
South Africa itself has an ongoing problem with various stem rust outbreaks which requires better response, including an indigenous
Taxonomy
There is considerable genetic diversity within the species P. graminis, and several special forms, forma specialis, which vary in host range have been identified.
- Puccinia graminis f. sp. avenae, oat
- Puccinia graminis f. sp. dactylis
- Puccinia graminis f. sp. hordei, barley
- Puccinia graminis f. sp. lolii
- Puccinia graminis f. sp. poae
- Puccinia graminis f. sp. secalis, rye and barley
- Puccinia graminis f. sp. tritici / Pgt, wheat and barley[13][14][15]
P. graminis is a member of the phylum
The rust is sometimes termed "red rust" or "red dust"[3][16] owing to the spores on the leaf surfaces that range from orange to dark-red in color. Later, the spores change and become dark in color, which gives rise to another common name, "black rust".[17][2]
Puccinia graminis f. sp. tritici
This section needs expansion. You can help by adding to it. (April 2021) |
The North American race nomenclature system[18][19][20] was introduced in 1988 by Roelfs and Martens.[21] This nomenclature is a series of letters each of which indicate virulence/avirulence against one resistance gene, as diagnosed by performance against a group of cultivars known to bear that gene.
Ug99
Pgt contains many races of wheat diseases, including some of the most significant in the world. Ug99 began as a race (TTKSK) of Pgt and now has proliferated into a large number of races of its own.[citation needed]
The virulent new race, against which most current wheat strains show no resistance, was identified in 1999. The race was named TTKSK (e.g. isolate Ug99), named after the country where it was identified (
JRCQC
JRCQC is a race affecting Durum in Ethiopia.[23]
MCC
Affects barley.[14]
QCC
Successfully overwintered in Kansas in 1989/90, and in Texas and Kansas in 1990/91, and so was expected to thereafter be a permanent part of the North American Pg population. Further pathogen adaptation, resulting in widening of the host range, is expected.[24]
QCCJ
Synonymous with QCCJB[15] or known as QCC-2 by some classifications.
Most common Pg race in 1991 in the
QCCJB
The first QCC race (since renamed QCCJ or QCCJB) was detected in the northwest Great Plains in 1988, and by 1990 was over 90% of Pgs on barley in the United States.[15] Also afflicted wheat until a mass switch away from vulnerable cultivars resulted in complete absence in 1997 or 1998.[25][15] Barley virulence is temperature-sensitive: from 18–20 °C (64–68 °F) rpg4 and Rpg5 are highly effective, but above 27 °C (81 °F) they are ineffective. Not necessarily distinguishable from QCCJ, used synonymously by some practitioners.[15]
QCCS
Found in the US on wheat in 1997 and 1998 – but only in the West across both years. On barley in 1997 but not 1998.[25]
QFCS
25% of Pgs on wheat in 1991. Traces found growing in northwest Illinois fields, also in 1991.[24] 8% of all Pgs on wheat, barley, and oat in the US in 1997, and 31% in 1998. Displaced the previously dominant TPMK suddenly in 1998.[25]
TPMK
36% of Pg samples from wheat in 1991 in the United States. Unusually severe in
Synonyms
As listed by
- Aecidium berberidis Pers. ex J.F. Gmel., Syst. Nat., Edn 13 2(2): 1473 (1792)
- Aecidium berberidis var. cyathiforme Rebent., Prodr. fl. neomarch. (Berolini): 352 (1804)
- Aecidium berberidis var. cylindricum Rebent., Prodr. fl. neomarch. (Berolini): 352, tab. 3:11a-b (1804)
- Caeoma berberidis (Pers. ex J.F. Gmel.) Schltdl., Fl. berol. (Berlin) 2: 112 (1824)
- Dicaeoma anthistiriae (Barclay) Syd., Annls mycol. 20(3/4): 117 (1922)
- Dicaeoma anthoxanthi (Fuckel) Kuntze, Revis. gen. pl. (Leipzig) 3(3): 467 (1898)
- Dicaeoma graminis (Pers.) Gray, Nat. Arr. Brit. Pl. (London) 1: 542 (1821)
- Dicaeoma phlei-pratensis (Erikss. & Henn.) Kuntze, Revis. gen. pl. (Leipzig) 3(3): 470 (1898)
- Dicaeoma vilis (Arthur) Arthur, Résult. Sci. Congr. Bot. Wien 1905: 344 (1906)
- Epitea dactylidis G.H. Otth, Mitt. naturf. Ges. Bern 531-552: 88 (1864)
- Lycoperdon berberidis C.-J. Duval, in Hoppe, Bot. Taschenb.: 257 (1793)
- Puccinia albigensis Mayor, Revue Mycol., Paris 22(3): 278 (1957)
- Puccinia anthistiriae Barclay, J. Asiat. Soc. Bengal, Pt. 2, Nat. Sci. 58: 246 (1889)
- Puccinia anthoxanthi Fuckel, Jb. nassau. Ver. Naturk. 27-28: 15 (1874)
- Puccinia brizae-maximae T.S. Ramakr., Indian Phytopath. 6: 30 (1954)
- Puccinia cerealis H. Mart., Prodr. Fl. Mosq., Edn 2: 227 (1817)
- Puccinia culmorum Schumach., Enum. pl. (Kjbenhavn) 2: 233 (1801)
- Puccinia dactylidis G.H. Otth, Mitt. naturf. Ges. Bern 531-552: 88 (1864)
- Puccinia dactylidis Gäum., Ber. schweiz. bot. Ges. 55: 79 (1945)
- Puccinia elymina Miura, Flora of Manchuria and East Mongolia, III Cryptogams, Fungi (Industr. Contr. S. Manch. Rly 27): 283 (1928)
- Puccinia favargeri Mayor, Revue Mycol., Paris 22(3): 273 (1957)
- Puccinia graminis f. macrospora Baudyš, Lotos 64: 29 (1916)
- Puccinia graminis subsp. graminicola Z. Urb., Česká Mykol. 21(1): 14 (1967)
- Puccinia graminis subsp. major A.L. Guyot, Massenot & Saccas, Annales de l'École Nationale d'Agriculture de Grignon, sér. 3 5: 142 (1946)
- Puccinia graminis var. phlei-pratensis (Erikss. & Henn.) Stakman & Piem., J. Agric. Res., Washington 10: 433 (1917)
- Puccinia graminis var. stakmanii A.L. Guyot, Massenot & Saccas, Ann. Ec. Agric. Grignon 5: 145 (1946)
- Puccinia graminis var. stakmanii A.L. Guyot, Massenot & Saccas ex Z. Urb., Česká Mykol. 21(1): 14 (1967)
- Puccinia graminis var. tritici A.L. Guyot, Massenot & Saccas, Annales de l'École Nationale d'Agriculture de Grignon, sér. 3 5: 145 (1946)
- Puccinia jubata Ellis & Barthol., Erythea 4: 2 (1896)
- Puccinia linearis Röhl., Deutschl. Fl. (Frankfurt) 3(3): 132 (1813)
- Puccinia megalopotamica Speg., Anal. Mus. nac. Hist. nat. B. Aires 6: 224 (1898)
- Puccinia phlei-pratensis Erikss. & Henn., Z. PflKrankh. 4: 140 (1894)
- Puccinia vilis Arthur, Bull. Torrey bot. Club 28: 663 (1901)
- Roestelia berberidis (Pers. ex J.F. Gmel.) Gray, Nat. Arr. Brit. Pl. (London) 1: 534 (1821)
- Uredo frumenti Sowerby, Col. fig. Engl. Fung. Mushr. (London) 2(no. 13): tab. 140 (1799)
Pathology
The stem rust fungus attacks the parts of the plant that are above ground. Spores that land on green wheat plants form a pustule that invades the outer layers of the stalk.[7] Infected plants produce fewer tillers and set fewer seed, and in cases of severe infection the plant may die. Infection can reduce what is an apparently healthy crop about three weeks before harvest into a black tangle of broken stems and shriveled grains by harvest.[1]
Stem rust of cereals causes yield losses in several ways:[2]
- Fungus absorbs nutrients that would otherwise be used for grain development.
- Pustules break through epidermis, which disrupt the plant's control of transpiration and can lead to desiccation and infection by other fungi.
- Interference with plant vascular tissue leads to shriveled grains.
- The fungus weakens the stems, which can lead to lodging (falling over). In severe cases lodging can make mechanical harvesting impossible.
Signs and symptoms
On wheat
Stem rust on wheat is characterized by the presence of uredinia on the plant, which are brick-red, elongated, blister-like pustules that are easily shaken off. They most frequently occur on the leaf sheaths, but are also found on stems, leaves, glumes and awns. On leaves they develop mostly on the underside but may penetrate to the upperside. On leaf sheaths and glumes pustules rupture the epidermis, giving a ragged appearance.[1]
Towards the end of the growing season black telia are produced. For this reason stem rust is also known as "black rust". The telia are firmly attached to the plant tissue.[1]
The site of infection is a visible symptom of the disease.
On barberry
Pycnia appear on
Life cycle
Like other
P. graminis can complete its life cycle either with or without barberry (the alternate host).[2]
P. g. tritici's
Life cycle on barberry
Due to its cyclical nature, there is no true 'start point' for this process. Here, the production of urediniospores is arbitrarily chosen as a start point.
Urediniospores are formed in structures called uredinia, which are produced by fungal mycelia on the cereal host 1–2 weeks after infection. The urediniospores are dikaryotic (contain two un-fused, haploid nuclei in one cell) and are formed on individual stalks within the uredinium. They are spiny and brick-red. Urediniospores are the only type of spores in the rust fungus life cycle that are capable of infecting the host on which they are produced, and this is therefore referred to as the 'repeating stage' of the life cycle. It is the spread of urediniospores that allows infection to spread from one cereal plant to another.[2] This phase can rapidly spread the infection over a wide area.
Towards the end of the cereal host's growing season, the mycelia produce structures called telia. Telia produce a type of spore called teliospores. These black, thick-walled spores are dikaryotic. They are the only form in which Puccinia graminis is able to overwinter independently of a host.[2]
Each teliospore undergoes karyogamy (fusion of nuclei) and meiosis to form four haploid spores called basidiospores. This is an important source of genetic recombination in the life cycle. Basidiospores are thin-walled and colourless. They cannot infect the cereal host, but can infect the alternative host (barberry).[2] They are usually carried to the alternative host by wind.
Once basidiospores arrive on a leaf of the alternative host, they germinate to produce a
This dikaryotic mycelium then forms structures called aecia, which produce a type of dikaryotic spores called aeciospores. These have a worty appearance and are formed in chains – unlike the urediniospores that are spiny and are produced on individual stalks. The chains of aeciospores are surrounded by a bell-like enclosure of fungal cells. The aeciospores are able to germinate on the cereal host but not on the alternative host (they are produced on the alternative host, which is usually barberry). They are carried by wind to the cereal host where they germinate and the germ tubes penetrate into the plant. The fungus grows inside the plant as a dikaryotic mycelium. Within 1–2 weeks the mycelium produces uredinia and the cycle is complete.[2]
Life cycle without barberry
Since the urediniospores are produced on the cereal host and can infect the cereal host, it is possible for the infection to pass from one year's crop to the next without infecting the alternate host (barberry). For example, infected volunteer wheat plants can serve as a bridge from one growing season to another. In other cases the fungus passes between winter wheat and spring wheat, meaning that it has a cereal host all year round. Since the urediniospores are wind dispersed, this can occur over large distances.[2] Note that this cycle consists simply of vegetative propagation – urediniospores infect one wheat plant, leading to the production of more urediniospores that then infect other wheat plants.
Spore dispersal
Puccinia graminis produces all five of the spore types that are known for rust fungi.[2]
Spores are typically deposited close to the source, but long-distance dispersal is also well documented[1] commonly out to hundreds of kilometres/miles.[30] The following three categories of long-distance dispersal are known to occur:[1]
- Extremely long-distance dispersal
This can occur unassisted (the robust nature of the spores allows them to be carried long distances in the air and then deposited by rain-scrubbing) or assisted (typically on human clothing or infected plant material that is transported between regions).[1] This type of dispersal is rare and is very difficult to predict.[1] This is especially known to rarely occur across thousands of km/mi from South Africa to Western Australia.[31][32]
- Step-wise range expansion
This is probably the most common mode of long-distance dispersal and usually occurs within a country or region.[1]
- Extinction and recolonisation
This occurs in areas that have unsuitable conditions for year-round survival of Puccinia graminis – typically temperate regions where hosts are absent during either the winter or summer.[1] Spores overwinter or oversummer in another region and then recolonise when conditions are favorable.[1]
Wheat stem rust resistance genes
A number of stem rust
None of the Sr genes provide resistance to all races of stem rust. For instance many of them are ineffective against the
There has been significant uptake of resistant wheat varieties among Ethiopian farmers since 2014[35][36] – a great deal of which is thanks to CGIAR and CIMMYT (the International Maize and Wheat Improvement Center).[37][36]
Although Sr5, Sr21, Sr9e, Sr7b, Sr11, Sr6, Sr8a, Sr9g, Sr9b, Sr30, Sr17, Sr9a, Sr9d, Sr10, SrTmp, Sr38, and SrMcN are no longer effective in Lebanon, Sr11, Sr24, and Sr31 still are which is diagnostic for the presence of various races of stem rust – but the complete absence of Ug99 specifically – from Lebanon.[38]
Sr9h
Discovered and found to provide
Sr14
Sr14 does not protect seedlings against TTKSK[40] but does provide moderate resistance at later stages.[40] It is effective against TTKST.[40]
Sr22
There is considerable variation among Sr22 alleles, with some conferring resistance and some susceptibility.[41]
Sr27
Sr27
Sr31
Ug99 is virulent against Sr31, which was effective against all previous stem rust races.[34]
Sr33
An
Sr35
Sr35 is an
Sr59
Recently, a new stem rust resistance gene Sr59 from
Sr62
An NLR (or NB-LRR, or
SrTmp
Originally from the widespread Ethiopian 'Digalu'.[52] Resistant to Ug99, susceptible to § TKTTF.[52]
Weaponization
In the 1950s, the United States Air Force developed Operation Steelyard, a plan to drop wheat stem rust mixed with feathers over wheat farms in the Soviet Union. If the plan were enacted, Boeing B-29 Superfortress bombers would drop 500-pound M115 bombs over Soviet farms, with the intention of destroying up to 50% of the Soviet winter wheat harvest.[53]
Future
Alone amongst cereals,
See also
References
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- ^ a b c Vincent, Michael (1 March 2011). "Wheat disease a threat to global food security". ABC News. Australian Broadcasting Corporation. Retrieved 27 May 2021.
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- ^ "Wheat Rust Diseases". Crop Science US. Retrieved 27 May 2021.
- ^ "North American Stem Rust Nomenclature Code Sheet" (PDF). CIMMYT (International Maize and Wheat Improvement Center). Archived from the original (PDF) on 2021-04-03.
- ^ "Rust - Stem: Race Nomenclature". United Nations Food and Agriculture Organization. Archived from the original on 2021-04-03. Retrieved 2021-04-03.
- ^ "Race identification". Agricultural Research Service. United States Department of Agriculture. Archived from the original on 2021-04-03. Retrieved 2021-04-03.
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- ^ a b c d Hawkins, Nichola (2021-01-20). "Ug99 Stem Rust – Breaching Wheat's Defences". Plant Pandemic Studies. British Society for Plant Pathology: 1–35.
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- ^ "Species Fungorum - GSD Species". www.speciesfungorum.org. Retrieved 19 August 2023.
- ^ Animated video of the life cycle of stem rust
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- ^ Sounding the Alarm on Global Stem Rust (PDF) (Report). Borlaug Global Rust Initiative. p. 4:
Like all...
- ^ "Watson, I A.; Cass Smith, W. P.; and Shipton, W. A. (1966) "Strains of stem and leaf rust on wheat in Western Australia since 1951," Journal of the Department of Agriculture, Western Australia, Series 4: Vol. 7 : No. 8 , Article 7". p. 365:
It may be...
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- ^ a b "Press release: Rust-resistant bread wheat varieties widely adopted in Ethiopia, study shows » CGIAR Research Program on WHEAT". CGIAR WHEAT. 10 November 2020. Retrieved 2020-11-18.
- ^ Yasabu, Simret (2020-12-15). "Shining a brighter light on adoption and diffusion". CIMMYT (the International Maize and Wheat Improvement Center). Retrieved 2021-01-01.
- ^ Rola El Amil (Lebanese Agricultural Research Institute, Lebanon) (2020-11-09). (DAY 2) - Phytosanitary Safety for Transboundary pest prevention - Yellow and Black rust population variability. CGIAR Germplasm Health Webinar series. Vol. Phytosanitary Awareness Week. International Institute of Tropical Agriculture / CGIAR. Slide at 00:44:37. Archived from the original on 2021-12-14.
- ^
- • Wessels, Elsabet; Prins, Renee; Boshoff, Willem; Zurn, Jason; Acevedo, Maricelis; Pretorius, Zacharias (2019), "Mapping a Resistance Gene to Puccinia graminis f. sp. tritici in the Bread Wheat Cultivar 'Matlabas'", S2CID 233672540
- • Wessels, Elsabet; Prins, Renee; Boshoff, Willem; Zurn, Jason; Acevedo, Maricelis; Pretorius, Zacharias (2019), "Mapping a Resistance Gene to Puccinia graminis f. sp. tritici in the Bread Wheat Cultivar 'Matlabas'",
- ^ .
- .
- ^ a b "Sr27". Borlaug Global Rust Initiative (BGRI). Retrieved 2021-07-24.
- ^ PMID 22920559.
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- • Yu, Guotai; Matny, Oadi; Champouret, Nicolas; Steuernagel, Burkhard; Moscou, Matthew J.; Hernández-Pinzón, Inmaculada; Green, Phon; Hayta, Sadiye; Smedley, Mark; Harwood, Wendy; Kangara, Ngonidzashe; Yue, Yajuan; Gardener, Catherine; Banfield, Mark J.; Olivera, Pablo D.; Welchin, Cole; Simmons, Jamie; Millet, Eitan; Minz-Dub, Anna; Ronen, Moshe; Avni, Raz; Sharon, Amir; Patpour, Mehran; Justesen, Annemarie F.; Jayakodi, Murukarthick; Himmelbach, Axel; Stein, Nils; Wu, Shuangye; Poland, Jesse; Ens, Jennifer; Pozniak, Curtis; Karafiátová, Miroslava; Molnár, István; Doležel, Jaroslav; Ward, Eric R.; Reuber, T. Lynne; Jones, Jonathan D. G.; Mascher, Martin; Steffenson, Brian J.; Wulff, Brande B. H. (2022). "Aegilops sharonensis genome-assisted identification of stem rust resistance gene Sr62". .
- ^ S2CID 205345605.
- ^ Kirby; Carus (2020). "Agroterrorism Perspectives". In Mauroni; Norton (eds.). Agroterrorism: National Defense Assessment, Strategies, and Capabilities (PDF). U.S. Air Force Center for Strategic Deterrence Studies. p. 9.
- Rice Today. Vol. 10, no. 1. International Rice Research Institute (IRRI) (CGIAR's Research Program on Rice). pp. 38–39.
- Centre for Agriculture and Bioscience International). Retrieved 2021-04-10.
Further reading
- Bhardwaj, S.C.; Nyar, S.K.; Prashar, M.; Kumar, J; Menon, M.K.; Singh, S.B. (1990). "A pathotype of Puccinia graminis f. sp. tritici on Sr24 in India". Cereal Rusts and Powdery Mildews Bulletin: 35–38.
External links
- Borlaug Global Rust Initiative
- Stem rust by FAO (the Food and Agriculture Organization of the United Nations
- Animation of stem rust life cycle
- "Norwich Rust Group". Norwich Rust Group. Retrieved 2020-12-18.
- "Stem Rust". Retrieved 2021-05-26.