Stem rust - Knowledge

424: 1337: 393:. 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. 1618:. 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. 51: 1425: 606: 315:), the rust god. They would perform this ritual in the spring during a festival known as the Robigalia in hopes of the wheat crop being spared from the destruction caused by the rust. Weather records from that time have been reexamined and it has been speculated that the fall of the Roman Empire was due to a string of rainy seasons in which the rust would have been more harsh, resulting in reduced wheat harvests. Laws banning barberry were established in 1660 in 38: 4871:

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).

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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. 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.

1549:. 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. This phase can rapidly spread the infection over a wide area. 3622:

A.; Lucas, S. M.; Mago, R.; Mauceli, E.; Morin, E.; Murat, C.; Pangilinan, J. L.; Park, R.; Pearson, M.; Quesneville, H.; Rouhier, N.; Sakthikumar, S.; Salamov, A. A.; Schmutz, J.; Selles, B.; Shapiro, H.; Tanguay, P.; Tuskan, G. A.; Henrissat, B.; Van de Peer, Y.; Rouze, P.; Ellis, J. G.; Dodds, P. N.; Schein, J. E.; Zhong, S.; Hamelin, R. C.; Grigoriev, I. V.; Szabo, L. J.; Martin, F. (2011-05-02).

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Hu, Zhaorong; Guo, Weilong; Li, Guoqiang; Ma, Xin; Li, Junming; Han, Fangpu; Fu, Xiangdong; Ma, Zhengqiang; Wang, Daowen; Zhang, Xueyong; Ling, Hong-Qing; Xia, Guangmin; Tong, Yiping; Liu, Zhiyong; He, Zhonghu; Jia, Jizeng; Chong, Kang (2022). "Wheat genomic study for genetic improvement of traits in China".

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Xiao, Jun; Liu, Bao; Yao, Yingyin; Guo, Zifeng; Jia, Haiyan; Kong, Lingrang; Zhang, Aimin; Ma, Wujun; Ni, Zhongfu; Xu, Shengbao; Lu, Fei; Jiao, Yuannian; Yang, Wuyun; Lin, Xuelei; Sun, Silong; Lu, Zefu; Gao, Lifeng; Zhao, Guangyao; Cao, Shuanghe; Chen, Qian; Zhang, Kunpu; Wang, Mengcheng; Wang, Meng;

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lifestyle involves the dramatic up-regulation of particular gene transcriptions, constituting its biotrophy genomic features. These genomic regions have parallels in other eukaryotic plant pathogens. These parallels – between these independently evolved and unrelated sets of genes – show a strong and

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million towards research into Ug99, to be spent on critical infrastructure in Africa. Scientists are working on breeding strains of wheat that are resistant to UG99. However, wheat is grown in a broad range of environments. This means that breeding programs would have extensive work remaining to get

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later conducted experiments to observe the beliefs of European farmers regarding the relationship between the rust and barberry plants, and after connecting the basidiospores of the basidia stage to barberry, he also identified that the aeciospores in the aecia stage reinfect the wheat host. Upon de

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Olivera, Pablo; Newcomb, Maria; Szabo, Les J.; Rouse, Matthew; Johnson, Jerry; Gale, Samuel; Luster, Douglas G.; Hodson, David; Cox, James A.; Burgin, Laura; Hort, Matt; Gilligan, Christopher A.; Patpour, Mehran; Justesen, Annemarie F.; Hovmøller, Mogens S.; Woldeab, Getaneh; Hailu, Endale; Hundie,

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Duplessis, S.; Cuomo, C. A.; Lin, Y.-C.; Aerts, A.; Tisserant, E.; Veneault-Fourrey, C.; Joly, D. L.; Hacquard, S.; Amselem, J.; Cantarel, B. L.; Chiu, R.; Coutinho, P. M.; Feau, N.; Field, M.; Frey, P.; Gelhaye, E.; Goldberg, J.; Grabherr, M. G.; Kodira, C. D.; Kohler, A.; Kues, U.; Lindquist, E.

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discovered the characteristic five-spore stage that is known in some stem rust species. The brothers were also able to make a connection between the red (urediniospore) and black (teliospore) spores as different stages within the life cycle of the same organism, but the rest of the stages remained

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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). This type of dispersal is rare and is very

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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,

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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

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Upadhyaya, Narayana M.; Mago, Rohit; Panwar, Vinay; Hewitt, Tim; Luo, Ming; Chen, Jian; Sperschneider, Jana; Nguyen-Phuc, Hoa; Wang, Aihua; Ortiz, Diana; Hac, Luch; Bhatt, Dhara; Li, Feng; Zhang, Jianping; Ayliffe, Michael; Figueroa, Melania; Kanyuka, Kostya; Ellis, Jeffrey G.; Dodds, Peter N.

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s on wheat in 1997 in the United States – being absent only from the southern Great Plains and the west, but then was down to 10% in 1998. In the upper Great Plains it was already declining – to 26% of samples in 1997, and 12% in 1998. In the most fertile areas of the eastern US it was 96% of

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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

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Singh, Ravi P.; Hodson, David P.; Huerta-Espino, Julio; Jin, Yue; Bhavani, Sridhar; Njau, Peter; Herrera-Foessel, Sybil; Singh, Pawan K.; Singh, Sukhwinder; Govindan, Velu (2011-09-08). "The Emergence of Ug99 Races of the Stem Rust Fungus is a Threat to World Wheat Production".

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Singh, Ravi P.; Hodson, David P.; Huerta-Espino, Julio; Jin, Yue; Bhavani, Sridhar; Njau, Peter; Herrera-Foessel, Sybil; Singh, Pawan K.; Singh, Sukhwinder; Govindan, Velu (2011). "The Emergence of Ug99 Races of the Stem Rust Fungus is a Threat to World Wheat Production".

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and the receptive hyphae. The pycniospores are produced in a sticky honeydew that attracts insects. The insects carry pycniospores from one leaf to another. Splashing raindrops can also spread pycniospores. A pycniospore can fertilise a receptive hypha of the opposite

2027:, Australia, rapidly produced virulence between 1982 and 1984 – the first virulence on this gene in the world. (This was especially associated with the cultivar Coorong.) Therefore, CIMMYT's triticale offerings were tested and many were found to depend solely on 4291:

Hatta, Muhammad Asyraf Muhammad; Ghosh, Sreya; Athiyannan, Naveenkumar; Richardson, Terese; Steuernagel, Burkhard; Yu, Guotai; Rouse, Matthew N.; Ayliffe, Michael; Lagudah, Evans S.; Radhakrishnan, Guru V.; Periyannan, Sambasivam K.; Wulff, Brande B. H. (2020).

1579:. 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). They are usually carried to the alternative host by wind. 236:. These diseases have affected cereal farming throughout history. The annual recurrence of stem rust of wheat in North Indian plains was discovered by K.C. Mehta. Since the 1950s, wheat strains bred to be resistant to stem rust have become available. 4697:

Periyannan, Sambasivam; Moore, John; Ayliffe, Michael; Bansal, Urmil; Wang, Xiaojing; Huang, Li; Deal, Karin; Luo, Mingcheng; Kong, Xiuying; Bariana, Harbans; Mago, Rohit; McIntosh, Robert; Dodds, Peter; Dvorak, Jan; Lagudah, Evans (2013-06-27).

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The rust is sometimes termed "red rust" or "red dust" 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".

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Salcedo, Andres; Rutter, William; Wang, Shichen; Akhunova, Alina; Bolus, Stephen; Chao, Shiaoman; Anderson, Nickolas; De Soto, Monica Fernandez; Rouse, Matthew; Szabo, Les; Bowden, Robert L.; Dubcovsky, Jorge; Akhunov, Eduard (2017-12-21).

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Kumar, Sachin; Fetch, Tom G.; Knox, Ron E.; Singh, Asheesh K.; Clarke, John M.; Depauw, Ron M.; Cuthbert, Richard D.; Campbell, Heather L.; Singh, Davinder; Bhavani, Sridhar; Pozniak, Curtis J.; Meyer, Brad; Clarke, Fran R. (2020-11-19).

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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

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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.

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s on barley in the United States. Also afflicted wheat until a mass switch away from vulnerable cultivars resulted in complete absence in 1997 or 1998. Barley virulence is temperature-sensitive: from 18–20 °C (64–68 °F)

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Spores are typically deposited close to the source, but long-distance dispersal is also well documented commonly out to hundreds of kilometres/miles. The following three categories of long-distance dispersal are known to occur:

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was introgressed into wheat, which provides an additional asset for wheat improvement to mitigate yield losses caused by stem rust. Singh et al. (2011) provide a list of known Sr genes and their effectiveness against Ug99.

319:, France. This was due to the fact that European farmers noticed a correlation between barberry and stem rust epidemics in wheat. The law banned the planting of barberry near wheat fields and was the first of its kind. 5087:

Singh, Ravi; Hodson, David; Jin, Yue; Lagudah, Evans; Ayliffe, Michael; Bhavani, Sridhar; Rouse, Matthew; Pretorius, Zacharias; Szabo, Les; Huerta-Espino, Julio; Basnet, Bhoja; Lan, Caixia; Hovmoller, Mogens (2015).

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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

1590:) that directly penetrates the epidermis and colonises the leaf. Once inside the leaf the mycelium produces specialised infection structures called pycnia. The pycnia produce two types of haploid gametes, the 1296:

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. Infected plants produce fewer

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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

297:, "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. 292:

The fungal ancestors of stem rust have infected grasses for millions of years and wheat crops for as long as they have been grown. According to Jim Peterson, professor of wheat breeding and genetics at

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Hodson, D. P.; Jaleta, M.; Tesfaye, K.; Yirga, C.; Beyene, H.; Kilian, A.; Carling, J.; Disasa, T.; Alemu, S. K.; Daba, T.; Alemayehu, Y.; Badebo, A.; Abeyo, B.; Erenstein, O. (2020-10-28).

1681:– typically temperate regions where hosts are absent during either the winter or summer. Spores overwinter or oversummer in another region and then recolonise when conditions are favorable. 4241: 4170: 956:

s in 1997 but then suddenly fell to 29% in 1998. In a few other locations in the US, and overall across the US, this race declined 97–98 in favor of other races, and not because of overall

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The North American race nomenclature system was introduced in 1988 by Roelfs and Martens. This nomenclature is a series of letters each of which indicate virulence/avirulence against one

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resistance into regionally adapted germplasms even after resistance is identified. Similarly, in 2014, a Ug99 race called "Digalu" emerged and devastated the Digalu variety in Ethiopia.

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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

508: 3178: 1368:. 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. 2661:

Saunders, Diane G. O.; Wulff, Brande B. H.; Thomas, Jane; Fenwick, Paul M.; Visser, Botma; Steffenson, Brian; Singh, Ravi P.; Sela, Hanan; Roohparvar, Ramin (2018-02-08).

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samples, and 67% in 1990. Afflicted spring-sown barley in the northern Great Plains in 1990. Was the first barley stem rust to show up in the United States in 1991, in

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are highly effective, but above 27 °C (81 °F) they are ineffective. Not necessarily distinguishable from QCCJ, used synonymously by some practitioners.

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on stems and leaves is typical of a general stem rust as well as any variation of this type of fungus. Different from most fungi, the rust variations have five

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Steffenson, B. J.; Case, A. J.; Pretorius, Z. A.; Coetzee, V.; Kloppers, F. J.; Zhou, H.; Chai, Y.; Wanyera, R.; Macharia, G.; Bhavani, S.; Grando, S. (2017).

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Visser, Botma; Meyer, Marcel; Park, Robert F.; Gilligan, Christopher A.; Burgin, Laura E.; Hort, Matthew C.; Hodson, David P.; Pretorius, Zacharias A. (2019).

2189: 4294:"Extensive Genetic Variation at the Sr22 Wheat Stem Rust Resistance Gene Locus in the Grasses Revealed Through Evolutionary Genomics and Functional Analyses" 2370: 243:

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

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Fetch, Thomas; Park, Robert; Pretorius, Zacharias; Depauw, Ronald (2021), "Stem rust: its history in Kenya and research to combat a global wheat threat",

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Raccolta di tutte le opere del chiarissimo cavaliere Giuseppe Maria Giovene con note dell'editore Luigi Marinelli Giovene: Memorie fisico agrarie

1908: 1964: 4006: 304:'s time (384–322 BC). An early ancient practice by the Romans was one where they would sacrifice red animals such as foxes, dogs, and cows to 5526: 4107:

Wessels, Elsabet; Prins, Renee; Boshoff, Willem; Zurn, Jason; Acevedo, Maricelis; Pretorius, Zacharias (2019), "Mapping a Resistance Gene to

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still are which is diagnostic for the presence of various races of stem rust – but the complete absence of Ug99 specifically – from Lebanon.

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The fungus weakens the stems, which can lead to lodging (falling over). In severe cases lodging can make mechanical harvesting impossible.

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Towards the end of the cereal host's growing season, the mycelia produce structures called telia. Telia produce a type of spore called

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s afflicting wheat in the US in 1995, 1% in 1996, and not at all in 1997 or 1998. Not found on barley in 1997 but found again in 1998.

4007:"Press release: Rust-resistant bread wheat varieties widely adopted in Ethiopia, study shows » CGIAR Research Program on WHEAT" 2173:

was introgressed into wheat, which provides an additional asset for wheat improvement to mitigate yield losses caused by stem rust.

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South Africa itself has an ongoing problem with various stem rust outbreaks which requires better response, including an indigenous

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Watson, I A.; Cass Smith, W. P.; Shipton, W. A. (1966). "Strains of stem and leaf rust on wheat in Western Australia since 1951".

5474: 5090:"Emergence and Spread of New Races of Wheat Stem Rust Fungus: Continued Threat to Food Security and Prospects of Genetic Control" 5027: 4890: 3220: 3189: 1657:

difficult to predict. This is especially known to rarely occur across thousands of km/mi from South Africa to Western Australia.

2487: 1408: in) from the leaf surface. So important is its role in maintenance of prevalence that since the near extermination of the 1907:, 2019 find the Ug99 resistance of cv. 'Matlabas' is probably due to this gene. Wessels finds it is present in less than 5% of 282:, combined with plant pathology and climate data, has pointed out the potential of the re-emergence of stem wheat rust in UK. 5182: 5479: 1607:. This is the sexual stage of the life cycle and cross-fertilisation provides an important source of genetic recombination. 4298: 3161: 423: 1971:

and rarely in hexaploid wheat. Located on the 3A chromosome arm, originally from 3R. Virulence has been observed in field

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are produced. For this reason stem rust is also known as "black rust". The telia are firmly attached to the plant tissue.

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None of the Sr genes provide resistance to all races of stem rust. For instance many of them are ineffective against the

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break through the lower leaf surface. The aecial cups are yellow and sometimes elongate to extend up to 5 millimetres (

707:, causing major concern due to the large numbers of people dependent on wheat for sustenance, thus threatening global 263:, causing major concern due to the large numbers of people dependent on wheat for sustenance, thus threatening global 5272: 4544: 3184: 2394: 903:

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.

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This is probably the most common mode of long-distance dispersal and usually occurs within a country or region.

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Miura, Flora of Manchuria and East Mongolia, III Cryptogams, Fungi (Industr. Contr. S. Manch. Rly 27): 283 (1928)

456: 4375: 4073:(DAY 2) - Phytosanitary Safety for Transboundary pest prevention - Yellow and Black rust population variability 3782: 1229:

A.L. Guyot, Massenot & Saccas, Annales de l'École Nationale d'Agriculture de Grignon, sér. 3 5: 145 (1946)

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A.L. Guyot, Massenot & Saccas, Annales de l'École Nationale d'Agriculture de Grignon, sér. 3 5: 142 (1946)

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Sharma Poudel, Roshan; Richards, Jonathan; Shrestha, Subidhya; Solanki, Shyam; Brueggeman, Robert (2019).

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developed Operation Steelyard, a plan to drop wheat stem rust mixed with feathers over wheat farms in the

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McIntosh, R. A.; Brown, G. N. (1997). "Anticipatory Breeding for Resistance to Rust Diseases in Wheat".

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in a sticky honeydew. Five to ten days later, cup-shaped structures filled with orange-yellow, powdery

345: 327: 161: 1760:, which was effective against all previous stem rust races. Recently, a new stem rust resistance gene 5541: 5414: 2279: 1530:

Due to its cyclical nature, there is no true 'start point' for this process. Here, the production of

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plants in the spring, usually in the upper leaf surfaces. They are often in small clusters and exude

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Pustules break through epidermis, which disrupt the plant's control of transpiration and can lead to

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Bhardwaj, S.C.; Nyar, S.K.; Prashar, M.; Kumar, J; Menon, M.K.; Singh, S.B. (1990). "A pathotype of

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Jin, Y; Steffenson, BJ; Miller, JD (1994). "Inheritance of resistance to pathotypes QCC and MCC of

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and Kansas in 1990/91, and so was expected to thereafter be a permanent part of the North American

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over Soviet farms, with the intention of destroying up to 50% of the Soviet winter wheat harvest.

1903:, both in 2010, already had virulence when retested against this new gene. Both Rouse and Wessels 1545:(contain two un-fused, haploid nuclei in one cell) and are formed on individual stalks within the 923:

s on wheat, barley, and oat in the US in 1997, and 31% in 1998. Displaced the previously dominant

19:"Red rust" redirects here. For the genus of parasitic thalloid green algae known as red rust, see 4117: 4089: 3485: 3419: 2271: 412: 294: 4061: 617: 5453: 5323: 4350: 2718: 2667: 1741: 847: 771:

population. Further pathogen adaptation, resulting in widening of the host range, is expected.

331: 4452:"Genomics accelerated isolation of a new stem rust avirulence gene–wheat resistance gene pair" 4881: 4400:

Park, Robert F.; Wellings, Colin R. (2012-09-08). "Somatic Hybridization in the Uredinales".

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contains many races of wheat diseases, including some of the most significant in the world.

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Due to the useful nature of both barberry and wheat plants, they were eventually brought to

349: 5383: 4968: 4894: 4797: 4780:"Variation in the AvrSr35gene determines Sr35 resistance against wheat stem rust race Ug99" 4725: 4472: 4183: 3723: 3587: 2553: 1512: 855: 3942:"Ethiopia's transforming wheat landscape: tracking variety use through DNA fingerprinting" 3310: 2298:

is naturally immune to rusts. If a genetic source of this resistance could be identified,

8: 4536: 4179: 2787:"Microsatellite Analysis and Urediniospore Dispersal Simulations Support the Movement of 2315: 1717: 1582:

Once basidiospores arrive on a leaf of the alternative host, they germinate to produce a

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Bhattacharya, Shaoni (2017-02-09). "Deadly new wheat disease threatens Europe's crops".

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The parasitic nature of stem rust was discovered in the 1700s. Two Italian scientists,

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Urediniospores are formed in structures called uredinia, which are produced by fungal

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on barley was QCCJ. On wheat, QCCJ was still the most common race but only at 38% of

439: 3761: 3607: 3528: 3452: 3390: 3288: 1639: 1193:(Erikss. & Henn.) Stakman & Piem., J. Agric. Res., Washington 10: 433 (1917) 640:, as diagnosed by performance against a group of cultivars known to bear that gene. 5375: 5107: 5031: 4976: 4918: 4902: 4823: 4805: 4733: 4671: 4623: 4619: 4575: 4480: 4415: 4311: 4254: 4191: 4130: 3975: 3959: 3905: 3844: 3731: 3671: 3653: 3595: 3498: 3432: 3360: 3268: 3245:

Roelfs, A. P.; Martens, J. W. (1988). "An International System of Nomenclature for

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Hovmøller, Mogens S.; Pretorius, Zacharias A.; Saunders, Diane G. O. (2019-02-04).

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can complete its life cycle either with or without barberry (the alternate host).

340:, also thoroughly studied the stem rust. Thirty years later it received its name, 5549: 4468: 3955: 3892: 3710: 2206: 2020: 1884: 1298: 944: 712: 448: 364:

Bary's discovery of all five spore stages and their need for barberry as a host,

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in barley line Q21861 and temperature effects on the expression of resistance".

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This occurs in areas that have unsuitable conditions for year-round survival of

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The first QCC race (since renamed QCCJ or QCCJB) was detected in the northwest

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Kirby; Carus (2020). "Agroterrorism Perspectives". In Mauroni; Norton (eds.).

5035: 4981: 4950: 4316: 4293: 3736: 3705: 3624:"Obligate biotrophy features unraveled by the genomic analysis of rust fungi, 3566: 3096: 2926: 2731: 2680: 2565: 2368: 1928:

but does provide moderate resistance at later stages. It is effective against

5594: 5121: 5043: 4990: 4914: 4819: 4747: 4683: 4631: 4587: 4492: 4459: 4427: 4325: 4268: 4205: 4144: 3971: 3856: 3745: 3667: 3512: 3444: 3374: 3333:

Bekele; Tadesse, Kebede; Pumphrey, Michael; Singh, Ravi P.; Jin, Yue (2015).

3280: 3065: 2935: 2880: 2826: 2109: 2104: 2064: 1531: 1424: 801: 789: 708: 564: 386: 376: 360: 264: 102: 82: 4810: 4738: 4699: 3658: 2595:

Wanyera, R.; Macharia, J. K.; Kilonzo, S. M.; Kamundia, J. W. (2009-08-06).

1309:

Fungus absorbs nutrients that would otherwise be used for grain development.

5466: 5129: 5051: 4998: 4932: 4837: 4755: 4639: 4500: 4435: 4333: 4152: 3989: 3917: 3864: 3798: 3753: 3685: 3520: 3382: 3073: 2953: 2908: 2834: 2749: 2698: 2622: 2613: 2596: 2573: 2458:

Schumann, G.L.; Leonard, K. J. (2011) . "Stem rust of wheat (black rust)".

2275: 2202: 2032: 1896: 1576: 1471: 1453:, requiring two hosts to complete its life cycle – the cereal host and the 1450: 1413: 1365: 1217:

A.L. Guyot, Massenot & Saccas ex Z. Urb., Česká Mykol. 21(1): 14 (1967)

879: 835: 572: 405: 369: 330:, first explained the stem rust fungus in wheat in 1767. Italian scientist 274:

in 2016, suggesting that the disease is returning to Europe. Comprehensive

112: 2597:"Foliar Fungicides to Control Wheat Stem Rust, Race TTKS (Ug99), in Kenya" 5554: 5500: 5435: 5355: 4080:

Germplasm Health Webinar series. Vol. Phytosanitary Awareness Week.

3272: 2999: 2369:

Singh, Ravi P.; Hodson, David; Huerta-Espino, Julio; et al. (2008).

2141:

came first, followed by the selective pressure of widespread adoption of

1696: 1596: 1591: 1445:(it colonizes living plant cells) and has a complex life cycle featuring 1391: 1313: 1285:

Sowerby, Col. fig. Engl. Fung. Mushr. (London) 2(no. 13): tab. 140 (1799)

851: 704: 390: 381: 260: 221: 20: 5154:

Agroterrorism: National Defense Assessment, Strategies, and Capabilities

4560:

Roelfs, A P (1988). "Genetic Control of Phenotypes in Wheat Stem Rust".

1205:

A.L. Guyot, Massenot & Saccas, Ann. Ec. Agric. Grignon 5: 145 (1946)

1041:(Erikss. & Henn.) Kuntze, Revis. gen. pl. (Leipzig) 3(3): 470 (1898) 605: 270:

An outbreak of another virulent race of stem rust, TTTTF, took place in

5427: 5177: 4343: 4278: 3927: 3872: 3719: 3599: 3436: 3343:

that Caused a Wheat Stem Rust Epidemic in Southern Ethiopia in 2013–14"

3083: 2303: 2122: 2024: 1615: 1553: 1395: 1353: 579:

stages and alternate between two hosts. Wheat is the primary host, and

5440: 1277:(Pers. ex J.F. Gmel.) Gray, Nat. Arr. Brit. Pl. (London) 1: 534 (1821) 5492: 5396: 5278: 2663:"Potential for re-emergence of wheat stem rust in the United Kingdom" 2283: 2265: 2008: 1968: 1774:

There has been significant uptake of resistant wheat varieties among

1753: 1723: 1568: 1546: 1541:

on the cereal host 1–2 weeks after infection. The urediniospores are

939:

samples from wheat in 1991 in the United States. Unusually severe in

859: 843: 839: 397: 301: 237: 233: 62: 37: 5317: 5160:. U.S. Air Force Center for Strategic Deterrence Studies. p. 9. 2893: 2872: 5487: 5401: 5340: 5284: 4964: 2648: 2073: 2004: 1996: 1900: 1775: 1604: 1600: 1583: 1557: 1542: 1538: 1459: 1433: 1387: 916: 737: 692: 680: 580: 248: 122: 4366: 1946:

alleles, with some conferring resistance and some susceptibility.

1319:

Interference with plant vascular tissue leads to shriveled grains.

1001:

Rebent., Prodr. fl. neomarch. (Berolini): 352, tab. 3:11a-b (1804)

4250: 4067: 1929: 1925: 1858: 1587: 1572: 1465: 1009:(Pers. ex J.F. Gmel.) Schltdl., Fl. berol. (Berlin) 2: 112 (1824) 806: 781:

Synonymous with QCCJB or known as QCC-2 by some classifications.

664: 656: 305: 275: 5388: 4656:

Johnson, R (1984). "A Critical Analysis of Durable Resistance".

2714:"Tackling the re-emergence of wheat stem rust in Western Europe" 2039:

has become less common in CIMMYT triticales since the mid-'80s.

1695:(Sr genes) have been identified in wheat. Some of them arose in 4346: 2184: 2081: 1692: 1611: 1372: 1081:

Barclay, J. Asiat. Soc. Bengal, Pt. 2, Nat. Sci. 58: 246 (1889)

1049:(Arthur) Arthur, Résult. Sci. Congr. Bot. Wien 1905: 344 (1906) 760: 696: 672: 554: 533: 492: 401: 312: 271: 252: 229: 217: 208: 72: 4290: 4237:"Mapping of Ug99 stem rust resistance in Canadian durum wheat" 3023: 2594: 404:

origin, and in 2018 DNA analysis confirmed that, specifically

5186: 4339: 4274: 4085: 4077: 4011: 3923: 3868: 3413:

Roelfs, A. P.; Long, D. L.; Roberts, J. J. (1993). "Races of

3079: 2016: 1779: 1708: 1361: 764: 733: 688: 684: 676: 576: 568: 550: 428: 316: 225: 220:

crops. Crop species that are affected by the disease include

4869: 4696: 3335:"Phenotypic and Genotypic Characterization of Race TKTTF of 2457: 2053:, which was effective against all previous stem rust races. 4001: 3999: 3827: 3801:

Journal of the Department of Agriculture, Western Australia

3408: 3406: 3404: 3402: 3400: 2295: 2249: 2085: 1749: 1416:

in the United States, epidemics in crops have become rare.

1378:

The site of infection is a visible symptom of the disease.

1357: 1025:(Fuckel) Kuntze, Revis. gen. pl. (Leipzig) 3(3): 467 (1898) 700: 652: 256: 244: 5263: 3578:

Peterson, Ronald H. (1974). "The rust fungus life cycle".

3470: 3468: 3466: 3464: 3462: 2711: 2660: 1999:

and slight degree of sporulation commonly allowed by most

1305:

Stem rust of cereals causes yield losses in several ways:

4861: 4859: 4857: 4855: 4776: 4448: 4027: 1960: 1732: 1483:) is considered to be the most important alternate host. 981:

Pers. ex J.F. Gmel., Syst. Nat., Edn 13 2(2): 1473 (1792)

695:

of stem rust on wheat caused by race TTKSK spread across

529: 466: 368:, a Canadian pathologist, identified the function of the 251:

of stem rust on wheat caused by race TTKSK spread across

5233: 4167: 3996: 3939: 3397: 3331: 3019: 3017: 3015: 3013: 3011: 3009: 2904:

that modulate barley rpg4-mediated stem rust resistance"

1786:(the International Maize and Wheat Improvement Center). 1778:

farmers since 2014 – a great deal of which is thanks to

1253:

Speg., Anal. Mus. nac. Hist. nat. B. Aires 6: 224 (1898)

1033:(Pers.) Gray, Nat. Arr. Brit. Pl. (London) 1: 542 (1821) 4949:

Sinha, Arunima; Singh, Lovepreet; Rawat, Nidhi (2022).

4708:

Genes, Encodes Resistance to Wheat Stem Rust Race Ug99"

4106: 3459: 2853:

Park, R. F. (2007). "Stem rust of wheat in Australia".

2784: 2762: 2646:

Kaplan, Karen (July 22, 2009) "A red alert for wheat".

2145:

wheat races, followed by the evolution of virulence on

2084:. Confers broad resistance to multiple races including 4852: 4599: 4597: 4526: 4524: 4522: 4520: 4518: 4055: 4043:(the International Maize and Wheat Improvement Center) 3620: 1610:

This dikaryotic mycelium then forms structures called

915:

s on wheat in 1991. Traces found growing in northwest

5086: 4690: 4395: 4393: 3006: 1121:

G.H. Otth, Mitt. naturf. Ges. Bern 531-552: 88 (1864)

1057:

G.H. Otth, Mitt. naturf. Ges. Bern 531-552: 88 (1864)

675:) and the year of its discovery (1999). It spread to 300:

Stem rust has been an ongoing problem dating back to

4530: 4442: 4036:"Shining a brighter light on adoption and diffusion" 3614: 2031:. Four years later, in 1988, virulence was found in 992:

Rebent., Prodr. fl. neomarch. (Berolini): 352 (1804)

5210:

Centre for Agriculture and Bioscience International

5082: 5080: 4794:

American Association for the Advancement of Science

4722:

American Association for the Advancement of Science

4594: 4515: 4233: 4229: 4227: 4225: 3823: 3821: 3819: 3817: 3238: 2971: 1686: 1614:, which produce a type of dikaryotic spores called 590: 240:effective against stem rust are available as well. 4770: 4390: 4361: 4359: 3475:McVey, D. V.; Long, D. L.; Roberts, J. J. (2002). 3325: 3158:(International Maize and Wheat Improvement Center) 3148:"North American Stem Rust Nomenclature Code Sheet" 3026:"Vulnerability of Barley to African Pathotypes of 854:before the 1991 season resulted in an epidemic in 337:Lettera al dottor Cosimo Moschettini sulla ruggine 3933: 3792: 3412: 2967: 2965: 2963: 2887: 1245:Röhl., Deutschl. Fl. (Frankfurt) 3(3): 132 (1813) 1089:Fuckel, Jb. nassau. Ver. Naturk. 27-28: 15 (1874) 1065:C.-J. Duval, in Hoppe, Bot. Taschenb.: 257 (1793) 452:, which vary in host range have been identified. 400:were found and for decades one hypothesis was an 5592: 5077: 4284: 4222: 3814: 3474: 3203: 2488:"Wheat disease a threat to global food security" 2371:"Will Stem Rust Destroy The World's Wheat Crop?" 2121:gene so named because it was discovered causing 1261:Erikss. & Henn., Z. PflKrankh. 4: 140 (1894) 1017:(Barclay) Syd., Annls mycol. 20(3/4): 117 (1922) 822:in 1991. 67% of QCCJ was from barley and 95% of 804:. Thought to be responsible for rusting of wild 663:and now has proliferated into a large number of 4948: 4651: 4649: 4356: 4082:International Institute of Tropical Agriculture 3641:Proceedings of the National Academy of Sciences 3304: 3302: 3300: 3298: 1564:is able to overwinter independently of a host. 1343:(Infected and uninfected depending on specific 691:, and becomes more virulent as it spreads. An 5193: 3768: 3171: 2960: 2780: 2778: 2776: 2015:virulent on wheat with this gene was found in 1621: 1113:Schumach., Enum. pl. (Kjbenhavn) 2: 233 (1801) 5189:'s Research Program on Rice). pp. 38–39. 5014: 4603: 4531:McIntosh, RA; Wellings, CR; Park, RF (1995). 3699: 3697: 3695: 3567:Animated video of the life cycle of stem rust 3311:"Ug99 Stem Rust – Breaching Wheat's Defences" 3244: 2481: 2479: 2477: 2453: 2451: 2449: 2447: 2445: 2443: 2441: 2439: 2437: 1707:), while others have been bred in from other 1475:) that are susceptible to stem rust, but the 1269:Arthur, Bull. Torrey bot. Club 28: 663 (1901) 1105:H. Mart., Prodr. Fl. Mosq., Edn 2: 227 (1817) 335: 5068:National Natural Science Foundation of China 4646: 4399: 3295: 3140: 2848: 2846: 2844: 2543: 2435: 2433: 2431: 2429: 2427: 2425: 2423: 2421: 2419: 2417: 2149:by way of nonfunctionalization mutations of 1423: 1371:Towards the end of the growing season black 1145:Mayor, Revue Mycol., Paris 22(3): 273 (1957) 1129:Gäum., Ber. schweiz. bot. Ges. 55: 79 (1945) 1097:T.S. Ramakr., Indian Phytopath. 6: 30 (1954) 1073:Mayor, Revue Mycol., Paris 22(3): 278 (1957) 4553: 3115: 2856:Australian Journal of Agricultural Research 2773: 2642: 2640: 2638: 2636: 2634: 2632: 5170: 5150: 4533:Wheat Rusts - An Atlas of Resistance Genes 3888: 3886: 3884: 3882: 3692: 3571: 3481:in the United States During 1997 and 1998" 2474: 2364: 2362: 2360: 2242:Originally from the widespread Ethiopian ' 2163:Recently, a new stem rust resistance gene 36: 5249:Cereal Rusts and Powdery Mildews Bulletin 5111: 4980: 4922: 4827: 4809: 4737: 4315: 4258: 4195: 4134: 4115:in the Bread Wheat Cultivar 'Matlabas'", 3979: 3735: 3675: 3657: 3502: 3364: 3055: 2943: 2925: 2841: 2816: 2739: 2688: 2612: 2414: 2358: 2356: 2354: 2352: 2350: 2348: 2346: 2344: 2342: 2340: 1995:s that does not allow the underdeveloped 1640:spore types that are known for rust fungi 1525: 1487:is macrocyclic (exhibits all five of the 1237:Ellis & Barthol., Erythea 4: 2 (1896) 814:and Great Plains, and overall was 94% of 396:In 1969 two races not detected before in 4064:Lebanese Agricultural Research Institute 3577: 2629: 2197:from that species. Was discovered by Yu 1534:is arbitrarily chosen as a start point. 1335: 947:, in 1991. TPMK was the worst at 69% of 422: 5199: 5164: 5028:Springer Science and Business Media LLC 4891:Springer Science and Business Media LLC 4655: 3879: 3308: 3221:United States Department of Agriculture 2918:Springer Science and Business Media LLC 2485: 1556:. These black, thick-walled spores are 943:in the first week of June, and in west 5593: 5202:"Does rust-free rice hold the secret?" 4559: 4033: 3776:Sounding the Alarm on Global Stem Rust 2337: 1942:There is considerable variation among 1169:Z. Urb., Česká Mykol. 21(1): 14 (1967) 830:. Continues to threaten barley in the 216:, which causes significant disease in 5322: 5321: 5183:International Rice Research Institute 3703: 3089: 1721:) or from other members of the tribe 1326: 882:in 1988, and by 1990 was over 90% of 5542:155d2162-bd25-401d-8b82-00b695e834a6 4299:Molecular Plant-Microbe Interactions 2852: 2539: 2537: 2019:in 1972. Deployment in triticale in 1967:), now (as of 2021) widely found in 998:Aecidium berberidis var. cylindricum 711:. In 2011, after it had spread into 600: 4676:10.1146/annurev.py.22.090184.001521 4580:10.1146/annurev.py.26.090188.002031 4420:10.1146/annurev-phyto-072910-095405 4242:Canadian Journal of Plant Pathology 4171:Canadian Journal of Plant Pathology 3910:10.1146/annurev-phyto-072910-095423 3849:10.1146/annurev-phyto-072910-095423 3417:in the United States During 1991". 3319:British Society for Plant Pathology 2496:Australian Broadcasting Corporation 1924:does not protect seedlings against 1575:to form four haploid spores called 13: 5227: 5104:American Phytopathological Society 4308:American Phytopathological Society 4127:American Phytopathological Society 4096: 3495:American Phytopathological Society 3429:American Phytopathological Society 3357:American Phytopathological Society 3265:American Phytopathological Society 3048:American Phytopathological Society 2992:American Phytopathological Society 2809:American Phytopathological Society 2795:from Southern Africa to Australia" 2228:which are common among plants. Yu 1630: 1560:. They are the only form in which 14: 5622: 5279:Animation of stem rust life cycle 5273:Food and Agriculture Organization 5257: 5200:Cameron, Katherine (2011-02-28). 3185:Food and Agriculture Organization 2534: 2486:Vincent, Michael (1 March 2011). 2129:– is the ancestral allele to all 1651:Extremely long-distance dispersal 1599:, leading to the production of a 4956:Current Opinion in Plant Biology 4092:from the original on 2021-12-14. 3543:"Species Fungorum - GSD Species" 3179:"Rust - Stem: Race Nomenclature" 2763:Giovene, Giuseppe Maria (1839). 2259: 1883:Discovered and found to provide 1687:Wheat stem rust resistance genes 1491:that are known for rust fungi). 919:fields, also in 1991. 8% of all 604: 49: 5144: 4659:Annual Review of Phytopathology 4607:Annual Review of Phytopathology 4563:Annual Review of Phytopathology 4403:Annual Review of Phytopathology 3897:Annual Review of Phytopathology 3832:Annual Review of Phytopathology 3560: 3535: 3309:Hawkins, Nichola (2021-01-20). 2756: 5264:Borlaug Global Rust Initiative 4624:10.1146/annurev.phyto.35.1.311 4376:Borlaug Global Rust Initiative 3783:Borlaug Global Rust Initiative 2705: 2654: 2588: 2509: 1381: 16:Fungus disease of cereal crops 1: 4260:10.1080/07060661.2020.1843073 4197:10.1080/07060661.2021.1902860 4034:Yasabu, Simret (2020-12-15). 3807:(8): Article 7. p. 365: 3216:Agricultural Research Service 2522:Plant diseases identification 2515: 2387:10.1016/S0065-2113(08)00205-8 2330: 2282:bombers would drop 500-pound 2133:alleles that are virulent on 1672:Extinction and recolonisation 1419: 1316:and infection by other fungi. 759:Successfully overwintered in 446:, and several special forms, 179: 5181:. Vol. 10, no. 1. 3650:National Academy of Sciences 2278:. If the plan were enacted, 2113:conferring some resistance. 1457:. There are many species in 1291: 7: 5113:10.1094/phyto-01-15-0030-fi 5064:Chinese Academy of Sciences 5019:Science China Life Sciences 3366:10.1094/phyto-11-14-0302-fi 2460:The Plant Health Instructor 2309: 1857:are no longer effective in 1622:Life cycle without barberry 1331: 1157:Baudyš, Lotos 64: 29 (1916) 963: 433:Botanical Museum Greifswald 418: 10: 5627: 5171:McCandless, Linda (2011). 4907:10.1038/s41467-022-29132-8 4485:10.1038/s41477-021-00971-5 4136:10.1094/pdis-10-18-1731-re 3964:10.1038/s41598-020-75181-8 3626:Melampsora larici-populina 3504:10.1094/pdis.2002.86.6.568 3057:10.1094/phyto-11-16-0400-r 3034:and Sources of Resistance" 2818:10.1094/phyto-04-18-0110-r 2468:10.1094/PHI-I-2000-0721-01 2263: 2253: 1752:lineage. Notably Ug99 has 1567:Each teliospore undergoes 1447:alternation of generations 732:JRCQC is a race affecting 563:is a member of the phylum 287: 18: 5330: 5036:10.1007/s11427-022-2178-7 4982:10.1016/j.pbi.2022.102247 4317:10.1094/mpmi-01-20-0018-r 3737:10.1016/j.cub.2013.09.034 2927:10.1186/s12864-019-6369-7 2769:. Cannone. pp. 152–. 2732:10.1038/s42003-019-0294-9 2681:10.1038/s42003-018-0013-y 2566:10.1038/nature.2017.21424 2289: 2280:Boeing B-29 Superfortress 2049:Ug99 is virulent against 1662:Step-wise range expansion 1638:produces all five of the 924: 858:and northwestern Kansas. 850:and the adjacent part of 334:(1753–1837), in his work 176: 169: 151: 144: 46:Scientific classification 44: 35: 30: 4704:, an Ortholog of Barley 2235: 1469:(and their hybrid genus 1258:Puccinia phlei-pratensis 1038:Dicaeoma phlei-pratensis 873: 812:Midwestern United States 727: 5611:Fungi described in 1794 4811:10.1126/science.aao7294 4739:10.1126/science.1239028 3704:Gross, Michael (2013). 3659:10.1073/pnas.1019315108 3547:www.speciesfungorum.org 2294:Alone amongst cereals, 2272:United States Air Force 2176: 2156: 2093: 2056: 2042: 1949: 1935: 1914: 1876: 1571:(fusion of nuclei) and 1250:Puccinia megalopotamica 1094:Puccinia brizae-maximae 930: 906: 898: 862:against barley bearing 776: 643: 583:is the alternate host. 413:breeding for resistance 348:, and in 1854 brothers 295:Oregon State University 4088:. Slide at 00:44:37. 3315:Plant Pandemic Studies 2719:Communications Biology 2668:Communications Biology 2614:10.1094/PDIS-93-9-0929 2183:An NLR (or NB-LRR, or 1742:Thinopyrum intermedium 1691:A number of stem rust 1526:Life cycle on barberry 1428: 1348: 848:South Central Oklahoma 751: 743: 438:There is considerable 435: 336: 332:Giuseppe Maria Giovene 5275:of the United Nations 4882:Nature Communications 4253:UK Limited: 599–611. 3211:"Race identification" 3123:"Wheat Rust Diseases" 2306:could be the future. 2224:which is composed of 2222:tandem protein kinase 2003:s. Instead there are 1977:and in an artificial 1891:, 2014. However Ug99 1508:obligately biotrophic 1427: 1339: 1078:Puccinia anthistiriae 1062:Lycoperdon berberidis 1014:Dicaeoma anthistiriae 842:. Higher than normal 717:Bill Gates Foundation 571:. The characteristic 431:, late 19th century, 426: 5285:"Norwich Rust Group" 4180:Taylor & Francis 3580:The Botanical Review 3273:10.1094/phyto-78-526 3000:10.1094/Phyto-84-452 2375:Advances in Agronomy 2193:, one of only three 2190:Aegilops sharonensis 1513:convergent evolution 1274:Roestelia berberidis 1086:Puccinia anthoxanthi 1022:Dicaeoma anthoxanthi 788:race in 1991 in the 5013: • 4973:2022COPB...6802247S 4947: • 4899:2022NatCo..13.1607Y 4868: • 4802:2017Sci...358.1604S 4796:(AAAS): 1604–1606. 4730:2013Sci...341..786P 4477:2021NatPl...7.1220U 4344:0000-0002-8952-5684 4279:0000-0002-4091-2608 4188:2021CaJPP..43S.275F 4166: • 4105: • 3928:0000-0002-4091-2608 3873:0000-0002-4091-2608 3728:2013CBio...23.R855G 3706:"Pests on the move" 3592:1974BotRv..40..453P 3097:"Red Rust of Wheat" 3084:0000-0002-4091-2608 2558:2017Natur.542..145B 2316:Chilean wheat cycle 2110:Triticum monococcum 1987:. When successful, 1959:is originally from 1126:Puccinia dactylidis 1118:Puccinia dactylidis 1070:Puccinia albigensis 986:Aecidium berberidis 978:Aecidium berberidis 763:in 1989/90, and in 567:within the kingdom 442:within the species 207:, is caused by the 5290:Norwich Rust Group 5206:The Plantwise Blog 5173:"No rust for rice" 4129:(APS): 2337–2344, 4016:. 10 November 2020 3947:Scientific Reports 3600:10.1007/BF02860021 3437:10.1094/pd-77-0129 3101:Rainy River Record 2270:In the 1950s, the 2067:from a wild wheat 1429: 1412:from the northern 1349: 1327:Signs and symptoms 1142:Puccinia favargeri 927:suddenly in 1998. 616:. You can help by 436: 5588: 5587: 5578:puccinia-graminis 5563:Open Tree of Life 5362:Puccinia graminis 5332:Puccinia graminis 5324:Taxon identifiers 5236:Puccinia graminis 4109:Puccinia graminis 3630:Puccinia graminis 3479:Puccinia graminis 3415:Puccinia graminis 3337:Puccinia graminis 3247:Puccinia graminis 3028:Puccinia graminis 2974:Puccinia graminis 2898:Puccinia graminis 2789:Puccinia graminis 2552:(7640): 145–146. 2302:with rice as the 2300:transgenic wheats 2252:, susceptible to 2070:Aegilops tauschii 1991:is among the few 1679:Puccinia graminis 1636:Puccinia graminis 1562:Puccinia graminis 1511:broad pattern of 1443:obligate biotroph 1386:Pycnia appear on 1341:Race differential 1242:Puccinia linearis 1222:Puccinia graminis 1210:Puccinia graminis 1198:Puccinia graminis 1186:Puccinia graminis 1174:Puccinia graminis 1162:Puccinia graminis 1150:Puccinia graminis 1110:Puccinia culmorum 1102:Puccinia cerealis 1054:Epitea dactylidis 1030:Dicaeoma graminis 1006:Caeoma berberidis 941:southern Illinois 866:. Down to 26% of 655:began as a race ( 634: 633: 592:Puccinia graminis 539:Puccinia graminis 521:Puccinia graminis 510:Puccinia graminis 499:Puccinia graminis 484:Puccinia graminis 473:Puccinia graminis 458:Puccinia graminis 440:genetic diversity 342:Puccinia graminis 280:Puccinia graminis 213:Puccinia graminis 186: 185: 155:Puccinia graminis 5618: 5581: 5580: 5571: 5570: 5558: 5557: 5545: 5544: 5535: 5534: 5522: 5521: 5519:NHMSYS0001495612 5509: 5508: 5496: 5495: 5483: 5482: 5470: 5469: 5457: 5456: 5444: 5443: 5431: 5430: 5418: 5417: 5405: 5404: 5392: 5391: 5379: 5378: 5366: 5365: 5364: 5351: 5350: 5349: 5319: 5318: 5314: 5312: 5311: 5300: 5298: 5297: 5253: 5221: 5220: 5218: 5217: 5197: 5191: 5190: 5168: 5162: 5161: 5159: 5148: 5142: 5141: 5115: 5106:(APS): 872–884. 5084: 5075: 5071: 5010: 4984: 4944: 4926: 4863: 4850: 4849: 4831: 4813: 4774: 4768: 4767: 4741: 4694: 4688: 4687: 4653: 4644: 4643: 4601: 4592: 4591: 4557: 4551: 4550: 4528: 4513: 4512: 4456: 4446: 4440: 4439: 4397: 4388: 4387: 4385: 4384: 4363: 4354: 4337: 4319: 4288: 4282: 4272: 4262: 4231: 4220: 4216: 4199: 4163: 4138: 4100: 4094: 4093: 4070:) (2020-11-09). 4059: 4053: 4052: 4050: 4049: 4031: 4025: 4024: 4022: 4021: 4003: 3994: 3993: 3983: 3937: 3931: 3921: 3890: 3877: 3876: 3825: 3812: 3811: 3796: 3790: 3789: 3780: 3772: 3766: 3765: 3739: 3701: 3690: 3689: 3679: 3661: 3618: 3612: 3611: 3575: 3569: 3564: 3558: 3557: 3555: 3553: 3539: 3533: 3532: 3506: 3472: 3457: 3456: 3410: 3395: 3394: 3368: 3329: 3323: 3322: 3306: 3293: 3292: 3242: 3236: 3235: 3233: 3232: 3223:. Archived from 3207: 3201: 3200: 3198: 3197: 3188:. Archived from 3175: 3169: 3168: 3166: 3160:. Archived from 3152: 3144: 3138: 3137: 3135: 3133: 3119: 3113: 3112: 3110: 3108: 3093: 3087: 3077: 3059: 3021: 3004: 3003: 2969: 2958: 2957: 2947: 2929: 2891: 2885: 2884: 2865:CSIRO Publishing 2850: 2839: 2838: 2820: 2782: 2771: 2770: 2760: 2754: 2753: 2743: 2709: 2703: 2702: 2692: 2658: 2652: 2644: 2627: 2626: 2616: 2592: 2586: 2585: 2541: 2532: 2531: 2529: 2528: 2513: 2507: 2506: 2504: 2502: 2483: 2472: 2471: 2455: 2412: 2411: 2409: 2408: 2399:. Archived from 2366: 2248:'. Resistant to 2247: 2246: 2201:, 2017 and then 2195:resistance genes 1983: 1693:resistance genes 1517:plant pathogenic 1505: 1485:P. graminis 1449:. The fungus is 1439:P. graminis 1407: 1406: 1402: 1345:resistance genes 1286: 1278: 1270: 1262: 1254: 1246: 1238: 1230: 1218: 1206: 1194: 1182: 1170: 1158: 1146: 1138: 1134:Puccinia elymina 1130: 1122: 1114: 1106: 1098: 1090: 1082: 1074: 1066: 1058: 1050: 1042: 1034: 1026: 1018: 1010: 1002: 993: 982: 970:Species Fungorum 832:Red River Valley 756:Affects barley. 748:Affects barley. 665:races of its own 629: 626: 608: 601: 444:P. graminis 339: 191:, also known as 157: 137:P. graminis 54: 53: 40: 28: 27: 5626: 5625: 5621: 5620: 5619: 5617: 5616: 5615: 5601:Cereal diseases 5591: 5590: 5589: 5584: 5576: 5574: 5566: 5561: 5553: 5550:Observation.org 5548: 5540: 5538: 5530: 5525: 5517: 5512: 5504: 5499: 5491: 5486: 5478: 5473: 5465: 5460: 5452: 5447: 5439: 5434: 5426: 5421: 5413: 5408: 5400: 5395: 5387: 5382: 5374: 5369: 5360: 5359: 5354: 5345: 5344: 5339: 5326: 5309: 5307: 5303: 5295: 5293: 5283: 5260: 5230: 5228:Further reading 5225: 5224: 5215: 5213: 5198: 5194: 5169: 5165: 5157: 5149: 5145: 5085: 5078: 5074: 4864: 4853: 4775: 4771: 4695: 4691: 4654: 4647: 4602: 4595: 4558: 4554: 4547: 4529: 4516: 4469:Nature Research 4454: 4447: 4443: 4398: 4391: 4382: 4380: 4365: 4364: 4357: 4289: 4285: 4232: 4223: 4219: 4101: 4097: 4060: 4056: 4047: 4045: 4032: 4028: 4019: 4017: 4005: 4004: 3997: 3956:Springer Nature 3938: 3934: 3891: 3880: 3826: 3815: 3797: 3793: 3778: 3774: 3773: 3769: 3711:Current Biology 3702: 3693: 3619: 3615: 3576: 3572: 3565: 3561: 3551: 3549: 3541: 3540: 3536: 3473: 3460: 3411: 3398: 3330: 3326: 3307: 3296: 3243: 3239: 3230: 3228: 3209: 3208: 3204: 3195: 3193: 3183:United Nations 3177: 3176: 3172: 3164: 3150: 3146: 3145: 3141: 3131: 3129: 3127:Crop Science US 3121: 3120: 3116: 3106: 3104: 3095: 3094: 3090: 3022: 3007: 2970: 2961: 2892: 2888: 2873:10.1071/ar07117 2851: 2842: 2783: 2774: 2761: 2757: 2710: 2706: 2659: 2655: 2645: 2630: 2593: 2589: 2542: 2535: 2526: 2524: 2514: 2510: 2500: 2498: 2484: 2475: 2456: 2415: 2406: 2404: 2397: 2367: 2338: 2333: 2312: 2292: 2268: 2262: 2244: 2243: 2240: 2181: 2161: 2098: 2061: 2047: 2021:New South Wales 1981: 1954: 1940: 1919: 1885:Ug99 resistance 1881: 1689: 1633: 1631:Spore dispersal 1624: 1528: 1503: 1477:common barberry 1422: 1404: 1400: 1399: 1384: 1334: 1329: 1294: 1289: 1284: 1276: 1268: 1260: 1252: 1244: 1236: 1234:Puccinia jubata 1228: 1216: 1204: 1192: 1190:phlei-pratensis 1180: 1168: 1156: 1144: 1136: 1128: 1120: 1112: 1104: 1096: 1088: 1080: 1072: 1064: 1056: 1048: 1040: 1032: 1024: 1016: 1008: 1000: 991: 980: 966: 945:central Indiana 933: 909: 901: 876: 779: 754: 746: 730: 713:southern Africa 646: 638:resistance gene 630: 624: 621: 614:needs expansion 599: 449:forma specialis 421: 354:Charles Tulasne 290: 180:§ Synonyms 165: 159: 153: 140: 93:Pucciniomycetes 48: 24: 17: 12: 11: 5: 5624: 5614: 5613: 5608: 5603: 5586: 5585: 5583: 5582: 5572: 5559: 5546: 5536: 5523: 5510: 5497: 5484: 5471: 5458: 5445: 5432: 5419: 5406: 5393: 5380: 5367: 5352: 5336: 5334: 5328: 5327: 5316: 5315: 5301: 5281: 5276: 5266: 5259: 5258:External links 5256: 5255: 5254: 5229: 5226: 5223: 5222: 5192: 5163: 5143: 5095:Phytopathology 5076: 5073: 5072: 5011: 4945: 4865: 4851: 4769: 4689: 4668:Annual Reviews 4645: 4616:Annual Reviews 4593: 4572:Annual Reviews 4552: 4545: 4514: 4441: 4412:Annual Reviews 4389: 4355: 4283: 4221: 4218: 4217: 4164: 4102: 4095: 4062:Rola El Amil ( 4054: 4026: 3995: 3932: 3904:(1): 465–481. 3878: 3841:Annual Reviews 3813: 3791: 3767: 3691: 3613: 3586:(4): 453–513. 3570: 3559: 3534: 3458: 3396: 3348:Phytopathology 3324: 3294: 3256:Phytopathology 3237: 3202: 3170: 3167:on 2021-04-03. 3139: 3114: 3088: 3039:Phytopathology 3005: 2983:Phytopathology 2959: 2886: 2840: 2800:Phytopathology 2772: 2755: 2704: 2653: 2628: 2607:(9): 929–932. 2587: 2533: 2508: 2473: 2413: 2395: 2335: 2334: 2332: 2329: 2328: 2327: 2318: 2311: 2308: 2291: 2288: 2264:Main article: 2261: 2258: 2239: 2234: 2180: 2175: 2170:Secale cereale 2160: 2155: 2097: 2092: 2060: 2055: 2046: 2041: 1953: 1948: 1939: 1934: 1918: 1913: 1909:breeding lines 1880: 1875: 1767:Secale cereale 1711:species (e.g. 1688: 1685: 1675: 1674: 1665: 1664: 1654: 1653: 1632: 1629: 1623: 1620: 1532:urediniospores 1527: 1524: 1472:x Mahoberberis 1455:alternate host 1421: 1418: 1410:alternate host 1383: 1380: 1333: 1330: 1328: 1325: 1324: 1323: 1320: 1317: 1310: 1293: 1290: 1288: 1287: 1282:Uredo frumenti 1279: 1271: 1266:Puccinia vilis 1263: 1255: 1247: 1239: 1231: 1219: 1207: 1195: 1183: 1171: 1159: 1147: 1139: 1131: 1123: 1115: 1107: 1099: 1091: 1083: 1075: 1067: 1059: 1051: 1046:Dicaeoma vilis 1043: 1035: 1027: 1019: 1011: 1003: 995: 983: 974: 965: 962: 932: 929: 908: 905: 900: 897: 875: 872: 846:production in 798:southern Texas 778: 775: 753: 750: 745: 742: 729: 726: 645: 642: 632: 631: 611: 609: 598: 589: 558: 557: 536: 517: 506: 495: 480: 469: 420: 417: 289: 286: 184: 183: 174: 173: 167: 166: 160: 149: 148: 142: 141: 134: 132: 128: 127: 120: 116: 115: 110: 106: 105: 100: 96: 95: 90: 86: 85: 80: 76: 75: 70: 66: 65: 60: 56: 55: 42: 41: 33: 32: 15: 9: 6: 4: 3: 2: 5623: 5612: 5609: 5607: 5604: 5602: 5599: 5598: 5596: 5579: 5573: 5569: 5564: 5560: 5556: 5551: 5547: 5543: 5537: 5533: 5528: 5524: 5520: 5515: 5511: 5507: 5502: 5498: 5494: 5489: 5485: 5481: 5476: 5472: 5468: 5463: 5459: 5455: 5450: 5446: 5442: 5437: 5433: 5429: 5424: 5420: 5416: 5411: 5407: 5403: 5398: 5394: 5390: 5385: 5381: 5377: 5372: 5368: 5363: 5357: 5353: 5348: 5342: 5338: 5337: 5335: 5333: 5329: 5325: 5320: 5306: 5302: 5292: 5291: 5286: 5282: 5280: 5277: 5274: 5270: 5267: 5265: 5262: 5261: 5251: 5250: 5245: 5241: 5237: 5232: 5231: 5211: 5207: 5203: 5196: 5188: 5184: 5180: 5179: 5174: 5167: 5156: 5155: 5147: 5139: 5135: 5131: 5127: 5123: 5119: 5114: 5109: 5105: 5101: 5097: 5096: 5091: 5083: 5081: 5069: 5065: 5061: 5057: 5053: 5049: 5045: 5041: 5037: 5033: 5030:: 1718–1775. 5029: 5025: 5021: 5020: 5012: 5008: 5004: 5000: 4996: 4992: 4988: 4983: 4978: 4974: 4970: 4966: 4962: 4958: 4957: 4952: 4946: 4942: 4938: 4934: 4930: 4925: 4920: 4916: 4912: 4908: 4904: 4900: 4896: 4892: 4888: 4884: 4883: 4878: 4876: 4867: 4866: 4862: 4860: 4858: 4856: 4847: 4843: 4839: 4835: 4830: 4825: 4821: 4817: 4812: 4807: 4803: 4799: 4795: 4791: 4787: 4786: 4781: 4773: 4765: 4761: 4757: 4753: 4749: 4745: 4740: 4735: 4731: 4727: 4723: 4719: 4715: 4714: 4709: 4707: 4703: 4693: 4685: 4681: 4677: 4673: 4669: 4665: 4661: 4660: 4652: 4650: 4641: 4637: 4633: 4629: 4625: 4621: 4617: 4613: 4609: 4608: 4600: 4598: 4589: 4585: 4581: 4577: 4573: 4569: 4565: 4564: 4556: 4548: 4546:9789401040419 4542: 4538: 4534: 4527: 4525: 4523: 4521: 4519: 4510: 4506: 4502: 4498: 4494: 4490: 4486: 4482: 4478: 4474: 4471:: 1220–1228. 4470: 4466: 4462: 4461: 4460:Nature Plants 4453: 4445: 4437: 4433: 4429: 4425: 4421: 4417: 4413: 4409: 4405: 4404: 4396: 4394: 4379: 4377: 4372: 4370: 4362: 4360: 4352: 4348: 4345: 4341: 4335: 4331: 4327: 4323: 4318: 4313: 4310:: 1286–1298. 4309: 4305: 4301: 4300: 4295: 4287: 4280: 4276: 4270: 4266: 4261: 4256: 4252: 4248: 4244: 4243: 4238: 4230: 4228: 4226: 4215: 4211: 4207: 4203: 4198: 4193: 4189: 4185: 4182:: S275–S297, 4181: 4177: 4173: 4172: 4165: 4162: 4158: 4154: 4150: 4146: 4142: 4137: 4132: 4128: 4124: 4120: 4119: 4118:Plant Disease 4114: 4110: 4104: 4103: 4099: 4091: 4087: 4083: 4079: 4075: 4074: 4069: 4065: 4058: 4044: 4042: 4037: 4030: 4015: 4013: 4008: 4002: 4000: 3991: 3987: 3982: 3977: 3973: 3969: 3965: 3961: 3957: 3953: 3949: 3948: 3943: 3936: 3929: 3925: 3919: 3915: 3911: 3907: 3903: 3899: 3898: 3889: 3887: 3885: 3883: 3874: 3870: 3866: 3862: 3858: 3854: 3850: 3846: 3842: 3838: 3834: 3833: 3824: 3822: 3820: 3818: 3810: 3806: 3802: 3795: 3788: 3785:. p. 4: 3784: 3777: 3771: 3763: 3759: 3755: 3751: 3747: 3743: 3738: 3733: 3729: 3725: 3722:: R855–R857. 3721: 3717: 3713: 3712: 3707: 3700: 3698: 3696: 3687: 3683: 3678: 3673: 3669: 3665: 3660: 3655: 3652:: 9166–9171. 3651: 3647: 3643: 3642: 3637: 3635: 3631: 3627: 3617: 3609: 3605: 3601: 3597: 3593: 3589: 3585: 3581: 3574: 3568: 3563: 3548: 3544: 3538: 3530: 3526: 3522: 3518: 3514: 3510: 3505: 3500: 3496: 3492: 3488: 3487: 3486:Plant Disease 3482: 3480: 3471: 3469: 3467: 3465: 3463: 3454: 3450: 3446: 3442: 3438: 3434: 3430: 3426: 3422: 3421: 3420:Plant Disease 3416: 3409: 3407: 3405: 3403: 3401: 3392: 3388: 3384: 3380: 3376: 3372: 3367: 3362: 3358: 3354: 3350: 3349: 3344: 3342: 3338: 3328: 3320: 3316: 3312: 3305: 3303: 3301: 3299: 3290: 3286: 3282: 3278: 3274: 3270: 3266: 3262: 3258: 3257: 3252: 3248: 3241: 3227:on 2021-04-03 3226: 3222: 3218: 3217: 3212: 3206: 3192:on 2021-04-03 3191: 3187: 3186: 3180: 3174: 3163: 3159: 3157: 3149: 3143: 3128: 3124: 3118: 3103:. 15 May 2012 3102: 3098: 3092: 3085: 3081: 3075: 3071: 3067: 3063: 3058: 3053: 3049: 3045: 3041: 3040: 3035: 3033: 3029: 3020: 3018: 3016: 3014: 3012: 3010: 3001: 2997: 2993: 2989: 2985: 2984: 2979: 2975: 2968: 2966: 2964: 2955: 2951: 2946: 2941: 2937: 2933: 2928: 2923: 2919: 2915: 2911: 2910: 2905: 2903: 2899: 2890: 2882: 2878: 2874: 2870: 2866: 2862: 2858: 2857: 2849: 2847: 2845: 2836: 2832: 2828: 2824: 2819: 2814: 2810: 2806: 2802: 2801: 2796: 2794: 2790: 2781: 2779: 2777: 2768: 2767: 2759: 2751: 2747: 2742: 2737: 2733: 2729: 2725: 2721: 2720: 2715: 2708: 2700: 2696: 2691: 2686: 2682: 2678: 2674: 2670: 2669: 2664: 2657: 2651: 2650: 2643: 2641: 2639: 2637: 2635: 2633: 2624: 2620: 2615: 2610: 2606: 2602: 2601:Plant Disease 2598: 2591: 2583: 2579: 2575: 2571: 2567: 2563: 2559: 2555: 2551: 2547: 2540: 2538: 2523: 2519: 2512: 2497: 2493: 2489: 2482: 2480: 2478: 2469: 2465: 2461: 2454: 2452: 2450: 2448: 2446: 2444: 2442: 2440: 2438: 2436: 2434: 2432: 2430: 2428: 2426: 2424: 2422: 2420: 2418: 2403:on 2020-11-08 2402: 2398: 2396:9780123743558 2392: 2388: 2384: 2380: 2376: 2372: 2365: 2363: 2361: 2359: 2357: 2355: 2353: 2351: 2349: 2347: 2345: 2343: 2341: 2336: 2326: 2324: 2319: 2317: 2314: 2313: 2307: 2305: 2301: 2297: 2287: 2285: 2281: 2277: 2273: 2267: 2260:Weaponization 2257: 2255: 2251: 2238: 2233: 2231: 2227: 2223: 2220:for a unique 2219: 2216: 2212: 2208: 2204: 2200: 2196: 2192: 2191: 2186: 2179: 2174: 2172: 2171: 2166: 2159: 2154: 2152: 2148: 2144: 2140: 2136: 2132: 2128: 2124: 2120: 2116: 2112: 2111: 2106: 2105:introgression 2102: 2096: 2091: 2089: 2087: 2083: 2079: 2075: 2072: 2071: 2066: 2065:introgression 2059: 2054: 2052: 2045: 2040: 2038: 2034: 2030: 2026: 2022: 2018: 2014: 2010: 2006: 2002: 1998: 1994: 1990: 1986: 1980: 1976: 1975: 1970: 1966: 1962: 1958: 1952: 1947: 1945: 1938: 1933: 1931: 1927: 1923: 1917: 1912: 1910: 1906: 1902: 1898: 1894: 1890: 1886: 1879: 1874: 1872: 1868: 1864: 1860: 1856: 1852: 1848: 1844: 1840: 1836: 1832: 1828: 1824: 1820: 1816: 1812: 1808: 1804: 1800: 1796: 1792: 1787: 1785: 1781: 1777: 1772: 1769: 1768: 1763: 1759: 1755: 1751: 1746: 1744: 1743: 1738: 1734: 1730: 1726: 1725: 1720: 1719: 1718:T. monococcum 1714: 1710: 1706: 1702: 1698: 1694: 1684: 1682: 1680: 1673: 1670: 1669: 1668: 1663: 1660: 1659: 1658: 1652: 1649: 1648: 1647: 1643: 1641: 1637: 1628: 1619: 1617: 1613: 1608: 1606: 1602: 1598: 1593: 1589: 1585: 1580: 1578: 1577:basidiospores 1574: 1570: 1565: 1563: 1559: 1555: 1550: 1548: 1544: 1540: 1535: 1533: 1523: 1521: 1518: 1514: 1509: 1502: 1501:P. g. tritici 1498: 1496: 1492: 1490: 1486: 1482: 1478: 1474: 1473: 1468: 1467: 1462: 1461: 1456: 1452: 1448: 1444: 1440: 1436: 1435: 1426: 1417: 1415: 1411: 1397: 1393: 1389: 1379: 1376: 1374: 1369: 1367: 1363: 1359: 1355: 1346: 1342: 1338: 1321: 1318: 1315: 1311: 1308: 1307: 1306: 1303: 1300: 1283: 1280: 1275: 1272: 1267: 1264: 1259: 1256: 1251: 1248: 1243: 1240: 1235: 1232: 1227: 1223: 1220: 1215: 1211: 1208: 1203: 1199: 1196: 1191: 1187: 1184: 1179: 1175: 1172: 1167: 1163: 1160: 1155: 1151: 1148: 1143: 1140: 1135: 1132: 1127: 1124: 1119: 1116: 1111: 1108: 1103: 1100: 1095: 1092: 1087: 1084: 1079: 1076: 1071: 1068: 1063: 1060: 1055: 1052: 1047: 1044: 1039: 1036: 1031: 1028: 1023: 1020: 1015: 1012: 1007: 1004: 999: 996: 994: 987: 984: 979: 976: 975: 973: 971: 968:As listed by 961: 959: 955: 950: 946: 942: 938: 928: 926: 922: 918: 914: 904: 896: 894: 890: 885: 881: 871: 869: 865: 861: 857: 856:North Central 853: 849: 845: 841: 837: 833: 829: 825: 821: 817: 813: 809: 808: 803: 799: 795: 792:, 68% of all 791: 790:United States 787: 782: 774: 772: 770: 766: 762: 757: 749: 741: 739: 735: 725: 722: 718: 714: 710: 709:food security 706: 702: 698: 694: 690: 686: 682: 678: 674: 668: 666: 662: 658: 654: 650: 641: 639: 628: 619: 615: 612:This section 610: 607: 603: 602: 597: 593: 588: 584: 582: 578: 574: 570: 566: 565:Basidiomycota 562: 556: 552: 548: 544: 540: 537: 535: 531: 527: 526: 522: 518: 516: 515: 511: 507: 505: 504: 500: 496: 494: 490: 489: 485: 481: 479: 478: 474: 470: 468: 464: 463: 459: 455: 454: 453: 451: 450: 445: 441: 434: 430: 425: 416: 414: 409: 407: 406:South African 403: 399: 394: 392: 388: 387:United States 383: 378: 377:North America 373: 371: 367: 362: 361:Anton de Bary 358: 355: 351: 347: 343: 338: 333: 329: 325: 320: 318: 314: 311: 307: 303: 298: 296: 285: 283: 281: 277: 273: 268: 266: 265:food security 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