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Phytophthora infestans
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{{Short description|Species of single-celled organism}}{{Speciesbox| image = Late_blight_on_potato_leaf_2.jpg| image_alt = Symptom of late blight on the underside of a potato leaf| image_caption = Symptom of late blight on the underside of a potato leaf| genus = Phytophthora| species = infestans- the content below is remote from Wikipedia
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Jean Pierre François Camille Montagne>Mont.) de Bary| synonyms = | subdivision_ranks = | subdivision = }}Phytophthora infestans is an oomycete or water mold, a fungus-like microorganism that causes the serious potato and tomato disease known as late blight or potato blight. Early blight, caused by Alternaria solani, is also often called "potato blight". Late blight was a major culprit in the 1840s European, the 1845–1852 Irish, and the 1846 Highland potato famines. The organism can also infect some other members of the Solanaceae.{{Citation|url=http://news.bbc.co.uk/1/hi/sci/tech/8246944.stm|title=Killer genes cause potato famine |last=Chand|first=Sudeep |date=9 September 2009|publisher=BBC News|access-date=26 September 2009}}{{Citation|title=Potato and tomato late blight caused by Phytophthora infestans: An overview of pathology and resistance breeding |last=Nowicki|first=Marcin|date=17 August 2011|doi= 10.1094/PDIS-05-11-0458|pmid=30731850|display-authors=etal|volume=96|issue=1|journal=Plant Disease|publisher=American Phytopathological Society|pages=4–17|doi-access=}}{{Citation |last=Nowicki|first=Marcin |chapter= Late Blight of Tomato|title=Translational Genomics for Crop Breeding|date=11 October 2013|doi=10.1002/9781118728475.ch13|display-authors=etal |pages=241–65|isbn=978-1-118-72847-5|s2cid=83142160 }} The pathogen is favored by moist, cool environments: sporulation is optimal at {{convert|12–18|C|F|0}} in water-saturated or nearly saturated environments, and zoospore production is favored at temperatures below {{convert|15|C|F|0}}. Lesion growth rates are typically optimal at a slightly warmer temperature range of {{convert|20 to 24|C|F|0}}.{{Citation |last1=Haverkort |first1=A. J. |last2=Struik |first2=P. C. |last3=Visser |first3=R. G. F.|last4=Jacobsen |first4= E |year=2009 |title=Applied biotechnology to combat late blight in potato caused by Phytophthora infestans|journal=Potato Research|volume=52 |issue=3 |pages=249–64 |doi= 10.1007/s11540-009-9136-3|s2cid=2850128 |url=http://library.wur.nl/WebQuery/wurpubs/381896 |type=Submitted manuscript }}EtymologyThe genus name Phytophthora comes from the Greek ({{transliteration|el|phyto}}), meaning "plant" – plus the Greek ({{transliteration|el|phthora}}), meaning "decay, ruin, perish". The species name infestans is the present participle of the Latin verb , meaning "attacking, destroying", from which the word "to infest" is derived. The name Phytophthora infestans was coined in 1876 by the German mycologist Heinrich Anton de Bary (1831–1888).JOURNAL, Bary, A. de, Researches into the nature of the potato fungus Phytophthora infestans, Journal of the Royal Society of Agriculture of England, Royal Society of Agriculture of England, 1876, 12, 239–269,weblink 2nd series, WEB,weblink Taxonomy browser (Phytophthora infestans), www.ncbi.nlm.nih.gov,Life cycle, signs and symptomsmissing image!
- Tomato with Phytophthora infestans (late blight).jpg - frameless|alt=Infected ripe tomato missing image!
- Tomato late blight stem lesion 3 (5816739322).jpg - frameless|alt=Infected tomato plant missing image!
- Tomato late blight fruit cluster (5816739612).jpg - frameless|alt=Infected unripe tomatoes missing image!
File:07 08 life cycle, Phytophthora infestans on potato, Peronosporales, Oomycota (M. Piepenbring).png|right|thumb|Life cycleLife cycleThe asexual life cycle of Phytophthora infestans is characterized by alternating phases of hyphal growth, sporulation, sporangia germination (either through zoospore release or direct germination, i.e. germ tube emergence from the sporangium), and the re-establishment of hyphal growth.{{Citation|title=A simple dual stain for detailed investigations of plant-fungal pathogen interactions|last=Nowicki|first=Marcin|date=15 May 2013|doi= 10.2478/v10032-012-0016-z|display-authors=etal|volume=77|pages=61–74|journal=Vegetable Crops Research Bulletin|doi-access=free}} There is also a sexual cycle, which occurs when isolates of opposite mating type (A1 and A2, see {{Slink||Mating types}} below) meet. Hormonal communication triggers the formation of the sexual spores, called oospores.Judelson HS, Blanco FA (2005) The spores of Phytophthora: weapons of the plant destroyer. Nature Microbiology Reviews 3: 47–58. The different types of spores play major roles in the dissemination and survival of P. infestans. Sporangia are spread by wind or water and enable the movement of P. infestans between different host plants. The zoospores released from sporangia are biflagellated and chemotactic, allowing further movement of P. infestans on water films found on leaves or soils. Both sporangia and zoospores are short-lived, in contrast to oospores which can persist in a viable form for many years.People can observe P. infestans produce dark green, then brown then black spots on the surface of potato leaves and stems, often near the tips or edges, where water or dew collects.WEB, May 2017, Late Blight in Potato,weblink 2021-11-30, North Dakota State University Agriculture Department, en, The sporangia and sporangiophores appear white on the lower surface of the foliage. As for tuber blight, the white mycelium often shows on the tubers' surface.{{Citation |last1=Fry |first1=W. E. |last2=Grünwald |first2=N. J. |year=2010|title=Introduction to Oomycetes|journal=The Plant Health Instructor |doi=10.1094/PHI-I-2010-1207-01}}Under ideal conditions, P. infestans completes its life cycle on potato or tomato foliage in about five days. Sporangia develop on the leaves, spreading through the crop when temperatures are above {{convert|10|°C|°F}} and humidity is over 75–80% for 2 days or more. Rain can wash spores into the soil where they infect young tubers, and the spores can also travel long distances on the wind. The early stages of blight are easily missed. Symptoms include the appearance of dark blotches on leaf tips and plant stems. White mold will appear under the leaves in humid conditions and the whole plant may quickly collapse. Infected tubers develop grey or dark patches that are reddish brown beneath the skin, and quickly decay to a foul-smelling mush caused by the infestation of secondary soft bacterial rots. Seemingly healthy tubers may rot later when in store.P. infestans survives poorly in nature apart from on its plant hosts. Under most conditions, the hyphae and asexual sporangia can survive for only brief periods in plant debris or soil, and are generally killed off during frosts or very warm weather. The exceptions involve oospores, and hyphae present within tubers. The persistence of viable pathogen within tubers, such as those that are left in the ground after the previous year's harvest or left in cull piles is a major problem in disease management. In particular, volunteer plants sprouting from infected tubers are thought to be a major source of inoculum (or propagules) at the start of a growing season.{{citation |last1=Koepsell |first1=Paul A. |last2=Pscheidt |first2=Jay W. |title=1994 Pacific Northwest Plant Disease Control Handbook |year=1994|publisher=Oregon State University Press|location=Corvallis |page=165 }} This can have devastating effects by destroying entire crops.{{anchor|Mating type|Ia|IIa|Ib}}- Phytophtora infestans-effects.jpg - Infected potato|alt=Infected potatoes are shrunken on the outside, and corky as well as rotted on the inside. Mating typesThe mating types are broadly divided into A1 and A2.JOURNAL, Judelson, Howard S., Metabolic Diversity and Novelties in the Oomycetes, Annual Review of Microbiology, Annual Reviews (publisher), Annual Reviews, 71, 1, 2017-09-08, 0066-4227, 10.1146/annurev-micro-090816-093609, 21–39, 28504899, Until the 1980s populations could only be distinguished by virulence assays and mating types, but since then more detailed analysis has shown that mating type and genotype are substantially decoupled. These types each produce a mating hormone of their own. Pathogen populations are grouped into clonal lineages of these mating types and includes:A1A1 produces a mating hormone, a diterpene α1. Clonal lineages of A1 include:
A2Discovered by John Niederhauser in the 1950s, in the Toluca Valley in Central Mexico, while working for the Rockefeller Foundation's Mexican Agriculture Program. Published in Niederhauser 1956. A2 produces a mating hormone α2. Clonal lineages of A2 include:
Self-fertileA self-fertile type was present in China between 2009 and 2013.Physiology{{Vanchor|PiINF1|text=PiINF1}} is the {{Vanchor|INF1|text=INF1}} in P. infestans. Hosts respond with autophagy upon detection of this elicitor, Liu et al. 2005 finding this to be the only alternative to mass hypersensitivity leading to mass programmed cell death.JOURNAL, Yang, Meng, Ismayil, Asigul, Liu, Yule, Autophagy in Plant-Virus Interactions, Annual Review of Virology, Annual Reviews (publisher), Annual Reviews, 7, 1, 2020-09-29, 2327-056X, 10.1146/annurev-virology-010220-054709, 403–419, 32530794, 219621432,GeneticsP. infestans is diploid, with about 8–10 chromosomes, and in 2009 scientists completed the sequencing of its genome. The genome was found to be considerably larger (240 Mbp) than that of most other Phytophthora species whose genomes have been sequenced; P. sojae has a 95 Mbp genome and P. ramorum had a 65 Mbp genome. About 18,000 genes were detected within the P. infestans genome. It also contained a diverse variety of transposons and many gene families encoding for effector proteins that are involved in causing pathogenicity. These proteins are split into two main groups depending on whether they are produced by the water mold in the symplast (inside plant cells) or in the apoplast (between plant cells). Proteins produced in the symplast included RXLR proteins, which contain an arginine-X-leucine-arginine (where X can be any amino acid) sequence at the amino terminus of the protein. Some RXLR proteins are avirulence proteins, meaning that they can be detected by the plant and lead to a hypersensitive response which restricts the growth of the pathogen. P. infestans was found to encode around 60% more of these proteins than most other Phytophthora species. Those found in the apoplast include hydrolytic enzymes such as proteases, lipases and glycosylases that act to degrade plant tissue, enzyme inhibitors to protect against host defence enzymes and necrotizing toxins. Overall the genome was found to have an extremely high repeat content (around 74%) and to have an unusual gene distribution in that some areas contain many genes whereas others contain very few.JOURNAL, 2009, free, Nature Portfolio, Nature (journal), Nature, 0028-0836, 461, 7262, Haas, Brian J., Kamoun, Sophien, Sophien Kamoun, Zody, Michael, Jiang, Rays, Handsaker, Robert, Cano, Liliana, Grabherr, Manfred, Kodira, Chinnappa, Raffaele, Sylvain, Torto-Alalibo, Trudy, Bozkurt, Tolga, Ah-Fong, Audrey, Alvarado, Lucia, Anderson, Vicky, Armstrong, Miles, Avrova, Anna, Baxter, Laura, Beynon, Jim, Boevink, Petra, Bollmann, Stephanie, Bos, Jorunn, Bulone, Vincent, Cai, Guohong, Cakir, Cahid, Carrington, James, Chawner, Megan, Conti, Lucio, Costanzo, Stefano, Ewan, Richard, Fahlgren, Noah, Fischbach, Michael, Fugelstad, Johanna, Gilroy, Eleanor, Gnerre, Sante, Green, Pamela, Grenville-Briggs, Laura, Griffith, John, Grünwald, Niklaus, Horn, Karolyn, Horner, Neil, Hu, Chia-Hui, Huitema, Edgar, Jeong, Dong-Hoon, Jones, Alexandra, Jones, Jonathan, Jones, Richard, Karlsson, Elinor, Kunjeti, Sridhara, Lamour, Kurt, Liu, Zhenyu, Ma, LiJun, MacLean, Daniel, Chibucos, Marcus, McDonald, Hayes, McWalters, Jessica, Meijer, Harold, Morgan, William, Morris, Paul, Munro, Carol, O'Neill, Keith, Ospina-Giraldo, Manuel, Pinzón, Andrés, Pritchard, Leighton, Ramsahoye, Bernard, Ren, Qinghu, Restrepo, Silvia, Roy, Sourav, Sadanandom, Ari, Savidor, Alon, Schornack, Sebastian, Schwartz, David, Schumann, Ulrike, Schwessinger, Ben, Seyer, Lauren, Sharpe, Ted, Silvar, Cristina, Song, Jing, Studholme, David, Sykes, Sean, Thines, Marco, van de Vondervoort, Peter, Phuntumart, Vipaporn, Wawra, Stephan, Weide, Rob, Win, Joe, Young, Carolyn, Zhou, Shiguo, Fry, William, Meyers, Blake, van West, Pieter, Ristaino, Jean, Govers, Francine, Birch, Paul, Whisson, Stephen, Judelson, Howard, Nusbaum, Chad, 393–398, Genome sequence and analysis of the Irish potato famine pathogen Phytophthora infestans, 10.1038/nature08358, 19741609,weblink 2009Natur.461..393H, 4385549, The pathogen shows high allelic diversity in many isolates collected in Europe. This may be due to widespread trisomy or polyploidy in those populations.ResearchStudy of P. infestans presents sampling difficulties in the United States. It occurs only sporadically and usually has significant founder effects due to each epidemic starting from introduction of a single genotype.Origin and diversity{{See also|Plant disease epidemiology}}File:Modell eines Querschnitts durch ein Kartoffelblatt mit Phytophthora infestans (Kartoffelpilz) -Osterloh Nr. 55- -Brendel 10 k, 1- (2).jpg|thumb|Histological model of a potato leaf cross-section, Botanical Museum GreifswaldBotanical Museum GreifswaldThe highlands of central Mexico are considered by many to be the center of origin of P. infestans, although others have proposed its origin to be in the Andes, which is also the origin of potatoes.JOURNAL, Gomez-Alpizar, L, Carbone, I, Ristaino, JB, 2007, An Andean origin of Phytophthora infestans inferred from mitochondrial and nuclear gene genealogies, Proceedings of the National Academy of Sciences, National Academy of Sciences of the United States, 104, 9, 3306–11, 10.1073/pnas.0611479104, 17360643, 1805513, 2007PNAS..104.3306G, free, JOURNAL, 10.1146/annurev.phyto.43.040204.135906, 16078881, The Biology of Phytophthora infestans at Its Center of Origin, Annual Review of Phytopathology, 43, 171–90, 2005, Grünwald, N. J., Flier, W. G., A recent study evaluated these two alternate hypotheses and found conclusive support for central Mexico being the center of origin.JOURNAL, 10.1073/pnas.1401884111, The Irish potato famine pathogen Phytophthora infestans originated in central Mexico rather than the Andes, Proceedings of the National Academy of Sciences, 2014, Goss, E. M., Tabima, J. F., Cooke, D. E. L., Restrepo, S., Fry, W. E., Forbes, G. A., Fieland, V. J., Cardenas, M., Grünwald, N. J., 111, 24, 8791–96, 24889615, 4066499 | doi-access = free, Support for Mexico specifically the Toluca Valley2015>LOCATION=ST. PAUL, MINNESOTA, US | TITLE=POPULATION BIOLOGY OF PLANT PATHOGENS : GENETICS, ECOLOGY, AND EVOLUTION | APS PRESS THE AMERICAN PHYTOPATHOLOGICAL SOCIETY>LAST=MILGROOM | LCCN=2014943809, comes from multiple observations including the fact that populations are genetically most diverse in Mexico, late blight is observed in native tuber-bearing Solanum species, populations of the pathogen are in Hardy–Weinberg equilibrium, the two mating (see #Mating types | above) types occur in a 1:1 ratio, and detailed phylogeographic and evolutionary studies.10.1094/PHYTO.2001.91.9.882 | TITLE = POPULATION STRUCTURE OF PHYTOPHTHORA INFESTANS IN THE TOLUCA VALLEY REGION OF CENTRAL MEXICO | PHYTOPATHOLOGY (JOURNAL)>PHYTOPATHOLOGY | AMERICAN PHYTOPATHOLOGICAL SOCIETY > VOLUME = 91 | PAGES = 882–90 | LAST1 = GRüNWALD | LAST2 = FLIER | LAST3 = STURBAUM | LAST4 = GARAY-SERRANO | LAST5 = VAN DEN BOSCH | LAST6 = SMART | LAST7 = MATUSZAK | LAST8 = LOZOYA-SALDAñA | LAST9 = TURKENSTEEN | LAST10 = FRY | DOI-ACCESS =, 10.1094/PHYTO.2003.93.4.382> PMID = 18944351 | PHYTOPHTHORA INFESTANS FROM THE TOLUCA VALLEY OF CENTRAL MEXICO SUGGESTS GENETIC DIFFERENTIATION BETWEEN POPULATIONS FROM CULTIVATED POTATO AND WILD SOLANUM SPP.>JOURNAL=PHYTOPATHOLOGY (JOURNAL) | >PUBLISHER=AMERICAN PHYTOPATHOLOGICAL SOCIETY | ISSUE = 4 | YEAR = 2003 | FIRST1 = W. G. | FIRST2 = N. J. | FIRST3 = L. P. N. M. | FIRST4 = A. K. | FIRST5 = T. B. M. | FIRST6 = E. | FIRST7 = H. | FIRST8 = W. E. | FIRST9 = L. J. | PHYTOPHTHORA INFESTANS FROM POTATO AND SOLANUM DEMISSUM IN THE TOLUCA VALLEY, CENTRAL MEXICO> JOURNAL = MYCOLOGICAL RESEARCH | ISSUE = 8 | YEAR = 2001 | FIRST1 = W. G. | FIRST2 = N. J. | FIRST3 = W. E. | FIRST4 = L. J., 10.1094/PHYTO.1997.87.4.375> PMID = 18945115 | PHYTOPHTHORA INFESTANS IN ECUADOR>JOURNAL=PHYTOPATHOLOGY (JOURNAL) | >PUBLISHER=AMERICAN PHYTOPATHOLOGICAL SOCIETY | ISSUE = 4 | YEAR = 1997 | FIRST1 = G. A. | FIRST2 = X. C. | FIRST3 = C. C. | FIRST4 = J. | FIRST5 = M. E. | FIRST6 = B. A. | FIRST7 = K. | FIRST8 = W. E. | PHYTOPHTHORA INFESTANS?>JOURNAL=PLANT PATHOLOGY (JOURNAL) | >PUBLISHER=WILEY-BLACKWELL (BRITISH SOCIETY FOR PLANT PATHOLOGY) | ISSUE = 3 | YEAR = 1991 | FIRST1 = L. J. | FIRST2 = A. | FIRST3 = L. C. | FIRST4 = L. J. | FIRST5 = W. | FIRST6 = P. W. | FIRST7 = W. E., Furthermore, the closest relatives of P. infestans, namely ''Phytophthora mirabilis | and Phytophthora ipomoeae>P. ipomoeae'' are endemic to central Mexico.10.1017/S0953756202006123> TITLE = PHYTOPHTHORA IPOMOEAE SP. NOV., A NEW HOMOTHALLIC SPECIES CAUSING LEAF BLIGHT ON IPOMOEA LONGIPEDUNCULATA IN THE TOLUCA VALLEY OF CENTRAL MEXICO | MYCOLOGICAL RESEARCH> VOLUME = 106 | PAGES = 848–56 | LAST1 = FLIER | LAST2 = GRüNWALD | LAST3 = KROON | LAST4 = VAN DEN BOSCH | LAST5 = GARAY-SERRANO | LAST6 = LOZOYA-SALDAñA | LAST7 = BONANTS | LAST8 = TURKENSTEEN | S2CID = 20294303, On the other hand, the only close relative found in South America, namely ''Phytophthora andina | , is a hybrid that does not share a single common ancestor with P. infestans. Finally, populations of P. infestans'' in South America lack genetic diversity and are Vegetative reproduction>clonal.10.1371/JOURNAL.PONE.0024543> PMID = 21949727 | PHYTOPHTHORA ANDINA EMERGED VIA HYBRIDIZATION OF AN UNKNOWN PHYTOPHTHORA SPECIES AND THE IRISH POTATO FAMINE PATHOGEN, P. INFESTANS> JOURNAL = PLOS ONE | ISSUE = 9 | YEAR = 2011 | FIRST1 = E. M. | FIRST2 = M. E. | FIRST3 = K. | FIRST4 = G. A. | FIRST5 = W. E. | FIRST6 = S. | FIRST7 = N. J. | BIBCODE = 2011PLOSO...624543G | TITLE = GENETIC DIVERSITY OF PHYTOPHTHORA INFESTANS IN THE NORTHERN ANDEAN REGION | BMC GENETICS> VOLUME = 12 | YEAR = 2011 | FIRST1 = M. | FIRST2 = A. | FIRST3 = R. | FIRST4 = A. | FIRST5 = A. | FIRST6 = A. | FIRST7 = M. | FIRST8 = G. | FIRST9 = L. E. | FIRST10 = N. J. | FIRST11 = A. | FIRST12 = C. | FIRST13 = S. | DOI-ACCESS = FREE, Migrations from Mexico to North America or Europe have occurred several times throughout history, probably linked to the movement of tubers.JOURNAL, 10.1073/pnas.91.24.11591, 7972108, Panglobal distribution of a single clonal lineage of the Irish potato famine fungus, Proceedings of the National Academy of Sciences, 91, 24, 11591–95, 1994, Goodwin, S. B., Cohen, B. A., Fry, W. E., 45277, 1994PNAS...9111591G, free, JOURNAL, 10.7554/eLife.00731, 23741619, The rise and fall of the Phytophthora infestans lineage that triggered the Irish potato famine, eLife, 2, e00731, 2013, Yoshida, K., Schuenemann, V. J., Cano, L. M., Pais, M., Mishra, B., Sharma, R., Lanz, C., Martin, F. N., Kamoun, Sophien, Sophien Kamoun, Krause, J., Thines, M., Weigel, D., Burbano, H. A., 3667578, 1305.4206, free, Until the 1970s, the A2 mating type was restricted to Mexico, but now in many regions of the world both A1 and A2 isolates can be found in the same region.JOURNAL, 10.1111/j.1364-3703.2007.00465.x, 18705878, Phytophthora infestans: The plant (and R gene) destroyer, Molecular Plant Pathology, 9, 3, 385–402, 2008, Fry, William E., 6640234, The co-occurrence of the two mating types is significant due to the possibility of sexual recombination and formation of oospores, which can survive the winter. Only in Mexico and Scandinavia, however, is oospore formation thought to play a role in overwintering.JOURNAL, 10.2137/1239099043633332, Oospores of Phytphthora infestans in soil provide an important new source of primary inoculum in Finland, 2008, Hannukkala, A., Lehtinen, A., Agricultural and Food Science, 13, 4, 399, free, In other parts of Europe, increasing genetic diversity has been observed as a consequence of sexual reproduction.{{Citation|title=Appraisal of artificial screening techniques of tomato to accurately reflect field performance of the late blight resistance|last=Nowakowska|first=Marzena|date=3 October 2014|doi=10.1371/journal.pone.0109328|display-authors=etal|volume=9|issue=10|journal=PLOS ONE|pages=e109328|pmid=25279467|pmc=4184844|bibcode=2014PLoSO...9j9328N|doi-access=free}} This is notable since different forms of P. infestans vary in their aggressiveness on potato or tomato, in sporulation rate, and sensitivity to fungicides.NEWS,weblink Study provides evidence on movement of potato famine pathogen, 29 December 2016, Variation in such traits also occurs in North America, however importation of new genotypes from Mexico appears to be the predominant cause of genetic diversity, as opposed to sexual recombination within potato or tomato fields. In 1976 – due to a summer drought in Europe – there was a potato production shortfall and so eating potatoes were imported to fill the shortfall. It is thought that this was the vehicle for mating type A2 to reach the rest of the world. In any case, there had been little diversity, consisting of the US-1 strain, and of that only one type of: mating type, mtDNA, restriction fragment length polymorphism, and di-locus{{clarify|date=June 2021}} isozyme. Then in 1980 suddenly greater diversity and A2 appeared in Europe. In 1981 it was found in the Netherlands, United Kingdom, 1985 in Sweden, the early 1990s in Norway and Finland, 1996 in Denmark, and 1999 in Iceland. In the UK new A1 lineages only replaced the old lineage by end of the '80s, and A2 spread even more slowly, with Britain having low levels and Ireland (north and Republic) having none-to-trace detections through the '90s.JOURNAL, Cooke, L. R., Schepers, H. T. A. M., Hermansen, A., Bain, R. A., Bradshaw, N. J., Ritchie, F., Shaw, D. S., Evenhuis, A., Kessel, G. J. T., Wander, J. G. N., Andersson, B., Hansen, J. G., Hannukkala, A., Nærstad, R., Nielsen, B. J., Epidemiology and Integrated Control of Potato Late Blight in Europe, Potato Research, European Association for Potato Research (Springer Science and Business Media, Springer), 54, 2, 2011-05-24, 0014-3065, 10.1007/s11540-011-9187-0, 183–222, 21251642, free, Many of the strains that appeared outside of Mexico since the 1980s have been more aggressive, leading to increased crop losses. In Europe since 2013 the populations have been tracked by the EuroBlight network (see links below). Some of the differences between strains may be related to variation in the RXLR effectors that are present.Disease management{{See also|Plant pathology}}P. infestans is still a difficult disease to control. There are many chemical options in agriculture for the control of damage to the foliage as well as the fruit (for tomatoes) and the tuberWEB,weblink PlantVillage, 2015-12-16, 2015-12-22,weblink dead, (for potatoes). A few of the most common foliar-applied fungicides are Ridomil, a Gavel/SuperTin tank mix, and Previcur Flex. All of the aforementioned fungicides need to be tank mixed with a broad-spectrum fungicide such as mancozeb or chlorothalonil not just for resistance management but also because the potato plants will be attacked by other pathogens at the same time.If adequate field scouting occurs and late blight is found soon after disease development, localized patches of potato plants can be killed with a desiccant (e.g. paraquat) through the use of a backpack sprayer. This management technique can be thought of as a field-scale hypersensitive response similar to what occurs in some plant-viral interactions whereby cells surrounding the initial point of infection are killed in order to prevent proliferation of the pathogen.If infected tubers make it into a storage bin, there is a very high risk to the storage life of the entire bin. Once in storage, there isn't much that can be done besides emptying the parts of the bin that contain tubers infected with Phytophthora infestans. To increase the probability of successfully storing potatoes from a field where late blight was known to occur during the growing season, some products can be applied just prior to entering storage (e.g. Phostrol)weblink {{Bare URL PDF|date=March 2022}}Around the world the disease causes around $6 billion of damage to crops each year.Resistant plantsmissing image!
File:Älskade knöl – utflykt med SLU till deras potatisförsöksodlingar – 20190825 09 (cropped).jpg|thumb|Genetically modified King Edward (right) next to King Edward which has not been genetically modified (left). Research field, Swedish University of Agricultural SciencesSwedish University of Agricultural SciencesBreeding for resistance, particularly in potato plants, has had limited success in part due to difficulties in crossing cultivated potato to its wild relatives, which are the source of potential resistance genes. In addition, most resistance genes only work against a subset of P. infestans isolates, since effective plant disease resistance only results when the pathogen expresses a RXLR effector gene that matches the corresponding plant resistance (R) gene; effector-R gene interactions trigger a range of plant defenses, such as the production of compounds toxic to the pathogen.Potato and tomato varieties vary in their susceptibility to blight. Most early varieties are very vulnerable; they should be planted early so that the crop matures before blight starts (usually in July in the Northern Hemisphere). Many old crop varieties, such as King Edward potato are also very susceptible but are grown because they are wanted commercially. Maincrop varieties which are very slow to develop blight include Cara, Stirling, Teena, Torridon, Remarka, and Romano. Some so-called resistant varieties can resist some strains of blight and not others, so their performance may vary depending on which are around. These crops have had polygenic resistance bred into them, and are known as "field resistant". New varieties such as Sarpo Mira and Sarpo Axona show great resistance to blight even in areas of heavy infestation. Defender is an American cultivar whose parentage includes Ranger Russet and Polish potatoes resistant to late blight. It is a long white-skinned cultivar with both foliar and tuber resistance to late blight. Defender was released in 2004.{{Citation |last1=Novy |first1=R. G. |last2=Love |first2=S. L. |year=2006 |title=Defender: A high-yielding, processing potato cultivar with foliar and tuber resistance to late blight |journal=American Journal of Potato Research|volume=83 |issue=1 |pages=9–19 |doi=10.1007/BF02869605 |s2cid=41454585 |display-authors=etal}}Genetic engineering may also provide options for generating resistance cultivars. A resistance gene effective against most known strains of blight has been identified from a wild relative of the potato, Solanum bulbocastanum, and introduced by genetic engineering into cultivated varieties of potato.{{Citation |last1=Song |first1=Junqi |last2=Bradeen |first2=James M. |last3=Naess |first3=S. Kristine |last4=Raasch |first4=John A. |last5=Wielgus |first5=Susan M. |last6=Haberlach |first6=Geraldine T. |last7=Liu |first7=Jia |last8=Kuang |first8=Hanhui |last9=Austin-Phillips |first9=Sandra |last10=Jiang |first10=Jiming |year=2003 |title=Gene RB cloned from Solanum bulbocastanum confers broad spectrum resistance to potato late blight |journal=Proceedings of the National Academy of Sciences |volume=100 |issue=16 |pages=9128–33 |doi=10.1073/pnas.1533501100 |pmid=12872003 |pmc=170883 |bibcode=2003PNAS..100.9128S |doi-access=free }} This is an example of cisgenic genetic engineering.JOURNAL, Jacobsen, E., Schouten, H. J., Cisgenesis, a New Tool for Traditional Plant Breeding, Should be Exempted from the Regulation on Genetically Modified Organisms in a Step by Step Approach, Potato Research, 51, 75–88, 2008, 10.1007/s11540-008-9097-y, 38742532, Free version {{Webarchive|url=https://web.archive.org/web/20150923203713weblink |date=2015-09-23 }}Melatonin in the plant/P. infestans co-environment reduces the stress tolerance of the parasite.JOURNAL, Socaciu, Andreea Iulia, Ionuţ, Răzvan, Socaciu, Mihai Adrian, Ungur, Andreea Petra, Bârsan, Maria, Chiorean, Angelica, Socaciu, Carmen, Râjnoveanu, Armand Gabriel, Melatonin, an ubiquitous metabolic regulator: functions, mechanisms and effects on circadian disruption and degenerative diseases, Reviews in Endocrine and Metabolic Disorders- Cisgenicpotatoes.JPG - Potatoes after exposure. The normal potatoes have blight but the cisgenic potatoes are healthy. | Springer Science+Business Media>Springer, 21, 4, 2020-07-21, 1389-9155, 10.1007/s11154-020-09570-9, 465–478, 32691289, 220657247, Reducing inoculumBlight can be controlled by limiting the source of (wikt:inoculum|inoculum). Only good-quality seed potatoes and tomatoes obtained from certified suppliers should be planted. Often discarded potatoes from the previous season and self-sown tubers can act as sources of inoculum.{{Citation |last1=Zwankhuizen |first1=Maarten J. |last2=Govers |first2=Francine |last3=Zadoks |first3=Jan C. |author-link3=Jan Zadoks |year=1998 |title=Development of potato late blight epidemics: Disease foci, disease gradients, and infection sources |journal=Phytopathology|publisher=American Phytopathological Society |volume=88 |issue=8 |pages=754–63 |doi=10.1094/PHYTO.1998.88.8.754 |pmid=18944880 |doi-access= }}Compost, soil or potting medium can be heat-treated to kill oomycetes such as Phytophthora infestans. The recommended sterilisation temperature for oomycetes is {{convert|120|F|C}} for 30 minutes.WEB,weblink Phytophthora in nursery stock and restoration plantings, 4 September 2017, BOOK, Baker, K., The U.C. System for Producing Healthy Container Grown Plants, Manual 23, 1957, University of California, Division of Agricultural Sciences, Agricultural Experiment Station Extension Service,Environmental conditionsThere are several environmental conditions that are conducive to P. infestans. An example of such took place in the United States during the 2009 growing season. As colder than average for the season and with greater than average rainfall, there was a major infestation of tomato plants, specifically in the eastern states.{{Citation |last=Moskin |first=Julia |date=17 July 2009 |title=Outbreak of Fungus Threatens Tomato Crop |newspaper=The New York Times |url=https://www.nytimes.com/2009/07/18/nyregion/18tomatoes.html }} By using weather forecasting systems, such as BLITECAST, if the following conditions occur as the canopy of the crop closes, then the use of fungicides is recommended to prevent an epidemic.{{Citation |doi=10.1094/PD-65-394 |last=MacKenzie |first=D. R. |year=1981 |title=Scheduling fungicide applications for potato late blight with Blitecast |journal=Plant Disease|publisher=American Phytopathological Society |volume=65 |issue= 5|pages=394–99 }}
Use of fungicidesFile:Potato blight spraying system.jpg|thumb|Spraying potato, NottinghamshireNottinghamshireFungicides for the control of potato blight are normally only used in a preventative manner, optionally in (wikt:conjunction|conjunction) with disease forecasting. In susceptible varieties, sometimes fungicide applications may be needed weekly. An early spray is most effective. The choice of fungicide can depend on the nature of local strains of P. infestans. Metalaxyl is a fungicide that was marketed for use against P. infestans, but suffered serious resistance issues when used on its own. In some regions of the world during the 1980s and 1990s, most strains of P. infestans became resistant to metalaxyl, but in subsequent years many populations shifted back to sensitivity. To reduce the occurrence of resistance, it is strongly advised to use single-target fungicides such as metalaxyl along with carbamate compounds. A combination of other compounds are recommended for managing metalaxyl-resistant strains. These include mandipropamid, chlorothalonil, fluazinam, triphenyltin, mancozeb, and others. In the United States, the Environmental Protection Agency has approved oxathiapiprolin for use against late blight.WEB,weblink Oxathiapiprolin, New Active Ingredient Review, October 2015, Minnesota Department of Agriculture, 2017-11-09,weblink" title="web.archive.org/web/20171107025015weblink">weblink 2017-11-07, In African smallholder production fungicide application can be necessary up to once every three days.WEB, Daba, Tadessa, Why a new potato variety could be a game-changed for farmers in East Africa, Alliance for Science, 2020-12-11,weblink 2021-09-01,In organic productionIn the past, copper(II) sulfate solution (called 'bluestone') was used to combat potato blight. Copper pesticides remain in use on organic crops, both in the form of copper hydroxide and copper sulfate. Given the dangers of copper toxicity, other organic control options that have been shown to be effective include horticultural oils, phosphorous acids, and rhamnolipid biosurfactants, while sprays containing "beneficial" microbes such as Bacillus subtilis or compounds that encourage the plant to produce defensive chemicals (such as knotweed extract) have not performed as well.Gevens, Amanda, University of Wisconsin Madison Extension. Managing Late Blight in Organic Tomato & Potato Crops {{Webarchive|url=https://web.archive.org/web/20140618023311weblink |date=2014-06-18 }}.During the crop year 2008, many of the certified organic potatoes produced in the United Kingdom and certified by the Soil Association as organic were sprayed with a copper pesticideWEB,weblinkweblink" title="web.archive.org/web/20210901204043weblink">weblink 2021-09-01, Soil Association organic standards for producers, January 2009, 16; section 4.11.11, 103, Soil Association, to control potato blight. According to the Soil Association, the total copper that can be applied to organic land is {{convert|6|kg/ha|lb/acre}}/year.WEB,weblink Links to forms permitting application of copper fungicide on the website of the Soil Association, Soilassociation.org,weblink" title="web.archive.org/web/20091015055455weblink">weblink 15 October 2009, 16 July 2010,Control of tuber blightRidging is often used to reduce tuber contamination by blight. This normally involves piling soil or mulch around the stems of the potato blight meaning the pathogen has farther to travel to get to the tuber.{{Citation |last1=Glass |first1=J. R. |last2=Johnson |first2=K. B. |last3=Powelson |first3=M. L. |year=2001 |title=Assessment of Barriers to Prevent the Development of Potato Tuber Blight |journal=Plant Disease|publisher=American Phytopathological Society|volume=85 |issue=5 |pages=521–28 |doi=10.1094/PDIS.2001.85.5.521 |pmid=30823129 |doi-access= }} Another approach is to destroy the canopy around five weeks before harvest, using a contact herbicide or sulfuric acid to burn off the foliage. Eliminating infected foliage reduces the likelihood of tuber infection.Historical impact{{See also|Timeline of plant pathology}}File:Suggested paths of migration and diversification of P. infestans lineages HERB-1 and US-1.jpg|thumb|right|Suggested paths of migration and diversification of lineages HERB-1 and US-1 ]]The effect of Phytophthora infestans in Ireland in 1845–52 was one of the factors which caused over one million to starve to deathJOURNAL, Bourouiba, Lydia, The Fluid Dynamics of Disease Transmission, Annual Review of Fluid Mechanics, Annual Reviews (publisher), Annual Reviews, 53, 1, 2021-01-05, 0066-4189, 10.1146/annurev-fluid-060220-113712, 473–508, 2021AnRFM..53..473B, 225114407, free, and forced another two million to emigrate from affected countries. Most commonly referenced is the Great Irish Famine, during the late 1840s, from which the Irish population has still not fully recouped. The first recorded instances of the disease were in the United States, in Philadelphia and New York City in early 1843. Winds then spread the spores, and in 1845 it was found from Illinois to Nova Scotia, and from Virginia to Ontario. It crossed the Atlantic Ocean with a shipment of seed potatoes for Belgian farmers in 1845.{{Citation |first=John |last=Reader |title=The Fungus That Conquered Europe |url=https://www.nytimes.com/2008/03/17/opinion/17reader.html |work=New York Times |date=17 March 2008 |access-date=18 March 2008}}WEB,weblink Between two worlds: The Hungry Forties in Europe, RTÉ.ie, 27 January 2022, All of the potato-growing countries in Europe were affected, but the potato blight hit Ireland the hardest. Implicated in Ireland's fate was the island's disproportionate dependency on a single variety of potato, the Irish Lumper. The lack of genetic variability created a susceptible host population for the organismWEB, Great Famine potato makes a comeback after 170 years
During the First World War, all of the copper in Germany was used for shell casings and electric wire and therefore none was available for making copper sulfate to spray potatoes. A major late blight outbreak on potato in Germany therefore went untreated, and the resulting scarcity of potatoes led to the deaths of 700,000 German civilians from starvation.WEB,weblink The Origin of Plant Pathology and The Potato Famine, and Other Stories of Plant Diseases., 2018-09-01,weblink" title="web.archive.org/web/20160304001917weblink">weblink 2016-03-04, Carefoot, G.L. and E.R. Sprott. 1967. Famine on the Wind: Man's Battle Against Plant Disease. Rand McNally.{{ISBN?}}{{page needed|date=July 2023}}Since 1941, Eastern Africa has been suffering potato production losses because of strains of P. infestans from Europe.JOURNAL, Genotyping of Phytophthora infestans in Eastern Africa Reveals a Dominating Invasive European Lineage, Phytopathology (journal), Phytopathology, 109, 4, 670–680, American Phytopathological Society, 10.1094/PHYTO-07-18-0234-R, 2019, Njoroge, Anne W., Andersson, Björn, Lees, Alison K., Mutai, Collins, Forbes, Gregory A., Yuen, Jonathan E., Pelle, Roger, 30253119, free, France, Canada, the United States, and the Soviet Union researched P. infestans as a biological weapon in the 1940s and 1950s.{{Citation |last1=Suffert |first1=Frédéric |last2=Latxague |first2=Émilie |last3=Sache |first3=Ivan |year=2009 |title=Plant pathogens as agroterrorist weapons: assessment of the threat for European agriculture and forestry |journal=Food Security|volume=1 |issue=2 |pages=221–232 |doi=10.1007/s12571-009-0014-2 |s2cid=23830595 }} Potato blight was one of more than 17 agents that the United States researched as potential biological weapons before the nation suspended its biological weapons program."Chemical and Biological Weapons: Possession and Programs Past and Present", James Martin Center for Nonproliferation Studies, Middlebury College, April 9, 2002, accessed November 14, 2008. Whether a weapon based on the pathogen would be effective is questionable, due to the difficulties in delivering viable pathogen to an enemy's fields, and the role of uncontrollable environmental factors in spreading the disease.{{Citation needed|date = June 2020}}Late blight (A2 type) has not yet been detected in Australia and strict biosecurity measures are in place. The disease has been seen in China, India and south-east Asian countries.A large outbreak of P. infestans occurred on tomato plants in the Northeast United States in 2009.WEB, The 2009 Late Blight Pandemic in Eastern USA,weblink 2023-11-24, The 2009 Late Blight Pandemic in Eastern USA, en-US, In light of the periodic epidemics of P. infestans ever since its first emergence, it may be regarded as a periodically emerging pathogen – or a periodically "re-emerging pathogen".JOURNAL, Fones, Helen N., Bebber, Daniel P., Chaloner, Thomas M., Kay, William T., Steinberg, Gero, Gurr, Sarah J., Threats to global food security from emerging fungal and oomycete crop pathogens, Nature Food, Nature Research, 1, 6, 2020-06-02, 2662-1355, 10.1038/s43016-020-0075-0, 332–342, 37128085, 219924805, {{snd}}(JOURNAL, Fones, Helen N., Bebber, Daniel P., Chaloner, Thomas M., Kay, William T., Steinberg, Gero, Gurr, Sarah J., 0, Author Correction: Threats to global food security from emerging fungal and oomycete crop pathogens, Nature Food, Nature Research, 1, 7, 2020, 2662-1355, 10.1038/s43016-020-0111-0, 455–456, free, )JOURNAL, Fry, W. E., Birch, P. R. J., Judelson, H. S., Grünwald, N. J., Danies, G., Everts, K. L., Gevens, A. J., Gugino, B. K., Johnson, D. A., Johnson, S. B., McGrath, M. T., Myers, K. L., Ristaino, J. B., Roberts, P. D., Secor, G., Smart, C. D., Five Reasons to Consider Phytophthora infestans a Reemerging Pathogen, Phytopathology (journal), Phytopathology, American Phytopathological Society, 105, 7, 2015, 0031-949X, 10.1094/phyto-01-15-0005-fi, 966–981, 25760519, 27658217, free, , IrishCentral ,weblink , 2013-03-05, 2013-03-03 , after the blight strains originating in Chiloé Archipelago replaced earlier potatoes of Peruvian origin in Europe.NEWS, Mash hits: the land that spawned the supermarket spud,weblink Johanson, Mark, August 28, 2020, September 1, 2020, The Economist, ReferencesFurther reading
| title=Rise and fall of the Phytophthora infestans lineage that triggered the Irish potato famine, eLife, in press, 10.7554/elife.00731 | display-authors=etal, 23741619, 3667578, 2, 2013 | doi-access=free,
| doi-access = free,
| title=Reconstructing genome evolution in historic samples of the Irish potato famine pathogen, Nature Communications, 4 | doi=10.1038/ncomms3172, 2013, etal, 23863894 | bibcode=2013NatCo...4.2172M,
| title=Genomic characterization of a South American Phytophthora hybrid mandates reassessment of the geographic origins of Phytophthora infestans | Molecular Biology and Evolution>volume=33 | doi=10.1093/molbev/msv241, 2016, etal, 478–91, 26576850, 4866541,
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