Editing Potato (section)

== Breeding ==

Potatoes, both ''S. tuberosum'' and most of its wild relatives, are [[self-incompatible]]: they bear no useful fruit when self-pollinated. This trait is problematic for crop breeding, as all sexually-produced plants must be [[hybrid (biology) |hybrid]]s. The gene responsible for self-incompatibility, as well as mutations to disable it, are now known. Self-compatibility has successfully been introduced both to diploid potatoes (including a special line of ''S. tuberosum'') by [[CRISPR-Cas9]].<ref name="Neofunctionalisation"/> Plants having a 'Sli' gene produce pollen which is compatible to its own parent and plants with similar S genes.<ref name="Hosaka Hanneman, Jr. 1998 pp. 191–197" >{{cite journal |last1=Hosaka |first1=Kazuyoshi |last2=Hanneman |first2=Robert E. Jr. |title=Genetics of self-compatibility in a self-incompatible wild diploid potato species ''Solanum chacoense''. 1. Detection of an S locus inhibitor (Sli) gene |journal=Euphytica |volume=99 |issue=3 |year=1998 |issn=0014-2336 |doi=10.1023/a:1018353613431 |pages=191–197 |s2cid=40678039 |url=https://link.springer.com/article/10.1023/A:1018353613431 }}</ref> This gene was cloned by [[Wageningen University & Research |Wageningen University]] and [[Solynta]] in 2021, which would allow for faster and more focused breeding.<ref name="Neofunctionalisation">{{cite journal |display-authors=3 |last1=Eggers |first1=Ernst-Jan |last2=Burgt |first2=van der |last3=Heusden |first3=van |last4=W. |first4=Sjaak A. |last5=Vries |first5=de |last6=E. |first6=Michiel |last7=Visser |first7=Richard G. F. |last8=Bachem |first8=Christian W. B. |last9=Lindhout |first9=Pim |title=Neofunctionalisation of the Sli gene leads to self-compatibility and facilitates precision breeding in potato |journal=Nature Communications |volume=12 |issue=1 |date=6 July 2021 |page=4141 |issn=2041-1723 |doi=10.1038/s41467-021-24267-6 |pmid=34230471 |pmc=8260583 |bibcode=2021NatCo..12.4141E}}</ref><ref>{{cite journal |last1=Ma |first1=Ling |last2=Zhang |first2=Chunzhi |last3=Zhang |first3=Bo |last4=Tang |first4=Fei |last5=Li |first5=Futing |last6=Liao |first6=Qinggang |last7=Tang |first7=Die |last8=Peng |first8=Zhen |last9=Jia |first9=Yuxin |last10=Gao |first10=Meng |last11=Guo |first11=Han |last12=Zhang |first12=Jinzhe |last13=Luo |first13=Xuming |last14=Yang |first14=Huiqin |last15=Gao |first15=Dongli |last16=Lucas |first16=William J. |last17=Li |first17=Canhui |last18=Huang |first18=Sanwen |last19=Shang |first19=Yi |display-authors=3 |title=A nonS-locus F-box gene breaks self-incompatibility in diploid potatoes |journal=Nature Communications |volume=12 |issue=1 |date=6 July 2021 |page=4142 |issn=2041-1723 |doi=10.1038/s41467-021-24266-7 |pmid=34230469 |pmc=8260799 |bibcode=2021NatCo..12.4142M}}
</ref>

[[Diploid]] hybrid potato breeding is a recent area of potato genetics supported by the finding that simultaneous [[homozygous |homozygosity]] and [[fixation (population genetics) |fixation]] of donor alleles is possible.<ref name="Lindhout Meijer Schotte Hutten 2011 pp. 301–312">{{cite journal |last1=Lindhout |first1=Pim |last2=Meijer |first2=Dennis |last3=Schotte |first3=Theo |last4=Hutten |first4=Ronald C. B. |last5=Visser |first5=Richard G. F. |last6=van Eck |first6=Herman J. |title=Towards F1 Hybrid Seed Potato Breeding |journal=Potato Research |publisher=Springer |volume=54 |issue=4 |year=2011 |issn=0014-3065 |doi=10.1007/s11540-011-9196-z |pages=301–312 |s2cid=39719359 |doi-access=free}}</ref> [[Crop wild relative |Wild potato species useful for breeding]] blight resistance include ''Solanum desmissum'' and ''S. stoloniferum'', among others.<ref name="Strategies">{{cite journal |year=2023 |publisher=Springer |first3=Anu |first2=Sat Pal |first1=Dechen |last3=Kalia |last2=Sharma |last1=Angmo |journal=Molecular Biology Reports |issn=0301-4851 |s2cid=260349512 |pmid=37526862 |doi=10.1007/s11033-023-08577-0 |title=Breeding strategies for late blight resistance in potato crop: recent developments |volume=50 |issue=9 |pages=7879–7891 }}</ref>

=== Varieties ===

{{further|List of potato cultivars}}
[[File:Potatoes for sale in a market in France.jpg|thumb|A variety of different potatoes for sale in a market in France]]

[[File:Papas de colores de Chiloe.jpg|thumb|Potatoes of different colors]] 

There are some 5,000 potato varieties worldwide, 3,000 of them in the [[Andes]] alone — mainly in Peru, Bolivia, Ecuador, Chile, and Colombia. Over 100 cultivars might be found in a single valley, and a dozen or more might be maintained by a single agricultural household.<ref>{{cite web |last=Theisen |first=K |date=1 January 2007 |title=History and overview |url=http://research.cip.cgiar.org/confluence/display/wpa/Peru |archive-url=https://web.archive.org/web/20080114015939/http://research.cip.cgiar.org/confluence/display/wpa/Peru |archive-date=14 January 2008 |access-date=10 September 2008 |work=World Potato Atlas: Peru |publisher=[[International Potato Center]]}}</ref><ref>{{cite web |url=http://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:821337-1#synonyms |title=''Solanum tuberosum'' L. |date=2017 |website=[[Plants of the World Online]] |publisher=Royal Botanic Gardens, Kew |access-date=7 September 2020}}</ref> 
The [[European Cultivated Potato Database]] is an online collaborative database of potato variety descriptions updated and maintained by the [[Scottish Agricultural Science Agency]] within the framework of the European Cooperative Programme for Crop Genetic Resources Networks—which is run by the [[International Plant Genetic Resources Institute]].<ref>{{cite web |title=Europotato.org |url=http://www.europotato.org/menu.php |url-status=dead |archive-url=https://web.archive.org/web/20091128021457/http://www.europotato.org/menu.php |archive-date=28 November 2009 |access-date=16 July 2010 |publisher=Europotato.org}}</ref>
Around 80 varieties are commercially available in the UK.<ref>{{cite web |author=Potato Council |title=Potato Varieties |url=http://www.britishpotatoes.co.uk/potato-varieties/ |url-status=dead |archive-url=https://web.archive.org/web/20090908212045/http://www.britishpotatoes.co.uk/potato-varieties |archive-date=8 September 2009 |access-date=13 September 2009 |publisher=Agriculture & Horticulture Development Board}}</ref> 

For culinary purposes, varieties are often differentiated by their waxiness: floury or mealy ''baking'' potatoes have more [[starch]] (20–22%) than waxy ''boiling'' potatoes (16–18%). The distinction may also arise from variation in the comparative ratio of two different potato starch compounds: [[amylose]] and [[amylopectin]]. Amylose, a long-chain molecule, diffuses from the starch granule when cooked in water, and lends itself to dishes where the potato is mashed. Varieties that contain a slightly higher amylopectin content, which is a highly branched molecule, help the potato retain its shape after being boiled in water.<ref>{{cite web |title=Potato Primer |url=http://www.cooksillustrated.com/images/document/howto/JF07_PotatoPrimer.pdf |url-status=dead |archive-url=https://web.archive.org/web/20081217030233/http://www.cooksillustrated.com/images/document/howto/JF07_PotatoPrimer.pdf |archive-date=17 December 2008 |access-date=8 December 2008 |publisher=[[Cook's Illustrated]]}}</ref> Potatoes that are good for making [[potato chip]]s or potato crisps are sometimes called "chipping potatoes", which means they meet the basic requirements of similar varietal characteristics, being firm, fairly clean, and fairly well-shaped.<ref>{{cite web |author=[[Agricultural Marketing Service]] |title=Potatoes for Chipping Grades and Standards |url=https://www.ams.usda.gov/grades-standards/potatoes-chipping-grades-and-standards |access-date=27 August 2018 }}</ref>

Immature potatoes may be sold fresh from the field as "{{Vanchor |creamer potatoes |text=creamer}}" or "{{Vanchor |new potatoes |text=new}}" potatoes and are particularly valued for their taste. They are typically small in size and tender, with a loose skin, and flesh containing a lower level of [[starch]] than other potatoes. In the United States they are generally either a [[Yukon Gold potato]] or a red potato, called gold creamers or red creamers respectively.<ref name="recipe tips">{{cite web |title=Creamer Potato |url=http://www.recipetips.com/glossary-term/t--35863/creamer-potato.asp |access-date=18 July 2008 |publisher=recipetips.com}}</ref><ref>{{cite news |date=12 August 2013 |title=What is a new potato? New guidelines issued |work=BBC News |url=https://www.bbc.co.uk/news/uk-scotland-glasgow-west-23667664 |access-date=13 June 2021}}</ref> In the UK, the [[Jersey Royal]] is a famous type of new potato.<ref>{{cite news |date=25 January 2010 |title=A look back at a Royal history |url=http://news.bbc.co.uk/local/jersey/hi/people_and_places/nature/newsid_8478000/8478833.stm |access-date=13 June 2021}}</ref> 

Dozens of potato [[cultivar]]s have been [[plant breeding|selectively bred]] specifically for their skin or flesh [[biological pigment |color]], including gold, red, and blue varieties.<ref>{{cite web |date=2017 |title=So many varieties, so many choices |url=http://wisconsinpotatoes.com/growing/varieties/ |publisher=Wisconsin Potato and Vegetable Growers Association}}</ref> These contain varying amounts of [[phytochemical]]s, including [[carotenoids]] for gold/yellow or [[polyphenol]]s for red or blue cultivars.<ref name="Hirsch">{{cite journal |last1=Hirsch |first1=C.N. |last2=Hirsch |first2=C.D. |last3=Felcher |first3=K |last4=Coombs |first4=J |last5=Zarka |first5=D |last6=Van Deynze |first6=A |last7=De Jong |first7=W |last8=Veilleux |first8=R.E. |last9=Jansky |first9=S |last10=Bethke |first10=P |last11=Douches |first11=D.S. |last12=Buell |first12=C.R. |year=2013 |title=Retrospective View of North American Potato (''Solanum tuberosum'' L.) Breeding in the 20th and 21st Centuries |journal=[[G3: Genes, Genomes, Genetics]] |volume=3 |issue=6 |pages=1003–13 |doi=10.1534/g3.113.005595 |pmc=3689798 |pmid=23589519}}</ref> Carotenoid compounds include [[provitamin A]] [[alpha-carotene]] and [[beta-carotene]], which are converted to the [[essential nutrient]], [[vitamin A]], during digestion. [[Anthocyanin]]s mainly responsible for red or blue pigmentation in potato cultivars do not have nutritional significance, but are used for visual variety and consumer appeal.<ref>{{cite journal |last1=Jemison |first1=John M. Jr. |last2=Sexton |first2=Peter |last3=Camire |first3=Mary Ellen |year=2008 |title=Factors Influencing Consumer Preference of Fresh Potato Varieties in Maine |journal=American Journal of Potato Research |volume=85 |issue=2 |page=140 |doi=10.1007/s12230-008-9017-3 |s2cid=34297429}}</ref> In 2010, potatoes were [[genetic engineering |bioengineered]] specifically for these pigmentation traits.<ref>{{cite book |last1=Mattoo |first1=A.K. |title=Bio-Farms for Nutraceuticals |last2=Shukla |first2=V |last3=Fatima |first3=T |last4=Handa |first4=A.K. |last5=Yachha |first5=S.K. |year=2010 |isbn=978-1-4419-7346-7 |series=Advances in Experimental Medicine and Biology |volume=698 |pages=122–43 |chapter=Genetic Engineering to Enhance Crop-Based Phytonutrients (Nutraceuticals) to Alleviate Diet-Related Diseases |doi=10.1007/978-1-4419-7347-4_10 |pmid=21520708}}</ref>

=== Genetic engineering ===

{{main|Genetically engineered potato}}

[[File:Auspflanzung Schweden 2.jpg|thumb|Amflora potatoes, modified to produce pure [[amylopectin]] starch]]

Genetic research has produced several [[genetically modified]] varieties. 'New Leaf', owned by [[Monsanto Company]], incorporates genes from ''[[Bacillus thuringiensis]]'' (source of most [[Bt toxin |''Bt'' toxins]] in [[Bt crops |transcrop use]]), which confers resistance to the [[Colorado potato beetle]]; 'New Leaf Plus' and 'New Leaf Y', approved by US regulatory agencies during the 1990s, also include resistance to [[potato virus |viruses]]. [[McDonald's]], [[Burger King]], [[Frito-Lay]], and [[Procter & Gamble]] announced they would not use [[genetically modified potato]]es, and Monsanto published its intent to discontinue the line in March 2001.<ref>{{cite web |title=Genetically Engineered Organisms Public Issues Education Project/Am I eating GE potatoes? |url=http://www.geo-pie.cornell.edu/crops/potato.html |access-date=16 December 2008 |publisher=[[Cornell University]] |archive-date=3 January 2009 |archive-url=https://web.archive.org/web/20090103033654/http://www.geo-pie.cornell.edu/crops/potato.html}}</ref>

Potato starch contains two types of [[glucan]], [[amylose]] and [[amylopectin]], the latter of which is most industrially useful. Waxy potato varieties produce [[waxy potato starch]], which is almost entirely amylopectin, with little or no amylose. [[BASF]] developed the '[[Amflora]]' potato, which was modified to express [[antisense RNA]] to inactivate the gene for [[NDP-glucose—starch glucosyltransferase |granule bound starch synthase]], an enzyme which catalyzes the formation of amylose.<ref>{{cite web |title=GMO compass database |url=http://www.gmo-compass.org/eng/gmo/popups/55.potato_eh92_527_1.html |url-status=dead |archive-url=https://web.archive.org/web/20141009210148/http://www.gmo-compass.org/eng/gmo/popups/55.potato_eh92_527_1.html |archive-date=9 October 2014 |access-date=6 October 2014}}</ref> 'Amflora' potatoes therefore produce starch consisting almost entirely of [[amylopectin]], and are thus more useful for the starch industry. In 2010, the European Commission cleared the way for 'Amflora' to be grown in the European Union for industrial purposes only—not for food. Nevertheless, under EU rules, individual countries have the right to decide whether they will allow this potato to be grown on their territory. Commercial planting of 'Amflora' was expected in the Czech Republic and Germany in the spring of 2010, and Sweden and the Netherlands in subsequent years.<ref>{{Cite web |date=2010-05-31 |title=GM potatoes: BASF at work |url=http://www.gmo-compass.org/eng/news/492.docu.html |archive-url=https://web.archive.org/web/20100531073525/http://www.gmo-compass.org/eng/news/492.docu.html |archive-date=31 May 2010 }}</ref> 

The 'Fortuna'<!-- see de:Fortuna (Kartoffel) fr:Fortuna (pomme de terre) --> GM potato variety developed by BASF was made resistant to [[Phytophthora infestans |late blight]] by [[introgression|introgressing]] two resistance genes, {{Visible anchor |blb1}} and {{Visible anchor |blb2}}, from ''[[Solanum bulbocastanum |S. bulbocastanum]]'', a wild potato native to Mexico.<ref name="Receptor-Mediated"/><ref>{{Cite web |date=2013-06-02 |title=Research in Germany: Business BASF applies for approval for another biotech potato |url=http://www.research-in-germany.de/84190/2011-11-17-business-basf-applies-for-approval-for-another-biotech-potato.html |archive-url=https://web.archive.org/web/20130602111343/http://www.research-in-germany.de/84190/2011-11-17-business-basf-applies-for-approval-for-another-biotech-potato.html |archive-date=2 June 2013 }}</ref><ref>{{Cite web |last=Burger |first=Ludwig |date=2015-11-10 |title=BASF applies for EU approval for Fortuna GM potato |website=[[Reuters]] |url=http://www.reuters.com/article/2011/10/31/us-basf-idUSTRE79U41Q20111031 |archive-url=https://web.archive.org/web/20151110105537/http://www.reuters.com/article/2011/10/31/us-basf-idUSTRE79U41Q20111031 |archive-date=10 November 2015 }}</ref> {{Vanchor|Rpi-blb1}} is a [[nucleotide-binding leucine-rich repeat]] (NB-LRR/NLR), an R-gene-produced immunoreceptor.<ref name="Receptor-Mediated"> {{Cite journal |last1=Oh |first1=Soohyun |last2=Choi |first2=Doil |year=2022 |title=Receptor-mediated nonhost resistance in plants |department=Review |journal=Essays in Biochemistry |publisher=[[Portland Press Limited]] ([[Biochemical Society]]) |volume=66 |issue=5 |pages=435–445 |doi=10.1042/EBC20210080 |pmc=9528085 |pmid=35388900 |s2cid=247999992}}</ref>

In October 2011, BASF requested cultivation and marketing approval as a feed and food from the EFSA. In 2012, GMO development in Europe was stopped by BASF.<ref>[http://www.dw.de/basf-stops-gm-crop-development-in-europe/a-15671900 BASF stops GM crop development in Europe], ''Deutsche Welle'', 17 January 2012</ref><ref>{{Cite news |last=Kanter |first=James |title=BASF to Stop Selling Genetically Modified Products in Europe |work=The New York Times |date=16 January 2012 |url=https://www.nytimes.com/2012/01/17/business/global/17iht-gmo17.html |archive-url=https://archive.today/20240123213954/https://www.nytimes.com/2012/01/17/business/global/17iht-gmo17.html |archive-date=23 January 2024 |access-date=23 January 2024}}</ref> In November 2014, the [[United States Department of Agriculture]] (USDA) approved a genetically modified potato developed by [[Simplot]], which contains genetic modifications that prevent bruising and produce less [[acrylamide]] when fried than conventional potatoes; the modifications do not cause new proteins to be made, but rather prevent proteins from being made via [[RNA interference]].<ref>{{Cite web |last=Pollack |first=Andrew |date=November 7, 2014 |title=U.S.D.A. Approves Modified Potato. Next Up: French Fry Fans |website=[[The New York Times]] |url=https://www.nytimes.com/2014/11/08/business/genetically-modified-potato-from-simplot-approved-by-usda.html |archive-url=https://archive.today/20141112021024/http://www.nytimes.com/2014/11/08/business/genetically-modified-potato-from-simplot-approved-by-usda.html?_r=0 |archive-date=12 November 2014}}</ref>

Genetically modified varieties have met public resistance in the U.S. and in the European Union.<ref>{{cite news |year=2002 |title=Consumer acceptance of genetically modified potatoes |journal=American Journal of Potato Research |url=http://www.agbioforum.org/v7n12/v7n12a13-mccluskey.pdf |url-status=dead |access-date=19 February 2012 |archive-url=https://web.archive.org/web/20121101142414/http://www.agbioforum.org/v7n12/v7n12a13-mccluskey.pdf |archive-date=1 November 2012 |id=cited through Bnet}}</ref><ref name="nytimes1">{{cite news |last=Rosenthal |first=Elisabeth |date=24 July 2007 |title=A genetically modified potato, not for eating, is stirring some opposition in Europe |work=[[The New York Times]] |url=https://www.nytimes.com/2007/07/24/business/worldbusiness/24spuds.html |access-date=15 November 2008}}</ref>

{{anchor|Growth}}