Editing Truffle

{{Short description|Fruiting body of a subterranean ascomycete fungus}}
{{hatgrp|{{distinguish|Chocolate truffle}}{{other uses}}}}
[[File:Truffe_noire_du_Périgord.jpg|thumb|[[Tuber melanosporum|Black truffle]] (''Tuber melanosporum'')]]
[[File:White truffles from San Miniato.jpg|thumb|300px|White truffles from San Miniato]]
[[File:Black truffles, San Miniato.jpg|thumb|300px|Black truffles from San Miniato]]

A '''truffle''' is the [[Sporocarp (fungi)|fruiting body]] of a subterranean [[ascomycete]] [[fungus]], one of the species of the genus ''[[Tuber (fungus)|Tuber]]''. More than one hundred other genera of fungi are classified as truffles including ''[[Geopora]]'', ''[[Peziza]]'', ''[[Choiromyces]]'', and ''[[Leucangium]]''.<ref name="Læssøe 2007">{{Cite journal|last1=Læssøe|first1=Thomas|last2=Hansen|first2=Karen|date=September 2007|title=Truffle trouble: what happened to the Tuberales?|journal=Mycological Research|volume=111|issue=9|pages=1075–1099|doi=10.1016/j.mycres.2007.08.004|issn=0953-7562|pmid=18022534}}</ref> These genera belong to the class [[Pezizomycetes]] and the [[Pezizales]] order. Several truffle-like [[basidiomycetes]] are excluded from Pezizales, including ''[[Rhizopogon]]'' and ''Glomus''. Truffles are [[ectomycorrhiza]]l fungi, so they are found in close association with tree roots. Spore dispersal is accomplished through [[fungivore]]s, animals that eat fungi.<ref>{{cite web|url=http://www.cpbr.gov.au/fungi/dispersal.html|title=Spore release and dispersal|last=Lepp|first=Heino|website=Australian National Botanic Gardens|access-date=5 December 2016|archive-date=14 November 2016|archive-url=https://web.archive.org/web/20161114222732/http://www.cpbr.gov.au/fungi/dispersal.html|url-status=live}}</ref> These fungi have ecological roles in [[nutrient cycling]] and drought tolerance.

Some truffle species are prized as food.<ref name="brillat">{{cite book |last=Brillat-Savarin |first=Jean Anthelme |url=https://archive.org/details/physiologiedugo02savgoog |title=Physiologie du goût |publisher=Charpentier |year=1838 |location=Paris |author-link=Jean Anthelme Brillat-Savarin |orig-year=1825}} [http://etext.library.adelaide.edu.au/b/b85p/part11.html English translation] {{webarchive|url=https://web.archive.org/web/20080706020126/http://etext.library.adelaide.edu.au/b/b85p/part11.html|date=2008-07-06}}</ref> Edible truffles are used in [[Italian cuisine|Italian]], [[French cuisine|French]]<ref>{{Cite news|url=http://www.traditionalfrenchfood.com/truffles.html|title=Truffles|date=2017|work=Traditional French Food Regional Recipes From Around France|access-date=2017-01-06|archive-date=2017-01-07|archive-url=https://web.archive.org/web/20170107102536/http://www.traditionalfrenchfood.com/truffles.html|url-status=live}}</ref> and other national {{lang|fr|[[haute cuisine]]s}}. Truffles are [[Fungiculture|cultivated]] and harvested from natural environments.

==Taxonomic details==

=== Species ===

==== Black ====
{{Main|Tuber melanosporum}}
[[File:Tuber brumale - Vue sur la tranche coupée.jpg|alt=|thumb|Black Périgord truffle, cross-section]]

The black truffle or black Périgord truffle (''Tuber melanosporum''), the second-most commercially valuable species, is named after the [[Périgord]] region in France.<ref name="Trappe-2009">{{Cite journal|last=Trappe|first=Jim|date=2009|title=Taming the truffle—the history, lore, and science of the ultimate mushroom |journal=Gastronomica|volume=9|issue=1|pages=116–117|doi=10.1525/gfc.2009.9.1.116|issn=1529-3262}}</ref> Black truffles associate with [[oak]]s, [[hazel]]nut, cherry, and other deciduous trees and are harvested in late autumn and winter.<ref name="Trappe-2009" /><ref name="Carluccio032">{{cite book|title=The Complete Mushroom Book|author=Carluccio|first=Antonio|publisher=Quadrille|year=2003|isbn=978-1-84400-040-1|author-link=Antonio Carluccio}}</ref> The genome sequence of the black truffle was published in March 2010.<ref name="truffle genome2">{{cite journal | last1 = Martin | first1 = F. | last2 = Kohler | first2 = A. | last3 = Murat | first3 = C. | last4 = Balestrini | first4 = R. | last5 = Coutinho | first5 = P.M. | last6 = Jaillon | first6 = O. | last7 = Montanini | first7 = B. | last8 = Morin | first8 = E. | last9 = Noel | first9 = B. | last10 = Percudani | first10 = R. | last11 = Porcel | first11 = B. | year = 2010 | title = Périgord black truffle genome uncovers evolutionary origins and mechanisms of symbiosis | journal = Nature | volume = 464 | issue = 7291| pages = 1033–1038 | doi=10.1038/nature08867 | pmid=20348908| bibcode = 2010Natur.464.1033M | doi-access = free | hdl = 2318/100278 | hdl-access = free }}</ref>

==== Summer or burgundy ====
{{Main|Tuber aestivum}}
[[File:Black.summer.truffle.arp.jpg|thumb|Summer truffles in a shop in Rome]]

The black summer truffle (''Tuber aestivum'') is found across Europe and is prized for its culinary value.<ref name="Paolocci-2004">{{Cite journal|last1=Paolocci|first1=Francesco|last2=Rubini|first2=Andrea|last3=Riccioni|first3=Claudia|last4=Topini|first4=Fabiana|last5=Arcioni|first5=Sergio|date=2004|title=Tuber aestivum and Tuber uncinatum: two morphotypes or two species?|journal=FEMS Microbiology Letters|volume=235|issue=1|pages=109–115|doi=10.1111/j.1574-6968.2004.tb09574.x|issn=0378-1097|pmid=15158269|doi-access=free}}</ref> Burgundy truffles (designated ''[[Tuber uncinatum]]'', but the same species) are harvested in autumn until December and have aromatic flesh of a darker colour. These are associated with various trees and shrubs.<ref name="Paolocci-2004" />

==== White ====
{{main|Tuber magnatum}}
[[File:Truffle washed and cut.jpg|alt=|thumb|A white truffle washed and with a corner cut to show the interior]] 
Tuber magnatum, the high-value ''white truffle'' ({{langx|it|tartufo bianco}}) is found mainly in the [[Langhe]] and [[Montferrat]] areas<ref>{{cite web|url=http://www.lifeinitaly.com/food/truffle.asp|title=White truffles from Alba|last=Demetri|first=Justin|date=2012|website=Life in Italy|publisher=Lifeinitaly.com|access-date=2012-06-16|archive-date=2009-10-16|archive-url=https://web.archive.org/web/20091016180401/http://www.lifeinitaly.com/food/truffle.asp|url-status=live}}</ref> of the [[Piedmont]] region in northern [[Italy]], and most famously, in the countryside around the cities of [[Alba, Piedmont|Alba]] and [[Asti]].<ref>{{Cite journal|last1=Bencivenga  |first1=M. |last2=Di Massimo |first2=G. |last3=Donnini |first3=D. |last4=Baciarelli Falini |first4=L. |date=2009|title=The cultivation of truffles in Italy |journal=Acta Botanica Yunnanica |volume=16 |issue=Suppl 16 |pages=100–102}}</ref> A large percentage of Italy's white truffles also come from [[Molise]].

==== Whitish ====
{{Main|Tuber borchii}}
The "[[whitish truffle]]" (''Tuber borchii'') is a similar species native to [[Tuscany]], [[Abruzzo]], [[Romagna]], [[Umbria]], the [[Marche]], and [[Molise]]. It is reportedly not as aromatic as those from Piedmont, although those from [[Città di Castello]] are said to come quite close.<ref name="Carluccio032" />

==== Other ''Tuber'' ====
A less common truffle is "garlic truffle" (''[[Tuber macrosporum]]'').

In the U.S. [[Pacific Northwest]], several species of truffle are harvested both recreationally and commercially, most notably, the ''[[Leucangium carthusianum]]'', Oregon black truffle; ''[[Tuber gibbosum]]'', Oregon spring white truffle; and ''[[Tuber oregonense]]'', Oregon winter white truffle. ''[[Kalapuya brunnea]]'', the Oregon brown truffle, has also been commercially harvested and is of culinary note. The Oregon white truffle is increasingly harvested due to its high quality and also exported to other countries. Oregon celebrates its traditional truffle harvesting with a 'truffle festival', combined with culinary shows and wine tastings.<ref>[https://knowyourforest.org/sites/default/files/documents/Oregon_Native_Truffles.pdf "Oregon Native Truffles: A primer for commercial production on small forestlands in the Pacific Northwest"]. ''Know Your Forest''. Accessed 8 October 2023.</ref>

The [[pecan truffle]] (''Tuber lyonii'')<ref>{{cite journal|first=Fred K. |last=Butters|year=1903|title=A Minnesota Species of Tuber|journal=Botanical Gazette|volume=35|issue=6|pages=427–431|doi=10.1086/328364|jstor=2556357|s2cid=84500806}}</ref> syn. ''texense''<ref>{{cite journal|first1=J. M. |last1=Trappe |first2=A. M. |last2=Jumpponen |first3=E. |last3=Cázares|name-list-style=amp|year=1996|title=NATS truffle and truffle-like fungi 5: ''Tuber lyonii'' (=''T. texense''), with a key to the spiny-spored ''Tuber'' species groups|url=http://www.cybertruffle.org.uk/cyberliber/59575/0060/0365.htm|journal=Mycotaxon|volume=60|pages=365–372|access-date=2010-06-04|archive-date=2015-09-23|archive-url=https://web.archive.org/web/20150923232916/http://www.cybertruffle.org.uk/cyberliber/59575/0060/0365.htm|url-status=live}}</ref> is found in the [[Southern United States]], usually associated with [[pecan]] trees. Chefs who have experimented with them agree "they are very good and have potential as a food commodity".<ref>{{cite web|url=http://www.caes.uga.edu/commodities/fruits/pecantruffles/index.html|title=Pecan Truffles|author=Tim Brenneman|date=2010|archive-url=https://web.archive.org/web/20100609195108/http://www.caes.uga.edu/commodities/fruits/pecantruffles/index.html|archive-date=2010-06-09|url-status=dead|access-date=2010-06-03}}</ref> Although pecan farmers used to find them along with pecans and discard them, considering them a nuisance, they sell for about $160 a pound and have been used in some gourmet restaurants.<ref name="pecanmag2">{{cite magazine|last1=Smith |first1=M. E. |display-authors=etal |date=2012|title=Pecan Truffles ( ''Tuber lyonii'' ): What We Know and What We Need to Know|magazine=Georgia Pecan Magazine|issue=Spring 2012|pages=52–58}}</ref>

==== Beyond ''Tuber'' {{anchor|Similar organisms}} ====
{{missing information|section|taste and flavor|date=February 2022}}
The term "truffle" has been applied to several other genera of similar underground fungi. The genera ''[[Terfezia]]'' and ''[[Tirmania]]'' of the family [[Terfeziaceae]] are known as the "desert truffles" of Africa and the Middle East. ''[[Pisolithus tinctorius]]'', which was historically eaten in parts of [[Germany]], is sometimes called "Bohemian truffle".<ref name="Ramsbottom53">{{cite book |last=Ramsbottom |first=J. |title=Mushrooms & Toadstools |publisher=Collins |year=1953}}</ref>

{{anchor|Geopora}}''[[Geopora]]'' spp. are important ectomycorrhizal partners of trees in woodlands and forests throughout the world.<ref name="Læssøe 2007" /> ''Pinus edulis'', a widespread pine species of the Southwest US, is dependent on ''Geopora'' for nutrient and water acquisition in arid environments.<ref name="Flores-Rentería-2014">{{Cite journal|last1=Flores-Rentería|first1=Lluvia|last2=Lau|first2=Matthew K.|last3=Lamit|first3=Louis J.|last4=Gehring|first4=Catherine A.|date=2014|title=An elusive ectomycorrhizal fungus reveals itself: a new species of Geopora (Pyronemataceae) associated with ''Pinus edulis'' |journal=Mycologia|volume=106|issue=3 |pages=553–563 |doi=10.3852/13-263|pmid=24871594|s2cid=207630013|issn=0027-5514}}</ref> Like other truffle fungi, ''Geopora'' produces subterranean [[Sporocarp (fungi)|sporocarps]] as a means of sexual reproduction.<ref name="Flores-Rentería-2014" /> ''[[Geopora cooperi]]'', also known as pine truffle or fuzzy truffle, is an edible species of this genus.<ref name="Læssøe 2007" />

[[File:Rhizopogon rubescens.jpg|thumb|''Rhizopogon'' truffle]]
''[[Rhizopogon]]'' spp. are ectomycorrhizal members of the Basidiomycota and the order [[Boletales]], a group of fungi that typically form mushrooms.<ref name="Molina-1994">{{Cite journal|last1=Molina |first1=Randy|last2=Trappe |first2=James M.|date=April 1994 |title=Biology of the ectomycorrhizal genus, Rhizopogon. I. Host associations, host-specificity and pure culture syntheses |journal=New Phytologist |volume=126 |issue=4 |pages=653–675|doi=10.1111/j.1469-8137.1994.tb02961.x |issn=0028-646X|doi-access=free }}</ref> Like their ascomycete counterparts, these fungi can create truffle-like fruiting bodies.<ref name="Molina-1994" /> ''Rhizopogon'' spp. are ecologically important in coniferous forests where they associate with various pines, firs, and [[Douglas fir]].<ref name="Griffiths">{{Cite journal |last1=Griffiths |first1=Robert P. |last2=Caldwell |first2=Bruce A. |last3=Cromack |first3=Kermit Jr. |last4=Morita |first4=Richard Y. |date=February 1990 |title=Douglas-fir forest soils colonized by ectomycorrhizal mats. I. Seasonal variation in nitrogen chemistry and nitrogen cycle transformation rates  |journal=Canadian Journal of Forest Research |volume=20 |issue=2 |pages=211–218 |doi=10.1139/x90-030 |bibcode=1990CaJFR..20..211G |issn=0045-5067}}</ref> In addition to their ecological importance, these fungi hold economic value, as well. ''Rhizopogon'' spp. are commonly used to inoculate coniferous seedlings in nurseries and during reforestation.<ref name="Molina-1994" />

''[[Hysterangium]]'' spp. are ectomycorrhizal members of the Basidiomycota and the order [[Hysterangiales]] that form sporocarps similar to true truffles.<ref name="Kluber-2010">{{Cite journal|last1=Kluber|first1=Laurel A.|last2=Tinnesand|first2=Kathryn M.|last3=Caldwell|first3=Bruce A.|last4=Dunham|first4=Susie M.|last5=Yarwood|first5=Rockie R.|last6=Bottomley|first6=Peter J.|last7=Myrold|first7=David D.|date=2010|title=Ectomycorrhizal mats alter forest soil biogeochemistry |journal=Soil Biology and Biochemistry|volume=42|issue=9|pages=1607–1613|doi=10.1016/j.soilbio.2010.06.001 |bibcode=2010SBiBi..42.1607K |s2cid=85720492 |issn=0038-0717}}</ref> These fungi form mycelial mats of vegetative hyphae that may cover 25–40% of the forest floor in Douglas fir forests, thereby contributing to a significant portion of the biomass present in soils.<ref name="Kluber-2010" /> Like other ectomycorrhizal fungi, ''Hysterangium'' spp. play a role in nutrient exchange in the nitrogen cycle by accessing nitrogen unavailable to host plants and acting as nitrogen sinks in forests.<ref name="Griffiths" />

''[[Glomus (fungus)|Glomus]]'' spp. are [[arbuscular mycorrhiza]]e of the phylum Glomeromycota within the order [[Glomerales]].<ref name="Trappe 2009">{{Cite work |last1=Trappe |first1=James M. |title=Diversity, ecology, and conservation of truffle fungi in forests of the Pacific Northwest |last2=Molina |first2=Randy |last3=Luoma |first3=Daniel L. |last4=Cázares |first4=Efren |last5=Pilz |first5=David |last6=Smith |first6=Jane E. |last7=Castellano |first7=Michael A. |last8=Miller |first8=Steven L. |last9=Trappe |first9=Matthew J. |date=2009 |work=General Technical Report PNW-GTR-772 |publisher=USDA Forest Service |location=Portland, Oregon |doi=10.2737/pnw-gtr-772 |hdl=2027/umn.31951d02938269i}}</ref> Members of this genus have low host specificity, associating with a variety of plants including hardwoods, forbs, shrubs, and grasses.<ref name="Trappe 2009" /> These fungi commonly occur throughout the Northern Hemisphere.<ref name="Trappe 2009" />

Members of the genus ''[[Elaphomyces]]'' are commonly mistaken for truffles.

=== Phylogeny ===
[[File:Truffle Evolution.jpg|thumb|upright=1.2|Evolution of subterranean fruiting bodies from above-ground mushrooms.]]
Phylogenetic analysis has demonstrated the [[convergent evolution]] of the ectomycorrhizal trophic mode in diverse fungi. The subphylum [[Pezizomycotina]], containing the order Pezizales, is approximately 400 million years old.<ref name="Kohler">{{Cite journal |last=Kohler |first=Annegre |year=2015 |title=Convergent losses of decay mechanisms and rapid turnover of symbiosis genes in mycorrhizal mutualists |journal=Nature Genetics |location=Nature Genetics |publisher=Lawrence Berkeley National Laboratory, United States Department of Energy |volume=47 |issue=4 |pages=410–5 |doi=10.1038/ng.3223 |oclc=946824824 |pmid=25706625 |s2cid=20914242 |doi-access=free|hdl=1942/18722 |hdl-access=free }}</ref> Within the order Pezizales, subterranean fungi evolved independently at least fifteen times.<ref name="Kohler" /> Contained within Pezizales are the families [[Tuberaceae]], [[Pezizaceae]], [[Pyronemataceae|Pyronematacae]], and [[Morchellaceae]]. All of these families contain lineages of subterranean or truffle fungi.<ref name="Læssøe 2007" />

The oldest ectomycorrhizal fossils are from the [[Eocene]] about 50 million years ago. The specimens are preserved [[permineralized]] in-situ in the [[Eocene Okanagan Highlands]] [[Princeton chert]] site. This indicates that the soft bodies of ectomycorrhizal fungi do not easily fossilize.<ref>{{cite journal |last1=LePage |first1=B. A. |last2=Currah |first2=R. S. |last3=Stockey |first3=R. A. |last4=Rothwell |first4=G. W. |year=1997 |title=Fossil ectomycorrhizae from the middle Eocene |journal=American Journal of Botany |volume=84 |issue=3 |pages=410–412 |doi=10.2307/2446014 |jstor=2446014 |pmid=21708594 |s2cid=29913925}}</ref> Molecular clockwork has suggested the evolution of ectomycorrhizal fungi occurred approximately 130 million years ago.<ref>{{Cite journal |last1=Berbee |first1=Mary L. |last2=Taylor |first2=John W. |date=August 1993 |title=Dating the evolutionary radiations of the true fungi |journal=Canadian Journal of Botany |volume=71 |issue=8 |pages=1114–1127 |doi=10.1139/b93-131 |bibcode=1993CaJB...71.1114B |issn=0008-4026}}</ref>

The evolution of subterranean fruiting bodies has occurred numerous times within the [[Ascomycota]], [[Basidiomycota]], and [[Glomeromycota]].<ref name="Læssøe 2007" /> For example, the genera ''[[Rhizopogon]]'' and ''[[Hysterangium]]'' of Basidiomycota both form subterranean fruiting bodies and play similar ecological roles as truffle forming ascomycetes. The ancestors of the Ascomycota genera ''Geopora'', ''Tuber'', and ''Leucangium'' originated in Laurasia during the Paleozoic era.<ref name="Trappe 2009" />

Phylogenetic evidence suggests that most subterranean fruiting bodies evolved from above-ground mushrooms. Over time mushroom stipes and caps were reduced, and caps began to enclose reproductive tissue. The dispersal of sexual spores then shifted from wind and rain to utilising animals.<ref name="Trappe 2009" />

The [[phylogeny]] and [[biogeography]] of the genus ''Tuber'' was investigated in 2008<ref>{{cite journal |last1=Jeandroz |first1=S. |last2=Murat |first2=C. |last3=Wang |first3=Y. |last4=Bonfante |first4=P. |last5=Le Tacon |first5=F. |year=2008 |title=Molecular phylogeny and historical biogeography of the genus Tuber, the true truffles |journal=Journal of Biogeography |volume=35 |issue=5 |pages=815–829 |doi=10.1111/j.1365-2699.2007.01851.x |bibcode=2008JBiog..35..815J |s2cid=84381208}}</ref> using [[Internal transcribed spacer|internal transcribed spacers (ITS)]] of [[nuclear DNA]] and revealed five major [[clade]]s (Aestivum, Excavatum, Rufum, Melanosporum and Puberulum); this was later improved and expanded in 2010 to nine major clades using 28S [[Large subunit|large subunits (LSU)]] rRNA of [[mitochondrial DNA]]{{citation needed|date=October 2023}}. The Magnatum and Macrosporum clades were distinguished as distinct from the Aestivum clade. The Gibbosum clade was resolved as distinct from all other clades, and the Spinoreticulatum clade was separated from the Rufum clade.<ref>{{cite journal |vauthors=Bonito GM, Gryganskyi AP, Trappe JM, Vilgalys R |year=2010 |title=A global meta-analysis of Tuber ITS rDNA sequences: species diversity, host associations and long-distance dispersal |journal=PLOS ONE |volume=8 |issue=1 |pages=e52765 |bibcode=2013PLoSO...852765B |doi=10.1371/journal.pone.0052765 |pmc=3534693 |pmid=23300990 |doi-access=free}}</ref>

The truffle habit has evolved independently among several basidiomycete genera.<ref name="Bruns-1989">{{Cite journal |last1=Bruns |first1=Thomas D. |last2=Fogel |first2=Robert |last3=White |first3=Thomas J. |last4=Palmer |first4=Jeffrey D. |date=1989 |title=Accelerated evolution of a false-truffle from a mushroom ancestor |url=https://deepblue.lib.umich.edu/bitstream/2027.42/62545/1/339140a0.pdf |url-status=live |journal=Nature |volume=339 |issue=6220 |pages=140–142 |bibcode=1989Natur.339..140B |doi=10.1038/339140a0 |issn=0028-0836 |pmid=2716834 |s2cid=4312286 |archive-url=https://web.archive.org/web/20210827065108/https://deepblue.lib.umich.edu/bitstream/handle/2027.42/62545/339140a0.pdf;jsessionid=6C2FB0F2AE5763E76E2D55E2708812C9?sequence=1 |archive-date=2021-08-27 |access-date=2019-09-01 |hdl-access=free |hdl=2027.42/62545}}</ref><ref>{{Cite journal |last=Hibbett |first=David S. |date=2007 |title=After the gold rush, or before the flood? Evolutionary morphology of mushroom-forming fungi (Agaricomycetes) in the early 21st century |journal=Mycological Research |volume=111 |issue=9 |pages=1001–1018 |doi=10.1016/j.mycres.2007.01.012 |issn=0953-7562 |pmid=17964768}}</ref><ref>{{Cite journal |last=Albee-Scott |first=Steven |date=2007 |title=The phylogenetic placement of the Leucogastrales, including Mycolevis siccigleba (Cribbeaceae), in the Albatrellaceae, using morphological and molecular data |journal=Mycological Research |volume=111 |issue=6 |pages=653–662 |doi=10.1016/j.mycres.2007.03.020 |issn=0953-7562 |pmid=17604150}}</ref> Phylogenetic analysis has revealed that basidiomycete subterranean fruiting bodies, like their ascomycete counterparts, evolved from above ground mushrooms. For example, ''Rhizopogon'' species likely arose from an ancestor shared with ''[[Suillus]]'', a mushroom-forming genus.<ref name="Bruns-1989" /> Studies have suggested that selection for subterranean fruiting bodies among ascomycetes and basidiomycetes occurred in water-limited environments.<ref name="Trappe 2009" /><ref name="Bruns-1989" />

=== Etymology ===
Most sources agree that the term "truffle" is derived from the [[Latin]] term {{lang|la|tūber}} by way of the Vulgar Latin {{lang|la|tufera}}, meaning "swelling" or "lump".<ref>{{cite web |date=3 July 2008 |title=Truffle – Podictionary Word of the Day, July 3rd 2008 |url=https://blog.oup.com/2008/07/truffle/ |access-date=12 March 2023 |website=OUP Blog |publisher=Oxford University Press}}</ref><ref>{{cite dictionary |title=truffle |url=https://www.merriam-webster.com/dictionary/truffle |access-date=12 March 2023 |dictionary=Merriam-Webster}}</ref><ref>{{cite dictionary |title=truffle |url=https://www.dictionary.com/browse/truffle |access-date=12 March 2023 |dictionary=Dictionary.com}}</ref><ref>{{cite dictionary |title=truffle |url=https://www.etymonline.com/word/truffle |access-date=12 March 2023 |dictionary=Online Etymology Dictionary}}</ref> This then entered other languages through [[Old French]] dialects.

== Ecology ==
The [[Mycelium|mycelia]] of truffles form [[symbiotic]], mycorrhizal relationships with the roots of several tree species, including [[beech]], [[birch]], [[hazel]], [[hornbeam]], [[oak]], [[pine]], and [[Populus|poplar]].<ref>{{cite web|url=https://www.angelfire.com/wizard/kimbrough/Textbook/NonCultivatedEdibleFungi_blue.htm|title=Non-cultivated Edible Fleshy Fungi|access-date=2008-05-17|quote=...it has been known for more than a century that truffles were mycorrhizal on various trees such as oak, beech, birch, hazels, and a few others|archive-date=2021-03-15|archive-url=https://web.archive.org/web/20210315143933/https://www.angelfire.com/wizard/kimbrough/Textbook/NonCultivatedEdibleFungi_blue.htm|url-status=live}}</ref> Mutualistic ectomycorrhizal fungi such as truffles provide valuable nutrients to plants in exchange for [[carbohydrate]]s.<ref name="Allen-2003">{{Cite journal|last1=Allen|first1=M.F.|last2=Swenson|first2=W.|last3=Querejeta|first3=J.I.|last4=Egerton-Warburton|first4=L.M.|last5=Treseder|first5=K.K.|author5-link= Kathleen Treseder |date=2003|title=Ecology of ycorrhizae: A conceptual framework for complex interactions among plants and fungi |journal=Annual Review of Phytopathology|volume=41 |issue=1|pages=271–303 |doi=10.1146/annurev.phyto.41.052002.095518 |pmid=12730396|bibcode=2003AnRvP..41..271A |issn=0066-4286}}</ref> Ectomycorrhizal fungi cannot survive in the soil without their plant hosts.<ref name="Kohler" /> In fact, many of these fungi have lost the [[enzyme]]s necessary for obtaining carbon through other means. For example, truffle fungi have lost their ability to degrade the cell walls of plants, limiting their capacity to decompose plant litter.<ref name="Kohler" /> Plant hosts can also depend on their associated truffle fungi. <!-- Studies have demonstrated that --> ''Geopora'', ''Peziza'', and ''Tuber'' spp. are vital in the establishment of oak communities.<ref name="Frank-2006">{{Cite journal|last1=Frank|first1=J.L.|last2=Barry|first2=S.|last3=Southworth|first3=D.|date=2006|title=Mammal mycophagy and dispersal of mycorrhizal inoculum in Oregon white oak woodlands |journal=Northwest Science|volume=80 |pages=264–273}}</ref>

''Tuber'' species prefer [[Argillaceous minerals|argillaceous]] or [[calcareous]] soils that are well drained and [[PH|neutral]] or [[alkaline]].<ref>{{cite journal|last1=Jaillard|first1=B|last2=Barry-Etienne|first2=D|last3=Colinas|first3=C|last4=de Miguel|first4=AM|last5=Genola|first5=L|last6=Libre|first6=A|last7=Oliach|first7=D|last8=Saenz|first8=W|last9=Saez|first9=M|date=2014|title=Alkalinity and structure of soils determine the truffle production in the Pyrenean Regions|url=http://arxiudigital.ctfc.cat/docs/upload/27_457_OliachColinas.pdf|journal=Forest Systems|volume=23|issue=2|pages=364–377|doi=10.5424/fs/2014232-04933|last10=Salducci|first10=X|last11=Souche|first11=G|last12=Sourzat|first12=P|last13=Villeneuve|first13=M|doi-access=free|access-date=2016-07-27|archive-date=2016-08-06|archive-url=https://web.archive.org/web/20160806102755/http://arxiudigital.ctfc.cat/docs/upload/27_457_OliachColinas.pdf|url-status=live}}</ref><ref>{{cite web|url=http://www.huh.harvard.edu/collections/fof/newsletters/newslspring2006.pdf|title=Basidiomycota truffles: Cup fungi go underground|author=Hansen|first=Karen|date=2006|website=Newsletter of the FRIENDS of the FARLOW|publisher=Harvard University|archive-url=https://web.archive.org/web/20081121001628/http://www.huh.harvard.edu/collections/fof/newsletters/newslSpring2006.pdf|archive-date=2008-11-21|url-status=dead|access-date=2008-05-17|quote=Generally, truffles seems to prefer warm, fairly dry climates and calcareous soils}}</ref><ref>{{Citation|last1=Chevalier|first1=Gérard|date=2012 |doi=10.1007/978-3-642-33823-6_10 |pages=163–189|publisher=Springer Berlin Heidelberg|isbn=9783642338229|last2=Sourzat|first2=Pierre|title=Edible Ectomycorrhizal Mushrooms |chapter=Soils and Techniques for Cultivating Tuber melanosporum and Tuber aestivum in Europe |series=Soil Biology |volume=34 }}</ref> ''Tuber'' truffles fruit throughout the year, depending on the species, and can be found buried between the [[Plant litter|leaf litter]] and the soil. Most fungal biomass is found in the [[humus]] and litter layers of soil.<ref name="Griffiths" />

[[File:Pezizales, Ascomycota Life Cycle.jpg|thumb|268x268px|The lifecycle of the order Pezizales in Ascomycota]]
Most truffle fungi produce both asexual spores (mitospores or [[Conidium|conidia]]) and sexual spores (meiospores or [[ascospore]]s/[[basidiospore]]s).<ref name="Tedersoo-2013">{{Cite journal|last1=Tedersoo|first1=Leho|last2=Arnold|first2=A. Elizabeth|last3=Hansen|first3=Karen|date=2013|title=Novel aspects in the life cycle and biotrophic interactions in Pezizomycetes (Ascomycota, Fungi) |journal=Molecular Ecology|volume=22|issue=6|pages=1488–1493|doi=10.1111/mec.12224|pmid=23599958|bibcode=2013MolEc..22.1488T |s2cid=45317735|issn=0962-1083|doi-access=free}}</ref> Conidia can be produced more readily and with less energy than ascospores and can disperse during disturbance events. Production of ascospores is energy intensive because the fungus must allocate resources to the production of large [[Sporocarp (fungi)|sporocarps]].<ref name="Tedersoo-2013" /> Ascospores are borne within sac-like structures called asci, which are contained within the sporocarp.

Because truffle fungi produce their sexual fruiting bodies underground, spores cannot be spread by wind and water. Therefore, nearly all truffles depend on [[Fungivore|mycophagous]] animal vectors for spore dispersal.<ref name="Læssøe 2007" /> This is analogous to the dispersal of seeds in fruit of angiosperms. When the ascospores are fully developed, the truffle exudes volatile compounds that attract animal vectors.<ref name="Læssøe 2007" /> For successful dispersal, these spores must survive passage through the digestive tracts of animals. Ascospores have thick walls composed of [[chitin]] to help them endure the environment of animal guts.<ref name="Tedersoo-2013" />

Animal vectors include birds, deer, and rodents such as [[vole]]s, [[squirrel]]s, and [[chipmunk]]s.<ref name="Læssøe 2007" /><ref name="Frank-2006" /><ref>{{Cite journal|last1=Ashkannejhad|first1=Sara|last2=Horton|first2=Thomas R.|date=2005|title=Ectomycorrhizal ecology under primary succession on coastal sand dunes: interactions involving Pinus contorta, suilloid fungi and deer |journal=New Phytologist |volume=169|issue=2|pages=345–354|doi=10.1111/j.1469-8137.2005.01593.x |issn=0028-646X |pmid=16411937|doi-access=free}}</ref> Many species of trees, such as ''[[Quercus garryana]]'', are dependent on the dispersal of sporocarps to inoculate isolated individuals. For example, the acorns of ''Q. garryana'' may be carried to new territory that lacks the necessary mycorrhizal fungi for establishment.<ref name="Frank-2006" />

Some mycophagous animals depend on truffles as their dominant food source. Flying squirrels, ''[[Northern flying squirrel|Glaucomys sabrinus]]'', of North America play a three-way symbiosis with truffles and their associated plants.<ref name="Læssøe 2007" /> ''G. sabrinus'' is particularly adapted to finding truffles using its refined sense of smell, visual clues, and long-term memory of prosperous populations of truffles.<ref name="Læssøe 2007" /> This intimacy between animals and truffles indirectly influences the success of mycorrhizal plant species.

After ascospores are dispersed, they remain dormant until germination is initiated by exudates excreted from host plant roots.<ref name="Pacioni-1989">{{Cite journal|last=Pacioni|first=G.|date=1989|title=Biology and ecology of the truffles |journal=Acta Medica Romana|volume=27|pages=104–117}}</ref> Following germination, hyphae form and seek out the roots of host plants. Arriving at roots, hyphae begin to form a mantle or sheath on the outer surface of root tips. Hyphae then enter the root cortex intercellularly to form the [[Hartig net]] for nutrient exchange. Hyphae can spread to other root tips colonising the entire root system of the host.<ref name="Pacioni-1989" /> Over time, the truffle fungus accumulates sufficient resources to form fruiting bodies.<ref name="Pacioni-1989" /><ref name="Frank-2006" /> Rate of growth is correlated with increasing photosynthetic rates in the spring as trees leaf out.<ref name="Frank-2006" />

=== Nutrient exchange ===
Truffle fungi receive carbohydrates from their host plants, providing them with valuable micro- and macronutrients. Plant macronutrients include [[potassium]], [[phosphorus]], [[nitrogen]], and [[sulfur]]. In contrast, micronutrients include [[iron]], [[copper]], [[zinc]], and [[chloride]].<ref name="Allen-2003" /> In truffle fungi, as in all ectomycorrhizae, the majority of nutrient exchange occurs in the Hartig net, the intercellular hyphal network between plant root cells. A unique feature of ectomycorrhizal fungi is the formation of the mantle on the outer surface of fine roots.<ref name="Allen-2003" />

Truffles have been suggested to co-locate with the orchid species ''[[Epipactis helleborine]]'' and ''[[Cephalanthera damasonium]]'',<ref>{{Cite web|url=https://www.researchgate.net/publication/254073638|title=Acta Biologica Szegediensis - Could orchids indicate truffle habitat? Mycorrhizal association between orchids and truffles|access-date=2018-12-19|archive-date=2021-08-27|archive-url=https://web.archive.org/web/20210827065109/https://www.researchgate.net/publication/254073638_Could_orchids_indicate_truffle_habitats_Mycorrhizal_association_between_orchids_and_truffles|url-status=live}}</ref> though this is not always the case.

=== Nutrient cycling ===
Truffle fungi are ecologically important in nutrient cycling. Plants obtain nutrients via their fine roots. Mycorrhizal fungi are much smaller than fine roots, so they have a higher surface area and a greater ability to explore soils for nutrients. Acquisition of nutrients includes the uptake of phosphorus, nitrate or ammonium, iron, [[magnesium]], and other [[ion]]s.<ref name="Allen-2003" /> Many ectomycorrhizal fungi form fungal mats in the upper layers of soils surrounding host plants. These mats have significantly higher carbon and fixed nitrogen concentrations than surrounding soils.<ref>{{Cite journal|last1=Cromack|first1=Kermit|last2=Fichter|first2=B.L.|last3=Moldenke|first3=A.M.|last4=Entry|first4=J.A.|last5=Ingham|first5=E.R.|date=1988|title=Interactions between soil animals and ectomycorrhizal fungal mats |journal=Agriculture, Ecosystems & Environment|volume=24|issue=1–3|pages=161–168|doi=10.1016/0167-8809(88)90063-1|bibcode=1988AgEE...24..161C |issn=0167-8809}}</ref> Because these mats are nitrogen sinks, leaching of nutrients is reduced.<ref name="Griffiths" />

Mycelial mats can also help maintain the structure of soils by holding organic matter in place and preventing [[erosion]].<ref name="Trappe 2009" /> Often, these networks of mycelium provide support for smaller organisms in the soil, such as bacteria and microscopic [[arthropod]]s. Bacteria feed on the exudates released by mycelium and colonise the soil surrounding them.<ref>{{cite book |last1=Reddy |first1=M. S. |last2=Satyanarayana |first2=T. |date=2006 |chapter=Interactions between ectomycorrhizal fungi and rhizospheric microbes |title=Microbial Activity in the Rhizoshere |series=Soil Biology |volume=7 |publisher=Springer |location=Berlin, Heidelberg |pages=245–263 |doi=10.1007/3-540-29420-1_13|isbn=3-540-29182-2 }}</ref> Microscopic arthropods such as mites feed directly on mycelium and release valuable nutrients for the uptake of other organisms.<ref>Moldenke, A.R., 1999. Soil-dwelling arthropods: their diversity and functional roles. United States Department of Agriculture Forest Service General Technical Report PNW. 33-44.</ref> Thus, truffle fungi and other ectomycorrhizal fungi facilitate a complex system of nutrient exchange between plants, animals, and microbes.

=== Importance in arid-land ecosystems ===
Plant community structure is often affected by the availability of compatible mycorrhizal fungi.<ref>{{Cite journal|last1=Hartnett|first1=David C.|last2=Wilson|first2=Gail W. T.|date=1999|title=Mycorrhizae influence plant community structure and diversity in tallgrass prairie |journal=Ecology|volume=80|issue=4|pages=1187|doi=10.2307/177066|jstor=177066|issn=0012-9658}}</ref><ref>{{Cite journal|last1=Haskins|first1=Kristin E.|last2=Gehring|first2=Catherine A.|date=2005|title=Evidence for mutualist limitation: the impacts of conspecific density on the mycorrhizal inoculum potential of woodland soils |journal=Oecologia|volume=145|issue=1|pages=123–131|doi=10.1007/s00442-005-0115-3|pmid=15891858|bibcode=2005Oecol.145..123H|s2cid=3102834|issn=0029-8549}}</ref> In arid-land ecosystems, these fungi become essential for the survival of their host plants by enhancing the ability to withstand drought.<ref>{{Cite journal|last1=Lehto|first1=Tarja|last2=Zwiazek|first2=Janusz J.|date=2010|title=Ectomycorrhizas and water relations of trees: a review |journal=Mycorrhiza|volume=21|issue=2|pages=71–90|doi=10.1007/s00572-010-0348-9|pmid=21140277|s2cid=20983637|issn=0940-6360}}</ref> A foundation species in arid-land ecosystems of the Southwest United States is ''[[Pinus edulis]]'', commonly known as pinyon pine. ''P. edulis'' associates with the subterranean fungi ''Geopora'' and ''Rhizopogon''.<ref name="Gehring-2017">{{Cite journal|last1=Gehring|first1=Catherine A.|last2=Sthultz|first2=Christopher M.|last3=Flores-Rentería|first3=Lluvia|last4=Whipple|first4=Amy V.|last5=Whitham|first5=Thomas G.|date=2017|title=Tree genetics defines fungal partner communities that may confer drought tolerance |journal=Proceedings of the National Academy of Sciences|volume=114|issue=42|pages=11169–11174|doi=10.1073/pnas.1704022114|issn=0027-8424|pmc=5651740|pmid=28973879|bibcode=2017PNAS..11411169G |doi-access=free}}</ref>

As global temperatures rise, so does the occurrence of severe droughts, detrimentally affecting the survival of aridland plants. This variability in climate has increased the mortality of ''P. edulis''.<ref>{{Cite journal|last1=Ogle|first1=Kiona|last2=Whitham|first2=Thomas G.|last3=Cobb|first3=Neil S.|date=2000|title=Tree-ring variation in pinyon predicts likelihood of death following severe drought |journal=Ecology|volume=81|issue=11|pages=3237|doi=10.2307/177414|jstor=177414|issn=0012-9658}}</ref> Therefore, the availability of compatible mycorrhizal inoculum can greatly affect the successful establishment of ''P. edulis'' seedlings.<ref name="Gehring-2017" /> Associated ectomycorrhizal fungi will likely play a significant role in the survival of ''P. edulis'' with continuing global [[climate change]].{{citation needed|date=June 2020}}

== Extraction ==
[[File:Cochon_truffier.JPG|thumb|A trained truffle hunting [[pig]] in [[Gignac, Lot]], France]]
[[File:Dog_smelling_for_truffles_in_Mons,_Var.jpg|thumb|A trained truffle hunting [[dog]] in [[Mons, Var]], France]]
{| class="wikitable"
|+Comparison of truffle dog and hog
!Truffle dog
!Truffle hog
|-
|Keen sense of smell
|Keen sense of smell
|-
|Must be trained
|Innate ability to sniff out truffles
|-
|Easier to control
|Tendency to eat truffles once found
|}
Because truffles are subterranean, they are often located with the help of an animal (sometimes called a '''truffler'''<ref>{{oed | truffler}}</ref>) possessing a refined sense of smell. Traditionally, [[pig]]s have been used to extract truffles.<ref name="Talou-1990">{{Cite journal|last1= Talou|first1= T.|last2=G aset|first2= A.|last3= Delmas|first3= M.|last4= Kulifaj |first4= M.|last5=Montant|first5=C.|date=1990|title=Dimethyl sulphide: the secret for black truffle hunting by animals? |journal= Mycological Research|volume= 94|issue= 2|pages= 277–278|doi= 10.1016/s0953-7562(09)80630-8|issn=0953-7562}}</ref> Both the female pig's natural truffle-seeking and her intent to eat the truffle were thought to be due to a compound within the truffle similar to [[androstenol]], the sex [[pheromone]] of boar saliva, to which the sow is keenly attracted. Studies in 1990 demonstrated that the compound actively recognised by both truffle pigs and dogs is [[dimethyl sulfide]].<ref name="Talou-1990" />

In Italy, the use of pigs to hunt truffles has been prohibited since 1985 because of damage caused by animals to truffle [[mycelia]] during the digging that dropped the production rate of the area for some years. An alternative to truffle pigs are dogs. Dogs offer an advantage because they do not have a strong desire to eat truffles, so they can be trained to locate sporocarps without digging them up. Pigs attempt to dig up truffles.<ref name="Talou-1990" />

Fly species of the genus ''[[Suillia tuberiperda|Suillia]]'' can also detect the volatile compounds associated with subterranean fruiting bodies. These flies lay their eggs above truffles to provide food for their young. At ground level, ''Suilla'' flies can be seen flying above truffles.<ref name="Talou-1990" />

== Volatile constituents ==
{{External media | width = 210px | float = right | headerimage= <!-- [[File:|210px]] --> | video1 = [https://www.youtube.com/watch?v=ltXd2BOrIHs "The Chemistry of Truffles, the Most Expensive Food in the World"], Sarah Everts, [[Chemical & Engineering News|CEN Online]]}}

The mycelia or fruiting bodies release the volatile constituents responsible for the natural aroma of truffles or derive from truffle-associated [[microbe]]s.  The [[chemical ecology]] of truffle volatiles is complex, interacting with plants, insects, and mammals, which contribute to [[spore]] dispersal. Depending on the truffle species, lifecycle, or location, they include:
* [[Sulfur]] [[volatility (chemistry)|volatiles]], which occur in all truffle species, such as dimethyl mono- (DMS), di- (DMDS) and tri- (DMTS) sulfides, as well as 2-methyl-4,5-dihydrothiophene, characteristic of the white truffle ''T. borchii'' and [[2,4-Dithiapentane]] occurring in all species but mostly characteristic of the white truffle ''T. magnatum''. Some very aromatic white truffles are notably pungent, even irritating the eye when cut or sliced.
* Metabolites of nonsulfur [[amino acid]] constituents (simple and branched-chain [[hydrocarbons]]) such as [[ethylene]] (produced by mycelia of white truffles affecting root architecture of host tree), as well as 2-methylbutanal, 2-methylpropanal, and [[2-phenylethanol]] (also common in baker's yeast).
* [[Fatty acid]]-derived volatiles (C8-alcohols and [[aldehyde]]s with a characteristic fungal odor, such as [[1-octen-3-ol]] and 2-octenal). The former is derived from [[linoleic acid]] and produced by mature white truffle ''T. borchii''.
* [[Thiophene]] derivatives appear to be produced by bacterial symbionts living in the truffle body. The most abundant of these, 3-methyl, 4-5 dihydrothiophene, contributes to the white truffle's aroma.<ref>{{cite journal | last1 = Splivallo | first1 = R | last2 = Ebeler | first2 = SE | date = Mar 2015 | title = Sulfur volatiles of microbial origin are key contributors to human-sensed truffle aroma | journal = Appl Microbiol Biotechnol | volume = 99 | issue = 6| pages = 2583–92 | doi = 10.1007/s00253-014-6360-9 | pmid = 2557347 | s2cid = 16749990 }}</ref><ref>{{cite journal | last1 = Splivallo | first1 = R | last2 = Deveau | first2 = A | last3 = Valdez | first3 = N | last4 = Kirchhoff | first4 = N | last5 = Frey-Klett | first5 = P | last6 = Karlovsky | first6 = P | date = Aug 2015 | title = Bacteria associated with truffle-fruiting bodies contribute to truffle aroma | journal = Environ Microbiol | volume = 17 | issue = 8| pages = 2647–60 | doi = 10.1111/1462-2920.12521 | pmid = 24903279 | bibcode = 2015EnvMi..17.2647S | doi-access = free }}</ref>
Several truffle species and varieties are differentiated based on their relative contents or absence of [[sulfide]]s, [[ether]]s or [[Alcohol (chemistry)|alcohol]]s, respectively. The sweaty-musky aroma of truffles is similar to that of the [[pheromone]] [[androstenol]] that also occurs in humans.<ref>[http://www.delectations.org/aroma.html Delectations]{{Webarchive|url=https://web.archive.org/web/20151113140927/http://www.delectations.org/aroma.html |date=2015-11-13 }}. Truffle Aroma. Retrieved December 19, 2015.</ref> {{As of|2010}}, the volatile profiles of seven black and six white truffle species have been studied.<ref>{{cite journal | last1 = Splivallo | first1 = R | display-authors = etal | year = 2010 | title = Truffle Volatiles: from chemical ecology to aroma biosynthesis | journal = New Phytologist | volume = 198 | issue = 3| pages = 688–699 | doi = 10.1111/j.1469-8137.2010.03523.x | pmid = 21287717 | doi-access =  | url = https://resolver.sub.uni-goettingen.de/purl?gro-2/24525 }}</ref>

== Cultivation ==
{{hatnote|This section focuses on the cultivation of truffles in the narrow sense, i.e. ''Tuber'' spp.}}

[[File:Joseph Talon.JPG|thumb|Statue of [[Joseph Talon]] in [[Saint-Saturnin-lès-Apt]]]]
[[File:Beaumont - truffières.jpg|thumb|Planted truffle groves near [[Beaumont-du-Ventoux]]]]

Truffles long eluded techniques of cultivation, as [[Jean-Anthelme Brillat-Savarin]] (1825) noted:

<blockquote>The most learned men have sought to ascertain the secret and fancied they discovered the seed. Their promises, however, were vain, and no planting was ever followed by a harvest. This perhaps is all right, for as one of the great values of truffles is their dearness, perhaps they would be less highly esteemed if they were cheaper.<ref name="brillat" /></blockquote>

Truffles can be cultivated. As early as 1808, attempts to cultivate truffles, known in French as {{lang|fr|trufficulture}}, were successful. People had long observed that truffles were growing among the roots of certain trees, and in 1808, [[Joseph Talon]], from [[Apt, Vaucluse|Apt]] ({{lang|fr|[[département in France|département]]}} of [[Vaucluse]]) in southern [[France]], had the idea of transplanting some seedlings that he had collected at the foot of oak trees known to host truffles in their root system.<ref name="Zambonelli-2013">{{Cite book |last1=Zambonelli |first1=Alessandra |url=https://link.springer.com/content/pdf/10.1007/978-3-642-33823-6.pdf |title=Edible Ectomycorrhizal Mushrooms: Current Knowledge and Future Prospects |last2=Bonito |first2=Gregory M. |date=2013-01-30 |publisher=Springer Science & Business Media |isbn=978-3-642-33823-6 |pages=3–16 |language=en |chapter=Laying the Foundations |doi=10.1007/978-3-642-33823-6 |s2cid=25454181}}</ref>

For discovering how to cultivate truffles, some sources now give priority to Pierre II Mauléon (1744–1831) of [[Loudun]] (in western France), who began to cultivate truffles around 1790. Mauléon saw an "obvious symbiosis" between the oak tree, the rocky soil, and the truffle and attempted to reproduce such an environment by taking acorns from trees known to have produced truffles and sowing them in chalky soil.<ref>See:  Thérèse Dereix de Laplane (2010) "Des truffes sauvages aux truffes cultivées en Loudunais" (From wild truffles to cultivated truffles in the area of Loudun), ''Mémoires de l'Académie des Sciences, Arts et Belles-Lettres de Touraine'', '''23''' :  215–241.  Available on-line at:  [http://academie-de-touraine.com/Tome_23_files/191_215-241.pdf Academy of Touraine] {{Webarchive|url=https://web.archive.org/web/20160506053635/http://academie-de-touraine.com/Tome_23_files/191_215-241.pdf|date=2016-05-06}}  From pp.&nbsp;224–225: "{{lang|fr|le paysan, a alors l'idée, vers 1790 — puisqu'il y a symbiose évidente entre le chêne, les galluches et la truffe — de provoquer la formation de truffières, en reproduisant leur environnement naturel par des semis de glands dans ses "terres galluches".  Avec "les glands venus sur les chênes donnant les truffes, des semis furent faits dans les terrains calcaires voisins"}}" (the farmer [viz, Pierre II Mauléon] then had the idea, around 1790 — because there is an obvious symbiosis between the oak tree, the rocky soil, and truffles — of inducing the formation of truffle patches, by reproducing their natural environment by sowing acorns in his rocky soils.  With "the acorns [that] came from the oak trees producing truffles, sowings were made in the neighbouring chalky plots")</ref><ref>[https://books.google.com/books?id=97oAAAAAYAAJ&pg=RA1-PA300 "Culture de la truffe à Loudun et à Richelieu,"] {{Webarchive|url=https://web.archive.org/web/20170403050208/https://books.google.com/books?id=97oAAAAAYAAJ&pg=RA1-PA300|date=2017-04-03}} ''Annales de la Société d'Agriculture Sciences, Arts, et Belles-lettres du Départment d'Indre-et-Loire'', 10th series, '''48''' : 300–302 (1869); see p.&nbsp;300.</ref> His experiment was successful, with truffles found in the soil around the newly grown oak trees years later. In 1847, [[Auguste Rousseau]] of [[Carpentras]] (in [[Vaucluse]]) planted {{convert|7|ha|acre|abbr=off}} of oak trees (again from acorns found on the soil around truffle-producing oak trees), and he subsequently obtained large harvests of truffles. He received a prize at the [[Exposition Universelle (1855)|1855 World's Fair]] in [[Paris]].<ref>Rousseau, "Truffes obtenues par la culture de chênes verts" (Truffles obtained by the cultivation of green oaks) in:  ''Exposition universelle de 1855 : Rapports du jury mixte international'', volume 1 (Paris, France:  Imprimerie Impériale, 1856), [https://books.google.com/books?id=m1ZAAAAAcAAJ&pg=PA173 pp. 173-174.] {{Webarchive|url=https://web.archive.org/web/20160518094525/https://books.google.com/books?id=m1ZAAAAAcAAJ&pg=PA173|date=2016-05-18}}</ref>

[[File:Truffle Market in Carpentras.jpg|thumb|A truffle market in [[Carpentras]], France]]
Others imitated these successful attempts in France and Italy.<ref name="Zambonelli-2013" /><ref name="perigord">{{Cite web |title=perigord.co // Truffes L&Co |url=https://www.perigord.co/en-gb/histoire-de-la-trufficulture |access-date=2022-08-18 |website=www.perigord.co |language=en}}</ref> In the late 19th century, an epidemic of [[phylloxera]] destroyed many of the vineyards in southern France. Another epidemic killed most of the [[silkworm]]s there, too, making the fields of [[mulberry]] trees useless. Trufficulture became an important source of income for those affected.<ref name="Zambonelli-2013" /><ref name="Van Vleet-2018">{{Cite thesis |title=Truffles Have Never Been Modern: An Actor-Network Theorization of 150 Years of French Trufficulture |publisher=Florida International University |first=Eric |last=Van Vleet |year=2018 |doi=10.25148/etd.fidc006559 |doi-access=free}}</ref> The calcareous and exposed vineyard soils were well-suited to the cultivation of truffles.<ref name="perigord" /> By 1890, truffières (truffle plantations) covered 750 km<sup>2</sup> of land in France, and 2,000 tonnes of truffles were produced in that year.<ref name="Zambonelli-2013" />

From the 19th century to the present, truffle production fell by 97–99% to 20–50 tonnes annually.<ref name="History and curiosities of the truffle-2015">{{Cite web |date=October 2015 |title=History and curiosities of the truffle – FoodsCross |url=https://foodscross.com/en/j/whats-cooking/history-and-curiosities-of-the-truffle/ |access-date=2022-08-18 |language=en-US}}</ref> Reasons given for this decline include the [[Industrial Revolution]], the subsequent [[rural flight]] and the multiple European wars of the 20th century, which reduced the rural population.<ref name="perigord" /><ref name="Van Vleet-2018" /><ref name="History and curiosities of the truffle-2015" /> For example, [[World War I]] resulted in the mobilisation of 65% of the agricultural workers from the region of [[Lot (department)|Lot]] alone.<ref name="Van Vleet-2018" /> Knowledge of truffle cultivation, the soil and the seasons was lost along with the people.<ref name="perigord" /> Another consequence was no more grazing sheep or shepherds who pruned trees for feed and fuelwood, so former truffle plantations turned into closed forests that no longer produced truffles.<ref name="Van Vleet-2018" /> Truffles were once sold at weekly markets (bi-weekly in the case of a market in [[Martel, Lot]]) and in quantities of two to six tonnes in good weeks, but only [[Lalbenque]] and [[Limogne-en-Quercy|Limogne]] today have weekly truffle markets.<ref name="Van Vleet-2018" /> Prices have increased so that truffles, once seen as a food of the middle class, have become a luxury.<ref name="Van Vleet-2018" />

The situation changed in the late 1960s and early 1970s, with researchers in France and Italy establishing mycorrhizas with truffle spores.<ref name="Zambonelli-2013" /> Beginning from the 1980s, truffle plantations are compensating for some of the decline in wild truffles, and exist in various countries including France, Italy, Spain and Australia.<ref name="Bungten-2015">{{cite journal |last1=Bungten |first1=Ulf |last2=Egli |first2=Simon |last3=Schneider |first3=Loic |last4=Von Arx |first4=Georg |last5=Rigling |first5=Andreas |last6=Camarero |first6=Julio |last7=Sangüesa |first7=Gabriel |last8=Fischer |first8=Christine |last9=Oliach |first9=Daniel |last10=Bonet |first10=Jose-Antonio |last11=Colinas |first11=Carlos |last12=Tegel |first12=Willy |last13=Ruiz |first13=Jose |last14=Martinez |first14=Fernando |date=2015 |title=Long-term irrigation effects on Spanish holm oak growth and its black truffle symbiont |url=https://repositori.udl.cat/bitstream/10459.1/65049/3/022489%20.pdf |url-status=live |journal=Agriculture, Ecosystems and Environment |volume=202 |pages=148–159 |doi=10.1016/j.agee.2014.12.016 |bibcode=2015AgEE..202..148B |archive-url=https://web.archive.org/web/20210827065120/https://repositori.udl.cat/bitstream/handle/10459.1/65049/022489%20.pdf;jsessionid=B40FA640177C921A181F8090B3D7B9D4?sequence=3 |archive-date=2021-08-27 |access-date=2019-09-24 |hdl-access=free |hdl=10261/113281}}</ref> About 80% of the truffles now produced in France come from specially planted truffle groves.{{citation needed|date=November 2020}} Investments in cultivated plantations are underway in many parts of the world using controlled irrigation for regular and resilient production.<ref name="Bungten-2015" /><ref>{{cite journal |last1=Olivera |first1=Antoni |last2=Fischer |first2=Christine |last3=Bonet |first3=Jose-Antonio |last4=Martinez de Aragon |first4=Juan |last5=Oliach |first5=Daniel |last6=Colinas |first6=Carlos |date=2011 |title=Weed management and irrigation are key treatments in emerging black truffle (Tuber melanosporum) cultivation |journal=New Forests |volume=42 |issue=2 |pages=227–239 |doi=10.1007/s11056-011-9249-9 |bibcode=2011NewFo..42..227O |s2cid=11586599}}</ref>

A critical phase of the cultivation is the quality control of the mycorrhizal plants. Between 7 and 10 years are needed for the truffles to develop their mycorrhizal network, and only after that do the host plants come into production. Complete soil analysis to avoid contamination by other dominant fungi and very strict control of the formation of mycorrhizae are necessary to ensure the success of a plantation. Total investment per hectare for an irrigated and barrier-sealed plantation (against wild boars) can cost up to €10,000.<ref>{{cite journal |last1=Oliach |first1=Daniel |last2=Muxi |first2=Pere |date=2012 |title=Estudi tècnic i econòmic del cultiu de la tòfona (in catalan) |url=http://cpf.gencat.cat/web/.content/or_organismes/or04_centre_propietat_forestal/01_organisme/publicacions/revista_silvicultura/numeros_publicats/silvicultura_66/silvi_66_edicio_digital.pdf |url-status=live |journal=Silvicultura |volume=66 |pages=8–10 |archive-url=https://web.archive.org/web/20160305025259/http://cpf.gencat.cat/web/.content/or_organismes/or04_centre_propietat_forestal/01_organisme/publicacions/revista_silvicultura/numeros_publicats/silvicultura_66/silvi_66_edicio_digital.pdf |archive-date=2016-03-05 |access-date=2016-07-27}}</ref> Considering the level of initial investment and the maturity delay, farmers who have not taken care of both soil conditions and seedling conditions are at high risk of failure.

=== New Zealand and Australia ===
The first black truffles (''[[Tuber melanosporum]]'') to be produced in the Southern Hemisphere were harvested in [[Gisborne, New Zealand]] in 1993.<ref name="NZ Truffle Association">{{cite web |title=Truffles in New Zealand |url=http://www.southern-truffles.co.nz/html/truffles_in_nz.html |url-status=dead |archive-url=https://web.archive.org/web/20130504065111/http://www.southern-truffles.co.nz/html/truffles_in_nz.html |archive-date=2013-05-04 |access-date=2012-07-19 |publisher=Southern_truffles.co.nz}}</ref>

New Zealand's first burgundy truffle was found in July 2012 at a Waipara truffle farm. It weighed 330 g and was found by the farm owner's beagle.<ref name="Stuff-burgundy truffle">{{cite web |date=18 July 2012 |title=Beagle digs up a New Zealand first |url=http://www.stuff.co.nz/life-style/food-wine/7305450/Beagle-digs-up-a-New-Zealand-first |url-status=live |archive-url=https://web.archive.org/web/20120721090701/http://www.stuff.co.nz/life-style/food-wine/7305450/Beagle-digs-up-a-New-Zealand-first |archive-date=2012-07-21 |access-date=2012-07-26 |publisher=stuff.co.nz}}</ref>

In 1999, the first Australian truffles were harvested in [[Tasmania]],<ref>{{cite book |url=https://books.google.com/books?id=vUo_AAAAQBAJ&pg=PA193 |title=Edible Ectomycorrhizal Mushrooms: Current Knowledge and Future Prospects |date=2013 |publisher=Springer |isbn=978-3-64233822-9 |editor1-last=Zambonelli |editor1-first=Alessandra |location=Germany |pages=193 |access-date=2020-09-05 |editor2-last=Bonito |editor2-first=Gregory M. |archive-url=https://web.archive.org/web/20210827065107/https://books.google.com/books?id=vUo_AAAAQBAJ&pg=PA193 |archive-date=2021-08-27 |url-status=live}}</ref> the result of eight years of work. Trees were [[Inoculation|inoculated]] with the truffle fungus to create a local truffle industry. Their success and the value of the resulting truffles has encouraged a small industry to develop.

Truffle production has expanded into the colder regions of [[Victoria (Australia)|Victoria]], [[New South Wales]] and [[Western Australia]].<ref>{{cite web |title=Growing Truffles |url=https://trufflegrowers.com.au/growing-truffles/ |access-date=2021-08-29 |publisher=Australian Truffle Industry Association}}</ref> In 2014, over {{convert|5000|kg|lb|abbr=off}} of truffles were harvested by Truffle Hill, Manjimup, Western Australia.

In June 2014, a grower harvested Australia's largest truffle from their property at [[Robertson, New South Wales|Robertson]], in the [[Southern Highlands (New South Wales)|Southern Highlands]] of New South Wales. It was a French black [[Périgord truffle|périgord]] fungus weighing in at {{convert|1.1172|kg|lboz|frac=16|abbr=on}} and was valued at over $2,000 per kilogram.<ref>{{cite web |date=30 June 2014 |title=Australia's largest truffle, worth $2,000 a kilo, grown in NSW town of Robertson |url=https://www.abc.net.au/news/2014-06-30/largest-truffle-ever-grown-in-australia-french-black-perigold/5559260 |archive-url=https://web.archive.org/web/20140630075053/http://www.abc.net.au/news/2014-06-30/largest-truffle-ever-grown-in-australia-french-black-perigold/5559260 |archive-date=2014-06-30 |work=ABC News (Australia)}}</ref>

=== United States ===

Périgord truffles were first farmed in Tennessee in 2007.<ref name="NPR">{{cite web |last=Burnham |first=Ted |date=February 29, 2012 |title=Truffles Take Root In Appalachian Soil |url=https://www.npr.org/sections/thesalt/2012/02/28/147586050/truffles-take-root-in-appalachian-soil |url-status=live |archive-url=https://web.archive.org/web/20180819051331/https://www.npr.org/sections/thesalt/2012/02/28/147586050/truffles-take-root-in-appalachian-soil |archive-date=August 19, 2018 |access-date=August 18, 2018 |publisher=[[NPR]]}}</ref> At its peak in the 2008–2009 season, one farm produced about 200 pounds of truffles, but [[Anisogramma anomala|Eastern filbert blight]] almost entirely wiped out the hazel trees by 2013 and production dropped, essentially ending the business.<ref name="Greeneville Sun">{{cite web |last=Jones |first=John M. Jr. |date=Mar 19, 2016 |title=Truffles Have Taken Tom Michaels On A Wild Ride |url=https://www.greenevillesun.com/news/local_news/truffles-have-taken-tom-michaels-on-a-wild-ride/article_df026ce0-29ce-11e6-a4ba-077074996893.html |url-status=live |archive-url=https://web.archive.org/web/20200809213511/https://www.greenevillesun.com/news/local_news/truffles-have-taken-tom-michaels-on-a-wild-ride/article_df026ce0-29ce-11e6-a4ba-077074996893.html |archive-date=August 9, 2020 |access-date=August 18, 2018 |publisher=[[The Greeneville Sun]]}}</ref> Eastern filbert blight similarly destroyed the orchards of other once-promising commercial farms in East Tennessee, while newer farms in California, North Carolina, Oregon, and Arkansas were started.<ref name="jacobsen">{{cite news |last1=Jacobsen |first1=Rowen |date=June 2021 |title=Has the American-Grown Truffle Finally Broken Through? |work=[[Smithsonian (magazine)|Smithsonian]] |url=https://www.smithsonianmag.com/innovation/american-grown-truffle-breakthrough-180977702/ |access-date=2 June 2021}}</ref><ref>{{cite news |last1=Krader |first1=Kate |date=25 October 2016 |title=How to Start Your Own Truffle Farm |newspaper=Bloomberg.com |publisher=Bloomberg |url=https://www.bloomberg.com/news/articles/2016-10-25/how-to-start-your-own-truffle-farm |url-status=live |access-date=10 April 2019 |archive-url=https://web.archive.org/web/20190410060139/https://www.bloomberg.com/news/articles/2016-10-25/how-to-start-your-own-truffle-farm |archive-date=2019-04-10}}</ref><ref name="Eater">{{cite web |last=Bamman |first=Mattie John |date=Jun 8, 2017 |title=Why Haven't American Truffles Taken Root Yet? |url=https://www.eater.com/2017/6/8/15757816/truffles-oregon-american-grown |url-status=live |archive-url=https://web.archive.org/web/20180819082750/https://www.eater.com/2017/6/8/15757816/truffles-oregon-american-grown |archive-date=August 19, 2018 |access-date=August 18, 2018 |publisher=[[Eater (website)|Eater]]}}</ref> {{As of|2022}}, the [[Appalachian Mountains|Appalachian]] truffle (''[[Tuber canaliculatum]]'') was being developed as a potential market.<ref>{{Cite magazine |last=Jacobson |first=Rowan |date=January 19, 2022 |title=America's Next Food Craze Is Buried in Appalachia |url=https://www.outsideonline.com/outdoor-adventure/environment/appalachian-truffle-hunters/ |website=Outside |language=en-US}}</ref>

== Uses ==
{{more citations needed section|date=July 2021}}
[[File:Shaved black truffles with pasta - Spago Las Vegas - March 2022 - Sarah Stierch.jpg|thumb|Shaved Périgord truffle with pasta at [[Spago]] in Las Vegas, Nevada.]]

Because of their high price<ref>{{Cite news|url=http://www.cbsnews.com/news/truffles-the-most-expensive-food-in-the-world/|title=Truffles: The Most Expensive Food in the World|access-date=2017-01-06|archive-date=2017-01-07|archive-url=https://web.archive.org/web/20170107100452/http://www.cbsnews.com/news/truffles-the-most-expensive-food-in-the-world/|url-status=live}}</ref> and their strong aroma, truffles are used sparingly. Supplies can be found commercially as unadulterated fresh produce or preserved, typically in a light [[brine]].

Their chemical compounds infuse well with fats such as butter, cream, cheeses, avocados, and coconut cream.

As the volatile aromas dissipate quicker when heated, truffles are generally served raw and shaved over warm, simple foods which highlight their flavour, such as buttered [[pasta]] or [[egg]]s. Thin truffle slices may be inserted into [[meat]]s, under the skins of roasted [[fowl]], in ''[[foie gras]]'' preparations, in ''[[pâté]]s'', or in [[stuffing]]s. Some speciality [[cheese]]s contain truffles, as well. Truffles are also used for producing truffle salt and truffle honey.

While chefs once peeled truffles, in modern times, most restaurants brush the truffle carefully and shave it or dice it with the skin on to make the most of the valuable ingredient. Some restaurants stamp out circular discs of truffle flesh and use the skins for sauces.
[[File:Truffle olive oil with Tuber melanosporum.JPG|thumb|upright=0.6|Truffle oil (olive oil with ''Tuber melanosporum'')]]

=== Oil ===
{{Main|Truffle oil}}

Truffle oil is used as a lower-cost and convenient substitute for truffles, to provide flavouring, or to enhance the flavour and aroma of truffles in cooking. Some products called "truffle oils" contain no truffles or include pieces of inexpensive, unprized [[List of Tuber (fungus) species|truffle varietals]], which have no culinary value, simply for show.<ref name="Patterson2007">{{cite news |url=https://www.nytimes.com/2007/05/16/dining/16truf.html |title=Hocus-Pocus, and a Beaker of Truffles |access-date=2008-05-17 |author=Daniel Patterson |date=16 May 2007 |newspaper=New York Times |quote=Most commercial truffle oils are concocted by mixing olive oil with one or more compounds like 2,4-dithiapentane |archive-date=2011-09-25 |archive-url=https://web.archive.org/web/20110925073741/http://www.nytimes.com/2007/05/16/dining/16truf.html |url-status=live }}</ref> The vast majority is oil that has been artificially flavoured using a synthetic agent such as [[2,4-dithiapentane]].<ref name="Patterson2007" />

The scientific name is included on the ingredient list of truffle oils infused with natural truffles.

=== Vodka ===
Because more aromatic molecules in truffles are soluble in alcohol, they can carry a more complex and accurate truffle flavour than oil without synthetic flavourings. Many commercial producers use 2,4-dithiapentane regardless, as it has become the dominant flavour most consumers, unexposed to fresh truffles but familiar with oils, associate with them. Because most Western nations do not have ingredient labelling requirements for spirits, consumers often do not know if artificial flavourings have been used.<ref>{{cite web |url=http://www.iard.org/resources/beverage-alcohol-labeling-requirements/ |title=Beverage Alcohol Labeling Requirements |publisher=International Alliance for Responsible Drinking |access-date=2018-12-04 |archive-date=2018-12-05 |archive-url=https://web.archive.org/web/20181205003335/http://www.iard.org/resources/beverage-alcohol-labeling-requirements/ |url-status=live }}</ref> It is used as a spirit in its own right, a [[cocktail]] mix or a food flavouring.<ref>{{cite web |url=http://www.mycorrhizalsystems.com/vodka-natural-way-taste-truffles |title=Truffle vodka article |publisher=Mycorrhizalsystems.com |date=2010-04-21 |access-date=2012-06-16 |archive-date=2013-11-04 |archive-url=https://web.archive.org/web/20131104214956/http://www.mycorrhizalsystems.com/vodka-natural-way-taste-truffles |url-status=live }}</ref>

== Cultural history ==

=== Antiquity ===
The first mention of truffles appears in the inscriptions of the [[Neo-Sumerian|neo-Sumerians]] regarding their [[Amorite]] enemy's eating habits ([[Third Dynasty of Ur]], 2nd century BCE)<ref>{{citation |last=Chiera |first=E. |title=Sumerian Epics and Myths |year=1934 |chapter=Nos. 58 and 112 |location=Chicago}}</ref> and later in writings of [[Theophrastus]] in the 4th century BCE. In classical times, their origins were a mystery that challenged many; [[Plutarch]] and others thought them to be the result of lightning, warmth, and water in the soil, while [[Juvenal]] thought thunder and rain to be instrumental in their origin. [[Cicero]] deemed them children of the earth, while [[Dioscorides]] thought they were tuberous roots.<ref name="Ramsbottom53" />

[[Ancient Rome|Rome]] and [[Thracia]] in the [[Ancient Rome|Classical]] period identified three kinds of truffles: ''[[Tuber melanosporum]]'', ''[[Tuber magnificanus|T. magnificus]]'', and ''[[Tuber magnatum|T. magnatum]]''. The Romans instead used a variety of fungus called [[Terfeziaceae|terfez]], also sometimes called a "desert truffle". Terfez used in Rome came from [[Lesbos]], [[Carthage]], and especially [[Libya]], where the coastal climate was less dry in ancient times.<ref name="Ramsbottom53" /> Their substance is pale, tinged with rose. Unlike truffles, terfez have little inherent flavour. The Romans used the terfez as a flavour carrier because the terfez tends to absorb surrounding flavours. Because [[Ancient Roman cuisine]] used many spices and flavourings, the terfez may have been appropriate in that context.

=== Middle Ages ===
Truffles were rarely used during the [[Middle Ages]]. Truffle hunting is mentioned by [[Bartolomeo Platina]], the papal historian, in 1481, when he recorded that the sows of [[Notza]] were without equal in hunting truffles, but they should be muzzled to prevent them from eating the prize.<ref name="benjamin95">{{Citation |last=Benjamin |first=D. R. |title=Mushrooms: Poisons and Panaceas — A Handbook for Naturalists, Mycologists and Physicians |pages=48–50 |year=1995 |chapter=Historical uses of truffles |location=New York |publisher=WH Freeman and Company |isbn=978-0716726005}}</ref>

=== Renaissance and modernity ===
During the [[Renaissance]], truffles regained popularity in Europe and were honoured at the court of King [[Francis I of France]]. They were popular in Parisian markets in the 1780s, imported seasonally from truffle grounds, where peasants had long enjoyed them. [[Jean Anthelme Brillat-Savarin|Brillat-Savarin]] (1825) noted that they were so expensive they appeared only at the dinner tables of great nobles and kept women. They were sometimes served with turkey.

== See also ==
{{Portal|Food|Fungi}}
* [[Chocolate truffle]] – a [[confection]] named for its similar appearance to a truffle
* [[List of Tuber (fungus) species|List of ''Tuber'' species]]
* [[Specialty foods]]

==References==
{{Reflist|30em}}

==Additional resources==
{{Commons|truffle}}
* {{cite book|last1= Morcillo |first1= M. |last2=Sanchez |first2=M. |last3=Vilanova |first3=X. |date=2015|title= Truffle Farming Today, a Comprehensive World Guide |publisher= Micología Forestal & Aplicada |isbn= 978-84-617-1307-3 |url=http://www.micofora.com}}
* {{cite book
 | title = Field Guide to North American Truffles: Hunting, Identifying, and Enjoying the World's Most Prized Fungi
 | series = Natural History Series
 | first1 = Matt
 | last1 = Trappe
 | first2 = Frank
 | author3-link = James Trappe
 | last2 = Evans
 | first3 = James M.
 | last3 = Trappe
 | publisher = Ten Speed Press
 | year = 2007
 | isbn = 9781580088626
}}
* {{cite book
 | title = Truffle: A Global History
 | series = The Edible Series
 | first1 = Zachary
 | last1 = Nowak
 | publisher = Reaktion
 | year = 2015
 | isbn = 978-1780234366
}}
* {{cite web|url=https://www.youtube.com/watch?v=RQ1nY51txoA |archive-url=https://ghostarchive.org/varchive/youtube/20211212/RQ1nY51txoA| archive-date=2021-12-12 |url-status=live|title=Why Real Truffles Are So Expensive|website=[[Business Insider]]|date=2019-10-26}}{{cbignore}}

{{Authority control}}

{{DEFAULTSORT:Truffle (Fungus)}}
[[Category:Truffles (fungi)| ]]
[[Category:Edible fungi]]
[[Category:French cuisine]]
[[Category:Fungi in cultivation]]
[[Category:Italian cuisine]]
[[Category:Mushroom types]]
[[Category:Fungus common names]]