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{{Short description|Species of mushroom, widely distributed in the Northern Hemisphere}}
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{{Redirect|Cep}}
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{{Featured article}}
{{Use British English|date=October 2015}}
{{Use British English|date=October 2015}}
{{stack begin}}
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{{Speciesbox
{{taxobox
| image = Boletus edulis EtgHollande 041031 091.jpg
| image = Boletus edulis IT.jpg
| image_caption = In a forest near [[Rambouillet]], France
| image_caption = In the northern [[Apennine Mountains]], Abetina Reale forest, Italy
| image_width = 234px
| image_alt = Two mushrooms with brown caps and light brown stems growing on the ground, surrounded by fallen leaves and other forest debris. One mushroom has been plucked and lies beside the other; its under-surface is visible, and is a light yellow colour.
| image_alt = Two mushrooms with brown caps and light brown stems growing on the ground, surrounded by fallen leaves and other forest debris. One mushroom has been plucked and lies beside the other; its under-surface is visible, and is a light yellow colour.
| regnum = [[Fungi]]
|status = LC
|status_system = IUCN3.1
| divisio = [[Basidiomycota]]
|status_ref = <ref name=iucn>{{cite iucn |author=Dahlberg, A. |year=2019 |errata=2022 |title=''Boletus edulis'' |volume=2019 |page=e.T122090234A222968388 |doi=10.2305/IUCN.UK.2019-3.RLTS.T122090234A222968388.en |access-date=2 July 2024}}</ref>
| classis = [[Agaricomycetes]]
| ordo = [[Boletales]]
| genus = Boletus
| familia = [[Boletaceae]]
| species = edulis
| authority = [[Jean Baptiste François Pierre Bulliard|Bull.]] (1782)
| genus = ''[[Boletus]]''
| synonyms_ref= <ref name="urlMycoBank: Boletus edulis">{{cite web |url=http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=356530 |title=''Boletus edulis'' Bull. 1782 |publisher=International Mycological Association |work=MycoBank |access-date=2010-10-21}}</ref>
| species = '''''B. edulis'''''
| binomial = ''Boletus edulis''
| binomial_authority = [[Jean Baptiste François Pierre Bulliard|Bull.]] (1782)
| synonyms_ref= <ref name="urlMycoBank: Boletus edulis">{{cite web |url=http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=356530 |title=''Boletus edulis'' Bull. 1782 |publisher=International Mycological Association |work=MycoBank |accessdate=2010-10-21}}</ref>
| synonyms =
| synonyms =
*''Ceriomyces crassus'' <small>[[Giovanni Antonio Battarra|Battarra]] (1775)</small>
*''Ceriomyces crassus'' <small>[[Giovanni Antonio Battarra|Battarra]] (1775)</small>
*''Boletus solidus'' <small>[[James Sowerby|Sowerby]] (1809)</small>
*''Boletus solidus'' <small>[[James Sowerby|Sowerby]] (1809)</small>
*''Leccinum edule'' <small>(Bull.) [[Samuel Frederick Gray|Gray]] (1821)</small>
*''Leccinum edule'' <small>(Bull.) [[Samuel Frederick Gray|Gray]] (1821)</small>
*''Dictyopus edulis'' <small>(Bull.) [[Lucien Forquignon|Forq.]] (1890)</small>
*''Dictyopus edulis'' <small>(Bull.) Forq. (1890)</small>
}}
}}
{{Mycomorphbox
{{mycomorphbox
| name = ''Boletus edulis''
| name = ''Boletus edulis''
| whichGills = adnate
| whichGills = adnate
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}}
}}
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'''''Boletus edulis''''' (English: '''cep''', '''penny bun''', '''porcino''' or '''porcini''') is a [[basidiomycete]] fungus, and the [[type species]] of the genus ''[[Boletus]]''.
'''''Boletus edulis''''' (English: '''cep''', '''porcino''' or '''porcini''') is a [[basidiomycete]] fungus, and the [[type species]] of the genus ''[[Boletus]]''. Widely distributed in the [[Northern Hemisphere]] across Europe, Asia, and North America, it does not occur naturally in the Southern Hemisphere, although it has been [[introduced species|introduced]] to southern Africa, Australia, and New Zealand. Several closely related European mushrooms formerly thought to be [[variety (botany)|varieties]] or [[form (botany)|forms]] of ''B.&nbsp;edulis'' have been shown using [[molecular phylogenetic]] analysis to be distinct species, and others previously classed as separate species are [[wikt:conspecific|conspecific]] with this species. The western North American species commonly known as the California king bolete (''Boletus edulis'' var. ''grandedulis'') is a large, darker-coloured variant first formally identified in 2007.


Prized as an ingredient in various culinary dishes, ''B.&nbsp;edulis'' is an [[edible mushroom]] held in high regard in many cuisines, and is commonly prepared and eaten in [[soups]], pasta, or [[risotto]]. The mushroom is low in fat and digestible [[carbohydrate]]s, and high in [[protein (nutrient)|protein]], [[vitamin]]s, [[dietary mineral|mineral]]s and [[dietary fibre]]. Although it is sold commercially, it is very difficult to [[fungiculture|cultivate]]. Available fresh in autumn throughout Europe and Russia, it is most often dried, packaged, and distributed worldwide. It keeps its flavour after drying, and it is then reconstituted and used in cooking. ''B.&nbsp;edulis'' is also one of the few fungi sold [[pickled]].
The fungus grows in [[deciduous forest|deciduous]] and [[coniferous forest]]s and tree [[plantations]], forming [[symbiotic]] [[mycorrhiza|ectomycorrhizal]] associations with living trees by enveloping the tree's underground roots with sheaths of fungal tissue. The fungus produces [[spore]]-bearing [[basidiocarp|fruit bodies]] above ground in summer and autumn. The fruit body has a large brown [[pileus (mycology)|cap]] which on occasion can reach {{convert|35|cm|in|0|abbr=on}} in diameter and {{convert|3|kg|lb|1|abbr=on}} in weight. Like other [[bolete]]s, it has tubes extending downward from the underside of the cap, rather than gills; spores escape at maturity through the tube openings, or pores. The pore surface of the ''B.&nbsp;edulis'' fruit body is whitish when young, but ages to a greenish-yellow. The stout [[Stipe (mycology)|stipe]], or stem, is white or yellowish in colour, up to {{convert|25|cm|in|0|abbr=on}} tall and {{convert|10|cm|in|0|abbr=on}} thick, and partially covered with a raised network pattern, or [[wikt:reticulate|reticulations]].


The fungus grows in [[deciduous forest|deciduous]] and [[coniferous forest]]s and tree [[plantation]]s, forming [[symbiotic]] [[Mycorrhiza#Ectomycorrhiza|ectomycorrhizal]] associations with living trees by enveloping the tree's underground roots with sheaths of fungal tissue. The fungus produces [[Basidiospore|spore]]-bearing [[basidiocarp|fruit bodies]] above ground in summer and autumn. The fruit body has a large brown [[pileus (mycology)|cap]] which on occasion can reach {{convert|30|cm|in|0|abbr=on}}, rarely {{convert|40|cm|in|0|abbr=on}} in diameter and {{convert|3|kg|lboz|abbr=on}} in weight. Like other [[bolete]]s, it has tubes extending downward from the underside of the cap, rather than gills; spores escape at maturity through the tube openings, or pores. The pore surface of the ''B.&nbsp;edulis'' fruit body is whitish when young, but ages to a greenish-yellow. The stout [[Stipe (mycology)|stipe]], or stem, is white or yellowish in colour, up to {{convert|20|cm|in|0|abbr=on}}, rarely {{convert|30|cm|in|0|abbr=on}} tall and {{convert|10|cm|in|0|abbr=on}} thick, and partially covered with a raised network pattern, or [[wikt:reticulate|reticulations]].
Prized as an ingredient in various foods, ''B.&nbsp;edulis'' is an [[edible mushroom]] held in high regard in many cuisines, and is commonly prepared and eaten in soups, pasta, or [[risotto]]. The mushroom is low in [[fat]] and digestible [[carbohydrate]]s, and high in [[protein (nutrient)|protein]], [[vitamin]]s, [[dietary mineral|mineral]]s and [[dietary fibre]]. Although it is sold commercially, it is very difficult to cultivate. Available fresh in autumn in [[Central Europe|Central]], [[Southern Europe|Southern]] and [[Northern Europe]], it is most often dried, packaged and distributed worldwide. Keeping its flavour after drying, it is then reconstituted and used in cooking. ''B.&nbsp;edulis'' is one of the few fungi sold pickled. The fungus also produces a variety of [[organic compound]]s with a diverse spectrum of [[biological activity]], including the steroid derivative [[ergosterol]], a [[lectin|sugar binding protein]], [[antiviral drug|antiviral]] compounds, [[antioxidant]]s, and [[phytochelatin]]s, which give the organism resistance to toxic [[heavy metal (chemistry)|heavy metals]].

Widely distributed in the [[Northern Hemisphere]] across Europe, Asia, and North America, it does not occur naturally in the Southern Hemisphere, although it has been [[introduced species|introduced]] to southern Africa, Australia, New Zealand, and Brazil. Several closely related European mushrooms formerly thought to be [[variety (botany)|varieties]] or [[form (botany)|forms]] of ''B.&nbsp;edulis'' have been shown using [[molecular phylogenetic]] analysis to be distinct species, and others previously classed as separate species are [[wikt:conspecific|conspecific]] with this species. The western North American species commonly known as the California king bolete (''Boletus edulis'' var. ''grandedulis'') is a large, darker-coloured variant first formally identified in 2007.


==Taxonomy==
==Taxonomy==
[[File:Bulliard00.jpg|thumb|right|Pierre Bulliard first described ''B.&nbsp;edulis'' in 1782.|alt=Side view of the head and upper body of a man wearing a dark jacket and white ruffled collar.]]
[[File:Bulliard00.jpg|thumb|right|Pierre Bulliard first described ''B.&nbsp;edulis'' in 1782.|alt=Side view of the head and upper body of a man wearing a dark jacket and white ruffled collar.]]
''Boletus edulis'' was first [[species description|described]] in 1782 by the French botanist [[Jean Baptiste François Pierre Bulliard|Pierre Bulliard]] and still bears its original name.<ref>{{cite book |title=Herbier de la France. Vol 2|author=Bulliard JBF.|year=1782|publisher=P.F. Didot|location=Paris, France|pages=49–96, plate 60|url=https://archive.org/stream/herbierdelafranc4996bull#page/22/mode/2up|accessdate=2009-11-24|language=French}}</ref> The starting date of fungal [[Taxonomy (biology)|taxonomy]] had been set as January&nbsp;1, 1821, to coincide with the date of the works of the 'father of mycology', Swedish naturalist [[Elias Magnus Fries]], which meant the name required sanction by Fries (indicated in the name by a colon) to be considered valid, as Bulliard's work preceded this date. It was thus written ''Boletus edulis'' Bull.:Fr. A 1987 revision of the [[International Code of Botanical Nomenclature]] set the starting date at May&nbsp;1, 1753, the date of publication of [[Carl Linnaeus|Linnaeus]]' work, the ''[[Species Plantarum]]''.<ref>{{cite book |author= Esser K, Lemke PA. |title= The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research |publisher = Springer|location=Heidelberg, Germany |year=1994|page=81|isbn = 3-540-66493-9}}</ref> Hence, the name no longer requires the ratification of Fries' authority. Early [[synonym (taxonomy)|alternate names]] include ''Boletus solidus'' by English naturalist [[James Sowerby]] in 1809,<ref>{{cite book |title=Coloured Figures of English Fungi |volume=Volume 4 |author=Sowerby J.|year=1809|publisher=J. Davis |location=London |page=199}} This entire work is [[commons:Coloured Figures of English Fungi or Mushrooms|available in Commons here]], but the reference is to plate 419 with textual description on page 697. Sowerby described the same modern species as ''B. edulis'' on plate 111 (description on page 57).</ref> and [[Samuel Frederick Gray|Gray's]] ''Leccinum edule''.<ref>{{cite book |title=A Natural Arrangement of British Plants |author=Gray SF. |year=1821 |publisher=Baldwin, Cradock, and Joy, Paternoster-Row |location=London |page=647 |url=https://books.google.com/?id=g-EYAAAAYAAJ&dq=A%20natural%20arrangement%20of%20British%20plants%20Gray%201821&pg=PA647#v=onepage&q=|accessdate=2009-11-24}}</ref> Gray's transfer of the species to ''Leccinum'' was later determined to be inconsistent with the rules of botanical [[biological classification|nomenclature]], and he apparently was unfamiliar with the earlier works of Fries when he published his arrangement of bolete species.<ref>{{cite journal|author= Šutara J. |year=1985 |title=''Leccinum'' and the question of superfluous names (Fungi: Boletaceae)|journal=Taxon |volume=34 |issue=4 |pages=678–86|doi= 10.2307/1222214|jstor= 1222214}}</ref>
''Boletus edulis'' was first [[species description|described]] in 1782 by the French botanist [[Jean Baptiste François Pierre Bulliard|Pierre Bulliard]] and still bears its original name.<ref>{{cite book |title=Herbier de la France. Vol 2|author=Bulliard JBF.|year=1782|publisher=P.F. Didot|location=Paris, France|pages=49–96, plate 60|url=https://archive.org/stream/herbierdelafranc4996bull#page/22/mode/2up|access-date=2009-11-24|language=fr}}</ref> The starting date of fungal [[Taxonomy (biology)|taxonomy]] had been set as January&nbsp;1, 1821, to coincide with the date of the works of the 'father of mycology', Swedish naturalist [[Elias Magnus Fries]], which meant the name required sanction by Fries (indicated in the name by a colon) to be considered valid, as Bulliard's work preceded this date. It was thus written ''Boletus edulis'' Bull.:Fr. A 1987 revision of the [[International Code of Botanical Nomenclature]] set the starting date at May&nbsp;1, 1753, the date of publication of [[Carl Linnaeus|Linnaeus]]' work, the ''[[Species Plantarum]]''.<ref>{{cite book |vauthors=Esser K, Lemke PA |title= The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research |url=https://archive.org/details/mossflorabritain00libg |url-access=limited |publisher = Springer|location=Heidelberg, Germany |year=1994|page=[https://archive.org/details/mossflorabritain00libg/page/n93 81]|isbn = 3-540-66493-9}}</ref> Hence, the name no longer requires the ratification of Fries' authority. Early [[synonym (taxonomy)|alternate names]] include ''Boletus solidus'' by English naturalist [[James Sowerby]] in 1809,<ref>{{cite book |title=Coloured Figures of English Fungi |volume=4 |author=Sowerby J.|year=1809|publisher=J. Davis |location=London |page=199}} This entire work is [[commons:Coloured Figures of English Fungi or Mushrooms|available in Commons here]], but the reference is to plate 419 with textual description on page 697. Sowerby described the same modern species as ''B. edulis'' on plate 111 (description on page 57).</ref> and [[Samuel Frederick Gray|Gray's]] ''Leccinum edule''.<ref>{{cite book |title=A Natural Arrangement of British Plants |author=Gray SF. |year=1821 |publisher=Baldwin, Cradock, and Joy, Paternoster-Row |location=London |page=647 |url=https://books.google.com/books?id=g-EYAAAAYAAJ&q=A%20natural%20arrangement%20of%20British%20plants%20Gray%201821&pg=PA647|access-date=2009-11-24}}</ref> Gray's transfer of the species to ''Leccinum'' was later determined to be inconsistent with the rules of botanical [[biological classification|nomenclature]], and he apparently was unfamiliar with the earlier works of Fries when he published his arrangement of bolete species.<ref>{{cite journal|author= Šutara J. |year=1985 |title=''Leccinum'' and the question of superfluous names (Fungi: Boletaceae) |journal=Taxon |volume=34 |issue=4 |pages=678–86|doi= 10.2307/1222214|jstor= 1222214}}</ref>


''B. edulis'' is the [[type species]] of the genus ''[[Boletus]]''. In [[Rolf Singer]]'s [[biological classification|classification]] of the [[Agaricales]] mushrooms, it is also the type species of [[Section (biology)#Ranks in botany|section]] ''Boletus'', a grouping of about 30 related boletes united by several characteristics: a mild-tasting, white [[trama (mycology)|flesh]] that does not change colour when exposed to air; a smooth to distinctly raised, netted pattern over at least the uppermost portion of the stem; a yellow-brown or olive-brown [[spore print]]; white tubes that later become yellowish then greenish, which initially appear to be stuffed with cotton; and [[cystidia]] that are not strongly coloured.<ref name=Singer1986>{{cite book |author=Singer R. |title=The Agaricales in Modern Taxonomy.|edition= 4th rev.|publisher=Koeltz Scientific Books|location=Koenigstein, Germany |year=1986|page=779|isbn=3-87429-254-1}}</ref><ref>{{cite book |author=Smith AH, Thiers HD. |year=1971 |title=The Boletes of Michigan |location=Ann Arbor, Michigan|publisher=University of Michigan Press|page=221|url=http://quod.lib.umich.edu/cgi/t/text/pageviewer-idx?c=fung1tc;cc=fung1tc;idno=agk0838.0001.001;frm=frameset;view=image;seq=229;page=root;size=s|accessdate=2010-12-02}}</ref> [[Molecular phylogenetics|Molecular]] analysis published in 1997 established that the bolete mushrooms are all [[monophyly|derived from a common ancestor]], and established the [[Boletales]] as an [[order (biology)|order]] separate from the Agaricales.<ref>{{cite journal |author=Binder M, Besl H, Bresinksy A. |year=1997 |title=Agaricales oder Boletales? Molekularbiologische Befunde zur Zuordnuhn einiger umstrittener Taxa. |journal=Z Mykol |volume=63 |issue= |pages=189–196 |doi= |pmid= }}</ref>
''Boletus edulis'' is the [[type species]] of the genus ''[[Boletus]]''. In [[Rolf Singer]]'s [[biological classification|classification]] of the [[Agaricales]] mushrooms, it is also the type species of [[Section (botany)#Ranks in botany|section]] ''Boletus'', a grouping of about 30 related boletes united by several characteristics: a mild-tasting, white [[trama (mycology)|flesh]] that does not change colour when exposed to air; a smooth to distinctly raised, netted pattern over at least the uppermost portion of the stem; a yellow-brown or olive-brown [[spore print]]; white tubes that later become yellowish then greenish, which initially appear to be stuffed with cotton; and [[cystidia]] that are not strongly coloured.<ref name=Singer1986>{{cite book |author=Singer R. |title=The Agaricales in Modern Taxonomy |edition= 4th rev.|publisher=Koeltz Scientific Books|location=Koenigstein, Germany |year=1986|page=779|isbn=3-87429-254-1}}</ref><ref>{{cite book |vauthors=Smith AH, Thiers HD |year=1971 |title=The Boletes of Michigan |location=Ann Arbor, Michigan|publisher=University of Michigan Press|page=221|url=http://quod.lib.umich.edu/cgi/t/text/pageviewer-idx?c=fung1tc;cc=fung1tc;idno=agk0838.0001.001;frm=frameset;view=image;seq=229;page=root;size=s|access-date=2010-12-02}}</ref> [[Molecular phylogenetics|Molecular]] analysis published in 1997 established that the bolete mushrooms are all [[monophyly|derived from a common ancestor]], and established the [[Boletales]] as an [[order (biology)|order]] separate from the Agaricales.<ref>{{cite journal |vauthors=Binder M, Besl H, Bresinsky A |year=1997 |title=Agaricales oder Boletales? Molekularbiologische Befunde zur Zuordnung einiger umstrittener Taxa |journal=Zeitschrift für Mykologie |volume=63 |pages=189–196 }}</ref>


The [[genus|generic]] name is derived from the [[Latin]] term ''bōlētus'' "mushroom", which was borrowed in turn from the [[Ancient Greek]] βωλίτης, "terrestrial fungus".<ref>{{cite book |author= Simpson DP. |title=Cassell's Latin Dictionary |publisher = Cassell Ltd |year = 1979 |origyear=1854 |edition = 5 |location = London |page=78 |isbn = 0-304-52257-0}}</ref> Ultimately, this last word derives from ''bōlos''/βῶλος "lump", "clod", and, metaphorically, "mushroom".<ref name="Liddell 1980">{{cite book |author = [[Henry George Liddell|Liddell HG]], [[Robert Scott (philologist)|Scott R.]] |year= 1980 |title = [[A Greek-English Lexicon]]|edition= Abridged |publisher = [[Oxford University Press]] | location= United Kingdom |isbn = 0-19-910207-4}}</ref> The βωλίτης of [[Galen]], like the ''boletus'' of Latin writers like [[Martial]], [[Seneca the Younger|Seneca]] and [[Petronius]],<ref>Peter Howell, ''A Commentary on Book One of the Epigrams of Martial,'' The Athlone Press, 1980 p.152-3. Howell doubts the identification, and mentions the view advanced by Augusta A. Imholtz Jr., 'Fungi and piace- names, thè origin of boletus,' in [[American Journal of Philology|AJP]] Vol.98, 1977 pp.71f., that the Latin word may derive from the Spanish town Boletum, modern-day [[Boltaña]], south of the [[Pyrenees]], which is still famous for its mushrooms.</ref> is often identified as the much prized ''[[Amanita caesarea]]''.<ref name = "Ramsbottom53">{{cite book |author=Ramsbottom J. |year=1953 |title=Mushrooms & Toadstools |page=6 |publisher=Collins |location=London, England|isbn= 1-870630-09-2}}</ref> The [[specific name (botany)|specific epithet]] ''edulis'' in Latin means "eatable" or "edible".<ref>{{cite book|author = Jamieson A, Ainsworth R, Morell T.| title = Latin Dictionary: Morell's Abridgment | publisher = Moon, Boys & Graves| year = 1828|edition = |location = London|pages = 121, 596| url=https://books.google.com/?id=U3oTAAAAYAAJ&printsec=frontcover&dq=latin+dictionary|accessdate=2009-11-02}}</ref>
The [[genus|generic]] name is derived from the [[Latin]] term ''bōlētus'' "mushroom", which was borrowed in turn from the [[Ancient Greek]] βωλίτης, "terrestrial fungus".<ref>{{cite book |author= Simpson DP. |title=Cassell's Latin Dictionary |publisher = Cassell Ltd |year = 1979 |orig-date=1854 |edition = 5 |location = London |page=78 |isbn = 0-304-52257-0}}</ref> Ultimately, this last word derives from ''bōlos''/βῶλος "lump", "clod", and, metaphorically, "mushroom".<ref name="Liddell 1980">{{cite book |author = [[Henry Liddell|Liddell HG]], [[Robert Scott (philologist)|Scott R.]] |year= 1980 |title = A Greek–English Lexicon|edition= Abridged |publisher = [[Oxford University Press]] | location= United Kingdom |isbn = 0-19-910207-4|title-link= A Greek–English Lexicon }}</ref> The βωλίτης of [[Galen]], like the ''boletus'' of Latin writers like [[Martial]], [[Seneca the Younger|Seneca]] and [[Petronius]],<ref>Peter Howell, ''A Commentary on Book One of the Epigrams of Martial,'' The Athlone Press, 1980 p.152-3. Howell doubts the identification, and mentions the view advanced by Augusta A. Imholtz Jr., 'Fungi and piace- names, thè origin of boletus,' in [[American Journal of Philology|AJP]] Vol.98, 1977 pp.71f., that the Latin word may derive from the Spanish town Boletum, modern-day [[Boltaña]], south of the [[Pyrenees]], which is still famous for its mushrooms.</ref> is often identified as the much prized ''[[Amanita caesarea]]''.<ref name = "Ramsbottom53">{{cite book |author=Ramsbottom J. |year=1953 |title=Mushrooms & Toadstools |page=6 |publisher=Collins |location=London, England|isbn= 1-870630-09-2}}</ref> The [[specific name (botany)|specific epithet]] ''edulis'' in Latin means "eatable" or "edible".<ref>{{cite book|vauthors=Jamieson A, Ainsworth R, Morell T | title = Latin Dictionary: Morell's Abridgment | publisher = Moon, Boys & Graves| year = 1828|location = London|pages = [https://archive.org/details/bub_gb_U3oTAAAAYAAJ/page/n550 121], 596| url=https://archive.org/details/bub_gb_U3oTAAAAYAAJ|access-date=2009-11-02}}</ref>


===Common names===
===Common names===
[[Common name]]s for ''B. edulis'' vary by region. The standard Italian name, ''porcino'' (pl. ''porcini''), means ''[[wikt:porcine|porcine]];''<ref>[http://www.treccani.it/vocabolario/porcino/ Treccani dictionary, "porcino"]</ref> ''fungo porcino'', in [[Italian language|Italian]], echoes the term ''suilli'', literally "hog mushrooms," a term used by the Ancient Romans<ref>[[Pliny the Elder|Pliny]], ''Natural History'', Bk.16, 11, 31: 'Such is the multiplicity of products in addition to the acorn that are borne by hard-oaks; but they also produce edible fungi (''boletos'') and hog mushrooms (''suillos'').' Pliny, ''Natural History'', 10 vols., tr.H. Rackham, Harvard University Press/Heinemann, (1945) 1968, vol.4, pp. 408–409.</ref> and still in use in southern Italian terms for this species.<ref>[[Neapolitan language|Neapolitan]] ''sillo'', and [[Calabrian dialects|Calabrian]] ''sillu''/''siddu''. See Glauco Sanga, Gherardo Ortalli, ''Nature knowledge: ethnoscience, cognition, and utility'', Berghahn Books, 2003 p. 78.</ref> The derivation has been ascribed to the resemblance of young fruit bodies to piglets, or to the fondness pigs have for eating them.<ref name="Carluccio03">Carluccio, pp. 36–38.</ref> It is also known as "king bolete".<ref name=Zeitl76>{{cite book|author=Zeitlmayr L.|year=1976|title=Wild Mushrooms: An Illustrated Handbook|publisher=Garden City Press|location= Hertfordshire, UK|isbn= 0-584-10324-7|page=96}}</ref> The English ''[[penny bun]]'' refers to its rounded brownish shape. The [[German language|German]] name ''Steinpilz'' (stone mushroom) refers to the species' firm flesh.<ref>{{cite book|author=[[Jacob Grimm|Grimm J.]], [[Wilhelm Grimm|Grimm W.]]|title=[[Deutsches Wörterbuch]]|publisher=Hirzel|location=Leipzig|date=1838–1961}} ([http://www.woerterbuchnetz.de/woerterbuecher/dwb/wbgui?lemid=GS42864 online])</ref> In Austria, it is called ''Herrenpilz'', the "noble mushroom",<ref name = "Carluccio03"/> while in Mexico, the [[Spanish language|Spanish]] name is ''panza'', meaning "belly".<ref>{{cite journal |author=Jarvis MC, Miller AM, Sheahan J, Ploetz K, Ploetz J, Watson RR, Ruiz MP, Villapan CAP, Alvarado JG, Ramirez AL, Orr B. |year=2004 |title=Edible wild mushrooms of the Cofre de Perote region, Veracruz, Mexico: An ethnomycological study of common names and uses |journal=Economic Botany |volume=58 |issue=Suppl. S |pages=S111–S115|doi=10.1663/0013-0001(2004)58[S111:EWMOTC]2.0.CO;2}}</ref> Another Spanish name, ''rodellon'', means "small round boulder", while the [[Dutch language|Dutch]] name ''eekhoorntjesbrood'' means "squirrel's bread".<ref name=Schalkwijk-Barendsen1991>{{cite book |author=Schalkwijk-Barendsen HME. |title=Mushrooms of Western Canada |publisher=Lone Pine Publishing |location=Edmonton |year=1991 |page=195 |isbn=0-919433-47-2}}</ref> Russian names are: "Belyy grib" ("white mushroom" as opposed to less valuable "black mushrooms") and "borovik" (from "bor" - "pine forest").
[[Common name]]s for ''B. edulis'' vary by region. The standard Italian name, ''porcino'' (pl. ''porcini''), means ''[[wikt:porcine|porcine]];''<ref>{{Cite web|url=https://www.treccani.it/vocabolario/porcino|title=porcino in Vocabolario – Treccani|website=www.treccani.it}}</ref> ''fungo porcino'', in Italian, echoes the term ''suilli'', literally "hog mushrooms", a term used by the Ancient Romans<ref>[[Pliny the Elder|Pliny]], ''Natural History'', Bk. 16, 11, 31: "Such is the multiplicity of products in addition to the acorn that are borne by hard-oaks; but they also produce edible fungi (''boletos'') and hog mushrooms (''suillos'')." Pliny, ''Natural History'', 10 vols., tr. H. Rackham, Harvard University Press/Heinemann, (1945) 1968, vol. 4, pp. 408–409.</ref> and still in use in southern Italian terms for this species.<ref>[[Neapolitan language|Neapolitan]] ''sillo'', and [[Calabrian dialects|Calabrian]] ''sillu''/''siddu''. See Glauco Sanga, Gherardo Ortalli, ''Nature knowledge: ethnoscience, cognition, and utility'', Berghahn Books, 2003 p. 78.</ref> The derivation has been ascribed to the resemblance of young fruit bodies to piglets, or to the fondness pigs have for eating them.<ref name="Carluccio03">Carluccio, pp. 36–38.</ref> It is also known as "king bolete".<ref name=Zeitl76>{{cite book|author=Zeitlmayr L.|year=1976|title=Wild Mushrooms: An Illustrated Handbook|publisher=Garden City Press|location= Hertfordshire, UK|isbn= 0-584-10324-7|page=96}}</ref> The English ''[[penny bun]]'' refers to its rounded brownish shape. The German name ''Steinpilz'' (stone mushroom) refers to the species' firm flesh.<ref>{{cite book|author=[[Jacob Grimm|Grimm J.]], [[Wilhelm Grimm|Grimm W.]]|title=Deutsches Wörterbuch|publisher=Hirzel|location=Leipzig|date=1838–1961|title-link=Deutsches Wörterbuch}} ([http://www.woerterbuchnetz.de/woerterbuecher/dwb/wbgui?lemid=GS42864 online] {{Webarchive|url=https://web.archive.org/web/20110719114321/http://www.woerterbuchnetz.de/woerterbuecher/dwb/wbgui?lemid=GS42864 |date=2011-07-19 }})</ref> In Austria, it is called ''Herrenpilz'', the "noble mushroom",<ref name = "Carluccio03"/> while in Mexico, the Spanish name is ''panza'', meaning "belly".<ref>{{cite journal |vauthors=Jarvis MC, Miller AM, Sheahan J, Ploetz K, Ploetz J, Watson RR, Ruiz MP, Villapan CA, Alvarado JG, Ramirez AL, Orr B |year=2004 |title=Edible wild mushrooms of the Cofre de Perote region, Veracruz, Mexico: An ethnomycological study of common names and uses |journal=Economic Botany |volume=58 |issue=Suppl. S |pages=S111–S115|doi=10.1663/0013-0001(2004)58[S111:EWMOTC]2.0.CO;2|s2cid=27729532 }}</ref> Another Spanish name, ''rodellon'', means "small round boulder", while the [[Dutch language|Dutch]] name ''eekhoorntjesbrood'' means "squirrel's bread".<ref name=Schalkwijk-Barendsen1991>{{cite book |author=Schalkwijk-Barendsen H. M. E. |title=Mushrooms of Western Canada |publisher=Lone Pine Publishing |location=Edmonton |year=1991 |page=[https://archive.org/details/mushroomsofweste0000scha/page/195 195] |isbn=0-919433-47-2 |url=https://archive.org/details/mushroomsofweste0000scha/page/195 }}</ref> Russian names are ''belyy grib'' ([[:ru:белый гриб]]; "white mushroom" as opposed to less valuable "black mushrooms") and ''borovik'' ([[:ru:боровик]]; from ''bor''—"pine forest").
The vernacular name ''cep'' is derived from the [[Catalan language|Catalan]] ''cep'' or its [[French language|French]] name ''cèpe'', although the latter is a generic term applying to several related species. In France, it is more fully ''cèpe de Bordeaux'', derived from the [[Gascon language|Gascon]] ''cep'' "trunk" for its fat stalk,<ref>{{cite book |author=Grigson J.|authorlink=|title=The Mushroom Feast |page=8|year=1975 |publisher=Penguin |location=London |isbn=0-14-046273-2}}</ref> ultimately from the [[Latin]] ''cippus'' "stake".<ref>{{Cite encyclopedia| title=cepe|encyclopedia=Oxford English Dictionary|editor=J. Simpson, E. Weiner (eds)| year=1989 |edition= 2nd| location=Oxford |publisher=Clarendon Press|isbn= 0-19-861186-2}}</ref> ''Ceppatello'', ''ceppatello buono'', ''ceppatello bianco'', ''giallo leonato'', ''ghezzo'', and ''moreccio'' are names from Italian dialects,<ref>{{cite book|author=Naccari NL.|title=Flora veneta|publisher=L. Bonvecchiato|year=1827|volume=4–6|page=10}}</ref><ref>{{cite book|author=Zeitlmayr L.|title=I funghi|publisher=Edizioni Studio Tesi|year=1977|page=180}}</ref> and ''ciurenys'' or ''surenys'' is another term in Catalan.<ref>{{cite book|last=Andrews|first=Colman |title=Catalan cuisine: vivid flavors from Spain's Mediterranean coast|publisher=Harvard Common Press|location=Boston, MA|year=1999|origyear=1988|edition=2nd|page=88|isbn=1-55832-329-5}}</ref> The French-born King [[Charles XIV John of Sweden|Charles XIV John]] popularised ''B.&nbsp;edulis'' in Sweden after 1818,<ref>{{cite book|author=Spoerke DG, Rumack BH.|title=Handbook of Mushroom Poisoning: Diagnosis and Treatment|publisher=CRC Press|location=Boca Raton, Florida|year=1994|page=11|isbn=0-8493-0194-7}}</ref> and is honoured in the local vernacular name ''Karljohanssvamp'' as well as the Danish name Karl Johan svamp. The monarch cultivated the fungus about his residence, [[Rosersberg Palace]].<ref>{{cite book|author=Stensaas M, Sonstegard J.|title=Canoe Country Flora: Plants and Trees of the North Woods and Boundary Waters |publisher=University of Minnesota Press|location=Minneapolis, Minnesota|year=2004|page=189|isbn=1-57025-121-5}}The Finnish name is "Herkkutatti", "herkku" from "herkullinen" which means delicious, and "tatti" describing the type of mushroom.</ref> It is known as ''hed tab tao'' {{lang|th|เห็ดตับเต่า}} in [[Thai language|Thai]].<ref name=Solomon/>
The vernacular name ''cep'' is derived from the [[Catalan language|Catalan]] ''cep'' or its French name ''cèpe'', although the latter is a generic term applying to several related species. In France, it is more fully ''cèpe de Bordeaux'', derived from the [[Gascon language|Gascon]] ''cep'' "trunk" for its fat stalk,<ref>{{cite book |author=Grigson J. |title=The Mushroom Feast |page=[https://archive.org/details/mushroomfeast00grig/page/8 8] |year=1975 |publisher=Penguin |location=London |isbn=0-14-046273-2 |url=https://archive.org/details/mushroomfeast00grig/page/8 }}</ref> ultimately from the [[Latin]] ''cippus'' "stake".<ref>{{Cite encyclopedia| title=cepe|encyclopedia=Oxford English Dictionary|editor1=J. Simpson |editor2=E. Weiner | year=1989 |edition= 2nd| location=Oxford |publisher=Clarendon Press|isbn= 0-19-861186-2}}</ref> ''Ceppatello'', ''ceppatello buono'', ''ceppatello bianco'', ''giallo leonato'', ''ghezzo'', and ''moreccio'' are names from Italian dialects,<ref>{{cite book|author=Naccari N. L.|title=Flora veneta|publisher=L. Bonvecchiato|year=1827|volume=4–6|page=10}}</ref><ref>{{cite book|author=Zeitlmayr L.|title=I funghi|publisher=Edizioni Studio Tesi|year=1977|page=180}}</ref> and ''ciurenys'' or ''surenys'' is another term in Catalan.<ref>{{cite book|last=Andrews|first=Colman |title=Catalan cuisine: vivid flavors from Spain's Mediterranean coast|publisher=Harvard Common Press|location=Boston, MA|year=1999|orig-date=1988|edition=2nd|page=88|isbn=1-55832-329-5}}</ref> The French-born King [[Charles XIV John of Sweden|Charles XIV John]] popularised ''B.&nbsp;edulis'' in Sweden after 1818,<ref>{{cite book|vauthors=Spoerke DG, Rumack BH |title=Handbook of Mushroom Poisoning: Diagnosis and Treatment|publisher=CRC Press|location=Boca Raton, Florida|year=1994|page=11|isbn=0-8493-0194-7}}</ref> and is honoured in the local vernacular name ''Karljohanssvamp'', as well as the Danish name ''Karl Johan svamp''. The monarch cultivated the fungus about his residence, [[Rosersberg Palace]].<ref>{{cite book|vauthors=Stensaas M, Sonstegard J |title=Canoe Country Flora: Plants and Trees of the North Woods and Boundary Waters |publisher=University of Minnesota Press|location=Minneapolis, Minnesota|year=2004|page=189|isbn=1-57025-121-5}}</ref> The Finnish name is ''herkkutatti'', from ''herkku'' 'delicacy', and ''tatti'', 'bolete'.{{Cn|date=December 2024}}


==Description==
==Description==
[[File:Boletus edulis herkkutatti halki.jpg|thumb|alt=Both halves of a thick-stemmed bisected mushroom.|Cross-section showing white flesh, broad stem, and spore tubes on the underside of the cap]]
[[File:Boletus edulis herkkutatti halki.jpg|thumb|alt=Both halves of a thick-stemmed bisected mushroom.|Cross-section showing white flesh, broad stem, and spore tubes on the underside of the cap]]
The [[Pileus (mycology)|cap]] of this mushroom is {{convert|7|–|30|cm|in|abbr=on}} broad at maturity. Slightly sticky to touch, it is convex in shape when young and flattens with age. The colour is generally reddish-brown fading to white in areas near the margin, and continues to darken as it matures. The [[Stipe (mycology)|stipe]], or stem, is {{convert|8|–|25|cm|in|abbr=on}} in height, and up to {{convert|7|cm|in|1|abbr=on}} thick—rather large in comparison to the cap;<ref name=Kozikowski1996/> it is club-shaped, or bulges out in the middle. It is finely [[wikt:reticulate|reticulate]] on the upper portion, but smooth or irregularly ridged on the lower part. The under surface of the cap is made of thin tubes, the site of [[spore]] production; they are {{convert|1|to|2|cm|in|1|abbr=on}} deep, and whitish in colour when young, but mature to a greenish-yellow.<ref name=Grund1976/> The angular pores, which do not stain when bruised, are small — roughly 2 to 3 pores per millimetre.<ref name=Tylutki1987II>{{cite book|title=Mushrooms of Idaho and the Pacific Northwest. Vol. 2. Non-gilled Hymenomycetes|author=Tylukti EE. |year=1987 |publisher=The [[University of Idaho Press]] |location=Moscow, Idaho|pages=9–10 |isbn= 0-89301-097-9}}</ref> In youth, the pores are white and appear as if stuffed with cotton (which are actually [[mycelium|mycelia]]); as they age, they change colour to yellow and later to brown. The [[spore print]] is olive brown. The [[trama (mycology)|flesh]] of the fruit body is white, thick and firm when young, but becomes somewhat spongy with age. When bruised or cut, it either does not change colour, or turns a very light brown or light red.<ref name=Wang1995/> Fully mature specimens can weigh about {{convert|1|kg|lb|1|abbr=on}}; a huge specimen collected on the [[Isle of Skye]], Scotland, in 1995 bore a cap of {{convert|42|cm|in|1|abbr=on}}, with a stipe {{convert|18|cm|in|1|abbr=on}} in height and {{convert|14|cm|in|1|abbr=on}} wide, and weighed {{convert|3.2|kg|lb|1|abbr=on}}.<ref name=Kozikowski1996>{{cite journal|author=Kozikowski GR.|year=1996 |title=Foray Report from Skye |journal=Mycologist |volume=10 |pages=183–84 |doi=10.1016/S0269-915X(96)80022-X |issue=4}}</ref> A similarly sized specimen found in Poland in 2013 made international news.<ref>http://www.bbc.co.uk/news/world-europe-24292021</ref>
{{double image|left|Boletus edulis (7).jpg|72|Boletus edulis JPG9.jpg|128|Stem shape can range from club-shaped to centrally bulbous||A brown-capped mushroom lying flat on the grass with a white or light-brown coloured stem that gradually gets thicker, so as to roughly resemble the shape of a club.|A brown-capped mushroom with a short, stout stem that is thickest in the middle, and whose thickness approaches the width of the cap it supports.}}
[[File:Young Boletus Edulis.png|thumb|This is what a young Boletus edulis will look like: a nice fat stem, and a small cap.
Those young ones are generally better to take than old ones due to common infections by maggots and the absorption of harmful gases
]]
''B. edulis'' is considered one of the safest wild mushrooms to pick for the table, as no poisonous species closely resemble it.<ref name="Carluccio03" /> The most similar poisonous mushroom may be the devil's bolete (''[[Rubroboletus satanas]]''), which has a similar shape, but has a red stem and stains blue on bruising.<ref name="Carluccio03" /> It is often confused with the very bitter and unpalatable ''[[Tylopilus felleus]]'', but can be distinguished by the reticulation on the stalk; in porcini, it is a whitish, net-like pattern on a brownish stalk, whereas it is a dark pattern on white in the latter. Porcini have whitish pores while the other has pink. If in doubt, tasting a tiny bit of flesh will yield a bitter taste.<ref name="Carluccio03" /> It can also resemble the "bolete-like" ''[[Gyroporus castaneus]]'', which is generally smaller, and has a browner stem.


The [[Pileus (mycology)|cap]] of this mushroom is {{convert|7|–|30|cm|in|frac=2|abbr=on}} broad at maturity. Slightly sticky to touch, it is convex in shape when young and flattens with age. The colour is generally reddish-brown fading to white in areas near the margin, and continues to darken as it matures. The [[Stipe (mycology)|stipe]], or stem, is {{convert|8|–|25|cm|in|frac=2|abbr=on}} in height, and up to {{convert|7|cm|in|frac=2|abbr=on}} thick—rather large in comparison to the cap;<ref name=Kozikowski1996/> it is club-shaped, or bulges out in the middle. It is finely [[wikt:reticulate|reticulate]] on the upper portion, but smooth or irregularly ridged on the lower part. The under surface of the cap is made of thin tubes, the site of [[Basidiospore|spore]] production; they are {{convert|1|to|2|cm|in|frac=4|abbr=on}} deep, and whitish in colour when young, but mature to a greenish-yellow.<ref name=Grund1976/> The angular pores, which do not stain when bruised, are small—roughly 2 to 3 pores per millimetre.<ref name=Tylutki1987II>{{cite book|title=Mushrooms of Idaho and the Pacific Northwest. Vol. 2. Non-gilled Hymenomycetes|author=Tylukti EE. |year=1987 |publisher=The [[University of Idaho Press]] |location=Moscow, Idaho|pages=9–10 |isbn= 0-89301-097-9}}</ref> In youth, the pores are white and appear as if stuffed with cotton (which are actually [[mycelia]]); as they age, they change colour to yellow and later to brown. The spore print is olive brown. The flesh of the fruit body is white, thick and firm when young, but becomes somewhat spongy with age. When bruised or cut, it either does not change colour, or turns a very light brown or light red.<ref name=Wang1995/> Fully mature specimens can weigh about {{convert|1|kg|lboz|abbr=on}}; a huge specimen collected on the [[Isle of Skye]], Scotland, in 1995 bore a cap of {{convert|42|cm|in|frac=2|abbr=on}}, with a stipe {{convert|18|cm|in|frac=2|abbr=on}} in height and {{convert|14|cm|in|frac=2|abbr=on}} wide, and weighed {{convert|3.2|kg|lboz|abbr=on}}.<ref name=Kozikowski1996>{{cite journal|author=Kozikowski GR.|year=1996 |title=Foray Report from Skye |journal=Mycologist |volume=10 |pages=183–84 |doi=10.1016/S0269-915X(96)80022-X |issue=4}}</ref> A similarly sized specimen found in Poland in 2013 made international news.<ref>{{Cite news | url=https://www.bbc.co.uk/news/world-europe-24292021 | title=Giant mushroom found in Poland| work=BBC News}}</ref>
The [[basidiospore|spores]] are elliptical to spindle-shaped, with dimensions of 12–17 by 5–7&nbsp;[[micrometre|µm]]. The [[basidia]], the spore-bearing cells, are produced in a layer lining the tubes, and arrange themselves so their ends are facing the center of the tube; this layer of cells is known technically as a [[hymenium]]. The basidia are thin-walled, mostly attached to four spores, and measure 25–30 by 8–10&nbsp;µm. Another cell type present in the hymenium is the [[cystidia]], larger sterile cells that protrude beyond the basidia into the [[lumen (anatomy)|lumen]] of the hymenium, and act as air traps, regulating humidity.<ref>{{cite book|author=Alexopoulos CJ, Mims CW, Blackwell M.| title=Introductory Mycology | year=1996 | publisher=John Wiley and Sons |location=New York, New York|page=495 | isbn=0-471-52229-5}}</ref> ''B.&nbsp;edulis'' has pleurocystidia (cystidia located on the face of a pore) that are thin-walled, roughly spindle-shaped to [[ventricose]], and measure 30–45 by 7–10&nbsp;µm; the "stuffed" feature of the hymenium is caused by cheilocystidia — cells found on the edges of the pores.<ref name=Grund1976>{{cite book |title=Nova Scotian Boletes |author=Grund DW, Harrison AK. |year=1976 |publisher=J. Cramer |location=Lehre, Germany |isbn=3-7682-1062-6|pages=73–75}}</ref> The [[hypha]]e of ''B.&nbsp;edulis'' do not have [[clamp connection]]s.<ref name=Tylutki1987II/>
{{multiple image
| align = left
| image1 = Boletus edulis (7).jpg
| width1 = 72
| alt1 = A brown-capped mushroom lying flat on the grass with a white or light-brown coloured stem that gradually gets thicker, so as to roughly resemble the shape of a club.
| caption1 =
| image2 = Boletus edulis JPG9.jpg
| width2 = 128
| alt2 = A brown-capped mushroom with a short, stout stem that is thickest in the middle, and whose thickness approaches the width of the cap it supports.
| caption2 =
| footer = Stem shape can range from club-shaped to centrally bulbous
}}
''Boletus edulis'' is considered one of the safest wild mushrooms to pick for the table, as few poisonous species closely resemble it, and those that do may be easily distinguished by careful examination.<ref name = "Carluccio03"/> The most similar poisonous mushroom may be the devil's bolete (''[[Rubroboletus satanas]]''), which has a similar shape, but has a red stem and stains blue on bruising.<ref name="Carluccio03" /> It is often confused with the very bitter and unpalatable ''[[Tylopilus felleus]]'', but can be distinguished by the reticulation on the stalk; in porcini, it is a whitish, net-like pattern on a brownish stalk, whereas it is a dark pattern on white in the latter. Porcini have whitish pores while the other has pink. If in doubt, tasting a tiny bit of flesh will yield a bitter taste.<ref name = "Carluccio03"/> It can also resemble the "bolete-like" ''[[Gyroporus castaneus]]'', which is generally smaller, and has a browner stem. ''[[Boletus huronensis]],'' an uncommon mushroom of northeastern North America, is another recognized look-alike known to cause severe [[gastrointestinal disorder]]s.<ref>{{cite web|url=https://namyco.org/boletus_huronensis.php|title=''Boletus huronensis'': Comments on its toxicity with diagnostic images of its field characteristics and staining reactions|author=Bill Bakaitis|publisher=North American Mycological Association|date=2019}}</ref>

The [[basidiospore|spores]] are elliptical to spindle-shaped, with dimensions of 12–17 by 5–7&nbsp;[[micrometre|μm]]. The [[basidia]], the spore-bearing cells, are produced in a layer lining the tubes, and arrange themselves so their ends are facing the center of the tube; this layer of cells is known technically as a [[hymenium]]. The basidia are thin-walled, mostly attached to four spores, and measure 25–30 by 8–10&nbsp;μm. Another cell type present in the hymenium is the cystidia, larger sterile cells that protrude beyond the basidia into the [[lumen (anatomy)|lumen]] of the hymenium, and act as air traps, regulating humidity.<ref>{{cite book|vauthors=Alexopoulos CJ, Mims CW, Blackwell M | title=Introductory Mycology | year=1996 | publisher=John Wiley and Sons |location=New York City|page=495 | isbn=0-471-52229-5}}</ref> ''B.&nbsp;edulis'' has pleurocystidia (cystidia located on the face of a pore) that are thin-walled, roughly spindle-shaped to [[ventricose]], and measure 30–45 by 7–10&nbsp;μm; the "stuffed" feature of the hymenium is caused by cheilocystidia—cells found on the edges of the pores.<ref name=Grund1976>{{cite book |title=Nova Scotian Boletes |vauthors=Grund DW, Harrison AK |year=1976 |publisher=J. Cramer |location=Lehre, Germany |isbn=3-7682-1062-6|pages=73–75}}</ref> The [[hypha]]e of ''B.&nbsp;edulis'' do not have [[clamp connection]]s.<ref name=Tylutki1987II/>


===Related species===
===Related species===
{{multiple image
{{double image|right|Boletus edulis var. grandedulis 27911.jpg|100|Boletus regineus 28082.jpg|100|''B. edulis'' var. ''grandedulis''|''B. regineus''|A mushroom with an orange-brown cap and a yellowish underside that somewhat resembles a sponge. The light-yellow stem is about half the thickness of the caps diameter. This mushroom is growing on the ground, surrounded by twigs, leaves, log and other forest floor debris.|A mushroom with an reddish-brown cap that is curled upwards to reveal a cream-coloured porous underside that somewhat resembles a sponge. The thick stipe has a pinkish hue, and its thickness is a little less than half of the cap's diameter. The mushroom has been pulled from the ground and the end of its stipe is a whitish colour caused by hyphal tufts, and is embedded with dirt and other small twigs.}}
| align = right
| image1 = Boletus edulis var. grandedulis 27911.jpg
| width1 = 100
| alt1 = A mushroom with an orange-brown cap and a yellowish underside that somewhat resembles a sponge. The light-yellow stem is about half the thickness of the caps diameter. This mushroom is growing on the ground, surrounded by twigs, leaves, log and other forest floor debris.
| caption1 = ''B. edulis'' var. ''grandedulis''
| image2 = Boletus regineus 28082.jpg
| width2 = 100
| alt2 = A mushroom with a reddish-brown cap that is curled upwards to reveal a cream-coloured porous underside that somewhat resembles a sponge. The thick stipe has a pinkish hue, and its thickness is a little less than half of the cap's diameter. The mushroom has been pulled from the ground and the end of its stipe is a whitish colour caused by hyphal tufts, and is embedded with dirt and other small twigs.
| caption2 = ''B. regineus''
| footer =
}}
Several similar brownish-coloured species are sometimes considered subspecies or forms of this mushroom. In Europe, in addition to ''B.&nbsp;edulis'' (or ''cèpe de Bordeaux''), the most popular are:
Several similar brownish-coloured species are sometimes considered subspecies or forms of this mushroom. In Europe, in addition to ''B.&nbsp;edulis'' (or ''cèpe de Bordeaux''), the most popular are:
*''Tête de nègre'' ("negro's head"; ''[[Boletus aereus]]''), much rarer than ''B.&nbsp;edulis'', is more highly regarded by gourmets, and more expensive. Usually smaller than ''B.&nbsp;edulis'', it is also distinctively darker in colour.<ref name = "Carluccio03"/> It is especially suited to drying.<ref name=Zeitl76/>
*''Cèpe bronzé'' ("dark cep"; ''[[Boletus aereus]]''), much rarer than ''B.&nbsp;edulis'', is more highly regarded by gourmets, and consequently more expensive. Usually smaller than ''B.&nbsp;edulis'', it is also distinctively darker in colour.<ref name = "Carluccio03"/> It is especially suited to drying.<ref name=Zeitl76/>
*''Cèpe des pins'' ("pine tree cep"; ''[[Boletus pinophilus]]'' or ''Boletus pinicola'') grows among pine trees. Rarer than ''B.&nbsp;edulis'', it is less appreciated by gourmets than the two other kinds of porcini, but remains a mushroom rated above most others.<ref name=Zeitl76/>
*''Cèpe des pins'' ("pine tree cep"; ''[[Boletus pinophilus]]'' or ''Boletus pinicola'') grows among pine trees. Rarer than ''B.&nbsp;edulis'', it is less appreciated by gourmets than the two other kinds of porcini, but remains a mushroom rated above most others.<ref name=Zeitl76/>
*''Cèpe d'été'' ("summer cep"; ''[[Boletus reticulatus]]''), also less common and found earlier.<ref name = "Carluccio03"/>
*''Cèpe d'été'' ("summer cep"; ''[[Boletus reticulatus]]''), also less common and found earlier.<ref name = "Carluccio03"/>


[[Molecular phylogenetic]] analyses have proven these three are all distinctive and separate species;<ref>{{cite journal |author=Vizzini A, Mello A, Ghignone S, Sechi C, Ruiu P, Bonfante P. |year=2008 |title=''Boletus edulis'' complex: from phylogenetic relationships to specific primers |journal=Pagine di Micologia |volume= |issue=30 |pages=49–52 |issn=1122-8911}}</ref> other taxa formerly believed to be unique species or subspecies, such as ''B.&nbsp;betulicola'', ''B.&nbsp;chippewaensis'', ''B.&nbsp;persoonii'', ''B.&nbsp;quercicola'' and ''B.&nbsp;venturii'', are now known to be part of a ''B.&nbsp;edulis'' [[species complex]] with a wide [[morphology (biology)|morphological]], ecological and geographic range,<ref>{{cite journal |author= Beugelsdijk DCM, van der Linde S, Zuccarello GC, den Bakker, Draisma SGA, Noordeloos ME.|year=2008 |title= A phylogenetic study of ''Boletus'' section ''Boletus'' in Europe |journal=Persoonia |pmid= 20467482 |volume=20 |issue=20 |pmc= 2865352 |pages=1–7|doi=10.3767/003158508X283692 }}</ref><ref name=dent10>{{cite journal| author=Dentinger, Bryn T.M. ''et. al''| title=Molecular phylogenetics of porcini mushrooms (''Boletus'' section ''Boletus'')| journal=Molecular Phylogenetics and Evolution| volume=57| year=2010| pages=1276–1292| url=http://static.msi.umn.edu/rreports/2010/211.pdf| doi=10.1016/j.ympev.2010.10.004| issue=3| pmid=20970511}}</ref> and that the genetic variability in this complex is low.<ref>{{cite journal|author=Beugelsdijk DC, van der Linde S, Zuccarello GC, den Bakker HC, Draisma SG, Noordeloos ME|year=2008|title=A phylogenetic study of ''Boletus'' section ''Boletus'' in Europe|journal=Persoonia|volume=20|pages=1–7|pmid=20467482|doi=10.3767/003158508X283692|pmc=2865352}}</ref> Similar molecular technology has been developed to rapidly and accurately identify ''B.&nbsp;edulis'' and other commercially important fungi.<ref>{{cite journal |author=Mello A, Ghignone S, Vizzini A, Sechi C, Ruiu P, Bonfante P. |year=2006 |title=ITS primers for the identification of marketable boletes |journal=Journal of Biotechnology |volume=121 |issue=3 |pages=318–29 |doi=10.1016/j.jbiotec.2005.08.022 |pmid=16213623}}</ref><ref>{{cite journal |author=Lian B, Zang J-P, Hou W-G, Yuan S, Smith DL.|year=2008 |title=PCR-based sensitive detection of the edible fungus ''Boletus edulis'' from rDNA ITS sequences |journal=Electronic Journal of Biotechnology |volume=11 |issue=3 |pages=1–8 |doi=10.2225/vol11-issue3-fulltext-4 |issn=0717-3458}}</ref>
[[Molecular phylogenetic]] analyses have proven these three are all distinctive and separate species;<ref>{{cite journal |vauthors=Vizzini A, Mello A, Ghignone S, Sechi C, Ruiu P, Bonfante P |year=2008 |title=''Boletus edulis'' complex: from phylogenetic relationships to specific primers |journal=Pagine di Micologia |issue=30 |pages=49–52 |issn=1122-8911}}</ref> other taxa formerly believed to be unique species or subspecies, such as ''B.&nbsp;betulicola'', ''B.&nbsp;chippewaensis'', ''B.&nbsp;persoonii'', ''B.&nbsp;quercicola'' and ''B.&nbsp;venturii'', are now known to be part of a ''B.&nbsp;edulis'' [[species complex]] with a wide [[morphology (biology)|morphological]], ecological and geographic range,<ref>{{cite journal |author1=Beugelsdijk DCM |author2=van der Linde S |author3=Zuccarello GC |author4=den Bakker |author5=Draisma SGA |author6=Noordeloos ME. |year=2008 |title= A phylogenetic study of ''Boletus'' section ''Boletus'' in Europe |journal=Persoonia |pmid= 20467482 |volume=20 |issue=1 |pmc= 2865352 |pages=1–7|doi=10.3767/003158508X283692 }}</ref><ref name=dent10>{{cite journal|vauthors=Dentinger BT, Ammirati JF, Both EE, Desjardin DE, Halling RE, Henkel TW, Moreau PA, Nagasawa E, Soytong K, Taylor AF, Watling R, Moncalvo JM, McLaughlin DJ |title=Molecular phylogenetics of porcini mushrooms (''Boletus'' section ''Boletus'') |journal=Molecular Phylogenetics and Evolution |volume=57 |year=2010 |pages=1276–1292 |url=http://static.msi.umn.edu/rreports/2010/211.pdf |doi=10.1016/j.ympev.2010.10.004 |issue=3 |pmid=20970511 |archive-url=https://web.archive.org/web/20130523094609/http://static.msi.umn.edu/rreports/2010/211.pdf |archive-date=2013-05-23 }}</ref> and that the genetic variability in this complex is low.<ref>{{cite journal|vauthors=Beugelsdijk DC, van der Linde S, Zuccarello GC, den Bakker HC, Draisma SG, Noordeloos ME |year=2008|title=A phylogenetic study of ''Boletus'' section ''Boletus'' in Europe|journal=Persoonia|volume=20|pages=1–7|pmid=20467482|doi=10.3767/003158508X283692|pmc=2865352}}</ref> Similar molecular technology has been developed to rapidly and accurately identify ''B.&nbsp;edulis'' and other commercially important fungi.<ref>{{cite journal |vauthors=Mello A, Ghignone S, Vizzini A, Sechi C, Ruiu P, Bonfante P |year=2006 |title=ITS primers for the identification of marketable boletes |journal=Journal of Biotechnology |volume=121 |issue=3 |pages=318–29 |doi=10.1016/j.jbiotec.2005.08.022 |pmid=16213623}}</ref><ref>{{cite journal |vauthors=Lian B, Zang JP, Hou WG, Yuan S, Smith DL |year=2008 |title=PCR-based sensitive detection of the edible fungus ''Boletus edulis'' from rDNA ITS sequences |journal=Electronic Journal of Biotechnology |volume=11 |issue=3 |pages=1–8 |doi=10.2225/vol11-issue3-fulltext-4 |doi-broken-date=2024-11-10 |issn=0717-3458|url=http://www.bioline.org.br/pdf?ej08040 |hdl=1807/48975 |hdl-access=free }}</ref>


Three divergent lineages found in Yunnan province in China that are commonly marketed and sold as ''B. edulis'' (and are actually more closely related to ''B. aereus'') were described in 2013 as ''[[Boletus bainiugan|B. bainiugan]]'', ''[[Boletus meiweiniuganjun|B. meiweiniuganjun]]'' and ''[[Boletus shiyong|B. shiyong]]''.<ref name="Dentinger 2013">{{cite journal|author=Dentinger B|date=12 October 2013|title=Nomenclatural novelties|journal=Index Fungorum|issue=29|pages=1|issn=2049-2375|url=http://www.indexfungorum.org/Publications/Index%20Fungorum%20no.29.pdf}}</ref><ref name="Dentinger 2014">{{cite journal|author=Dentinger B, Suz LM|date=2014|title=What’s for dinner? Undescribed species of porcini in a commercial packet|journal=PeerJ |issue=2:e570|doi=10.7717/peerj.570}}</ref>
Three divergent lineages found in Yunnan province in China that are commonly marketed and sold as ''B. edulis'' (and are actually more closely related to ''B. aereus'') were described in 2013 as ''[[Boletus bainiugan|B. bainiugan]]'', ''[[Boletus meiweiniuganjun|B. meiweiniuganjun]]'' and ''[[Boletus shiyong|B. shiyong]]''.<ref name="Dentinger 2013">{{cite journal|author=Dentinger B|date=12 October 2013|title=Nomenclatural novelties|journal=Index Fungorum|issue=29|page=1|issn=2049-2375|url=http://www.indexfungorum.org/Publications/Index%20Fungorum%20no.29.pdf}}</ref><ref name="Dentinger 2014">{{cite journal|vauthors=Dentinger B, Suz LM |date=2014|title=What's for dinner? Undescribed species of porcini in a commercial packet|journal=PeerJ |volume=2|issue=2:e570|pages=e570|doi=10.7717/peerj.570|pmid=25279259|pmc=4179395 |doi-access=free }}</ref> The classification has since been updated and expanded. All lineages are still members of ''Boletus'' sect. ''Boletus'', the sensu sticto "porcini clade" of the genus.<ref name=Cui2016>{{cite journal |last1=Cui |first1=Yang-Yang |last2=Feng |first2=Bang |last3=Wu |first3=Gang |last4=Xu |first4=Jianping |last5=Yang |first5=Zhu L. |title=Porcini mushrooms (Boletus sect. Boletus) from China |journal=Fungal Diversity |date=November 2016 |volume=81 |issue=1 |pages=189–212 |doi=10.1007/s13225-015-0336-7 |s2cid=12866685 |url=http://groups.kib.cas.cn/klpb/yzl/tnh_fblw/201509/P020150928547013572278.pdf}}</ref>


[[File:Boletus rex-veris 42967.jpg|thumb|left|''B. rex-veris''|alt=A group of three mushrooms with reddish-brown caps, bright yellow porous undersides, and thick white stipes. They are growing on the ground in soil covered with pine needles.]]
[[File:Boletus rex-veris 42967.jpg|thumb|left|''B. rex-veris''|alt=A group of three mushrooms with reddish-brown caps, bright yellow porous undersides, and thick white stipes. They are growing on the ground in soil covered with pine needles.]]
Western North America has several species closely related to ''B.&nbsp;edulis''. The white king bolete (''[[Boletus barrowsii]]''), found in parts of [[Colorado]], [[New Mexico]], [[Arizona]], and [[California]] (and possibly elsewhere), is named after its discoverer Chuck Barrows.<ref>{{cite web|url=http://botit.botany.wisc.edu/Toms_fungi/aug2004.html|accessdate=2008-09-23|title=''Boletus barrowsii'', Chuck Barrows' bolete|author=Volk T.|date=August 2004|work=Tom Volk's Fungus of the Month|publisher=Department of Biology, University of Wisconsin-La Crosse|archiveurl=https://web.archive.org/web/20081003220326/http://botit.botany.wisc.edu/toms_fungi/aug2004.html <!--Added by H3llBot-->|archivedate=2008-10-03}}</ref> It is lighter in colour than ''B.&nbsp;edulis'', having a cream-coloured cap with pink tones;<ref name=Miller2006>{{cite book |author=Miller HR, Miller OK. |title=North American Mushrooms: a Field Guide to Edible and Inedible Fungi |publisher=Falcon Guide |location=Guilford, Conn |year=2006 |page=392 |isbn=0-7627-3109-5 |url=https://books.google.com/?id=zjvXkLpqsEgC&lpg=PA381&dq=boletus%20edulis&pg=PA392#v=onepage&q=boletus%20edulis |accessdate=2009-11-04}}</ref> often mycorrhizal with [[Ponderosa pine]], it tends to grow in areas where there is less rainfall. Some find its flavour as good as if not better than ''B.&nbsp;edulis''.<ref name=Evenson>{{cite book |title=Mushrooms of Colorado and the Southern Rocky Mountains |author=Evenson VS. |year=1997 |publisher=Westcliffe Publishers |location= Boulder, Colorado|isbn=978-1-56579-192-3 |page=157|url=https://books.google.com/?id=EAeDeyqZLq0C&lpg=PA156&dq=boletus%20edulis&pg=PA157#v=onepage&q=boletus%20edulis|accessdate=2009-10-20}}</ref> The California king bolete (''Boletus edulis var. grandedulis'') can reach massive proportions, and is distinguished from ''B.&nbsp;edulis'' by a mature pore surface that is brown to slightly reddish. The cap colour appears to be affected by the amount of light received during its development, and may range from white in young specimens grown under thick [[forest canopy|canopy]], to dark-brown, red-brown or yellow brown in those specimens receiving more light.<ref name=eb1>{{cite journal |author=Arora D. |year=2008 |title=California porcini: three new taxa, observations on their harvest, and the tragedy of no commons |journal=Economic Botany |volume=62 |issue=3 |pages=356–75 |url=http://www.davidarora.com/uploads/arora_california_porcini.pdf |format=PDF |doi=10.1007/s12231-008-9050-7}}</ref> The queen bolete (''[[Boletus regineus]]''), formerly considered a variety of ''B.&nbsp;aereus'', is also a choice edible. It is generally smaller than ''B.&nbsp;edulis'', and unlike that species, is typically found in [[mixed forest]]s.<ref name="urlCalifornia Fungi: Boletus regineus">{{cite web |url=http://www.mykoweb.com/CAF/species/Boletus_regineus.html |title=California Fungi: ''Boletus regineus'' |author=Wood M, Stevens F |publisher=MykoWeb |accessdate=2009-11-02}}</ref> The spring king bolete (''[[Boletus rex-veris]]''), formerly considered a variety of ''B.&nbsp;edulis'' or ''B.&nbsp;pinophilus'', is found throughout western North America. In contrast to ''B.&nbsp;edulis'', ''B.&nbsp;rex-veris'' tends to fruit in clusters, and, as its common name suggests, appears in the spring.<ref name="urlCalifornia Fungi: Boletus rex-veris">{{cite web |url=http://www.mykoweb.com/CAF/species/Boletus_rex-veris.html |title=California Fungi: ''Boletus rex-veris'' |author=Wood M, Stevens F |publisher= MykoWeb|accessdate=2009-11-03}}</ref>
Western North America has several species closely related to ''B.&nbsp;edulis''. The white king bolete (''[[Boletus barrowsii]]''), found in parts of [[Colorado]], [[New Mexico]], [[Arizona]], and California (and possibly elsewhere), is named after its discoverer Chuck Barrows.<ref>{{cite web|url=http://botit.botany.wisc.edu/Toms_fungi/aug2004.html|access-date=2008-09-23|title=''Boletus barrowsii'', Chuck Barrows' bolete|author=Volk T.|date=August 2004|work=Tom Volk's Fungus of the Month|publisher=Department of Biology, University of Wisconsin-La Crosse|archive-url=https://web.archive.org/web/20081003220326/http://botit.botany.wisc.edu/toms_fungi/aug2004.html <!--Added by H3llBot-->|archive-date=2008-10-03}}</ref> It is lighter in colour than ''B.&nbsp;edulis'', having a cream-coloured cap with pink tones;<ref name=Miller2006>{{cite book |vauthors=Miller HR, Miller OK |title=North American Mushrooms: a Field Guide to Edible and Inedible Fungi |publisher=Falcon Guide |location=Guilford, Conn |year=2006 |page=392 |isbn=0-7627-3109-5 }}</ref> often mycorrhizal with [[Ponderosa pine]], it tends to grow in areas where there is less rainfall. Some find its flavour as good as if not better than ''B.&nbsp;edulis''.<ref name=Evenson>{{cite book |title=Mushrooms of Colorado and the Southern Rocky Mountains |author=Evenson VS. |year=1997 |publisher=Westcliffe Publishers |location= Boulder, Colorado|isbn=978-1-56579-192-3 |page=157|url=https://books.google.com/books?id=EAeDeyqZLq0C&q=boletus%20edulis&pg=PA157|access-date=2009-10-20}}</ref> The California king bolete (''Boletus edulis var. grandedulis'') can reach massive proportions, and is distinguished from ''B.&nbsp;edulis'' by a mature pore surface that is brown to slightly reddish. The cap colour appears to be affected by the amount of light received during its development, and may range from white in young specimens grown under thick [[forest canopy|canopy]], to dark-brown, red-brown or yellow brown in those specimens receiving more light.<ref name=eb1>{{cite journal |author=Arora D. |year=2008 |title=California porcini: three new taxa, observations on their harvest, and the tragedy of no commons |journal=Economic Botany |volume=62 |issue=3 |pages=356–75 |url=http://www.davidarora.com/uploads/arora_california_porcini.pdf |doi=10.1007/s12231-008-9050-7 |s2cid=23176365 |access-date=2009-11-04 |archive-date=2012-03-12 |archive-url=https://web.archive.org/web/20120312213054/http://www.davidarora.com/uploads/arora_california_porcini.pdf }}</ref> The queen bolete (''[[Boletus regineus]]''), formerly considered a variety of ''B.&nbsp;aereus'', is also a choice edible. It is generally smaller than ''B.&nbsp;edulis'', and unlike that species, is typically found in [[mixed forest]]s.<ref name="urlCalifornia Fungi: Boletus regineus">{{cite web |url=http://www.mykoweb.com/CAF/species/Boletus_regineus.html |title=California Fungi: ''Boletus regineus'' |vauthors=Wood M, Stevens F |publisher=MykoWeb |access-date=2009-11-02}}</ref> The spring king bolete (''[[Boletus rex-veris]]''), formerly considered a variety of ''B.&nbsp;edulis'' or ''B.&nbsp;pinophilus'', is found throughout western North America. In contrast to ''B.&nbsp;edulis'', ''B.&nbsp;rex-veris'' tends to fruit in clusters, and, as its common name suggests, appears in the spring.<ref name="urlCalifornia Fungi: Boletus rex-veris">{{cite web |url=http://www.mykoweb.com/CAF/species/Boletus_rex-veris.html |title=California Fungi: ''Boletus rex-veris'' |vauthors=Wood M, Stevens F |publisher= MykoWeb|access-date=2009-11-03}}</ref> ''[[Boletus fibrillosus|B.&nbsp;fibrillosus]]'' is edible but considered inferior in taste.<ref>{{cite book|last1=Trudell|first1=Steve|url=https://books.google.com/books?id=WevHvt6Tr8kC|title=Mushrooms of the Pacific Northwest|last2=Ammirati|first2=Joe|publisher=Timber Press|year=2009|isbn=978-0-88192-935-5|series=Timber Press Field Guides|location=Portland, OR|page=217}}</ref>


==Habitat and distribution==
==Habitat and distribution==
[[File:Boletus edulis 11.jpg|thumb|right|upright|In Lithuania|alt=A mushroom with a brown smooth cap the shape of a halved sphere, atop a thick, dirty white stipe. The mushroom is growing on a sloping patch of ground amongst moss, twigs and other forest debris; trees can be faintly seen in the background.]]
[[File:Boletus edulis 11.jpg|thumb|right|upright|In Lithuania|alt=A mushroom with a brown smooth cap the shape of a halved sphere, atop a thick, dirty white stipe. The mushroom is growing on a sloping patch of ground amongst moss, twigs and other forest debris; trees can be faintly seen in the background.]]
The fruit bodies of ''Boletus edulis'' can grow singly or in small clusters of two or three specimens. The mushroom's habitat consists of areas dominated by pine (''[[Pinus]]'' spp.), spruce (''[[Picea]]'' spp.), hemlock (''[[Tsuga]]'' spp.) and fir (''[[Abies]]'' spp.) trees, although other hosts include [[chestnut]], [[Chrysolepis|chinquapin]], [[beech]], ''[[Keteleeria]]'' spp., ''[[Lithocarpus]]'' spp., and [[oak]]. In California, porcini have been collected in a variety of forests, such as coastal forests, dry interior oak forests and [[savanna]]s and interior high-elevation [[Montane forest|montane]] [[mixed forest]]s,<ref name=Hall1998/> to an altitude of {{convert|3,500|m|ft|abbr=on}}.<ref name=Hall2003>{{cite book|author=Hall IR, Stephenson SR, Buchanan PK, Yun W, Cole A.|title=Edible and Poisonous Mushrooms of the World|publisher=Timber Press|location=Portland, Oregon|year=2003|pages=224–25|isbn=0-88192-586-1}}</ref> In northwestern Spain, they are common in [[scrubland]]s dominated by the [[Cistaceae|rock rose]] species ''[[Cistus ladanifer]]'' and ''Halimium lasianthum''.<ref>{{cite journal|author=Oria-de-Rueda J, Martin-Pinto P, Olaizola J.|year=2008 |title=Bolete productivity of cistaceous scrublands in northwestern Spain |journal=Economic Botany |volume=62 |issue=3|pages=323–30 |doi=10.1007/s12231-008-9031-x}}</ref>
The fruit bodies of ''Boletus edulis'' can grow singly or in small clusters of two or three specimens. The mushroom's habitat consists of areas dominated by pine (''[[Pinus]]'' spp.), spruce (''[[Picea]]'' spp.), hemlock (''[[Tsuga]]'' spp.) and fir (''[[Abies]]'' spp.) trees, although other hosts include [[chestnut]], [[Chrysolepis|chinquapin]], [[beech]], ''[[Keteleeria]]'' spp., ''[[Lithocarpus]]'' spp., and [[oak]]. In California, porcini have been collected in a variety of forests, such as coastal forests, dry interior oak forests and [[savanna]]s and interior high-elevation [[Montane forest|montane]] [[mixed forest]]s,<ref name=Hall1998/> to an altitude of {{convert|3,500|m|ft|abbr=on}}.<ref name=Hall2003>{{cite book|vauthors=Hall IR, Stephenson SR, Buchanan PK, Yun W, Cole A |title=Edible and Poisonous Mushrooms of the World|url=https://archive.org/details/ediblepoisonousm00hall |url-access=limited |publisher=Timber Press|location=Portland, Oregon|year=2003|pages=[https://archive.org/details/ediblepoisonousm00hall/page/n224 224]–25|isbn=0-88192-586-1}}</ref> In northwestern Spain, they are common in [[scrubland]]s dominated by the [[Cistaceae|rock rose]] species ''[[Cistus ladanifer]]'' and ''Halimium lasianthum''.<ref>{{cite journal|vauthors=Oria-de-Rueda J, Martin-Pinto P, Olaizola J |year=2008 |title=Bolete productivity of cistaceous scrublands in northwestern Spain |journal=Economic Botany |volume=62 |issue=3|pages=323–30 |doi=10.1007/s12231-008-9031-x|s2cid=20095996 }}</ref> In [[Southern France|the Midi region of south-west France]], they are especially favoured and locally called ''cèpe de Bordeaux'' after the town from which they are traded to the north and abroad.<ref>{{cite book|first=Colin Duncan|last=Taylor|title=Menu from the Midi: A Gastronomic Journey through the South of France|year=2021|publisher=Matador |language=English| isbn= 978-1-80046-496-4}}</ref>


''Boletus edulis'' has a [[cosmopolitan distribution]], concentrated in cool-[[temperate]] to [[subtropical]] regions.<ref name=Hall1998/> It is common in [[Europe]]—from northern Scandinavia, south to the extremities of Greece and Italy—and [[North America]], where its southern range extends as far south as Mexico.<ref name=Wang1995>{{cite journal |doi=10.1080/01140671.1995.9513892 |author=Wang Y, Sinclair L, Hall IR, Cole ALJ. |year=1995 |title=''Boletus edulis'' sensu lato: a new record for New Zealand|journal=New Zealand Journal of Crop and Horticultural Science |volume=23 |pages=227–31 |url=http://pdfserve.informaworld.com/915037__920077166.pdf |accessdate=2009-12-06 |issue=2}}</ref> It is well known from the [[Borgotaro]] area of [[Parma]], [[Italy]], and has [[Protected designation of origin|PGI]] status there. The European distribution extends north to Scandinavia and south to southern Italy and Morocco.<ref name=Hall1998>{{cite journal |author=Hall IR, Lyon AJE, Wang Y, Sinclair L. |year=1998 |title=Ectomycorrhizal fungi with edible fruiting bodies 2. ''Boletus edulis'' |journal=Economic Botany |volume=52 |issue=1 |pages=44–56 |doi=10.1007/BF02861294}}</ref> In China, the mushroom can be found from the northeastern [[Heilongjiang Province]] to the [[Yunnan-Guizhou Plateau]] and [[Tibet]].<ref name=Wang1995/> It has been recorded growing under ''Pinus'' and ''Tsuga'' in [[Sagarmatha National Park]] in [[Nepal]],<ref>{{cite journal |author=Giri A, Rana P.|year=2007|title=Some Higher Fungi from Sagarmatha National Park (SNP) and its adjoining areas, Nepal|journal=Scientific World |volume=5 |issue=5 |pages=67–74 |doi=10.3126/sw.v5i5.2659}}</ref> as well as in the [[India]]n forests of [[Arunachal Pradesh]].<ref>{{cite journal |author=Adhikary RK, Baruah P, Kalita P, Bordoloi D. |year=1999 |title=Edible mushrooms growing in forests of Arunachal Pradesh |journal=Advances in Horticulture and Forestry |volume=6 |pages=119–23}}</ref> In West Asia, the species has been reported from the northwest forests of Iran.<ref>{{cite journal |author=Asef MR. |year=2008 |title=Macrofungi of Arasbaran 2. Boletes (families Boletaceae and Suillaceae) |journal=Rostaniha |volume=9 |issue=2 |pages=210–29 |url=http://www.sid.ir/fa/VEWSSID/J_pdf/50313873207.pdf |format=PDF |language=Arabic}}</ref>
''Boletus edulis'' has a [[cosmopolitan distribution]], concentrated in cool-[[temperate]] to [[subtropical]] regions.<ref name=Hall1998/> It is common in Europe—from northern Scandinavia, south to the extremities of Greece and Italy—and North America, where its southern range extends as far south as Mexico.<ref name=Wang1995>{{cite journal |doi=10.1080/01140671.1995.9513892 |vauthors=Wang Y, Sinclair L, Hall IR, Cole AL |year=1995 |title=''Boletus edulis'' sensu lato: a new record for New Zealand|journal=New Zealand Journal of Crop and Horticultural Science |volume=23 |pages=227–31 |url=http://pdfserve.informaworld.com/915037__920077166.pdf |access-date=2009-12-06 |issue=2|doi-access=free }}</ref> It is well known from the [[Borgotaro]] area of [[Parma]], Italy, and has [[Protected designation of origin|PGI]] status there. The European distribution extends north to Scandinavia and south to southern Italy and Morocco.<ref name=Hall1998>{{cite journal |vauthors=Hall IR, Lyon AJ, Wang Y, Sinclair L |year=1998 |title=Ectomycorrhizal fungi with edible fruiting bodies 2. ''Boletus edulis'' |journal=Economic Botany |volume=52 |issue=1 |pages=44–56 |doi=10.1007/BF02861294|s2cid=20934023 }}</ref> In the American [[Pacific Northwest]], it can be found from May to October.<ref>{{Cite web |title=Seasonal Chart for Edible Mushrooms |url=https://www.centraloregonmushroomclub.com/Mushroom-Seasons.htm?m=102&s=866 |access-date=2024-03-31 |website=Central Oregon Mushroom Club |archive-date=2024-03-31 |archive-url=https://web.archive.org/web/20240331075937/https://www.centraloregonmushroomclub.com/Mushroom-Seasons.htm?m=102&s=866 |url-status=dead }}</ref> In China, the mushroom can be found from the northeastern [[Heilongjiang]] to the [[Yunnan–Guizhou Plateau]] and [[Tibet]].<ref name=Wang1995/> It has been recorded growing under ''Pinus'' and ''Tsuga'' in [[Sagarmatha National Park]] in Nepal,<ref>{{cite journal |vauthors=Giri A, Rana P |year=2007|title=Some Higher Fungi from Sagarmatha National Park (SNP) and its adjoining areas, Nepal|journal=Scientific World |volume=5 |issue=5 |pages=67–74 |doi=10.3126/sw.v5i5.2659|doi-access=free }}</ref> as well as in the Indian forests of [[Arunachal Pradesh]].<ref>{{cite journal |vauthors=Adhikary RK, Baruah P, Kalita P, Bordoloi D |year=1999 |title=Edible mushrooms growing in forests of Arunachal Pradesh |journal=Advances in Horticulture and Forestry |volume=6 |pages=119–23}}</ref> In West Asia, the species has been reported from the northwest forests of Iran.<ref>{{cite journal |author=Asef MR. |year=2008 |title=Macrofungi of Arasbaran 2. Boletes (families Boletaceae and Suillaceae) |journal=Rostaniha |volume=9 |issue=2 |pages=210–29 |url=http://www.sid.ir/fa/VEWSSID/J_pdf/50313873207.pdf |language=ar}}</ref>

===Cultivation===
Some steps have been made towards cultivating ''Boletus edulis'',<ref name="Wang2014">{{Citation |last1=Wang |first1=Yun |title=Recent Advances in Cultivation of Edible Mycorrhizal Mushrooms |date=2014 |url=https://www.researchgate.net/publication/270546094 |work=Mycorrhizal Fungi: Use in Sustainable Agriculture and Land Restoration |volume=41 |page=384 |editor-last=Solaiman |editor-first=Zakaria M. |place=Berlin, Heidelberg |publisher=Springer Berlin Heidelberg |doi=10.1007/978-3-662-45370-4_23 |isbn=978-3-662-45369-8 |access-date=2022-10-03 |last2=Chen |first2=Ying Long |editor2-last=Abbott |editor2-first=Lynette K. |editor3-last=Varma |editor3-first=Ajit}}</ref> including mycorrhization of [[Cistus ladanifer|rockrose]] shrubs enhanced by helper bacteria.<ref name="Mediavilla2015">{{Cite journal |last1=Mediavilla |first1=Olaya |last2=Olaizola |first2=Jaime |last3=Santos-del-Blanco |first3=Luis |last4=Oria-de-Rueda |first4=Juan Andrés |last5=Martín-Pinto |first5=Pablo |date=2015-07-26 |title=Mycorrhization between Cistus ladanifer L. and Boletus edulis Bull is enhanced by the mycorrhiza helper bacteria Pseudomonas fluorescens Migula |url=http://sostenible.palencia.uva.es/system/files/publicaciones/mediavilla_et_al._2015._mycorrhization_between_cistus_ladanifer_l._and_boletus_edulis.pdf |journal=Mycorrhiza |language=en |volume=26 |issue=2 |pages=161–168 |doi=10.1007/s00572-015-0657-0 |pmid=26208816 |s2cid=8967720 |issn=0940-6360}}</ref>


===Non-native introductions===
===Non-native introductions===
''Boletus edulis'' grows in some areas where it is not believed to be [[indigenous (ecology)|indigenous]]. It is often found underneath oak and [[Betula pendula|silver birch]] in [[Hagley Park, Christchurch|Hagley Park]] in central [[Christchurch]], New Zealand, where it is likely to have been [[introduced species|introduced]],<ref>{{cite journal |author = Orlovich D, Stringer A, Yun W, Hall I, Prime G, Danell E, Weden C, Bulman S.|year= 2004 |title = ''Boletus edulis'' Bull. Ex Fries in New Zealand|journal= Australasian Mycological Society Newsletter|volume=1 |issue=1 |page= 6}}</ref> probably on the roots of container-grown beech, birch, and oak in the mid-19th century—around the time exotic trees began to be planted in the Christchurch area.<ref name=Wang1995/> Similarly, it has been collected in [[Adelaide Hills]] region of Australia in association with three species of introduced trees.<ref name="Catcheside 2011">{{cite journal |author=Catcheside PS, Catcheside DEA. |title=''Boletus edulis'' (Boletaceae), a new record for Australia |journal=Journal of the Adelaide Botanical Garden |year=2011 |volume=25 |pages=5–10 |url=http://www.environment.sa.gov.au/Knowledge_Bank/Science_research/State_Herbarium/Resources/Publications/Journal_of_the_Adelaide_Botanic_Gardens/Journal_search/25/1 |format=PDF}}</ref> It has been growing plentifully in association with pine forests in the southern [[KwaZulu-Natal Midlands]] in South Africa for more than 50 years and is believed to have been introduced with the import of pine trees.<ref>{{cite journal |author=Eicker A.|year=1990|title=Commercial mushroom production in South Africa |journal=Bulletin|issue=418 | publisher = Department of Agricultural Development|location =Pretoria}}</ref><ref>{{cite journal |author=Marais LJ, Kotzé JM. |year=1977|title=Notes on ectotrophic mycorrhizae of ''Pinus patula'' in South Africa|journal=South African Forestry Journal |volume=100|pages=61–71 |doi=10.1080/00382167.1977.9629436}}</ref> It also grows in pine plantations in neighboring Zimbabwe.<ref name=Masuka1996/>
''Boletus edulis'' grows in some areas where it is not believed to be [[indigenous (ecology)|indigenous]]. It is often found underneath oak and [[Betula pendula|silver birch]] in [[Hagley Park, Christchurch|Hagley Park]] in central [[Christchurch]], New Zealand, where it is likely to have been [[introduced species|introduced]],<ref>{{cite journal |vauthors=Orlovich D, Stringer A, Yun W, Hall I, Prime G, Danell E, Weden C, Bulman S |year= 2004 |title = ''Boletus edulis'' Bull. Ex Fries in New Zealand|journal= Australasian Mycological Society Newsletter|volume=1 |issue=1 |page= 6}}</ref> probably on the roots of container-grown beech, birch, and oak in the mid-19th century—around the time exotic trees began to be planted in the Christchurch area.<ref name=Wang1995/> Similarly, it has been collected in [[Adelaide Hills]] region of Australia in association with three species of introduced trees.<ref name="Catcheside 2011">{{cite journal |vauthors=Catcheside PS, Catcheside DE |title=''Boletus edulis'' (Boletaceae), a new record for Australia |journal=Journal of the Adelaide Botanic Garden |year=2011 |volume=25 |pages=5–10 |url=http://www.environment.sa.gov.au/Knowledge_Bank/Science_research/State_Herbarium/Resources/Publications/Journal_of_the_Adelaide_Botanic_Gardens/Journal_search/25/1 |format=PDF |archive-url=https://web.archive.org/web/20140517151731/http://www.environment.sa.gov.au/Knowledge_Bank/Science_research/State_Herbarium/Resources/Publications/Journal_of_the_Adelaide_Botanic_Gardens/Journal_search/25/1 |archive-date=2014-05-17 }}</ref> It has been growing plentifully in association with pine forests in the southern [[KwaZulu-Natal Midlands]] in South Africa for more than 50 years and is believed to have been introduced with the import of pine trees.<ref>{{cite journal |author=Eicker A.|year=1990|title=Commercial mushroom production in South Africa |journal=Bulletin|issue=418 | publisher = Department of Agricultural Development|location =Pretoria}}</ref><ref>{{cite journal |vauthors=Marais LJ, Kotzé JM |year=1977|title=Notes on ectotrophic mycorrhizae of ''Pinus patula'' in South Africa|journal=South African Forestry Journal |volume=100|pages=61–71 |doi=10.1080/00382167.1977.9629436}}</ref> It also grows in pine plantations in neighboring Zimbabwe.<ref name=Masuka1996/>


==Ecology==
==Ecology==
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===Fruit body production===
===Fruit body production===
Italian folklore holds that porcini sprout up at the time of the new moon;<ref name = "Carluccio03"/> research studies have tried to investigate more scientifically the factors that influence the production of fruit bodies. Although fruit bodies may appear any time from summer to autumn (June to November in the UK), their growth is known to be triggered by rainfall during warm periods of weather followed by frequent autumn rain with a drop in soil temperature.<ref name=Hall1998/> Above average rainfall may result in the rapid appearance of large numbers of boletes, in what is known in some circles as a "bolete year".<ref name=Nilper>{{cite book | author = Nilson S, Persson O. | year = 1977 | title = Fungi of Northern Europe 1: Larger Fungi (Excluding Gill-Fungi) | publisher = Penguin |location=Harmondsworth, England|isbn=0-14-063005-8|page=100}}</ref> A 2004 [[field study]] indicated that fruit body production is enhanced by an open and sunny wood habitat,<ref>{{cite journal |author=Salerni E, Perini C.|year=2004 |title=Experimental study for increasing productivity of ''Boletus edulis'' s.l. in Italy |journal=Forest Ecology and Management |volume=201 |issue=2–3 |pages=161–70|doi=10.1016/j.foreco.2004.06.027}}</ref> corroborating an earlier observation made in a Zimbabwean study;<ref name=Masuka1996>{{cite journal |author=Masuka AJ. |year=1996 |title=Dynamics of mushroom (''Boletus edulis'') production in pine plantations in Zimbabwe |journal= JASSA, Journal of Applied Science in Southern Africa |volume=2 |issue=2 |pages=69–76 |doi=10.4314/jassa.v2i2.16877}}</ref> removal of the litter layer on the [[forest floor]] appeared to have a negative effect on fruit body production, but previous studies reported contradictory results.<ref>{{cite journal |author=Baar J, Ter Braak CJF.|year=1996 |title=Ectomycorrhizal sporocarp occurrence as affected by manipulation of litter and humus layers in Scots pine stands of different age |journal=Applied Soil Ecology |volume=4 |issue=1 |pages=61–73|doi= 10.1016/0929-1393(96)00097-2}}</ref><ref>{{cite journal |author=Baar J, de Vries FW. |year=1995 |title=Effects of manipulation of litter and humus layer on ectomycorrhizal colonization potential in Scots pine stands of different age |journal=Mycorrhiza |volume=5 |issue=4 |pages=267–72 |url=http://www.springerlink.com/content/k505pr50v0732860/fulltext.pdf?page=1 |doi=10.1007/BF00204960 |format=First page preview PDF}}</ref> A Lithuanian study conducted in 2001 concluded that the maximal daily growth rate of the cap (about 21&nbsp;mm or 0.8&nbsp;in) occurred when the [[relative air humidity]] was the greatest, and the fruit bodies ceased growing when the air humidity dropped below 40%. Factors most likely to inhibit the appearance of fruit bodies included prolonged drought, inadequate air and soil humidity, sudden decreases of night air temperatures, and the appearance of the first [[frost]].<ref>{{cite journal |author=Kasparavicius, J. |year=2001 |title=Influence of climatic conditions on the growth of fruit bodies of ''Boletus edulis'' |journal=Botanica Lithuanica |volume=7 |issue=1 |pages=73–78 |issn=1392-1665}}</ref> Plots facing north tend to produce more mushrooms compared to equivalent plots facing south.<ref>{{cite journal |author=Martín-Pinto P, Vaquerizo H, Peñalver F, Olaizola J, Oria-de-Rueda J.|year=2006 |title=Early effects of a wildfire on the diversity and production of fungal communities in Mediterranean vegetation types dominated by ''Cistus ladanifer'' and ''Pinus pinaster'' in Spain |journal=Forest Ecology and Management |volume=225 |issue=1–3 |pages=296–305|doi=10.1016/j.foreco.2006.01.006}}</ref>
Italian folklore holds that porcini sprout up at the time of the new moon;<ref name = "Carluccio03"/> research studies have tried to investigate more scientifically the factors that influence the production of fruit bodies. Although fruit bodies may appear any time from summer to autumn (June to November in the UK), their growth is known to be triggered by rainfall during warm periods of weather followed by frequent autumn rain with a drop in soil temperature.<ref name=Hall1998/> Above average rainfall may result in the rapid appearance of large numbers of boletes, in what is known in some circles as a "bolete year".<ref name=Nilper>{{cite book |vauthors=Nilson S, Persson O | year = 1977 | title = Fungi of Northern Europe 1: Larger Fungi (Excluding Gill-Fungi) | publisher = Penguin |location=Harmondsworth, England|isbn=0-14-063005-8|page=100}}</ref> A 2004 [[field study]] indicated that fruit body production is enhanced by an open and sunny wood habitat,<ref>{{cite journal |vauthors=Salerni E, Perini C |year=2004 |title=Experimental study for increasing productivity of ''Boletus edulis'' s.l. in Italy |journal=Forest Ecology and Management |volume=201 |issue=2–3 |pages=161–70|doi=10.1016/j.foreco.2004.06.027}}</ref> corroborating an earlier observation made in a Zimbabwean study;<ref name=Masuka1996>{{cite journal |author=Masuka AJ. |year=1996 |title=Dynamics of mushroom (''Boletus edulis'') production in pine plantations in Zimbabwe |journal= Journal of Applied Science in Southern Africa |volume=2 |issue=2 |pages=69–76 |doi=10.4314/jassa.v2i2.16877|url=http://opendocs.ids.ac.uk/opendocs/handle/123456789/8944 }}</ref> removal of the litter layer on the [[forest floor]] appeared to have a negative effect on fruit body production, but previous studies reported contradictory results.<ref>{{cite journal |vauthors=Baar J, Ter Braak CJ |year=1996 |title=Ectomycorrhizal sporocarp occurrence as affected by manipulation of litter and humus layers in Scots pine stands of different age |journal=Applied Soil Ecology |volume=4 |issue=1 |pages=61–73|doi= 10.1016/0929-1393(96)00097-2}}</ref><ref>{{cite journal |vauthors=Baar J, de Vries FW |year=1995 |title=Effects of manipulation of litter and humus layer on ectomycorrhizal colonization potential in Scots pine stands of different age |journal=Mycorrhiza |volume=5 |issue=4 |pages=267–72 |doi=10.1007/BF00204960 |s2cid=32618842 }}</ref> A Lithuanian study conducted in 2001 concluded that the maximal daily growth rate of the cap (about 21&nbsp;mm or 0.8&nbsp;in) occurred when the [[relative air humidity]] was the greatest, and the fruit bodies ceased growing when the air humidity dropped below 40%. Factors most likely to inhibit the appearance of fruit bodies included prolonged drought, inadequate air and soil humidity, sudden decreases of night air temperatures, and the appearance of the first [[frost]].<ref>{{cite journal |author=Kasparavicius, J. |year=2001 |title=Influence of climatic conditions on the growth of fruit bodies of ''Boletus edulis'' |journal=Botanica Lithuanica |volume=7 |issue=1 |pages=73–78 |issn=1392-1665}}</ref> Plots facing north tend to produce more mushrooms compared to equivalent plots facing south.<ref>{{cite journal |vauthors=Martín-Pinto P, Vaquerizo H, Peñalver F, Olaizola J, Oria-de-Rueda J |year=2006 |title=Early effects of a wildfire on the diversity and production of fungal communities in Mediterranean vegetation types dominated by ''Cistus ladanifer'' and ''Pinus pinaster'' in Spain |journal=Forest Ecology and Management |volume=225 |issue=1–3 |pages=296–305|doi=10.1016/j.foreco.2006.01.006}}</ref>


===Mycorrhizal associations===
===Mycorrhizal associations===
''Boletus edulis'' is [[mycorrhiza]]l—it is in a [[Mutualism (biology)|mutualistic]] relationship with the roots of plants (hosts), in which the fungus exchanges [[nitrogen]] and other nutrients extracted from the environment for [[carbon fixation|fixed carbon]] from the host. Other benefits for the plant are evident: in the case of the [[Castanea mollissima|Chinese chestnut]], the formation of mycorrhizae with ''B.&nbsp;edulis'' increases the ability of plant seedlings to resist [[water stress]], and increases leaf [[succulent plants|succulence]], leaf area, and water-holding ability.<ref>{{cite journal |author=Quan L, Lei Z-P. |year=2000 |title=A study on effect of ectomycorrhizae on promoting ''Castanea mollissima'' resistance to drought and its mechanism |journal=Forest Research |volume=13 |issue=3 |pages=249–56 |issn=1001-1498|language=Chinese}}</ref> The fungus forms a sheath of tissue around terminal, nutrient-absorbing root tips, often inducing a high degree of branching in the tips of the host, and penetrating into the root tissue, forming, to some mycologists, the defining feature of ectomycorrhizal relationships, a [[hartig net]].<ref name="ReferenceA">Smith, Sally; Read, David. Mycorrhizal Symbiosis. Academic press, 1996</ref> The ectomycorrhizal fungi are then able to exchange nutrients with the plant, effectively expanding the root system of the host plant to the furthest reaches of the symbiont fungi.<ref name="ReferenceA"/> Compatible hosts may belong to multiple families of [[vascular plant]]s that are widely distributed throughout the Northern Hemisphere; according to one 1995 estimate, there are at least 30 host plant species distributed over more than 15 genera.<ref name=Wang1995/> Examples of mycorrhizal associates include [[Pinus massoniana|Chinese red pine]],<ref>{{cite journal |author= Fu S-C, Tan Q, Chen MJ, Shang X-D, Cai LY, Zhang M-Y. |year=2009 |title=Mycorrhizal synthesis involving ''Boletus edulis'' and ''Pinus massoniana''|journal= Acta Edulis Fungi |volume=16 |issue=1 |pages=31–41 |issn=1005-9873}}</ref> [[Pinus patula|Mexican weeping pine]],<ref>{{cite journal |author=Gross E, Thomazini-Casagrande LI, Caetano FH. |year=1998 |title=A scanning electron microscopy study of ectomycorrhizae on ''Pinus patula'' Schiede and Deppe |journal=Naturalia (Rio Claro) |volume=23 |issue=1 |pages=93–101 |issn=0101-1944 }}</ref> [[Pinus sylvestris|Scots pine]], [[Picea abies|Norway spruce]],<ref>{{cite journal |author=Ceruti A, Tozzi M, Reitano G |year=1987–88 |title=Micorrize di sintesi tra ''Boletus edulis'', ''Pinus sylvestris'' e ''Picea excelsa''|trans_title=Mycorrhizal synthesis between ''Boletus edulis'', ''Pinus sylvestris'' and ''Picea excelsa''|journal=Allionia (Turin) |volume=28 |pages=117–24|issn=0065-6429|language=Italian}}</ref> [[Pseudotsuga menziesii|Coast Douglas-fir]],<ref>{{cite journal |author=Gobl F. |year=1977 |title=Mycorrhiza in Austrian Douglas fir stands |journal=Centralblatt fur das Gesamte Forstwesen |volume=94 |issue=4 |pages=185–94|issn=0379-5292|language=German}}</ref> [[Pinus mugo|mountain pine]],<ref>{{cite journal |author=Froidevaux L, Amiet R. |year=1975 |title= Ecto mycorrhizae endo mycorrhizae of ''Pinus mugo'' plus ''Boletus edulis'' ssp ''edulis'' and ''Pinus cembra'' plus ''Suillus variegatus'' formed in pure culture |journal=European Journal of Forest Pathology |volume=5 |issue=1 |pages=57–61 |issn=0300-1237 |doi=10.1111/j.1439-0329.1975.tb00935.x}}</ref> and [[Pinus virginiana|Virginia pine]].<ref>{{cite journal |author=Vozzo JA, Hackskaylo E. |year=1961 |title=Mycorrhizal fungi on ''Pinus virginiana'' |journal=Mycologia |volume=53 |issue=5 |pages=538–39 |doi=10.2307/3756310 |jstor=3756310}}</ref> The fungus has also been shown to associate with [[Cistus ladanifer|Gum rockrose]], a [[pioneer species|pioneer]] early stage shrub that is adapted for growth in degraded areas, such as burned forests.<ref name=Agueda2008/> These and other [[Cistus|Rockrose]] species are ecologically important as fungal reservoirs, maintaining an inoculum of mycorrhizal fungi for trees that appear later in the [[Ecological succession|forest regrowth cycle]].<ref>{{cite journal |author=Águeda B, Parladé J, de Miguel AM, Martínez-Peña F. |year=2006 |title=Characterization and identification of field ectomycorrhizae of ''Boletus edulis'' and ''Cistus ladanifer'' |journal=Mycologia |volume=98 |issue=1 |pages=23–30 |url=http://www.mycologia.org/cgi/content/full/98/1/23 |doi=10.3852/mycologia.98.1.23 |accessdate=2009-11-23 |pmid=16800301}}</ref>
''Boletus edulis'' is [[mycorrhiza]]l—it is in a [[Mutualism (biology)|mutualistic]] relationship with the roots of plants (hosts), in which the fungus exchanges [[nitrogen]] and other nutrients extracted from the environment for [[carbon fixation|fixed carbon]] from the host. Other benefits for the plant are evident: in the case of the [[Castanea mollissima|Chinese chestnut]], the formation of mycorrhizae with ''B.&nbsp;edulis'' increases the ability of plant seedlings to resist [[moisture stress|water stress]], and increases leaf [[succulent plants|succulence]], leaf area, and water-holding ability.<ref>{{cite journal |vauthors=Quan L, Lei ZP |year=2000 |title=A study on effect of ectomycorrhizae on promoting ''Castanea mollissima'' resistance to drought and its mechanism |journal=Forest Research |volume=13 |issue=3 |pages=249–56 |issn=1001-1498|language=zh}}</ref> The fungus forms a sheath of tissue around terminal, nutrient-absorbing root tips, often inducing a high degree of branching in the tips of the host, and penetrating into the root tissue, forming, to some mycologists, the defining feature of ectomycorrhizal relationships, a [[hartig net]].<ref name="ReferenceA">Smith, Sally; Read, David. Mycorrhizal Symbiosis. Academic press, 1996</ref> The ectomycorrhizal fungi are then able to exchange nutrients with the plant, effectively expanding the root system of the host plant to the furthest reaches of the symbiont fungi.<ref name="ReferenceA"/> Compatible hosts may belong to multiple families of [[vascular plant]]s that are widely distributed throughout the Northern Hemisphere; according to one 1995 estimate, there are at least 30 host plant species distributed over more than 15 genera.<ref name=Wang1995/> Examples of mycorrhizal associates include [[Pinus massoniana|Chinese red pine]],<ref>{{cite journal |vauthors=Fu SC, Tan Q, Chen MJ, Shang XD, Cai LY, Zhang MY |year=2009 |title=Mycorrhizal synthesis involving ''Boletus edulis'' and ''Pinus massoniana''|journal= Acta Edulis Fungi |volume=16 |issue=1 |pages=31–41 |issn=1005-9873}}</ref> [[Pinus patula|Mexican weeping pine]],<ref>{{cite journal |vauthors=Gross E, Thomazini-Casagrande LI, Caetano FH |year=1998 |title=A scanning electron microscopy study of ectomycorrhizae on ''Pinus patula'' Schiede and Deppe |journal=Naturalia (Rio Claro) |volume=23 |issue=1 |pages=93–101 |issn=0101-1944 }}</ref> [[Scots pine]], [[Picea abies|Norway spruce]],<ref>{{cite journal |vauthors=Ceruti A, Tozzi M, Reitano G |year=1987–88 |title=Micorrize di sintesi tra ''Boletus edulis'', ''Pinus sylvestris'' e ''Picea excelsa''|trans-title=Mycorrhizal synthesis between ''Boletus edulis'', ''Pinus sylvestris'' and ''Picea excelsa''|journal=Allionia (Turin) |volume=28 |pages=117–24|issn=0065-6429|language=it}}</ref> [[Pseudotsuga menziesii|Coast Douglas-fir]],<ref>{{cite journal |author=Gobl F. |year=1977 |title=Mycorrhiza in Austrian Douglas fir stands |journal=Centralblatt für das Gesamte Forstwesen |volume=94 |issue=4 |pages=185–94|issn=0379-5292|language=de}}</ref> [[Pinus mugo|mountain pine]],<ref>{{cite journal |vauthors=Froidevaux L, Amiet R |year=1975 |title= Ecto mycorrhizae endo mycorrhizae of ''Pinus mugo'' plus ''Boletus edulis'' ssp ''edulis'' and ''Pinus cembra'' plus ''Suillus variegatus'' formed in pure culture |journal=European Journal of Forest Pathology |volume=5 |issue=1 |pages=57–61 |issn=0300-1237 |doi=10.1111/j.1439-0329.1975.tb00935.x}}</ref> and [[Pinus virginiana|Virginia pine]].<ref>{{cite journal |vauthors=Vozzo JA, Hackskaylo E |year=1961 |title=Mycorrhizal fungi on ''Pinus virginiana'' |journal=Mycologia |volume=53 |issue=5 |pages=538–39 |doi=10.2307/3756310 |jstor=3756310}}</ref> The fungus has also been shown to associate with [[Cistus ladanifer|gum rockrose]], a [[pioneer species|pioneer]] early stage shrub that is adapted for growth in degraded areas, such as burned forests.<ref name=Agueda2008/> These and other [[Cistus|rockrose]] species are ecologically important as fungal reservoirs, maintaining an inoculum of mycorrhizal fungi for trees that appear later in the [[Ecological succession|forest regrowth cycle]].<ref>{{cite journal |vauthors=Águeda B, Parladé J, de Miguel AM, Martínez-Peña F |year=2006 |title=Characterization and identification of field ectomycorrhizae of ''Boletus edulis'' and ''Cistus ladanifer'' |journal=Mycologia |volume=98 |issue=1 |pages=23–30 |doi=10.3852/mycologia.98.1.23 |pmid=16800301|hdl=10171/18758 |hdl-access=free }}</ref>


The mushroom has been noted to commonly co-occur with ''[[Amanita muscaria]]'' or ''[[Amanita rubescens|A.&nbsp;rubescens]]'', although it is unclear whether this is due to a biological association between the species, or because of similarities in growing season, habitat, and ecological requirements.<ref name=Hall1998/> An association has also been reported between ''B.&nbsp;edulis'' and ''[[Amanita excelsa var. spissa|Amanita excelsa]]'' on ''[[Pinus radiata]]'' ectomycorrhizae in New Zealand, suggesting that other fungi may influence the life cycle of porcini.<ref>{{cite journal |author=Hall IR, Wang Y, Amicucci A. |year=2003 |title=Cultivation of edible ectomycorrhizal mushrooms |journal=Trends in Biotechnology |volume=21 |issue=10|pages=433–38 |doi=10.1016/S0167-7799(03)00204-X|pmid=14512229}}</ref> A 2007 field study revealed little correlation between the abundance of fruit bodies and presence of its mycelia below ground, even when soil samples were taken from directly beneath the mushroom; the study concluded that the triggers leading to formation of mycorrhizae and production of the fruit bodies were more complex.<ref>{{cite journal |author=Peintner U, Iotti M, Klotz P, Bonuso E, Zambonelli A. |year=2007 |title=Soil fungal communities in a ''Castanea sativa'' (chestnut) forest producing large quantities of ''Boletus edulis sensu lato'' (porcini): where is the mycelium of porcini? |journal=Environmental Microbiology |volume=9 |issue=4 |pages=880–89 |doi=10.1111/j.1462-2920.2006.01208.x|pmid=17359260}}</ref>
The mushroom has been noted to often co-occur with ''[[Amanita muscaria]]'' or ''[[Amanita rubescens|A.&nbsp;rubescens]]'', although it is unclear whether this is due to a biological association between the species, or because of similarities in growing season, habitat, and ecological requirements.<ref name=Hall1998/> An association has also been reported between ''B.&nbsp;edulis'' and ''[[Amanita excelsa var. spissa|Amanita excelsa]]'' on ''[[Pinus radiata]]'' ectomycorrhizae in New Zealand, suggesting that other fungi may influence the life cycle of porcini.<ref>{{cite journal |vauthors=Hall IR, Wang Y, Amicucci A |year=2003 |title=Cultivation of edible ectomycorrhizal mushrooms |journal=Trends in Biotechnology |volume=21 |issue=10|pages=433–38 |doi=10.1016/S0167-7799(03)00204-X|pmid=14512229}}</ref> A 2007 field study revealed little correlation between the abundance of fruit bodies and presence of its mycelia below ground, even when soil samples were taken from directly beneath the mushroom; the study concluded that the triggers leading to formation of mycorrhizae and production of the fruit bodies were more complex.<ref>{{cite journal |vauthors=Peintner U, Iotti M, Klotz P, Bonuso E, Zambonelli A |year=2007 |title=Soil fungal communities in a ''Castanea sativa'' (chestnut) forest producing large quantities of ''Boletus edulis sensu lato'' (porcini): where is the mycelium of porcini? |journal=Environmental Microbiology |volume=9 |issue=4 |pages=880–89 |doi=10.1111/j.1462-2920.2006.01208.x|pmid=17359260}}</ref>


===Heavy metal contamination===
===Heavy-metal contamination===
''Boletus edulis'' is known to be able to tolerate and even thrive on soil that is contaminated with toxic [[heavy metal (chemistry)|heavy metals]], such as soil that might be found near [[smelting|metal smelters]]. The mushroom's resistance to heavy metal toxicity is conferred by a biochemical called a [[phytochelatin]]—an [[oligopeptide]] whose production is induced after exposure to metal.<ref>{{cite journal |author=Collin-Hansen C, Pedersen SA, Andersen RA, Steinnes E. |year=2007 |title=First report of phytochelatins in a mushroom: induction of phytochelatins by metal exposure in ''Boletus edulis'' |journal=Mycologia |volume=99 |issue=2 |pages=161–74|doi=10.3852/mycologia.99.2.161 |pmid=17682769}}</ref> Phytochelatins are [[chelation|chelating agents]], capable of forming [[coordination complex|multiple bonds]] with the metal; in this state, the metal cannot normally react with other elements or ions and is stored in a detoxified form in the mushroom tissue.
''Boletus edulis'' is known to be able to tolerate and even thrive on soil that is contaminated with toxic heavy metals, such as soil that might be found near [[smelting|metal smelters]]. The mushroom's resistance to heavy-[[metal toxicity]] is conferred by a biochemical called a [[phytochelatin]]—an [[oligopeptide]] whose production is induced after exposure to metal.<ref>{{cite journal |vauthors=Collin-Hansen C, Pedersen SA, Andersen RA, Steinnes E |year=2007 |title=First report of phytochelatins in a mushroom: induction of phytochelatins by metal exposure in ''Boletus edulis'' |journal=Mycologia |volume=99 |issue=2 |pages=161–74|doi=10.3852/mycologia.99.2.161 |pmid=17682769}}</ref> Phytochelatins are [[chelation|chelating agents]], capable of forming [[coordination complex|multiple bonds]] with the metal; in this state, the metal cannot normally react with other elements or ions and is stored in a detoxified form in the mushroom tissue.{{Cn|date=December 2024}}


===Pests and predators===
===Pests and predators===
The fruit bodies of ''B.&nbsp;edulis'' can be infected by the parasitic mould-like fungus ''[[Hypomyces chrysospermus]]'', known as the bolete eater, which manifests itself as a white, yellow, or reddish-brown cottony layer over the surface of the mushroom.<ref name="urlHypomyces chrysospermus (MushroomExpert.Com)">{{cite web |url=http://www.mushroomexpert.com/hypomyces_chrysospermus.html |title=''Hypomyces chrysospermus'' |author=Kuo M |work= MushroomExpert.Com|accessdate=2009-11-02}}</ref> Some reported cases of [[Abdominal pain|stomach ache]] following consumption of dried porcini have been attributed to the presence of this mould on the fruit bodies.<ref>{{cite journal |author=Lunghini D, Onofri S, Zucconi L. |year=1984 |title= Some cases of intoxication probably caused by ''Sepedonium'' spp. infecting fruiting-bodies of some species of ''Boletus'' |journal=Micologia Italiana |volume=13 |issue=1 |pages=1/37–1/38 |issn=0390-0460|language=Italian}}</ref> The mushroom is also used as a food source by several species of mushroom flies,<ref name=Hall1998/> as well as other insects and their [[larvae]].<ref>{{cite book |title=Fungus-Insect Relationships: Perspectives in Ecology and Evolution |author=Bruns TD. |editor= Wheeler Q, Blackwell M|chapter=Insect mycophagy in the Boletales: fungivore diversity and the mushroom habitat |year=1984 |publisher=Columbia University Press |location=New York, NY |isbn=978-0-231-05695-3|page= |pages=91–129}}</ref> An unidentified species of [[virus]] was reported to have infected specimens found in the Netherlands and in Italy; fruit bodies affected by the virus had relatively thick stems and small or no caps, leading to the name "little-cap disease".<ref>{{cite journal |author=Huttinga H, Wichers HJ, Dieleman van Zaayen A. |year=1975 |title=Filamentous and polyhedral virus-like particle in ''Boletus edulis'' |journal=Netherlands Journal of Plant Pathology |volume=81 |pages=102–106|url=http://www.springerlink.com/content/t104136x1388073k/fulltext.pdf?page=1|format=PDF|accessdate=2009-11-02 |doi=10.1007/BF01999860 |issue=3}}</ref><ref>{{cite journal |author=Pisi A, Bellardi ME, Bernicchia A. |year=1988 |title=Virus-like particles in ''Boletus edulis'' Bull. ex Fr. in Italy |journal=Phytopathologia Mediterranea |volume=27 |issue=2 |pages=115–18 |issn=0031-9465|url=http://www.mycologia.org/cgi/reprint/94/5/757.pdf|format=PDF|accessdate=2009-11-03}}</ref>
The fruit bodies of ''B.&nbsp;edulis'' can be infected by the parasitic mould-like fungus ''[[Hypomyces chrysospermus]]'', known as the bolete eater, which manifests itself as a white, yellow, or reddish-brown cottony layer over the surface of the mushroom.<ref name="urlHypomyces chrysospermus (MushroomExpert.Com)">{{cite web |url=http://www.mushroomexpert.com/hypomyces_chrysospermus.html |title=''Hypomyces chrysospermus'' |author=Kuo M |work= MushroomExpert.Com|access-date=2009-11-02}}</ref> Some reported cases of [[Abdominal pain|stomach ache]] following consumption of dried porcini have been attributed to the presence of this mould on the fruit bodies.<ref>{{cite journal |vauthors=Lunghini D, Onofri S, Zucconi L |year=1984 |title= Some cases of intoxication probably caused by ''Sepedonium'' spp. infecting fruiting-bodies of some species of ''Boletus'' |journal=Micologia Italiana |volume=13 |issue=1 |pages=1/37–1/38 |issn=0390-0460|language=it}}</ref> The mushroom is also used as a food source by several species of mushroom flies,<ref name=Hall1998/> as well as other insects and their [[larvae]].<ref>{{cite book |title=Fungus-Insect Relationships: Perspectives in Ecology and Evolution |author=Bruns TD. |veditors=Wheeler Q, Blackwell M |chapter=Insect mycophagy in the Boletales: fungivore diversity and the mushroom habitat |year=1984 |publisher=Columbia University Press |location=New York, NY |isbn=978-0-231-05695-3|pages=91–129}}</ref> An unidentified species of virus was reported to have infected specimens found in the Netherlands and in Italy; fruit bodies affected by the virus had relatively thick stems and small or no caps, leading to the name "little-cap disease".<ref>{{cite journal |vauthors=Huttinga H, Wichers HJ, Dieleman van Zaayen A |year=1975 |title=Filamentous and polyhedral virus-like particle in ''Boletus edulis'' |journal=Netherlands Journal of Plant Pathology |volume=81 |pages=102–106|doi=10.1007/BF01999860 |issue=3|s2cid=35794592 }}</ref><ref>{{cite journal |vauthors=Pisi A, Bellardi ME, Bernicchia A |year=1988 |title=Virus-like particles in ''Boletus edulis'' Bull. ex Fr. in Italy |journal=Phytopathologia Mediterranea |volume=27 |issue=2 |pages=115–18 |issn=0031-9465|url=http://www.mycologia.org/cgi/reprint/94/5/757.pdf|access-date=2009-11-03}}</ref>


''Boletus edulis'' is a food source for animals such as the [[banana slug]] ''(Ariolimax columbianus)'',<ref>{{cite journal |author=Keller HW, Snell KL. |year=2002 |title=Feeding activities of slugs on Myxomycetes and macrofungi |journal=Mycologia |volume=94 |issue=5 |pages=757–60 |doi= 10.2307/3761690 |jstor=3761690 |pmid=21156549}}</ref> the [[long-haired grass mouse]],<ref>{{cite journal |author=Bozinovic F, Muñoz-Pedreros A. |year=1995 |title=Nutritional ecology and digestive responses of an omnivorous-insectivorous rodent (''Abrothrix longipilis'') feeding on fungus |journal=Physiological Zoology |volume=68 |issue=3 |pages=474–89}}</ref> the [[red squirrel]],<ref>{{cite journal |author=Grönwall O, Pehrson Å. |year=1984 |title=Nutrient content in fungi as a primary food of the Red Squirrel ''Sciurus vulgaris'' L |journal=Oecologia |volume=64 |issue=2 |pages=230–31 |doi= 10.1007/BF00376875}}</ref> and, as noted in one isolated report, the [[fox sparrow]].<ref>{{cite journal |author=Schiller AM, Larson KW. |year=2006 |title=Fox Sparrow foraging on a king bolete mushroom |journal=Northwestern Naturalist |volume=87 |issue=3 |page=252 |doi=10.1898/1051-1733(2006)87[252:FSFOAK]2.0.CO;2 |issn=1051-1733}}</ref>
''Boletus edulis'' is a food source for animals such as the [[banana slug]] ''(Ariolimax columbianus)'',<ref>{{cite journal |vauthors=Keller HW, Snell KL |year=2002 |title=Feeding activities of slugs on Myxomycetes and macrofungi |journal=Mycologia |volume=94 |issue=5 |pages=757–60 |doi= 10.2307/3761690 |jstor=3761690 |pmid=21156549}}</ref> the [[long-haired grass mouse]],<ref>{{cite journal |vauthors=Bozinovic F, Muñoz-Pedreros A |year=1995 |title=Nutritional ecology and digestive responses of an omnivorous-insectivorous rodent (''Abrothrix longipilis'') feeding on fungus |journal=Physiological Zoology |volume=68 |issue=3 |pages=474–89|doi=10.1086/physzool.68.3.30163780 |s2cid=87835970 }}</ref> the [[red squirrel]],<ref>{{cite journal |vauthors=Grönwall O, Pehrson Å |year=1984 |title=Nutrient content in fungi as a primary food of the Red Squirrel ''Sciurus vulgaris'' L |journal=Oecologia |volume=64 |issue=2 |pages=230–31 |doi= 10.1007/BF00376875|pmid=28312343 |bibcode=1984Oecol..64..230G |s2cid=28125328 }}</ref> and, as noted in one isolated report, the [[fox sparrow]].<ref>{{cite journal |vauthors=Schiller AM, Larson KW |year=2006 |title=Fox Sparrow foraging on a king bolete mushroom |journal=Northwestern Naturalist |volume=87 |issue=3 |page=252 |doi=10.1898/1051-1733(2006)87[252:FSFOAK]2.0.CO;2 |s2cid=86235892 |issn=1051-1733}}</ref>


==Culinary uses==
==Culinary uses==
[[File:Porcini mushroom soup in breadbowl poland 2010.JPG|thumb|left|A porcini mushroom and noodle soup served in a bread bowl at a Polish restaurant]]
''Boletus edulis'', as its name implies, is an [[edible mushroom]]. Italian chef and restaurateur [[Antonio Carluccio]] has described it as representing "the wild mushroom ''par excellence''", and hails it as the most rewarding of all fungi in the kitchen for its taste and versatility.<ref name = "Carluccio03"/> Considered a choice edible, particularly in France, Germany and Italy,<ref name=Zeitl76/> it was widely written about by the Roman writers [[Pliny the Elder]] and [[Martial]], although ranked below the esteemed ''[[Amanita caesarea]]''.
[[File:Penny bun sandwich at Nordiska Museet.jpg|thumb|left|A porcini mushroom sandwich in Stockholm, Sweden]]
{{quote|''sunt tibi boleti; fungos ego sumo suillos'' (Ep. iii. 60)<br />("You eat the choice boletus, I have mushrooms that swine grub up.")<ref>{{cite journal |author=Morford M. |year=1977 |title=Juvenal's Fifth Satire |journal=The American Journal of Philology |volume=98 |issue=3 |pages=219–45 |doi=10.2307/293772 |jstor=293772}}</ref>}}
wrote the disgruntled Martial when served ''suilli'' instead of ''boleti''.<ref>{{cite book |author=Brothwell DR, Brothwell P. |title=Food in Antiquity: a Survey of the Diet of Early Peoples |year=1998 |origyear=1969|page=92|publisher=The Johns Hopkins University Press|location=Baltimore, MD|isbn=0-8018-5740-6}}</ref> The term ''suilli'' was also thought to encompass the related ''[[Leccinum scabrum]]''.<ref>{{cite book|author=Rolfe RT, Rolfe EW.|title=The Romance of the Fungus World|publisher=Dover|location=New York|page=287|isbn=0-486-23105-4|year=1974}}</ref>
''Boletus edulis'', as the species [[Botanical name#Components of plant names|epithet]] ''edulis'' ({{langx|la|edible}}) indicates, is an edible mushroom. Italian chef and restaurateur [[Antonio Carluccio]] has described it as representing "the wild mushroom ''par excellence''", and hails it as the most rewarding of all fungi in the kitchen for its taste and versatility.<ref name = "Carluccio03"/> Considered a choice edible, particularly in France, Germany, Poland and Italy,<ref name=Zeitl76/> it was widely written about by the Roman writers [[Pliny the Elder]] and Martial, although ranked below the esteemed ''[[Amanita caesarea]]''. When he was served ''suilli''{{efn|The term ''suilli'' was also thought to encompass the related ''[[Leccinum scabrum]]''.<ref>{{cite book|vauthors=Rolfe RT, Rolfe EW |title=The Romance of the Fungus World|publisher=Dover|location=New York|page=287|isbn=0-486-23105-4|year=1974}}</ref>}} instead of ''boleti'',<ref>{{cite book |vauthors=Brothwell DR, Brothwell P |title=Food in Antiquity: a Survey of the Diet of Early Peoples |year=1998 |orig-date=1969|page=92|publisher=The Johns Hopkins University Press|location=Baltimore, MD|isbn=0-8018-5740-6}}</ref> the disgruntled Martial wrote: {{blockquote|''sunt tibi boleti; fungos ego sumo suillos'' (Ep. iii. 60)<br />("You eat the choice boletus, I have mushrooms that swine grub up.")<ref>{{cite journal |author=Morford M. |year=1977 |title=Juvenal's Fifth Satire |journal=The American Journal of Philology |volume=98 |issue=3 |pages=219–45 |doi=10.2307/293772 |jstor=293772}}</ref>}}


The flavour has been described as nutty and slightly meaty, with a smooth, creamy texture, and a distinctive aroma reminiscent of [[sourdough]]. Young, small porcini are most appreciated by gourmets, as the large ones often harbour [[maggot]]s (insect larvae), and become slimy, soft and less tasty with age. Fruit bodies are collected by holding the stipe near the base and twisting gently. Cutting the stipe with a knife may risk the part left behind rotting and the [[mycelium]] being destroyed. Peeling and washing are not recommended.<ref name="Carluccio03" /> The fruit bodies are highly [[decomposition|perishable]], due largely to the high water content (around 90%), the high level of [[enzyme]] activity, and the presence of a flora of microorganisms.<ref name=Jaworska2008>{{cite journal |author=Jaworska G, Bernaś E. |year=2008 |title=The effect of preliminary processing and period of storage on the quality of frozen ''Boletus edulis'' (Bull: Fr.) mushrooms |journal=Food Chemistry |volume=113 |issue=4 |pages=936–43 |doi=10.1016/j.foodchem.2008.08.023}}</ref> Caution should be exercised when collecting specimens from potentially polluted or [[Soil contamination|contaminated]] sites, as several studies have shown that the fruit bodies can [[bioaccumulation|bioaccumulate]] toxic [[heavy metal (chemistry)|heavy metals]] like [[mercury (element)|mercury]],<ref>{{cite journal |author=Melgar MJ, Alonson J, Garcia MA. |year=2009 |title=Mercury in edible mushrooms and underlying soil: Bioconcentration factors and toxicological risk |journal=Science of the Total Environment |volume=407 |issue=1 |pages=5328–34 |doi=10.1016/j.scitotenv.2009.07.001|pmid=19631362}}</ref> [[cadmium]],<ref>{{cite journal |author= Collin-Hansen C, Yttri KE, Andersen RA, Berthelsen BO, Steinnes E.|year=2002 |title=Mushrooms from two metal-contaminated areas in Norway: occurrence of metals and metallothionein-like proteins |journal=Geochemistry: Exploration, Environment, Analysis |volume=2 |issue=2 |pages= 121–30|doi=10.1144/1467-787302-015}}</ref> [[caesium]] and [[polonium]].<ref>{{cite journal |author=Collin-Hansen C, Andersen RA, Steinnes E. |year=2005 |title=Molecular defense systems are expressed in the king bolete (''Boletus edulis'') growing near metal smelters |journal=Mycologia |volume=97 |issue=5|pages=973–83|doi=10.3852/mycologia.97.5.973 |pmid=16596949 }}</ref><ref>{{cite journal |author=Collin-Hansen C, Andersen RA, Steinnes E. |year=2005 |title=Damage to DNA and lipids in ''Boletus edulis'' exposed to heavy metals |journal=Mycological Research |volume=109 |issue=12 |pages=1386–96 |doi= 10.1017/S0953756205004016|pmid=16353638}}</ref> Bioaccumulated metals or [[radionuclide|radioactive fission decay products]] are like chemical signatures: [[analytical chemistry|chemical]] and [[radiochemistry|radiochemical]] analysis can be used to identify the origin of imported specimens,<ref>{{cite journal |author=Marzano FN, Bracchi PG, Pizzetti P. |year=2001 |title=Radioactive and conventional pollutants accumulated by edible mushrooms (''Boletus'' sp.) are useful indicators of species origin |journal=Environmental Research |pmid=11237515 |volume=85 |issue=3 |pages=260–64 |doi=10.1006/enrs.2001.4233}}</ref> and for long-term [[radioecology|radioecological]] monitoring of polluted areas.<ref>{{cite journal |author=Grodzinskaya AA, Berreck M, Haselwandter K, Wasser SP. |year=2003 |title=Radiocesium contamination of wild-growing medicinal mushrooms in Ukraine |journal=International Journal of Medicinal Mushrooms |volume=5 |issue=1 |pages=61–86 |issn=1521-9437 |doi=10.1615/InterJMedicMush.v5.i1.90}}</ref>
The flavour of porcini has been described as nutty and slightly meaty, with a smooth, creamy texture, and a distinctive aroma reminiscent of [[sourdough]]. Young, small porcini are most appreciated, as the large ones often harbour [[maggot]]s (insect larvae), and become slimy, soft and less tasty with age. The fruit bodies are collected by holding the stipe near the base and twisting gently. Cutting the stipe with a knife may risk the part left behind rotting and the mycelium being destroyed. Peeling and washing are not recommended.<ref name="Carluccio03" /> The fruit bodies are highly [[decomposition|perishable]], due largely to the high water content (around 90%), the high level of [[enzyme]] activity, and the presence of a flora of microorganisms.<ref name=Jaworska2008>{{cite journal |vauthors=Jaworska G, Bernaś E |year=2008 |title=The effect of preliminary processing and period of storage on the quality of frozen ''Boletus edulis'' (Bull: Fr.) mushrooms |journal=Food Chemistry |volume=113 |issue=4 |pages=936–43 |doi=10.1016/j.foodchem.2008.08.023}}</ref> Caution should be exercised when collecting specimens from potentially polluted or [[Soil contamination|contaminated]] sites, as several studies have shown that the fruit bodies can [[bioaccumulation|bioaccumulate]] toxic heavy metals like [[mercury (element)|mercury]],<ref>{{cite journal |vauthors=Melgar MJ, Alonson J, Garcia MA |year=2009 |title=Mercury in edible mushrooms and underlying soil: Bioconcentration factors and toxicological risk |journal=Science of the Total Environment |volume=407 |issue=1 |pages=5328–34 |doi=10.1016/j.scitotenv.2009.07.001|pmid=19631362|bibcode=2009ScTEn.407.5328M }}</ref> [[cadmium]],<ref>{{cite journal |vauthors=Collin-Hansen C, Yttri KE, Andersen RA, Berthelsen BO, Steinnes E |year=2002 |title=Mushrooms from two metal-contaminated areas in Norway: occurrence of metals and metallothionein-like proteins |journal=Geochemistry: Exploration, Environment, Analysis |volume=2 |issue=2 |pages= 121–30|doi=10.1144/1467-787302-015|bibcode=2002GEEA....2..121C |s2cid=129662148 }}</ref> [[caesium]] and [[polonium]].<ref>{{cite journal |vauthors=Collin-Hansen C, Andersen RA, Steinnes E |year=2005 |title=Molecular defense systems are expressed in the king bolete (''Boletus edulis'') growing near metal smelters |journal=Mycologia |volume=97 |issue=5|pages=973–83|doi=10.3852/mycologia.97.5.973 |pmid=16596949 }}</ref><ref>{{cite journal |vauthors=Collin-Hansen C, Andersen RA, Steinnes E |year=2005 |title=Damage to DNA and lipids in ''Boletus edulis'' exposed to heavy metals |journal=Mycological Research |volume=109 |issue=12 |pages=1386–96 |doi= 10.1017/S0953756205004016|pmid=16353638}}</ref> Bioaccumulated metals or [[radionuclide|radioactive fission decay products]] are like chemical signatures: [[analytical chemistry|chemical]] and [[radiochemistry|radiochemical]] analysis can be used to identify the origin of imported specimens,<ref>{{cite journal |vauthors=Marzano FN, Bracchi PG, Pizzetti P |year=2001 |title=Radioactive and conventional pollutants accumulated by edible mushrooms (''Boletus'' sp.) are useful indicators of species origin |journal=Environmental Research |pmid=11237515 |volume=85 |issue=3 |pages=260–64 |doi=10.1006/enrs.2001.4233|bibcode=2001ER.....85..260M }}</ref> and for long-term [[radioecology|radioecological]] monitoring of polluted areas.<ref>{{cite journal |vauthors=Grodzinskaya AA, Berreck M, Haselwandter K, Wasser SP |year=2003 |title=Radiocesium contamination of wild-growing medicinal mushrooms in Ukraine |journal=International Journal of Medicinal Mushrooms |volume=5 |issue=1 |pages=61–86 |issn=1521-9437 |doi=10.1615/InterJMedicMush.v5.i1.90}}</ref>


Porcini are sold fresh in markets in summer and autumn in Europe and Russia, and dried or canned at other times of the year, and distributed worldwide to countries where they are not otherwise found.<ref name=Lamaison05>{{cite book |title=The Great Encyclopedia of Mushrooms |vauthors=Lamaison JL, Polese JM |year=2005 |publisher=Könemann |location=Koln, Germany |isbn=3-8331-1239-5|page=28}}</ref> They are eaten and enjoyed raw, sautéed with butter, ground into pasta, in soups, and in many other dishes. In France, they are used in recipes such as ''cèpes à la Bordelaise'', ''cèpe frits'' and ''cèpe aux tomates''.<ref>{{cite journal |author=Ramsbottom J. |year=1947 |title=Some edible fungi |journal=The British Medical Journal |volume=2 |issue=4250 |pages=304–305 |doi=10.1136/bmj.2.4520.304|pmid=20257565 |pmc=2055419 }}</ref> Porcini [[risotto]] is a traditional Italian autumn dish.<ref>Carluccio, p. 166.</ref> Porcini are a feature of many cuisines, including [[Provence|Provençal]],<ref>{{cite book |author=Olney R. |title= A Provençal Table|year=1995 |pages=31–32|publisher=Pavilion |location=London |isbn=1-85793-632-9}}</ref> and [[Vienna|Viennese]].<ref>{{cite book |author=Philpot R. |title= Viennese Cookery|year=1965 |pages=139–40|publisher=Hodder & Stoughton |location=London }}</ref> In [[Thailand]] they are used in soups and consumed blanched in salads.<ref name=Solomon>{{cite book |title=Charmaine Solomon's Encyclopedia of Asian Food |author=Solomon C. |year=1996 |publisher=William Heinemann Australia |location=Melbourne |isbn=0-85561-688-1|page=238}}</ref> Porcini can also be frozen, either while raw or after cooking in butter. The colour, aroma, and taste of porcini deteriorate noticeably after being frozen for four months. [[blanching (cooking)|Blanching]] (or soaking and blanching) as a processing step before freezing can extend the freezer life to 12 months.<ref name=Jaworska2008/> They are also one of the few species sold commercially as [[pickled mushrooms]].<ref>Carluccio, p. 99.</ref>
[[File:Porcini mushroom soup in breadbowl poland 2010.JPG|thumb|left|A porcini mushroom and noodle soup served in a bread bowl at a Polish restaurant]]
Porcini are sold fresh in markets in summer and autumn in [[Central Europe|Central]] and [[Southern Europe]], and dried or canned at other times of the year, and distributed worldwide to countries where they are not otherwise found.<ref name=Lamaison05>{{cite book |title=The Great Encyclopedia of Mushrooms |author=Lamaison J-L, Polese J-M. |year=2005 |publisher=Könemann |location=Koln, Germany |isbn=3-8331-1239-5|page=28}}</ref> They are eaten and enjoyed raw, sautéed with butter, ground into pasta, in soups, and in many other dishes. In France, they are used in recipes such as ''cèpes à la Bordelaise'', ''cèpe frits'' and ''cèpe aux tomates''.<ref>{{cite journal |author=Ramsbottom J. |year=1947 |title=Some edible fungi |journal=The British Medical Journal |volume=2 |issue=4250 |pages=304–305 |doi=10.1136/bmj.2.4520.304}}</ref> Porcini [[risotto]] is a traditional Italian autumn dish.<ref>Carluccio, p. 166.</ref> Porcini are a feature of many cuisines, including [[Provence|Provençal]],<ref>{{cite book |author=Olney R. |title= A Provençal Table|year=1995 |pages=31–32|publisher=Pavilion |location=London |isbn=1-85793-632-9}}</ref> and [[Vienna|Viennese]].<ref>{{cite book |author=Philpot R. |title= Viennese Cookery|year=1965 |pages=139–40|publisher=Hodder & Stoughton |location=London |isbn=}}</ref> They are used in soups and consumed blanched in salads in [[Thailand]].<ref name=Solomon>{{cite book |title=Charmaine Solomon's Encyclopedia of Asian Food |author=Solomon C. |year=1996 |publisher=William Heinemann Australia |location=Melbourne |isbn=0-85561-688-1|page=238}}</ref> Porcini can also be frozen — either raw or first cooked in butter. The colour, aroma, and taste of frozen porcini deteriorate noticeably if frozen longer than four months. [[blanching (cooking)|Blanching]] or soaking and blanching as a processing step before freezing can extend the freezer life up to 12 months.<ref name=Jaworska2008/> They are also one of the few mushroom species pickled and sold commercially.<ref>Carluccio, p. 99.</ref>


===Dried===
===Dried===
[[File:Pile of Porcini-2.jpg|thumb|right|alt=A heaped pile of dried sliced mushrooms on a large plate. A sign near the back of the plate reads "Fungo di Borgotaro I.G.P."; another sign at the front reads "Prezzo speciale di Fiera. 3 Etti 18,00". An electronic scale is partially visible on a table behind the mushrooms|A pile of dried porcini at the [[Borgotaro]] festival of the porcino, Italy]]
[[File:Pile of Porcini-2.jpg|thumb|right|alt=A heaped pile of dried sliced mushrooms on a large plate. A sign near the back of the plate reads "Fungo di Borgotaro I.G.P."; another sign at the front reads "Prezzo speciale di Fiera. 3 Etti 18,00". An electronic scale is partially visible on a table behind the mushrooms|A pile of dried porcini at the [[Borgotaro]] porcino festival in Italy]]
''Boletus edulis'' is well suited to drying—its flavour intensifies, it is easily reconstituted, and its resulting texture is pleasant.<ref>Carluccio, p. 96.</ref> Reconstitution is done by soaking in hot, but not boiling, water for about twenty minutes; the water used is infused with the mushroom aroma and it too can be used in subsequent cooking. Dried porcini have more protein than most other commonly consumed vegetables apart from [[soybean]]s. Some of this content is indigestible, though digestibility is improved with cooking.<ref name=Arora86>{{cite book |title=Mushrooms Demystified: A Comprehensive Guide to the Fleshy Fungi|author=Arora D.|year=1986|publisher=Ten Speed Press|location=Berkeley, California|isbn=0-89815-169-4|page=30 }}</ref>
''Boletus edulis'' is well suited to drying—its flavour intensifies, it is easily reconstituted, and its resulting texture is pleasant.<ref>Carluccio, p. 96.</ref> Reconstitution is done by soaking in hot, but not boiling, water for about twenty minutes; the water used is infused with the mushroom aroma and it too can be used in subsequent cooking. Dried porcini have more protein than most other commonly consumed vegetables, apart from [[soybean]]s. Some of their protein is indigestible, though digestibility is improved with cooking.<ref name=Arora86>{{cite book|title=Mushrooms Demystified: A Comprehensive Guide to the Fleshy Fungi|author=Arora D.|year=1986|publisher=Ten Speed Press|location=Berkeley, California|isbn=0-89815-169-4|page=[https://archive.org/details/mushroomsdemysti00aror_0/page/30 30]|url=https://archive.org/details/mushroomsdemysti00aror_0/page/30}}</ref>


Like other boletes, porcini can be dried by being strung separately on twine and hung close to the ceiling of a kitchen. Alternatively, the mushrooms can be dried by cleaning with a brush (washing is not recommended), and then placing them in a wicker basket or [[mushiki|bamboo steamer]] on top of a boiler or hot water tank.<ref name=Carluccio97>Carluccio, p. 97.</ref> Another method is drying in an oven at {{convert|25|to|30|C|F}} for two to three hours, then increasing the temperature to {{convert|50|C|F}} until crisp or brittle.<ref>{{cite book|author=Gates L, Delong D. |title=How to Dry Foods |publisher=HP Books |location=Los Angeles, California |year=1992 |page=61 |isbn=1-55788-050-6 |url=https://books.google.com/?id=aSBfsPQP_MQC&lpg=PA61&dq=cooking%20Boletus%20edulis&pg=PA61 |accessdate=2009-10-06}}</ref> Once dry, they are kept in an airtight jar.<ref name=Carluccio97/> Importantly for commercial production, porcini retain their flavour after industrial preparation in a [[pressure cooking|pressure cooker]] or after canning or bottling, and are thus useful for manufacturers of soups or stews. The addition of a few pieces of dried porcino can significantly add to flavour, and they are a major ingredient of the pasta sauce known as ''carrettiere'' (carter's sauce).<ref>{{cite book |author=Seed D. |title=The Top One Hundred Pasta Sauces |publisher=Ten Speed Press |location=Berkeley, California |year=1987 |page=25 |isbn=0-89815-232-1 |url=https://books.google.com/?id=5JfdNVZlwA0C&lpg=PA25&dq=cooking%20Boletus%20edulis&pg=PA25#v=onepage&q=cooking%20Boletus%20edulis |accessdate=2010-11-06}}</ref> The drying process is known to induce the formation of various [[volatile organic compound|volatile substances]] that contribute to the mushroom's aroma. Chemical analysis has shown that the odour of the dried mushroom is a complex mixture of 53 volatile compounds.<ref name="pmid19845295">{{cite journal |author=Misharina TA, Muhutdinova SM, Zharikova GG, Terenina MB, Krikunova NI. |title=The composition of volatile components of cepe (''Boletus edulis'') and oyster mushrooms (''Pleurotus ostreatus'') |journal=Applied Biochemistry and Microbiology |volume=45 |issue=2 |pages=187–93 |year=2009 |pmid=|doi=10.1134/S0003683809020124}}</ref>
Like other boletes, porcini can be dried by being strung separately on twine and hung close to the ceiling of a kitchen. Alternatively, the mushrooms can be dried by cleaning with a brush (washing is not recommended), and then placing them in a wicker basket or [[bamboo steamer]] on top of a boiler or hot water tank.<ref name=Carluccio97>Carluccio, p. 97.</ref> Another method is drying in an oven at {{convert|25|to|30|C|F}} for two to three hours, then increasing the temperature to {{convert|50|C|F}} until crisp or brittle.<ref>{{cite book|vauthors=Gates L, Delong D |title=How to Dry Foods |publisher=HP Books |location=Los Angeles, California |year=1992 |page=61 |isbn=1-55788-050-6 |url=https://books.google.com/books?id=aSBfsPQP_MQC&q=cooking%20Boletus%20edulis&pg=PA61 |access-date=2009-10-06}}</ref> Once dry, they must be kept in an airtight container.<ref name=Carluccio97/> Importantly for commercial production, porcini retain their flavour after industrial preparation in a [[pressure cooking|pressure cooker]] or after canning or bottling, and are thus useful for manufacturers of soups or stews. The addition of a few pieces of dried porcino can significantly add to flavour, and they are a major ingredient of the pasta sauce known as ''carrettiera'' (carter's sauce).<ref>{{cite book |author=Seed D. |title=The Top One Hundred Pasta Sauces |publisher=Ten Speed Press |location=Berkeley, California |year=1987 |page=25 |isbn=0-89815-232-1 |url=https://books.google.com/books?id=5JfdNVZlwA0C&q=cooking%20Boletus%20edulis&pg=PA25 |access-date=2010-11-06}}</ref> The drying process is known to induce the formation of various [[volatile organic compound|volatile substances]] that contribute to the mushroom's aroma. Chemical analysis has shown that the odour of the dried mushroom is a complex mixture of 53 volatile compounds.<ref name="pmid19845295">{{cite journal |vauthors=Misharina TA, Muhutdinova SM, Zharikova GG, Terenina MB, Krikunova NI |title=The composition of volatile components of cepe (''Boletus edulis'') and oyster mushrooms (''Pleurotus ostreatus'') |journal=Applied Biochemistry and Microbiology |volume=45 |issue=2 |pages=187–93 |year=2009 |pmid=19382709|doi=10.1134/S0003683809020124|s2cid=21525680 }}</ref>


===Commercial harvest===
===Commercial harvest===
[[File:Steinpilz 2006 08 3.jpg|thumb|right|alt=Approximately two dozen brown-capped, white or light-brown stemmed mushrooms of various sizes in a brown bowl.|Porcini can vary considerably in size.]]
[[File:Steinpilz 2006 08 3.jpg|thumb|right|alt=Approximately two dozen brown-capped, white or light-brown stemmed mushrooms of various sizes in a brown bowl.|Porcini can vary considerably in size.]]
A 1998 estimate suggests the total annual worldwide consumption of ''Boletus edulis'' and closely related species (''B.&nbsp;aereus'', ''B.&nbsp;pinophilus'', and ''B.&nbsp;reticulatus'') to be between 20,000 and 100,000 tons.<ref name=Hall1998/> Approximately 2,700&nbsp;tonnes (3,000&nbsp;[[Short ton|tons]]) were sold in France, Italy and Germany in 1988, according to official figures. The true amount consumed far exceeds this, as it does not account for informal sales or consumption by collectors.<ref name=Hall2003/> They are widely exported and sold in dried form, reaching countries where they do not occur naturally, such as Australia and New Zealand. The autonomous community of [[Castile and León]] in Spain produces 7,700&nbsp;tonnes (8,500&nbsp;tons) annually.<ref name=Agueda2008>{{cite journal |author=Agueda B, Parlade J, Fernandez-Toiran LM, Cisneros O, de Miguel AM, Modrego MP, Martinez-Pena F, Pera J.|year=2008 |title=Mycorrhizal synthesis between ''Boletus edulis'' species complex and rockroses (''Cistus'' sp.) |journal=Mycorrhiza |pmid=18695982 |volume=18 |issue=8 |pages=443–49 |doi=10.1007/s00572-008-0192-3}}</ref> In autumn, the price of porcini in the Northern Hemisphere typically ranges between $20 and $80&nbsp;dollars per kilogram, although in New York in 1997, the scarcity of fruit bodies elevated the wholesale price to over $200 per kilogram.<ref name=Hall2003/>
A 1998 estimate suggested that the total annual worldwide consumption of ''Boletus edulis'' and closely related species (''B.&nbsp;aereus'', ''B.&nbsp;pinophilus'', and ''B.&nbsp;reticulatus'') was between 20,000 and 100,000 tons.<ref name=Hall1998/> Approximately 2,700&nbsp;tonnes (3,000&nbsp;[[Short ton|tons]]) were sold in France, Italy and Germany in 1988, according to official figures. The true amount consumed far exceeds this, as the official sales figures did not account for informal sales or consumption by collectors.<ref name=Hall2003/> They are widely exported and sold in dried form, reaching countries where they do not occur naturally, such as Australia and New Zealand. The autonomous community of [[Castile and León]] in Spain produces 7,700&nbsp;tonnes (8,500&nbsp;tons) annually.<ref name=Agueda2008>{{cite journal |vauthors=Agueda B, Parlade J, Fernandez-Toiran LM, Cisneros O, de Miguel AM, Modrego MP, Martinez-Pena F, Pera J |year=2008 |title=Mycorrhizal synthesis between ''Boletus edulis'' species complex and rockroses (''Cistus'' sp.) |journal=Mycorrhiza |pmid=18695982 |volume=18 |issue=8 |pages=443–49 |doi=10.1007/s00572-008-0192-3|s2cid=11233267 }}</ref> In autumn, the price of porcini in the Northern Hemisphere typically ranges between $20 and $80&nbsp;per kilogram, although in New York in 1997 the wholesale price rose to more than $200 per kilogram due to scarcity.<ref name=Hall2003/>


In the vicinity of [[Borgo Val di Taro|Borgotaro]] in the [[Province of Parma]] of northern Italy, the four species ''Boletus edulis'', ''B.&nbsp;aereus'', ''B.&nbsp;aestivalis'' and ''B.&nbsp;pinophilus'' have been recognised for their superior taste and officially termed ''Fungo di Borgotaro''. Here, these mushrooms have been collected for centuries, and exported commercially. Due to the globalization of the mushroom trade, most of the porcini commercially available in Italy or exported by Italy no longer originate there. Porcini and other mushrooms are imported into Italy from various locations, especially China and eastern European countries; these are then often re-exported under the "Italian porcini" label.<ref>{{cite journal |author=Sitta N, Floriani M. |year=2008 |title=Nationalization and globalization trends in the wild mushroom commerce of Italy with emphasis on porcini (''Boletus edulis'' and allied species) |journal=Economic Botany |volume=62 |issue=3 |pages=307–22 |doi=10.1007/s12231-008-9037-4}}</ref><ref name=Boa2004>{{cite book |author= Boa E. |title=Wild Edible Fungi: A Global Overview of Their Use and Importance to People (Non-Wood Forest Products) |publisher=Food & Agriculture Organization of the UN |location=Rome, Italy |year=2004 |pages=96–97 |isbn=92-5-105157-7|url=https://books.google.com/?id=Zd2NlcNZgvcC&lpg=PA97&dq=boletus%20edulis&pg=PA97|accessdate=2009-10-30}}</ref>
In the vicinity of [[Borgo Val di Taro|Borgotaro]] in the [[Province of Parma]] of northern Italy, the four species ''Boletus edulis'', ''B.&nbsp;aereus'', ''B.&nbsp;aestivalis'' and ''B.&nbsp;pinophilus'' have been recognised for their superior taste and officially termed ''Fungo di Borgotaro''. Here these mushrooms have been collected for centuries and exported commercially. Owing to the globalisation of the mushroom trade most of the porcini commercially available in Italy or exported by Italy no longer originate there. Porcini and other mushrooms are also imported into Italy from various locations, especially China and eastern European countries; these are then often re-exported under the "Italian porcini" label.<ref>{{cite journal |vauthors=Sitta N, Floriani M |year=2008 |title=Nationalization and globalization trends in the wild mushroom commerce of Italy with emphasis on porcini (''Boletus edulis'' and allied species) |journal=Economic Botany |volume=62 |issue=3 |pages=307–22 |doi=10.1007/s12231-008-9037-4|s2cid=44274570 }}</ref><ref name=Boa2004>{{cite book |author= Boa E. |title=Wild Edible Fungi: A Global Overview of Their Use and Importance to People (Non-Wood Forest Products) |publisher=Food & Agriculture Organization of the UN |location=Rome, Italy |year=2004 |pages=96–97 |isbn=92-5-105157-7|url=https://books.google.com/books?id=Zd2NlcNZgvcC&q=boletus%20edulis&pg=PA97|access-date=2009-10-30}}</ref>


In Italy, the disconnect with local production has had an adverse effect on quality; for example, in the 1990s, some of the dried porcino mushrooms exported to Italy from China contained species of genus ''[[Tylopilus]]'', which are rather similar in appearance, and when dried, are difficult for both mushroom labourers and mycologists alike to distinguish from ''Boletus''. ''Tylopilus'' species typically have a very bitter taste, a bitterness that is imparted to the flavour of the porcini with which they are mixed.<ref>{{cite journal |author=Sitta N. |year=2000 |title=Presence of ''Tylopilus'' into dried "Porcini" mushrooms from China |journal=Micologia Italiana |volume=29 |issue=1 |pages=96–99 |issn=0390-0460|language=Italian}}</ref>
In Italy the disconnect with local production has had an adverse effect on quality; for example in the 1990s some of the dried porcino mushrooms exported to Italy from China contained species of genus ''[[Tylopilus]]'', which are rather similar in appearance and when dried are difficult for both mushroom labourers and mycologists alike to distinguish from ''Boletus''. ''Tylopilus'' species typically have a very bitter taste, which is imparted to the flavour of the porcini with which they are mixed.<ref>{{cite journal |author=Sitta N. |year=2000 |title=Presence of ''Tylopilus'' into dried "Porcini" mushrooms from China |journal=Micologia Italiana |volume=29 |issue=1 |pages=96–99 |issn=0390-0460|language=it}}</ref>


After the [[revolutions of 1989|fall of the Iron Curtain]] and the economic and political barriers that followed, central and eastern European countries with local mushroom harvesting traditions, such as Albania, Bulgaria, Macedonia, Romania, Serbia and Slovenia, developed into exporters of porcini, concentrating primarily on the Italian market.<ref name=Boa2004/> Exported porcini and other wild fungi are also destined for France, Germany and other western European markets, where demand for them exists, but collection on a commercial scale does not.<ref name=Boa2004/> Picking ''B.&nbsp;edulis'' has become an annual seasonal income earner and pastime in countries like [[Bulgaria]], especially for many [[Romani people|Roma]] communities and the [[unemployment|unemployed]].<ref name=belogradchik>{{cite web |url=http://www.belogradchik.info/1/content/view/390/2/lang,bg/ |language=Bulgarian |title=''По 1 тон гъби на ден се предават в изкупвателните пунктове в Белоградчик'' |trans_title=''One ton of mushrooms a day are being submitted to purchasing stations in Belogradchik''|author=Focus Information Agency, Bulgaria |work= Belogradchik.info|accessdate=2009-11-02}}</ref> A lack of control has led to heavy exploitation of the mushroom resource.<ref name=worldwildlife>{{cite web |url=http://www.worldwildlife.org/bsp/publications/europe/bulgaria/bulgaria1.html |title=The Macromycetes Fungi of Bulgaria |author=Drumeva M, Gyosheva M |work= World Wildlife Fund|accessdate=2009-11-27}}</ref>
After the [[revolutions of 1989|fall of the Iron Curtain]] and the subsequent reduction of economic and political barriers, central and eastern European countries with local mushroom harvesting traditions, such as Albania, Bulgaria, Macedonia, Romania, Serbia and Slovenia, developed into exporters of porcini, concentrating primarily on the Italian market.<ref name=Boa2004/> Porcini and other wild fungi from these countries are also destined for France, Germany and other western European markets, where demand for them exists but collection on a commercial scale does not.<ref name=Boa2004/> Picking ''B.&nbsp;edulis'' has become an annual seasonal income earner and pastime in countries like [[Bulgaria]], especially for many [[Romani people|Roma]] communities and the [[unemployment|unemployed]].<ref name=belogradchik>{{cite web |url=http://www.belogradchik.info/1/content/view/390/2/lang,bg/ |language=bg |title=''По 1 тон гъби на ден се предават в изкупвателните пунктове в Белоградчик'' |trans-title=One ton of mushrooms a day are being submitted to purchasing stations in Belogradchik |author=Focus Information Agency, Bulgaria |work=Belogradchik.info |access-date=2009-11-02 |archive-url=https://web.archive.org/web/20110721185827/http://www.belogradchik.info/1/content/view/390/2/lang%2Cbg/ |archive-date=2011-07-21 }}</ref> A lack of control of the harvest has led to heavy exploitation of the mushroom resource.<ref name=worldwildlife>{{cite web |url=http://www.worldwildlife.org/bsp/publications/europe/bulgaria/bulgaria1.html |title=The Macromycetes Fungi of Bulgaria |vauthors=Drumeva M, Gyosheva M |work= World Wildlife Fund|access-date=2009-11-27}}</ref>


Like many other strictly mycorrhizal fungi, ''B.&nbsp;edulis'' has to date eluded cultivation attempts.<ref name=Arora86/><ref>{{cite book |author=Chang S-T, Miles PG. |title=Mushrooms: Cultivation, Nutritional Value, Medicinal Effect, and Environmental Impact|edition=2nd |year=2004|publisher=CRC Press |location= Boca Raton, Florida|isbn=0-8493-1043-1|page=131}}</ref> The results of some studies suggest that unknown components of the soil microflora might be required for ''B.&nbsp;edulis'' to successfully establish a mycorrhizal relationship with the host plant.<ref>{{cite journal |author=Veselkov IM. |year=1975 |title=Artificial propagation of ''Boletus edulis'' in forests |journal=Растительньіе Ресурсы России|volume=11 |pages= 574–78}}</ref><ref>{{cite journal |author=Ceruti A, Tozzi M, Reitano G. |year=1988 |title=Micorize di sintesi tra ''Boletus edulis'', ''Pinus sylvestris'' e ''Picea excelsa'' |journal=Allionia (Turin)|volume=28 |pages= 117–24|language=Italian}}</ref><ref>{{cite book |author=Fitter AH, Garbaye J. |year=1994 |chapter=Interaction between mycorrhizal fungi and other organisms|title=Management of Mycorrhizas in Agriculture, Horticulture and Forestry |editor=Robson AD, Abbott LK, Malajczuk N |isbn=978-0-7923-2700-4 |publisher=Kluwer Academic Publishers |location=Dordrecht, Netherlands}}</ref>
Like many other strictly mycorrhizal fungi, ''B.&nbsp;edulis'' has eluded cultivation attempts for years.<ref name=Arora86/><ref>{{cite book |vauthors=Chang ST, Miles PG |title=Mushrooms: Cultivation, Nutritional Value, Medicinal Effect, and Environmental Impact|url=https://archive.org/details/mushroomscultiva00mile |url-access=limited |edition=2nd |year=2004|publisher=CRC Press |location= Boca Raton, Florida|isbn=0-8493-1043-1|page=[https://archive.org/details/mushroomscultiva00mile/page/n155 131]}}</ref><ref name="Wang2014" /> The results of some studies suggest that unknown components of the soil microflora might be required for ''B.&nbsp;edulis'' to establish a mycorrhizal relationship with the host plant.<ref>{{cite journal |author=Veselkov IM. |year=1975 |title=Artificial propagation of ''Boletus edulis'' in forests |journal=Растительньіе Ресурсы России|volume=11 |pages= 574–78}}</ref><ref>{{cite journal |vauthors=Ceruti A, Tozzi M, Reitano G |year=1988 |title=Micorize di sintesi tra ''Boletus edulis'', ''Pinus sylvestris'' e ''Picea excelsa'' |journal=Allionia (Turin)|volume=28 |pages= 117–24|language=it}}</ref><ref>{{cite book |vauthors=Fitter AH, Garbaye J |year=1994 |chapter=Interaction between mycorrhizal fungi and other organisms|title=Management of Mycorrhizas in Agriculture, Horticulture and Forestry |veditors=Robson AD, Abbott LK, Malajczuk N |isbn=978-0-7923-2700-4 |publisher=Kluwer Academic Publishers |location=Dordrecht, Netherlands}}</ref> Successful attempts at cultivating ''B.&nbsp;edulis'' have been made by Spanish scientists by mycorrhization of ''Cistus'' species,<ref name="Wang2014" /> with [[Pseudomonas fluorescens|''Pseudomonas&nbsp;fluorescens'']] bacteria helping the mycorrhiza.<ref name="Mediavilla2015" />


=== Nutritional composition ===
{{nutritionalvalue
{{nutritionalvalue
| name=''Boletus edulis'', fresh<ref name=Caglarlrmak2002>Nutritional values are based on [[food chemistry|chemical analysis]] of Turkish specimens, conducted by Çaglarlrmak and colleagues at the Agricultural Faculty, Food Engineering Department, [[Gaziosmanpasha University]]. Source: {{cite journal |author=Çaglarlrmak N, Ünal K, Ötles S.|year=2001 |title=Nutritional value of edible wild mushrooms collected from the Black Sea region of Turkey |journal=Micologia Aplicada International |volume= 14|issue=1 |pages=1–5 |url=http://www.micaplint.com/PDF/V14_1_01.pdf|format=PDF}}</ref>
| name=''Boletus edulis'', fresh<ref name=Caglarlrmak2002>Nutritional values are based on [[food chemistry|chemical analysis]] of Turkish specimens, conducted by Çaglarlrmak and colleagues at the Agricultural Faculty, Food Engineering Department, [[Gaziosmanpașa University]]. Source: {{cite journal |vauthors=Çaglarlrmak N, Ünal K, Ötles S |year=2001 |title=Nutritional value of edible wild mushrooms collected from the Black Sea region of Turkey |journal=Micologia Aplicada International |volume= 14|issue=1 |pages=1–5 |url=http://www.redalyc.org/pdf/685/68514101.pdf}}</ref>
| kJ=342.4
| kJ=342.4
| protein=7.39 g
| protein=7.39&nbsp;g
| fat=1.70 g
| fat=1.70&nbsp;g
| carbohydrates=9.23 g
| carbs=9.23&nbsp;g
| iron_mg=0.739
| iron_mg=0.739
| calcium_mg=1.195
| calcium_mg=1.195
Line 153: Line 175:
| riboflavin_mg=0.092
| riboflavin_mg=0.092
| niacin_mg=6.07
| niacin_mg=6.07
| right=1
| source_usda=1
| source_usda=1
}}
}}
''Boletus edulis'' constitutes a food source which, although not rich in easily absorbed [[carbohydrate]]s or [[dietary fat|fat]], contains [[vitamin]]s, [[dietary mineral|minerals]] and [[dietary fibre]]. Fresh mushrooms consist of over 80% moisture,<ref name=Ouzouni2006>{{cite journal |author=Ouzouni PK, Riganakos KA. |year=2006 |title=Nutritional value and metal content profile of Greek wild edible fungi |journal=Acta Alimentaria |volume=36 |issue=1 |pages=99–110 |doi=10.1556/AAlim.36.2007.1.11}}</ref> although reported values tend to differ somewhat as moisture content can be affected by environmental temperature and [[relative humidity]] during growth and storage, as well as the relative amount of water that may be produced as a result of normal metabolic processes during storage.<ref>{{cite book |title=The Biology and Cultivation of Edible Mushrooms |author=Crisan EV, Sands A|year=1978|editor=Chang ST, Hayes WA|chapter=Nutritional value |publisher=Academic Press |location=New York |isbn=978-0-12-168050-3|pages=727–93}}</ref>


== Nutrition==
Carbohydrates make up the bulk of the fruit bodies, comprising 9.23% of the fresh weight (see table), and 65.4% of the dry weight.<ref name=Ouzouni2006/> The carbohydrate component contains the [[monosaccharide]]s [[glucose]], [[mannitol]] and α,α-[[trehalose]], the [[polysaccharide]] [[glycogen]], and the water-insoluble structural polysaccharide [[chitin]], which accounts for up to 80–90% of dry matter in mushroom [[cell wall]]s. Chitin, [[hemicellulose]], and [[pectin]]-like carbohydrates—all indigestible by humans—contribute to the nutritionally desirable high proportion of insoluble fibre in ''B.&nbsp;edulis''.<ref name=Kalac2009/>
''Boletus edulis'' mushrooms are 9% [[carbohydrate]]s, 3% fat, and 7% [[protein]] (table). Fresh mushrooms consist of over 80% moisture,<ref name=Ouzouni2006>{{cite journal |vauthors=Ouzouni PK, Riganakos KA |year=2006 |title=Nutritional value and metal content profile of Greek wild edible fungi |journal=Acta Alimentaria |volume=36 |issue=1 |pages=99–110 |doi=10.1556/AAlim.36.2007.1.11}}</ref> although reported values tend to differ somewhat as moisture content can be affected by environmental temperature and [[relative humidity]] during growth and storage.<ref>{{cite book |title=The Biology and Cultivation of Edible Mushrooms |vauthors=Crisan EV, Sands A |year=1978|veditors=Chang ST, Hayes WA |chapter=Nutritional value |publisher=Academic Press |location=New York |isbn=978-0-12-168050-3|pages=727–93}}</ref> The carbohydrate component contains the [[monosaccharide]]s [[glucose]], [[mannitol]] and α,α-[[trehalose]], the [[polysaccharide]] [[glycogen]], and the water-insoluble structural polysaccharide [[chitin]], which accounts for up to 80–90% of dry matter in mushroom [[cell wall]]s. Chitin, [[hemicellulose]], and [[pectin]]-like carbohydrates—all indigestible by humans—contribute to the high proportion of insoluble fibre in ''B.&nbsp;edulis''.<ref name=Kalac2009>{{cite journal |author=Kalač P. |year=2009 |title=Chemical composition and nutritional value of European species of wild growing mushrooms: a review |journal=Food Chemistry |volume=113 |issue=1 |pages=9–16 |doi=10.1016/j.foodchem.2008.07.077}}</ref>


The total [[lipid]], or crude fat, content makes up 2.6% of the dry matter of the mushroom. The proportion of [[fatty acid]]s (expressed as a % of total fatty acids) are: [[palmitic acid]], 9.8%; [[stearic acid]], 2.7%; [[oleic acid]], 36.1%; [[linoleic acid]], 42.2%, and [[alpha-Linolenic acid|linolenic acid]], 0.2%.<ref>{{cite journal |author=Pedneault K, Angers P, Gosselin A, Tweddell RJ. |year=2006 |title=Fatty acid composition of lipids from mushrooms belonging to the family Boletaceae |journal=Mycological Research |volume=110|pages=1179–83 |doi=10.1016/j.mycres.2006.05.006 |pmid=16959482 |issue=Pt 10}}</ref><ref group="nb">A comparable Indian analysis produced somewhat different values: total lipids, 3.3% of dry matter; palmitic acid, 21.6%; stearic acid, 9.1%; oleic acid, 31.1%; linoleic acid, 33.8%, and linolenic acid, 1.7%. Source: Kavishree S, Hemavathy J, Lokesh BR, Shashirekha MN, Rajarathnam S. (2008). "Fat and fatty acids in Indian edible mushrooms". ''Food Chemistry'' '''106''': 597–602.</ref>
The total [[lipid]], or crude fat, content makes up 3% of the dry matter of the mushroom. The proportion of [[fatty acid]]s (expressed as a % of total fatty acids) are: [[linoleic acid]] 42%, [[oleic acid]] 36%, [[palmitic acid]] 10%, and [[stearic acid]] 3%.<ref>{{cite journal |vauthors=Pedneault K, Angers P, Gosselin A, Tweddell RJ |year=2006 |title=Fatty acid composition of lipids from mushrooms belonging to the family Boletaceae |journal=Mycological Research |volume=110|pages=1179–83 |doi=10.1016/j.mycres.2006.05.006 |pmid=16959482 |issue=Pt 10}}</ref>


A comparative study of the [[amino acid]] composition of eleven Portuguese wild edible mushroom species showed ''Boletus edulis'' to have the highest total amino acid content,<ref group="nb">Other species tested were ''[[Suillus bellinii]]'', ''[[Suillus luteus]]'', ''[[Suillus granulatus]]'', ''[[Tricholomopsis rutilans]]'', ''[[Hygrophorus agathosmus]]'', ''[[Amanita rubescens]]'', ''[[Russula cyanoxantha]]'', ''[[Tricholoma equestre]]'', ''[[Fistulina hepatica]]'', and ''[[Cantharellus cibarius]]''.</ref> about 2.3&nbsp;g per 100&nbsp;g of dried mushroom. This total includes a full complement of 20 [[essential amino acid|essential]] and nonessential amino acids.<ref name=Ribeiro2008b>{{cite journal |author= Ribeiro B, Andrade PB, Silva BM, Baptista P, Seabra RM, Valento P.|year=2008 |title=Comparative study on free amino acid composition of wild edible mushroom species |journal=Journal of Agricultural and Food Chemistry |volume=56 |issue=22 |pages=10973–79 |url=http://pubs.acs.org/doi/abs/10.1021/jf802076p |doi=10.1021/jf802076p |pmid=18942845}}</ref> Analysis of the free amino acids (that is, those not bound up in [[protein]]) revealed [[glutamine]] and [[alanine]] to be the principal amino acids (each about 25% of total compounds); a separate analysis concluded that [[lysine]] is another predominant compound.<ref>{{cite journal |author=Tsai S-Y, Tsai H-L, Mau J-L.|year=2008 |title=Non-volatile taste components of ''Agaricus blazei'', ''Agrocybe cylindracea'' and ''Boletus edulis''|journal=Food Chemistry |volume=107 |issue=3 |pages=977–83 |doi=10.1016/j.foodchem.2007.07.080}}</ref>
A comparative study of the [[amino acid]] composition of eleven Portuguese wild edible mushroom species showed ''Boletus edulis'' to have the highest total amino acid content.<ref name=Ribeiro2008b>{{cite journal |vauthors=Ribeiro B, Andrade PB, Silva BM, Baptista P, Seabra RM, Valento P |year=2008 |title=Comparative study on free amino acid composition of wild edible mushroom species |journal=Journal of Agricultural and Food Chemistry |volume=56 |issue=22 |pages=10973–79 |doi=10.1021/jf802076p |pmid=18942845}}</ref><ref>{{cite journal |vauthors=Tsai SY, Tsai HL, Mau JL |year=2008 |title=Non-volatile taste components of ''Agaricus blazei'', ''Agrocybe cylindracea'' and ''Boletus edulis''|journal=Food Chemistry |volume=107 |issue=3 |pages=977–83 |doi=10.1016/j.foodchem.2007.07.080}}</ref>


''B. edulis'' mushrooms are rich in the [[mineral (nutrient)|dietary minerals]], [[sodium]], iron, [[calcium]], and [[magnesium]], with amounts varying according to the mushroom component and to soil composition in the geographic region of China where they were sampled.<ref name=Kalac2009/><ref name="Su">{{cite journal | last1=Su | first1=Jiuyan | last2=Zhang | first2=Ji | last3=Li | first3=Jieqing | last4=Li | first4=Tao | last5=Liu | first5=Honggao | last6=Wang | first6=Yuanzhong | title=Determination of mineral contents of wild ''Boletus edulis'' mushroom and its edible safety assessment | journal=Journal of Environmental Science and Health, Part B | volume=53 | issue=7 | date=6 April 2018 | issn=0360-1234 | doi=10.1080/03601234.2018.1455361 | pages=454–463|pmid=29624491| s2cid=4665228 }}</ref> They also have high content of [[B vitamins]] and [[tocopherol]]s.<ref name="Jaworska">{{cite journal | last1=Jaworska | first1=Grażyna | last2=Pogoń | first2=Krystyna | last3=Skrzypczak | first3=Aleksandra | last4=Bernaś | first4=Emilia | title=Composition and antioxidant properties of wild mushrooms Boletus edulis and Xerocomus badius prepared for consumption | journal=Journal of Food Science and Technology | volume=52 | issue=12 | date=12 July 2015 | issn=0022-1155 | doi=10.1007/s13197-015-1933-x | pages=7944–7953|pmid=26604366|pmc=4648917}}</ref> ''B.&nbsp;edulis'' contains appreciable amounts of [[selenium]], a [[trace mineral]],<ref>{{cite journal |author=Falandysz J. |year=2008 |title= Selenium in edible mushrooms |journal= Journal of Environmental Science and Health, Part C|volume=26 |issue=3 |pages=256–99|pmid= 18781538 |doi=10.1080/10590500802350086|s2cid=205722725 |url=https://zenodo.org/record/918325 }}</ref> although the [[bioavailability]] of mushroom-derived selenium is low.<ref>{{cite journal |author=Mutanen M. |year=1986 |title= Bioavailability of selenium in mushrooms |journal=International Journal for Vitamin and Nutrition Research |volume=56 |issue=3 |pages=297–301|pmid=3781756}}</ref>
Reported values of the composition and concentrations of [[trace metal]]s and minerals in ''Boletus edulis'' tend to differ considerably, as the mushroom bioaccumulates different elements to varying degrees, and the element concentration in the fruit bodies is often a reflection of the element concentration of the soils from which they were picked.<ref name=Kalac2009/> In general, ''B.&nbsp;edulis'' contains appreciable amounts of [[selenium]] (13&ndash;17 ppm), a [[trace mineral]] essential for good health,<ref>{{cite journal |author=Falandysz J. |year=2008 |title= Selenium in edible mushrooms |journal= Journal of Environmental Science and Health Part C–Environmental Carcinogenesis & Ecotoxicology Reviews|volume=26 |issue=3 |pages=256–99|pmid= 18781538 |doi=10.1080/10590500802350086}}</ref> though the [[bioavailability]] of mushroom-derived selenium is low.<ref>{{cite journal |author=Mutanen M. |year=1986 |title= Bioavailability of selenium in mushrooms |journal=International Journal for Vitamin and Nutrition Research |volume=56 |issue=3 |pages=297–301|pmid=3781756}}</ref> Whole fruit bodies also contain 4.7&nbsp;[[microgram|μg]] of [[vitamin D2|vitamin D<sub>2</sub>]] per 100&nbsp;g dry weight.<ref name=Mattilla2002/> The relatively high [[ergosterol]] content (see next section) of the fruit bodies can make the mushroom nutritionally pragmatic for [[vegetarian]]s and [[vegan]]s, who would otherwise have a limited intake of vitamin D.<ref name=Kalac2009>{{cite journal |author=Kalač P. |year=2009 |title=Chemical composition and nutritional value of European species of wild growing mushrooms: a review |journal=Food Chemistry |volume=113 |issue=1 |pages=9–16 |doi=10.1016/j.foodchem.2008.07.077}}</ref>


== Phytochemicals and research==
== Bioactive compounds ==
[[File:Phytochelatin.svg|thumb|Phytochelatins give ''B. edulis'' resistance to toxic heavy metals like [[cadmium]].|alt=Skeletal formula depicting the amino acids cysteine and glycine joined in a peptide bond, with free carboxy and amino groups at either end of the peptide chain. Parentheses around the peptide bond indicate a variable number of amino acids are involved.]]
''Boletus edulis'' fruit bodies contain about 500&nbsp;mg of [[ergosterol]] per 100&nbsp;g of dried mushroom.<ref name=Mattilla2002>{{cite journal |author=Mattila P, Lampi A-M, Ronkainen R, Toivo J, Piironen V. |year=2002 |title=Sterol and vitamin D<sub>2</sub> contents in some wild and cultivated mushrooms |journal=Food Chemistry |volume=76 |issue=3 |pages=293–98|doi=10.1016/S0308-8146(01)00275-8}}</ref> Ergosterol is a [[sterol]] compound common in fungi. Additionally, the fruit bodies have about 30&nbsp;mg of [[ergosterol peroxide]] per 100&nbsp;g of dried mushroom. Ergosterol peroxide is a [[steroid]] [[Derivative (chemistry)|derivative]] with a wide spectrum of [[biological activity]], including [[antimicrobial]] and [[anti-inflammatory]] activity, and [[cytotoxicity]] to various tumor [[cell line]]s grown in [[in vitro|laboratory culture]].<ref>{{cite journal |author=Krzyczkowskia W, Malinowskaa E, Suchockib P, Klepsa J, Olejnikd M, Herold F. |year=2008 |title=Isolation and quantitative determination of ergosterol peroxide in various edible mushroom species |journal=Food Chemistry |volume=113 |issue=1 |pages=351–55 |doi=10.1016/j.foodchem.2008.06.075}}</ref>
[[File:Phytochelatin.svg|thumb|left|Phytochelatins give ''B. edulis'' resistance to toxic heavy metals like [[cadmium]].|alt=Skeletal formula depicting the amino acids cysteine and glycine joined in a peptide bond, with free carboxy and amino groups at either end of the peptide chain. Parentheses around the peptide bond indicate a variable number of amino acids are involved.]]
The mushroom also contains a sugar-binding [[protein]], or [[lectin]], that has affinity for the sugars [[xylose]] and [[melibiose]]. The lectin is [[mitogenic]]—that is, it can stimulate cells to begin the process of [[cell division]], resulting in [[mitosis]]. Further, the lectin has [[antiviral drug|antiviral]] properties: it inhibits the [[human immunodeficiency virus]] enzyme [[reverse transcriptase]].<ref>{{cite journal |author=Zheng S, Li C, Ng TB, Wang HX. |year=2007 |title=A lectin with mitogenic activity from the edible wild mushroom ''Boletus edulis'' |journal=Process Biochemistry |volume=42 |issue=12 |pages=1620–24|doi=10.1016/j.procbio.2007.09.004}}</ref> Other studies suggest that ''B.&nbsp;edulis'' also has antiviral activity against ''[[Vaccinia]]'' virus<ref>{{cite journal |author=Kandefer-Szersen M, Kawecki Z, Salata B, Witek M. |year=1980 |title=Mushrooms as a source of substances with antiviral activity |journal=Acta Mycologica |volume=16 |issue=2 |pages=215–20|issn=0001-625X|language=Polish}}</ref> and [[tobacco mosaic virus]] grown in culture.<ref>{{cite journal |author=Li D, Zhao W-H, Kong B-H, Ye M, Chen H-R. |year=2009 |title=Inhibition effects of the extract and polysaccharide in macrofungus on TMV |journal= Journal of Yunnan Agricultural University |volume=24 |issue=2 |pages=175–80|issn=1004-390X}}</ref> Antiviral compounds from mushrooms are a subject of interest in biomedical research for their potential to advance the knowledge of [[viral replication]], and as new drugs in the treatment of viral disease.<ref>{{cite journal |author= Piraino FF.|year=2006 |title=Emerging antiviral drugs from medicinal mushrooms |journal= International Journal of Medicinal Mushrooms |volume=8 |issue=2 |pages=101–14 |issn=1521-9437 |doi= 10.1615/IntJMedMushr.v8.i2.20}}</ref>


The fruit bodies have a high [[antioxidant|antioxidative]] capacity, due probably to a combination of various [[organic acid]]s (such as [[oxalic acid|oxalic]], [[citric acid|citric]], [[malic acid|malic]], [[succinic acid|succinic]] and [[fumaric acid]]s), [[tocopherol]]s, [[natural phenol|phenolic compounds]]<ref>{{cite journal |author=Shu-Yao T, Hui-Li T, Jeng-Leun Mau LWT. |title=Antioxidant properties of ''Agaricus blazei'', ''Agrocybe cylindracea'', and ''Boletus edulis'' |journal=Food Science and Technology |volume=40 |issue=8 |year=2007 |pages=1392–1402 |doi=10.1016/j.lwt.2006.10.001}}</ref> and [[alkaloid]]s; the highest antioxidant activity is in the mushroom caps.<ref>{{cite journal |author=Ribeiroa B, Lopesa R, Andradea PB, Seabraa RM, Gonçalvesa, Baptistab P, Quelhasa I, Valentão P. |year=2008 |title=Comparative study of phytochemicals and antioxidant potential of wild edible mushroom caps and stipes |journal=Food Chemistry |volume=110 |issue=1 |pages=47–56 |doi=10.1016/j.foodchem.2008.01.054}}</ref> Furthermore, fruit bodies were determined to have 528&nbsp;mg of the antioxidant compound [[ergothioneine]] per kilogram of fresh mushroom; this value was the highest among many food items tested in one study.<ref>{{cite journal |author=Ey J, Schömig E, Taubert D. |title=Dietary sources and antioxidant effects of ergothioneine |journal=Journal of Agricultural and Food Chemistry |volume=55 |issue=16 |pages=6466–74 |year=2007 |pmid=17616140 |doi=10.1021/jf071328f}}</ref> Porcini were thought to have anti-cancer properties according to Hungarian research conducted in the 1950s,<ref>{{cite journal |author=Byerrum RU, Clarke DA, Lucas EH, Ringler RL, Stevens JA, Stock CC. |year=1957 |title= Tumor inhibitors in ''Boletus edulis'' and other holobasidiomycetes |journal=Antibiotics and Chemotherapy |volume=7 |issue=1 |pages=1–4 |pmid=24544235}}</ref> but later investigations in the United States did not support this.<ref name=Lamaison05/>
''Boletus edulis'' fruit bodies contain diverse [[phytochemical]]s, including 500&nbsp;mg of [[ergosterol]] per 100&nbsp;g of dried mushroom,<ref name=Mattilla2002>{{cite journal |vauthors=Mattila P, Lampi AM, Ronkainen R, Toivo J, Piironen V |year=2002 |title=Sterol and vitamin D<sub>2</sub> contents in some wild and cultivated mushrooms |journal=Food Chemistry |volume=76 |issue=3 |pages=293–98|doi=10.1016/S0308-8146(01)00275-8}}</ref> and [[ergothioneine]].<ref>{{cite journal |vauthors=Ey J, Schömig E, Taubert D |title=Dietary sources and antioxidant effects of ergothioneine |journal=Journal of Agricultural and Food Chemistry |volume=55 |issue=16 |pages=6466–74 |year=2007 |pmid=17616140 |doi=10.1021/jf071328f}}</ref> The fruit bodies contain numerous [[polyphenol]]s, especially a high content of [[rosmarinic acid]],<ref name="Vamanu">{{cite journal | last1=Vamanu | first1=Emanuel | last2=Nita | first2=Sultana | title=Antioxidant capacity and the correlation with major phenolic compounds, anthocyanin, and tocopherol content in various extracts from the wild edible ''Boletus edulis'' mushroom | journal=BioMed Research International | volume=2013 | year=2013 | issn=2314-6133 | doi=10.1155/2013/313905 | pages=1–11|pmid=23509707|pmc=3591155| doi-access=free }}</ref> and [[organic acid]]s (such as [[oxalic acid|oxalic]], [[citric acid|citric]], [[malic acid|malic]], [[succinic acid|succinic]] and [[fumaric acid]]s),<ref>{{cite journal |vauthors=Shu-Yao T, Hui-Li T, Jeng-Leun Mau LW |title=Antioxidant properties of ''Agaricus blazei'', ''Agrocybe cylindracea'', and ''Boletus edulis'' |journal=Food Science and Technology |volume=40 |issue=8 |year=2007 |pages=1392–1402 |doi=10.1016/j.lwt.2006.10.001}}</ref> and [[alkaloid]]s.<ref>{{cite journal |author1=Ribeiroa B |author2=Lopesa R |author3=Andradea PB |author4=Seabraa RM |author5=Gonçalvesa, Baptistab P |author6=Quelhasa I |author7=Valentão P. |year=2008 |title=Comparative study of phytochemicals and antioxidant potential of wild edible mushroom caps and stipes |journal=Food Chemistry |volume=110 |issue=1 |pages=47–56 |doi=10.1016/j.foodchem.2008.01.054|pmid=26050164 |url=https://bibliotecadigital.ipb.pt/bitstream/10198/4404/1/18.pdf |hdl=10198/4404 |hdl-access=free }}</ref>

===Aroma===
[[Odor|Aroma]] compounds giving ''B. edulis'' mushrooms their characteristic fragrance include some 100 components, such as [[ester]]s and [[fatty acid]]s.<ref name="csoka">{{cite journal | last1=Csóka | first1=Mariann | last2=Geosel | first2=Andras | last3=Amtmann | first3=Maria | last4=Korany | first4=Kornel | title=Volatile Composition of Some Cultivated and Wild Culinary-Medicinal Mushrooms from Hungary | journal=International Journal of Medicinal Mushrooms | volume=19 | issue=5 | year=2017 | issn=1521-9437 | doi=10.1615/intjmedmushrooms.v19.i5.50 | pages=433–443|pmid=28845772}}</ref> In a study of aroma compounds, [[1-octen-3-one]] was the most prevalent chemical detected in raw mushrooms, with [[pyrazine]]s having increased aroma effect and elevated content after drying.<ref name="Zhang">{{cite journal | last1=Zhang | first1=Huiying | last2=Pu | first2=Dandan | last3=Sun | first3=Baoguo | last4=Ren | first4=Fazheng | last5=Zhang | first5=Yuyu | last6=Chen | first6=Haitao | title=Characterization and comparison of key aroma compounds in raw and dry porcini mushroom ( Boletus edulis ) by aroma extract dilution analysis, quantitation and aroma recombination experiments | journal=Food Chemistry | volume=258 | year=2018 | issn=0308-8146 | doi=10.1016/j.foodchem.2018.03.056 | pages=260–268|pmid= 29655732| s2cid=4890378 }}</ref>


==See also==
==See also==
{{Portal|Fungi}}
*[[List of Boletus species|List of ''Boletus'' species]]
*[[List of Boletus species|List of ''Boletus'' species]]
*[[List of North American boletes]]
*[[List of North American boletes]]

== Notes ==
<references group="nb" />


==References==
==References==
'''Footnotes'''
{{Reflist|30em}}
{{Notelist}}

'''Citations'''
{{Reflist}}


===Cited texts===
===Cited texts===
*{{cite book|author=Carluccio A.|authorlink=Antonio Carluccio|date= 2003-10-17 |title= The Complete Mushroom Book|publisher = Quadrille |location=London, England|isbn=978-1-84400-040-1}}
* {{cite book|author=Carluccio, A.|author-link=Antonio Carluccio|year= 2003 |title= The Complete Mushroom Book|publisher = Quadrille |location=London |isbn=978-1-84400-040-1}}
* {{Commons-inline}}
* {{Wikispecies-inline}}


{{Taxonbar|from=Q19740}}
== External links ==
{{Commons|Boletus edulis}}
{{Wikispecies}}
* {{indexFungorum|356530}}
* [http://mushroom-collecting.com/mushroomking.html Mushroom-Collecting.com] King bolete

{{Featured article}}


{{DEFAULTSORT:Boletus Edulis}}
[[Category:Boletus|edulis]]
[[Category:Boletus|edulis]]
[[Category:Fungi described in 1782]]
[[Category:Fungi described in 1782]]
[[Category:Fungi naturalized in Australia]]
[[Category:Fungi of Africa]]
[[Category:Fungi of Africa]]
[[Category:Fungi of Asia]]
[[Category:Fungi of Asia]]
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[[Category:Fungi of North America]]
[[Category:Fungi of North America]]
[[Category:Edible fungi]]
[[Category:Edible fungi]]
[[Category:French cuisine]]
[[Category:Italian cuisine]]
[[Category:Italian products with protected designation of origin]]
[[Category:Italian products with protected designation of origin]]
[[Category:Fungi of the Middle East]]
[[Category:Fungi of Western Asia]]
[[Category:Taxa named by Jean Baptiste François Pierre Bulliard]]
[[Category:Fungus species]]
[[Category:Fungi used for fiber dyes]]

Latest revision as of 04:23, 19 December 2024

Boletus edulis
Two mushrooms with brown caps and light brown stems growing on the ground, surrounded by fallen leaves and other forest debris. One mushroom has been plucked and lies beside the other; its under-surface is visible, and is a light yellow colour.
In the northern Apennine Mountains, Abetina Reale forest, Italy
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Boletales
Family: Boletaceae
Genus: Boletus
Species:
B. edulis
Binomial name
Boletus edulis
Bull. (1782)
Synonyms[2]
  • Ceriomyces crassus Battarra (1775)
  • Boletus solidus Sowerby (1809)
  • Leccinum edule (Bull.) Gray (1821)
  • Dictyopus edulis (Bull.) Forq. (1890)
Boletus edulis
View the Mycomorphbox template that generates the following list
Pores on hymenium
Cap is convex
Hymenium is adnate
Stipe is bare
Spore print is brown
Ecology is mycorrhizal
Edibility is choice

Boletus edulis (English: cep, penny bun, porcino or porcini) is a basidiomycete fungus, and the type species of the genus Boletus.

Prized as an ingredient in various culinary dishes, B. edulis is an edible mushroom held in high regard in many cuisines, and is commonly prepared and eaten in soups, pasta, or risotto. The mushroom is low in fat and digestible carbohydrates, and high in protein, vitamins, minerals and dietary fibre. Although it is sold commercially, it is very difficult to cultivate. Available fresh in autumn throughout Europe and Russia, it is most often dried, packaged, and distributed worldwide. It keeps its flavour after drying, and it is then reconstituted and used in cooking. B. edulis is also one of the few fungi sold pickled.

The fungus grows in deciduous and coniferous forests and tree plantations, forming symbiotic ectomycorrhizal associations with living trees by enveloping the tree's underground roots with sheaths of fungal tissue. The fungus produces spore-bearing fruit bodies above ground in summer and autumn. The fruit body has a large brown cap which on occasion can reach 30 cm (12 in), rarely 40 cm (16 in) in diameter and 3 kg (6 lb 10 oz) in weight. Like other boletes, it has tubes extending downward from the underside of the cap, rather than gills; spores escape at maturity through the tube openings, or pores. The pore surface of the B. edulis fruit body is whitish when young, but ages to a greenish-yellow. The stout stipe, or stem, is white or yellowish in colour, up to 20 cm (8 in), rarely 30 cm (12 in) tall and 10 cm (4 in) thick, and partially covered with a raised network pattern, or reticulations.

Widely distributed in the Northern Hemisphere across Europe, Asia, and North America, it does not occur naturally in the Southern Hemisphere, although it has been introduced to southern Africa, Australia, New Zealand, and Brazil. Several closely related European mushrooms formerly thought to be varieties or forms of B. edulis have been shown using molecular phylogenetic analysis to be distinct species, and others previously classed as separate species are conspecific with this species. The western North American species commonly known as the California king bolete (Boletus edulis var. grandedulis) is a large, darker-coloured variant first formally identified in 2007.

Taxonomy

[edit]
Side view of the head and upper body of a man wearing a dark jacket and white ruffled collar.
Pierre Bulliard first described B. edulis in 1782.

Boletus edulis was first described in 1782 by the French botanist Pierre Bulliard and still bears its original name.[3] The starting date of fungal taxonomy had been set as January 1, 1821, to coincide with the date of the works of the 'father of mycology', Swedish naturalist Elias Magnus Fries, which meant the name required sanction by Fries (indicated in the name by a colon) to be considered valid, as Bulliard's work preceded this date. It was thus written Boletus edulis Bull.:Fr. A 1987 revision of the International Code of Botanical Nomenclature set the starting date at May 1, 1753, the date of publication of Linnaeus' work, the Species Plantarum.[4] Hence, the name no longer requires the ratification of Fries' authority. Early alternate names include Boletus solidus by English naturalist James Sowerby in 1809,[5] and Gray's Leccinum edule.[6] Gray's transfer of the species to Leccinum was later determined to be inconsistent with the rules of botanical nomenclature, and he apparently was unfamiliar with the earlier works of Fries when he published his arrangement of bolete species.[7]

Boletus edulis is the type species of the genus Boletus. In Rolf Singer's classification of the Agaricales mushrooms, it is also the type species of section Boletus, a grouping of about 30 related boletes united by several characteristics: a mild-tasting, white flesh that does not change colour when exposed to air; a smooth to distinctly raised, netted pattern over at least the uppermost portion of the stem; a yellow-brown or olive-brown spore print; white tubes that later become yellowish then greenish, which initially appear to be stuffed with cotton; and cystidia that are not strongly coloured.[8][9] Molecular analysis published in 1997 established that the bolete mushrooms are all derived from a common ancestor, and established the Boletales as an order separate from the Agaricales.[10]

The generic name is derived from the Latin term bōlētus "mushroom", which was borrowed in turn from the Ancient Greek βωλίτης, "terrestrial fungus".[11] Ultimately, this last word derives from bōlos/βῶλος "lump", "clod", and, metaphorically, "mushroom".[12] The βωλίτης of Galen, like the boletus of Latin writers like Martial, Seneca and Petronius,[13] is often identified as the much prized Amanita caesarea.[14] The specific epithet edulis in Latin means "eatable" or "edible".[15]

Common names

[edit]

Common names for B. edulis vary by region. The standard Italian name, porcino (pl. porcini), means porcine;[16] fungo porcino, in Italian, echoes the term suilli, literally "hog mushrooms", a term used by the Ancient Romans[17] and still in use in southern Italian terms for this species.[18] The derivation has been ascribed to the resemblance of young fruit bodies to piglets, or to the fondness pigs have for eating them.[19] It is also known as "king bolete".[20] The English penny bun refers to its rounded brownish shape. The German name Steinpilz (stone mushroom) refers to the species' firm flesh.[21] In Austria, it is called Herrenpilz, the "noble mushroom",[19] while in Mexico, the Spanish name is panza, meaning "belly".[22] Another Spanish name, rodellon, means "small round boulder", while the Dutch name eekhoorntjesbrood means "squirrel's bread".[23] Russian names are belyy grib (ru:белый гриб; "white mushroom" as opposed to less valuable "black mushrooms") and borovik (ru:боровик; from bor—"pine forest"). The vernacular name cep is derived from the Catalan cep or its French name cèpe, although the latter is a generic term applying to several related species. In France, it is more fully cèpe de Bordeaux, derived from the Gascon cep "trunk" for its fat stalk,[24] ultimately from the Latin cippus "stake".[25] Ceppatello, ceppatello buono, ceppatello bianco, giallo leonato, ghezzo, and moreccio are names from Italian dialects,[26][27] and ciurenys or surenys is another term in Catalan.[28] The French-born King Charles XIV John popularised B. edulis in Sweden after 1818,[29] and is honoured in the local vernacular name Karljohanssvamp, as well as the Danish name Karl Johan svamp. The monarch cultivated the fungus about his residence, Rosersberg Palace.[30] The Finnish name is herkkutatti, from herkku 'delicacy', and tatti, 'bolete'.[citation needed]

Description

[edit]
Both halves of a thick-stemmed bisected mushroom.
Cross-section showing white flesh, broad stem, and spore tubes on the underside of the cap

The cap of this mushroom is 7–30 cm (3–12 in) broad at maturity. Slightly sticky to touch, it is convex in shape when young and flattens with age. The colour is generally reddish-brown fading to white in areas near the margin, and continues to darken as it matures. The stipe, or stem, is 8–25 cm (3–10 in) in height, and up to 7 cm (3 in) thick—rather large in comparison to the cap;[31] it is club-shaped, or bulges out in the middle. It is finely reticulate on the upper portion, but smooth or irregularly ridged on the lower part. The under surface of the cap is made of thin tubes, the site of spore production; they are 1 to 2 cm (12 to 34 in) deep, and whitish in colour when young, but mature to a greenish-yellow.[32] The angular pores, which do not stain when bruised, are small—roughly 2 to 3 pores per millimetre.[33] In youth, the pores are white and appear as if stuffed with cotton (which are actually mycelia); as they age, they change colour to yellow and later to brown. The spore print is olive brown. The flesh of the fruit body is white, thick and firm when young, but becomes somewhat spongy with age. When bruised or cut, it either does not change colour, or turns a very light brown or light red.[34] Fully mature specimens can weigh about 1 kg (2 lb 3 oz); a huge specimen collected on the Isle of Skye, Scotland, in 1995 bore a cap of 42 cm (16+12 in), with a stipe 18 cm (7 in) in height and 14 cm (5+12 in) wide, and weighed 3.2 kg (7 lb 1 oz).[31] A similarly sized specimen found in Poland in 2013 made international news.[35]

A brown-capped mushroom lying flat on the grass with a white or light-brown coloured stem that gradually gets thicker, so as to roughly resemble the shape of a club.
A brown-capped mushroom with a short, stout stem that is thickest in the middle, and whose thickness approaches the width of the cap it supports.
Stem shape can range from club-shaped to centrally bulbous

Boletus edulis is considered one of the safest wild mushrooms to pick for the table, as few poisonous species closely resemble it, and those that do may be easily distinguished by careful examination.[19] The most similar poisonous mushroom may be the devil's bolete (Rubroboletus satanas), which has a similar shape, but has a red stem and stains blue on bruising.[19] It is often confused with the very bitter and unpalatable Tylopilus felleus, but can be distinguished by the reticulation on the stalk; in porcini, it is a whitish, net-like pattern on a brownish stalk, whereas it is a dark pattern on white in the latter. Porcini have whitish pores while the other has pink. If in doubt, tasting a tiny bit of flesh will yield a bitter taste.[19] It can also resemble the "bolete-like" Gyroporus castaneus, which is generally smaller, and has a browner stem. Boletus huronensis, an uncommon mushroom of northeastern North America, is another recognized look-alike known to cause severe gastrointestinal disorders.[36]

The spores are elliptical to spindle-shaped, with dimensions of 12–17 by 5–7 μm. The basidia, the spore-bearing cells, are produced in a layer lining the tubes, and arrange themselves so their ends are facing the center of the tube; this layer of cells is known technically as a hymenium. The basidia are thin-walled, mostly attached to four spores, and measure 25–30 by 8–10 μm. Another cell type present in the hymenium is the cystidia, larger sterile cells that protrude beyond the basidia into the lumen of the hymenium, and act as air traps, regulating humidity.[37] B. edulis has pleurocystidia (cystidia located on the face of a pore) that are thin-walled, roughly spindle-shaped to ventricose, and measure 30–45 by 7–10 μm; the "stuffed" feature of the hymenium is caused by cheilocystidia—cells found on the edges of the pores.[32] The hyphae of B. edulis do not have clamp connections.[33]

[edit]
A mushroom with an orange-brown cap and a yellowish underside that somewhat resembles a sponge. The light-yellow stem is about half the thickness of the caps diameter. This mushroom is growing on the ground, surrounded by twigs, leaves, log and other forest floor debris.
B. edulis var. grandedulis
A mushroom with a reddish-brown cap that is curled upwards to reveal a cream-coloured porous underside that somewhat resembles a sponge. The thick stipe has a pinkish hue, and its thickness is a little less than half of the cap's diameter. The mushroom has been pulled from the ground and the end of its stipe is a whitish colour caused by hyphal tufts, and is embedded with dirt and other small twigs.
B. regineus

Several similar brownish-coloured species are sometimes considered subspecies or forms of this mushroom. In Europe, in addition to B. edulis (or cèpe de Bordeaux), the most popular are:

  • Cèpe bronzé ("dark cep"; Boletus aereus), much rarer than B. edulis, is more highly regarded by gourmets, and consequently more expensive. Usually smaller than B. edulis, it is also distinctively darker in colour.[19] It is especially suited to drying.[20]
  • Cèpe des pins ("pine tree cep"; Boletus pinophilus or Boletus pinicola) grows among pine trees. Rarer than B. edulis, it is less appreciated by gourmets than the two other kinds of porcini, but remains a mushroom rated above most others.[20]
  • Cèpe d'été ("summer cep"; Boletus reticulatus), also less common and found earlier.[19]

Molecular phylogenetic analyses have proven these three are all distinctive and separate species;[38] other taxa formerly believed to be unique species or subspecies, such as B. betulicola, B. chippewaensis, B. persoonii, B. quercicola and B. venturii, are now known to be part of a B. edulis species complex with a wide morphological, ecological and geographic range,[39][40] and that the genetic variability in this complex is low.[41] Similar molecular technology has been developed to rapidly and accurately identify B. edulis and other commercially important fungi.[42][43]

Three divergent lineages found in Yunnan province in China that are commonly marketed and sold as B. edulis (and are actually more closely related to B. aereus) were described in 2013 as B. bainiugan, B. meiweiniuganjun and B. shiyong.[44][45] The classification has since been updated and expanded. All lineages are still members of Boletus sect. Boletus, the sensu sticto "porcini clade" of the genus.[46]

A group of three mushrooms with reddish-brown caps, bright yellow porous undersides, and thick white stipes. They are growing on the ground in soil covered with pine needles.
B. rex-veris

Western North America has several species closely related to B. edulis. The white king bolete (Boletus barrowsii), found in parts of Colorado, New Mexico, Arizona, and California (and possibly elsewhere), is named after its discoverer Chuck Barrows.[47] It is lighter in colour than B. edulis, having a cream-coloured cap with pink tones;[48] often mycorrhizal with Ponderosa pine, it tends to grow in areas where there is less rainfall. Some find its flavour as good as if not better than B. edulis.[49] The California king bolete (Boletus edulis var. grandedulis) can reach massive proportions, and is distinguished from B. edulis by a mature pore surface that is brown to slightly reddish. The cap colour appears to be affected by the amount of light received during its development, and may range from white in young specimens grown under thick canopy, to dark-brown, red-brown or yellow brown in those specimens receiving more light.[50] The queen bolete (Boletus regineus), formerly considered a variety of B. aereus, is also a choice edible. It is generally smaller than B. edulis, and unlike that species, is typically found in mixed forests.[51] The spring king bolete (Boletus rex-veris), formerly considered a variety of B. edulis or B. pinophilus, is found throughout western North America. In contrast to B. edulis, B. rex-veris tends to fruit in clusters, and, as its common name suggests, appears in the spring.[52] B. fibrillosus is edible but considered inferior in taste.[53]

Habitat and distribution

[edit]
A mushroom with a brown smooth cap the shape of a halved sphere, atop a thick, dirty white stipe. The mushroom is growing on a sloping patch of ground amongst moss, twigs and other forest debris; trees can be faintly seen in the background.
In Lithuania

The fruit bodies of Boletus edulis can grow singly or in small clusters of two or three specimens. The mushroom's habitat consists of areas dominated by pine (Pinus spp.), spruce (Picea spp.), hemlock (Tsuga spp.) and fir (Abies spp.) trees, although other hosts include chestnut, chinquapin, beech, Keteleeria spp., Lithocarpus spp., and oak. In California, porcini have been collected in a variety of forests, such as coastal forests, dry interior oak forests and savannas and interior high-elevation montane mixed forests,[54] to an altitude of 3,500 m (11,500 ft).[55] In northwestern Spain, they are common in scrublands dominated by the rock rose species Cistus ladanifer and Halimium lasianthum.[56] In the Midi region of south-west France, they are especially favoured and locally called cèpe de Bordeaux after the town from which they are traded to the north and abroad.[57]

Boletus edulis has a cosmopolitan distribution, concentrated in cool-temperate to subtropical regions.[54] It is common in Europe—from northern Scandinavia, south to the extremities of Greece and Italy—and North America, where its southern range extends as far south as Mexico.[34] It is well known from the Borgotaro area of Parma, Italy, and has PGI status there. The European distribution extends north to Scandinavia and south to southern Italy and Morocco.[54] In the American Pacific Northwest, it can be found from May to October.[58] In China, the mushroom can be found from the northeastern Heilongjiang to the Yunnan–Guizhou Plateau and Tibet.[34] It has been recorded growing under Pinus and Tsuga in Sagarmatha National Park in Nepal,[59] as well as in the Indian forests of Arunachal Pradesh.[60] In West Asia, the species has been reported from the northwest forests of Iran.[61]

Cultivation

[edit]

Some steps have been made towards cultivating Boletus edulis,[62] including mycorrhization of rockrose shrubs enhanced by helper bacteria.[63]

Non-native introductions

[edit]

Boletus edulis grows in some areas where it is not believed to be indigenous. It is often found underneath oak and silver birch in Hagley Park in central Christchurch, New Zealand, where it is likely to have been introduced,[64] probably on the roots of container-grown beech, birch, and oak in the mid-19th century—around the time exotic trees began to be planted in the Christchurch area.[34] Similarly, it has been collected in Adelaide Hills region of Australia in association with three species of introduced trees.[65] It has been growing plentifully in association with pine forests in the southern KwaZulu-Natal Midlands in South Africa for more than 50 years and is believed to have been introduced with the import of pine trees.[66][67] It also grows in pine plantations in neighboring Zimbabwe.[68]

Ecology

[edit]
About half a dozen pine trees with upward-pointing branches 15 to 30 metres in height with green needles. The upper half of the background is blue sky.
B. edulis is ectomycorrhizal and may co-occur with Pinus radiata

Fruit body production

[edit]

Italian folklore holds that porcini sprout up at the time of the new moon;[19] research studies have tried to investigate more scientifically the factors that influence the production of fruit bodies. Although fruit bodies may appear any time from summer to autumn (June to November in the UK), their growth is known to be triggered by rainfall during warm periods of weather followed by frequent autumn rain with a drop in soil temperature.[54] Above average rainfall may result in the rapid appearance of large numbers of boletes, in what is known in some circles as a "bolete year".[69] A 2004 field study indicated that fruit body production is enhanced by an open and sunny wood habitat,[70] corroborating an earlier observation made in a Zimbabwean study;[68] removal of the litter layer on the forest floor appeared to have a negative effect on fruit body production, but previous studies reported contradictory results.[71][72] A Lithuanian study conducted in 2001 concluded that the maximal daily growth rate of the cap (about 21 mm or 0.8 in) occurred when the relative air humidity was the greatest, and the fruit bodies ceased growing when the air humidity dropped below 40%. Factors most likely to inhibit the appearance of fruit bodies included prolonged drought, inadequate air and soil humidity, sudden decreases of night air temperatures, and the appearance of the first frost.[73] Plots facing north tend to produce more mushrooms compared to equivalent plots facing south.[74]

Mycorrhizal associations

[edit]

Boletus edulis is mycorrhizal—it is in a mutualistic relationship with the roots of plants (hosts), in which the fungus exchanges nitrogen and other nutrients extracted from the environment for fixed carbon from the host. Other benefits for the plant are evident: in the case of the Chinese chestnut, the formation of mycorrhizae with B. edulis increases the ability of plant seedlings to resist water stress, and increases leaf succulence, leaf area, and water-holding ability.[75] The fungus forms a sheath of tissue around terminal, nutrient-absorbing root tips, often inducing a high degree of branching in the tips of the host, and penetrating into the root tissue, forming, to some mycologists, the defining feature of ectomycorrhizal relationships, a hartig net.[76] The ectomycorrhizal fungi are then able to exchange nutrients with the plant, effectively expanding the root system of the host plant to the furthest reaches of the symbiont fungi.[76] Compatible hosts may belong to multiple families of vascular plants that are widely distributed throughout the Northern Hemisphere; according to one 1995 estimate, there are at least 30 host plant species distributed over more than 15 genera.[34] Examples of mycorrhizal associates include Chinese red pine,[77] Mexican weeping pine,[78] Scots pine, Norway spruce,[79] Coast Douglas-fir,[80] mountain pine,[81] and Virginia pine.[82] The fungus has also been shown to associate with gum rockrose, a pioneer early stage shrub that is adapted for growth in degraded areas, such as burned forests.[83] These and other rockrose species are ecologically important as fungal reservoirs, maintaining an inoculum of mycorrhizal fungi for trees that appear later in the forest regrowth cycle.[84]

The mushroom has been noted to often co-occur with Amanita muscaria or A. rubescens, although it is unclear whether this is due to a biological association between the species, or because of similarities in growing season, habitat, and ecological requirements.[54] An association has also been reported between B. edulis and Amanita excelsa on Pinus radiata ectomycorrhizae in New Zealand, suggesting that other fungi may influence the life cycle of porcini.[85] A 2007 field study revealed little correlation between the abundance of fruit bodies and presence of its mycelia below ground, even when soil samples were taken from directly beneath the mushroom; the study concluded that the triggers leading to formation of mycorrhizae and production of the fruit bodies were more complex.[86]

Heavy-metal contamination

[edit]

Boletus edulis is known to be able to tolerate and even thrive on soil that is contaminated with toxic heavy metals, such as soil that might be found near metal smelters. The mushroom's resistance to heavy-metal toxicity is conferred by a biochemical called a phytochelatin—an oligopeptide whose production is induced after exposure to metal.[87] Phytochelatins are chelating agents, capable of forming multiple bonds with the metal; in this state, the metal cannot normally react with other elements or ions and is stored in a detoxified form in the mushroom tissue.[citation needed]

Pests and predators

[edit]

The fruit bodies of B. edulis can be infected by the parasitic mould-like fungus Hypomyces chrysospermus, known as the bolete eater, which manifests itself as a white, yellow, or reddish-brown cottony layer over the surface of the mushroom.[88] Some reported cases of stomach ache following consumption of dried porcini have been attributed to the presence of this mould on the fruit bodies.[89] The mushroom is also used as a food source by several species of mushroom flies,[54] as well as other insects and their larvae.[90] An unidentified species of virus was reported to have infected specimens found in the Netherlands and in Italy; fruit bodies affected by the virus had relatively thick stems and small or no caps, leading to the name "little-cap disease".[91][92]

Boletus edulis is a food source for animals such as the banana slug (Ariolimax columbianus),[93] the long-haired grass mouse,[94] the red squirrel,[95] and, as noted in one isolated report, the fox sparrow.[96]

Culinary uses

[edit]
A porcini mushroom and noodle soup served in a bread bowl at a Polish restaurant
A porcini mushroom sandwich in Stockholm, Sweden

Boletus edulis, as the species epithet edulis (Latin: edible) indicates, is an edible mushroom. Italian chef and restaurateur Antonio Carluccio has described it as representing "the wild mushroom par excellence", and hails it as the most rewarding of all fungi in the kitchen for its taste and versatility.[19] Considered a choice edible, particularly in France, Germany, Poland and Italy,[20] it was widely written about by the Roman writers Pliny the Elder and Martial, although ranked below the esteemed Amanita caesarea. When he was served suilli[a] instead of boleti,[98] the disgruntled Martial wrote:

sunt tibi boleti; fungos ego sumo suillos (Ep. iii. 60)
("You eat the choice boletus, I have mushrooms that swine grub up.")[99]

The flavour of porcini has been described as nutty and slightly meaty, with a smooth, creamy texture, and a distinctive aroma reminiscent of sourdough. Young, small porcini are most appreciated, as the large ones often harbour maggots (insect larvae), and become slimy, soft and less tasty with age. The fruit bodies are collected by holding the stipe near the base and twisting gently. Cutting the stipe with a knife may risk the part left behind rotting and the mycelium being destroyed. Peeling and washing are not recommended.[19] The fruit bodies are highly perishable, due largely to the high water content (around 90%), the high level of enzyme activity, and the presence of a flora of microorganisms.[100] Caution should be exercised when collecting specimens from potentially polluted or contaminated sites, as several studies have shown that the fruit bodies can bioaccumulate toxic heavy metals like mercury,[101] cadmium,[102] caesium and polonium.[103][104] Bioaccumulated metals or radioactive fission decay products are like chemical signatures: chemical and radiochemical analysis can be used to identify the origin of imported specimens,[105] and for long-term radioecological monitoring of polluted areas.[106]

Porcini are sold fresh in markets in summer and autumn in Europe and Russia, and dried or canned at other times of the year, and distributed worldwide to countries where they are not otherwise found.[107] They are eaten and enjoyed raw, sautéed with butter, ground into pasta, in soups, and in many other dishes. In France, they are used in recipes such as cèpes à la Bordelaise, cèpe frits and cèpe aux tomates.[108] Porcini risotto is a traditional Italian autumn dish.[109] Porcini are a feature of many cuisines, including Provençal,[110] and Viennese.[111] In Thailand they are used in soups and consumed blanched in salads.[112] Porcini can also be frozen, either while raw or after cooking in butter. The colour, aroma, and taste of porcini deteriorate noticeably after being frozen for four months. Blanching (or soaking and blanching) as a processing step before freezing can extend the freezer life to 12 months.[100] They are also one of the few species sold commercially as pickled mushrooms.[113]

Dried

[edit]
A heaped pile of dried sliced mushrooms on a large plate. A sign near the back of the plate reads "Fungo di Borgotaro I.G.P."; another sign at the front reads "Prezzo speciale di Fiera. 3 Etti 18,00". An electronic scale is partially visible on a table behind the mushrooms
A pile of dried porcini at the Borgotaro porcino festival in Italy

Boletus edulis is well suited to drying—its flavour intensifies, it is easily reconstituted, and its resulting texture is pleasant.[114] Reconstitution is done by soaking in hot, but not boiling, water for about twenty minutes; the water used is infused with the mushroom aroma and it too can be used in subsequent cooking. Dried porcini have more protein than most other commonly consumed vegetables, apart from soybeans. Some of their protein is indigestible, though digestibility is improved with cooking.[115]

Like other boletes, porcini can be dried by being strung separately on twine and hung close to the ceiling of a kitchen. Alternatively, the mushrooms can be dried by cleaning with a brush (washing is not recommended), and then placing them in a wicker basket or bamboo steamer on top of a boiler or hot water tank.[116] Another method is drying in an oven at 25 to 30 °C (77 to 86 °F) for two to three hours, then increasing the temperature to 50 °C (122 °F) until crisp or brittle.[117] Once dry, they must be kept in an airtight container.[116] Importantly for commercial production, porcini retain their flavour after industrial preparation in a pressure cooker or after canning or bottling, and are thus useful for manufacturers of soups or stews. The addition of a few pieces of dried porcino can significantly add to flavour, and they are a major ingredient of the pasta sauce known as carrettiera (carter's sauce).[118] The drying process is known to induce the formation of various volatile substances that contribute to the mushroom's aroma. Chemical analysis has shown that the odour of the dried mushroom is a complex mixture of 53 volatile compounds.[119]

Commercial harvest

[edit]
Approximately two dozen brown-capped, white or light-brown stemmed mushrooms of various sizes in a brown bowl.
Porcini can vary considerably in size.

A 1998 estimate suggested that the total annual worldwide consumption of Boletus edulis and closely related species (B. aereus, B. pinophilus, and B. reticulatus) was between 20,000 and 100,000 tons.[54] Approximately 2,700 tonnes (3,000 tons) were sold in France, Italy and Germany in 1988, according to official figures. The true amount consumed far exceeds this, as the official sales figures did not account for informal sales or consumption by collectors.[55] They are widely exported and sold in dried form, reaching countries where they do not occur naturally, such as Australia and New Zealand. The autonomous community of Castile and León in Spain produces 7,700 tonnes (8,500 tons) annually.[83] In autumn, the price of porcini in the Northern Hemisphere typically ranges between $20 and $80 per kilogram, although in New York in 1997 the wholesale price rose to more than $200 per kilogram due to scarcity.[55]

In the vicinity of Borgotaro in the Province of Parma of northern Italy, the four species Boletus edulis, B. aereus, B. aestivalis and B. pinophilus have been recognised for their superior taste and officially termed Fungo di Borgotaro. Here these mushrooms have been collected for centuries and exported commercially. Owing to the globalisation of the mushroom trade most of the porcini commercially available in Italy or exported by Italy no longer originate there. Porcini and other mushrooms are also imported into Italy from various locations, especially China and eastern European countries; these are then often re-exported under the "Italian porcini" label.[120][121]

In Italy the disconnect with local production has had an adverse effect on quality; for example in the 1990s some of the dried porcino mushrooms exported to Italy from China contained species of genus Tylopilus, which are rather similar in appearance and when dried are difficult for both mushroom labourers and mycologists alike to distinguish from Boletus. Tylopilus species typically have a very bitter taste, which is imparted to the flavour of the porcini with which they are mixed.[122]

After the fall of the Iron Curtain and the subsequent reduction of economic and political barriers, central and eastern European countries with local mushroom harvesting traditions, such as Albania, Bulgaria, Macedonia, Romania, Serbia and Slovenia, developed into exporters of porcini, concentrating primarily on the Italian market.[121] Porcini and other wild fungi from these countries are also destined for France, Germany and other western European markets, where demand for them exists but collection on a commercial scale does not.[121] Picking B. edulis has become an annual seasonal income earner and pastime in countries like Bulgaria, especially for many Roma communities and the unemployed.[123] A lack of control of the harvest has led to heavy exploitation of the mushroom resource.[124]

Like many other strictly mycorrhizal fungi, B. edulis has eluded cultivation attempts for years.[115][125][62] The results of some studies suggest that unknown components of the soil microflora might be required for B. edulis to establish a mycorrhizal relationship with the host plant.[126][127][128] Successful attempts at cultivating B. edulis have been made by Spanish scientists by mycorrhization of Cistus species,[62] with Pseudomonas fluorescens bacteria helping the mycorrhiza.[63]

Boletus edulis, fresh[129]
Nutritional value per 100 g (3.5 oz)
Energy342.4 kJ (81.8 kcal)
9.23 g
1.70 g
7.39 g
Vitamins and minerals
VitaminsQuantity
%DV
Thiamine (B1)
9%
0.105 mg
Riboflavin (B2)
7%
0.092 mg
Niacin (B3)
38%
6.07 mg
Pantothenic acid (B5)
53%
2.64 mg
Vitamin B6
3%
0.051 mg
Folate (B9)
73%
290 μg
Vitamin C
5%
4.21 mg
MineralsQuantity
%DV
Calcium
0%
1.195 mg
Copper
87%
0.786 mg
Iron
4%
0.739 mg
Phosphorus
2%
22.26 mg
Potassium
7%
203.3 mg
Zinc
38%
4.172 mg
Percentages estimated using US recommendations for adults,[130] except for potassium, which is estimated based on expert recommendation from the National Academies.[131]

Nutrition

[edit]

Boletus edulis mushrooms are 9% carbohydrates, 3% fat, and 7% protein (table). Fresh mushrooms consist of over 80% moisture,[132] although reported values tend to differ somewhat as moisture content can be affected by environmental temperature and relative humidity during growth and storage.[133] The carbohydrate component contains the monosaccharides glucose, mannitol and α,α-trehalose, the polysaccharide glycogen, and the water-insoluble structural polysaccharide chitin, which accounts for up to 80–90% of dry matter in mushroom cell walls. Chitin, hemicellulose, and pectin-like carbohydrates—all indigestible by humans—contribute to the high proportion of insoluble fibre in B. edulis.[134]

The total lipid, or crude fat, content makes up 3% of the dry matter of the mushroom. The proportion of fatty acids (expressed as a % of total fatty acids) are: linoleic acid 42%, oleic acid 36%, palmitic acid 10%, and stearic acid 3%.[135]

A comparative study of the amino acid composition of eleven Portuguese wild edible mushroom species showed Boletus edulis to have the highest total amino acid content.[136][137]

B. edulis mushrooms are rich in the dietary minerals, sodium, iron, calcium, and magnesium, with amounts varying according to the mushroom component and to soil composition in the geographic region of China where they were sampled.[134][138] They also have high content of B vitamins and tocopherols.[139] B. edulis contains appreciable amounts of selenium, a trace mineral,[140] although the bioavailability of mushroom-derived selenium is low.[141]

Phytochemicals and research

[edit]
Skeletal formula depicting the amino acids cysteine and glycine joined in a peptide bond, with free carboxy and amino groups at either end of the peptide chain. Parentheses around the peptide bond indicate a variable number of amino acids are involved.
Phytochelatins give B. edulis resistance to toxic heavy metals like cadmium.

Boletus edulis fruit bodies contain diverse phytochemicals, including 500 mg of ergosterol per 100 g of dried mushroom,[142] and ergothioneine.[143] The fruit bodies contain numerous polyphenols, especially a high content of rosmarinic acid,[144] and organic acids (such as oxalic, citric, malic, succinic and fumaric acids),[145] and alkaloids.[146]

Aroma

[edit]

Aroma compounds giving B. edulis mushrooms their characteristic fragrance include some 100 components, such as esters and fatty acids.[147] In a study of aroma compounds, 1-octen-3-one was the most prevalent chemical detected in raw mushrooms, with pyrazines having increased aroma effect and elevated content after drying.[148]

See also

[edit]

References

[edit]

Footnotes

  1. ^ The term suilli was also thought to encompass the related Leccinum scabrum.[97]

Citations

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