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Boletus edulis
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==Ecology== {{Multiple image|direction=vertical|align=right|image1=Pinus radiata HuckleberryHill1.jpg|image2=Amanita muscaria 3 vliegenzwammen op rij.jpg|width=200|caption1=''B. edulis'' is [[ectomycorrhizal]] and may co-occur with ''Pinus radiata''|caption2=and ''[[Amanita muscaria]]''.|alt1=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.|alt2=A group of three mushrooms, ranging in size from small to large, with red caps dotted with white warts, and white stems. The largest of the three has a droopy skirt hanging from the upper portion of its stem. The mushrooms are growing in the ground, surrounded by fallen brown leaves, green grass, and a dark green bush in the background.}} ===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 |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 |url-access=subscription }}</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 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]].<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=== ''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. 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 often co-occur with ''[[Amanita muscaria]]'' or ''[[Amanita rubescens|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.<ref name=Hall1998/> An association has also been reported between ''B. 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=== ''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=== 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.<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 |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>
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