A truffle is the fruiting body of a subterranean ascomycete

fungus A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...

, one of the species of the genus ''

Tuber Tubers are a type of enlarged structure that plants use as storage organs for nutrients, derived from stems or roots. Tubers help plants perennate (survive winter or dry months), provide energy and nutrients, and are a means of asexual reproduc ...

''. More than one hundred other genera of fungi are classified as truffles including '' Geopora'', '' Peziza'', '' Choiromyces'', and '' Leucangium''. These genera belong to the class Pezizomycetes and the Pezizales order. Several truffle-like

basidiomycetes Basidiomycota () is one of two large division (mycology), divisions that, together with the Ascomycota, constitute the subkingdom Dikarya (often referred to as the "higher fungi") within the kingdom Fungi. Members are known as basidiomycetes. Mor ...

are excluded from Pezizales, including '' Rhizopogon'' and ''Glomus''. Truffles are ectomycorrhizal fungi, so they are found in close association with tree roots. Spore dispersal is accomplished through fungivores, animals that eat fungi. These fungi have ecological roles in nutrient cycling and drought tolerance. Some truffle species are prized as food.English translation
Edible truffles are used in Italian, French and other national . Truffles are cultivated and harvested from natural environments.

Taxonomic details

Species

Black

The black truffle or black Périgord truffle (''Tuber melanosporum''), the second-most commercially valuable species, is named after the Périgord region in France. Black truffles associate with oaks, hazelnut, cherry, and other deciduous trees and are harvested in late autumn and winter. The genome sequence of the black truffle was published in March 2010.

Summer or burgundy

The black summer truffle (''Tuber aestivum'') is found across Europe and is prized for its culinary value. Burgundy truffles (designated '' Tuber uncinatum'', but the same species) are harvested in autumn until December and have aromatic flesh of a darker colour. These are associated with various trees and shrubs.

White

Tuber magnatum, the high-value ''white truffle'' () is found mainly in the Langhe and Montferrat areas of the

Piedmont Piedmont ( ; ; ) is one of the 20 regions of Italy, located in the northwest Italy, Northwest of the country. It borders the Liguria region to the south, the Lombardy and Emilia-Romagna regions to the east, and the Aosta Valley region to the ...

region in northern

Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...

, and most famously, in the countryside around the cities of Alba and Asti. A large percentage of Italy's white truffles also come from

Molise Molise ( , ; ; , ) is a Regions of Italy, region in Southern Italy. Until 1963, it formed part of the region of Abruzzi e Molise together with Abruzzo. The split, which did not become effective until 1970, makes Molise the newest region in Ital ...

Whitish

The " whitish truffle" (''Tuber borchii'') is a similar species native to

Tuscany Tuscany ( ; ) is a Regions of Italy, region in central Italy with an area of about and a population of 3,660,834 inhabitants as of 2025. The capital city is Florence. Tuscany is known for its landscapes, history, artistic legacy, and its in ...

Abruzzo Abruzzo (, ; ; , ''Abbrìzze'' or ''Abbrèzze'' ; ), historically also known as Abruzzi, is a Regions of Italy, region of Southern Italy with an area of 10,763 square km (4,156 sq mi) and a population of 1.3 million. It is divided into four ...

, Romagna,

Umbria Umbria ( ; ) is a Regions of Italy, region of central Italy. It includes Lake Trasimeno and Cascata delle Marmore, Marmore Falls, and is crossed by the Tiber. It is the only landlocked region on the Italian Peninsula, Apennine Peninsula. The re ...

, the

Marche Marche ( ; ), in English sometimes referred to as the Marches ( ) from the Italian name of the region (Le Marche), is one of the Regions of Italy, twenty regions of Italy. The region is located in the Central Italy, central area of the country, ...

, and

. It is reportedly not as aromatic as those from Piedmont, although those from Città di Castello are said to come quite close.

Other ''Tuber''

A less common truffle is "garlic truffle" ('' Tuber macrosporum''). In the U.S.

Pacific Northwest The Pacific Northwest (PNW; ) is a geographic region in Western North America bounded by its coastal waters of the Pacific Ocean to the west and, loosely, by the Rocky Mountains to the east. Though no official boundary exists, the most common ...

, several species of truffle are harvested both recreationally and commercially, most notably, the '' Leucangium carthusianum'', Oregon black truffle; '' Tuber gibbosum'', Oregon spring white truffle; and '' Tuber oregonense'', Oregon winter white truffle. '' Kalapuya brunnea'', the Oregon brown truffle, has also been commercially harvested and is of culinary note. The Oregon white truffle is increasingly harvested due to its high quality and also exported to other countries. Oregon celebrates its traditional truffle harvesting with a 'truffle festival', combined with culinary shows and wine tastings. The pecan truffle (''Tuber lyonii'') syn. ''texense'' is found in the

Southern United States The Southern United States (sometimes Dixie, also referred to as the Southern States, the American South, the Southland, Dixieland, or simply the South) is List of regions of the United States, census regions defined by the United States Cens ...

, usually associated with pecan trees. Chefs who have experimented with them agree "they are very good and have potential as a food commodity". Although pecan farmers used to find them along with pecans and discard them, considering them a nuisance, they sell for about $160 a pound and have been used in some gourmet restaurants.

Beyond ''Tuber''

The term "truffle" has been applied to several other genera of similar underground fungi. The genera ''

Terfezia ''Terfezia'' (Berber language, Berber: Tirfas) is a genus of truffle-like fungi within the Pezizaceae family. ''Terfezia'' species are common name, commonly known as desert truffles. Some authorities consider this the type (biology), type genus o ...

'' and '' Tirmania'' of the family

Terfeziaceae The Terfeziaceae, or desert truffles, is a family of truffles (, , , ) endemic to arid and semi-arid areas of the Mediterranean Region, North Africa, and the Middle East, where they live in ectomycorrhizal association with '' Helianthemum'' ...

are known as the "desert truffles" of Africa and the Middle East. '' Pisolithus tinctorius'', which was historically eaten in parts of

Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...

, is sometimes called "Bohemian truffle". '' Geopora'' spp. are important ectomycorrhizal partners of trees in woodlands and forests throughout the world. ''Pinus edulis'', a widespread pine species of the Southwest US, is dependent on ''Geopora'' for nutrient and water acquisition in arid environments. Like other truffle fungi, ''Geopora'' produces subterranean sporocarps as a means of sexual reproduction. '' Geopora cooperi'', also known as pine truffle or fuzzy truffle, is an edible species of this genus. Rhizopogon rubescens

'' Rhizopogon'' spp. are ectomycorrhizal members of the Basidiomycota and the order Boletales, a group of fungi that typically form mushrooms. Like their ascomycete counterparts, these fungi can create truffle-like fruiting bodies. ''Rhizopogon'' spp. are ecologically important in coniferous forests where they associate with various pines, firs, and Douglas fir. In addition to their ecological importance, these fungi hold economic value, as well. ''Rhizopogon'' spp. are commonly used to inoculate coniferous seedlings in nurseries and during reforestation. '' Hysterangium'' spp. are ectomycorrhizal members of the Basidiomycota and the order Hysterangiales that form sporocarps similar to true truffles. These fungi form mycelial mats of vegetative hyphae that may cover 25–40% of the forest floor in Douglas fir forests, thereby contributing to a significant portion of the biomass present in soils. Like other ectomycorrhizal fungi, ''Hysterangium'' spp. play a role in nutrient exchange in the nitrogen cycle by accessing nitrogen unavailable to host plants and acting as nitrogen sinks in forests. '' Glomus'' spp. are arbuscular mycorrhizae of the phylum Glomeromycota within the order Glomerales. Members of this genus have low host specificity, associating with a variety of plants including hardwoods, forbs, shrubs, and grasses. These fungi commonly occur throughout the Northern Hemisphere. Members of the genus '' Elaphomyces'' are commonly mistaken for truffles.

Phylogeny

Phylogenetic analysis has demonstrated the

convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...

of the ectomycorrhizal trophic mode in diverse fungi. The subphylum Pezizomycotina, containing the order Pezizales, is approximately 400 million years old. Within the order Pezizales, subterranean fungi evolved independently at least fifteen times. Contained within Pezizales are the families Tuberaceae,

Pezizaceae The Pezizaceae (commonly referred to as cup fungi) are a family of fungi in the Ascomycota which produce mushrooms that tend to grow in the shape of a "cup". Spores are formed on the inner surface of the fruit body ( ascoma). The cup shape typi ...

, Pyronematacae, and Morchellaceae. All of these families contain lineages of subterranean or truffle fungi. The oldest ectomycorrhizal fossils are from the

Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...

about 50 million years ago. The specimens are preserved permineralized in-situ in the Eocene Okanagan Highlands Princeton chert site. This indicates that the soft bodies of ectomycorrhizal fungi do not easily fossilize. Molecular clockwork has suggested the evolution of ectomycorrhizal fungi occurred approximately 130 million years ago. The evolution of subterranean fruiting bodies has occurred numerous times within the

Ascomycota Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...

Basidiomycota Basidiomycota () is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya (often referred to as the "higher fungi") within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Basi ...

, and Glomeromycota. For example, the genera '' Rhizopogon'' and '' Hysterangium'' of Basidiomycota both form subterranean fruiting bodies and play similar ecological roles as truffle forming ascomycetes. The ancestors of the Ascomycota genera ''Geopora'', ''Tuber'', and ''Leucangium'' originated in Laurasia during the Paleozoic era. Phylogenetic evidence suggests that most subterranean fruiting bodies evolved from above-ground mushrooms. Over time mushroom stipes and caps were reduced, and caps began to enclose reproductive tissue. The dispersal of sexual spores then shifted from wind and rain to utilising animals. The

phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...

and

biogeography Biogeography is the study of the species distribution, distribution of species and ecosystems in geography, geographic space and through evolutionary history of life, geological time. Organisms and biological community (ecology), communities o ...

of the genus ''Tuber'' was investigated in 2008 using internal transcribed spacers (ITS) of

nuclear DNA Nuclear DNA (nDNA), or nuclear deoxyribonucleic acid, is the DNA contained within each cell nucleus of a eukaryotic organism. It encodes for the majority of the genome in eukaryotes, with mitochondrial DNA and plastid DNA coding for the rest. ...

and revealed five major

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

s (Aestivum, Excavatum, Rufum, Melanosporum and Puberulum); this was later improved and expanded in 2010 to nine major clades using 28S large subunits (LSU) rRNA of

mitochondrial DNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...

. The Magnatum and Macrosporum clades were distinguished as distinct from the Aestivum clade. The Gibbosum clade was resolved as distinct from all other clades, and the Spinoreticulatum clade was separated from the Rufum clade. The truffle habit has evolved independently among several basidiomycete genera. Phylogenetic analysis has revealed that basidiomycete subterranean fruiting bodies, like their ascomycete counterparts, evolved from above ground mushrooms. For example, ''Rhizopogon'' species likely arose from an ancestor shared with '' Suillus'', a mushroom-forming genus. Studies have suggested that selection for subterranean fruiting bodies among ascomycetes and basidiomycetes occurred in water-limited environments.

Etymology

Most sources agree that the term "truffle" is derived from the

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

term by way of the Vulgar Latin , meaning "swelling" or "lump". This then entered other languages through

Old French Old French (, , ; ) was the language spoken in most of the northern half of France approximately between the late 8th

mycelia Mycelium (: mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Its normal form is that of branched, slender, entangled, anastomosing, hyaline threads. Fungal colonies composed of mycelium are fo ...

of truffles form symbiotic, mycorrhizal relationships with the roots of several tree species, including beech, birch, hazel, hornbeam, oak,

pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as cu ...

, and poplar. Mutualistic ectomycorrhizal fungi such as truffles provide valuable nutrients to plants in exchange for

carbohydrate A carbohydrate () is a biomolecule composed of carbon (C), hydrogen (H), and oxygen (O) atoms. The typical hydrogen-to-oxygen atomic ratio is 2:1, analogous to that of water, and is represented by the empirical formula (where ''m'' and ''n'' ...

s. Ectomycorrhizal fungi cannot survive in the soil without their plant hosts. In fact, many of these fungi have lost the

enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...

s necessary for obtaining carbon through other means. For example, truffle fungi have lost their ability to degrade the cell walls of plants, limiting their capacity to decompose plant litter. Plant hosts can also depend on their associated truffle fungi. ''Geopora'', ''Peziza'', and ''Tuber'' spp. are vital in the establishment of oak communities. ''Tuber'' species prefer argillaceous or calcareous soils that are well drained and neutral or

alkaline In chemistry, an alkali (; from the Arabic word , ) is a basic salt of an alkali metal or an alkaline earth metal. An alkali can also be defined as a base that dissolves in water. A solution of a soluble base has a pH greater than 7.0. The ...

. ''Tuber'' truffles fruit throughout the year, depending on the species, and can be found buried between the leaf litter and the soil. Most fungal biomass is found in the humus and litter layers of soil. Most truffle fungi produce both asexual spores (mitospores or

conidia A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...

) and sexual spores (meiospores or ascospores/ basidiospores). Conidia can be produced more readily and with less energy than ascospores and can disperse during disturbance events. Production of ascospores is energy intensive because the fungus must allocate resources to the production of large sporocarps. Ascospores are borne within sac-like structures called asci, which are contained within the sporocarp. Because truffle fungi produce their sexual fruiting bodies underground, spores cannot be spread by wind and water. Therefore, nearly all truffles depend on mycophagous animal vectors for spore dispersal. This is analogous to the dispersal of seeds in fruit of angiosperms. When the ascospores are fully developed, the truffle exudes volatile compounds that attract animal vectors. For successful dispersal, these spores must survive passage through the digestive tracts of animals. Ascospores have thick walls composed of

chitin Chitin (carbon, C8hydrogen, H13oxygen, O5nitrogen, N)n ( ) is a long-chain polymer of N-Acetylglucosamine, ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is the second most abundant polysaccharide in nature (behind only cell ...

to help them endure the environment of animal guts. Animal vectors include birds, deer, and rodents such as

vole Voles are small rodents that are relatives of lemmings and hamsters, but with a stouter body; a longer, hairy tail; a slightly rounder head; smaller eyes and ears; and differently formed molars (high-crowned with angular cusps instead of lo ...

s, squirrels, and chipmunks. Many species of trees, such as '' Quercus garryana'', are dependent on the dispersal of sporocarps to inoculate isolated individuals. For example, the acorns of ''Q. garryana'' may be carried to new territory that lacks the necessary mycorrhizal fungi for establishment. Some mycophagous animals depend on truffles as their dominant food source. Flying squirrels, '' Glaucomys sabrinus'', of North America play a three-way symbiosis with truffles and their associated plants. ''G. sabrinus'' is particularly adapted to finding truffles using its refined sense of smell, visual clues, and long-term memory of prosperous populations of truffles. This intimacy between animals and truffles indirectly influences the success of mycorrhizal plant species. After ascospores are dispersed, they remain dormant until germination is initiated by exudates excreted from host plant roots. Following germination, hyphae form and seek out the roots of host plants. Arriving at roots, hyphae begin to form a mantle or sheath on the outer surface of root tips. Hyphae then enter the root cortex intercellularly to form the Hartig net for nutrient exchange. Hyphae can spread to other root tips colonising the entire root system of the host. Over time, the truffle fungus accumulates sufficient resources to form fruiting bodies. Rate of growth is correlated with increasing photosynthetic rates in the spring as trees leaf out.

Nutrient exchange

Truffle fungi receive carbohydrates from their host plants, providing them with valuable micro- and macronutrients. Plant macronutrients include

potassium Potassium is a chemical element; it has Symbol (chemistry), symbol K (from Neo-Latin ) and atomic number19. It is a silvery white metal that is soft enough to easily cut with a knife. Potassium metal reacts rapidly with atmospheric oxygen to ...

phosphorus Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...

nitrogen Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...

, and

sulfur Sulfur ( American spelling and the preferred IUPAC name) or sulphur ( Commonwealth spelling) is a chemical element; it has symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms ...

. In contrast, micronutrients include

iron Iron is a chemical element; it has symbol Fe () and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, forming much of Earth's o ...

copper Copper is a chemical element; it has symbol Cu (from Latin ) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkish-orang ...

zinc Zinc is a chemical element; it has symbol Zn and atomic number 30. It is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodic tabl ...

, and

chloride The term chloride refers to a compound or molecule that contains either a chlorine anion (), which is a negatively charged chlorine atom, or a non-charged chlorine atom covalently bonded to the rest of the molecule by a single bond (). The pr ...

. In truffle fungi, as in all ectomycorrhizae, the majority of nutrient exchange occurs in the Hartig net, the intercellular hyphal network between plant root cells. A unique feature of ectomycorrhizal fungi is the formation of the mantle on the outer surface of fine roots. Truffles have been suggested to co-locate with the orchid species '' Epipactis helleborine'' and '' Cephalanthera damasonium'', though this is not always the case.

Nutrient cycling

Truffle fungi are ecologically important in nutrient cycling. Plants obtain nutrients via their fine roots. Mycorrhizal fungi are much smaller than fine roots, so they have a higher surface area and a greater ability to explore soils for nutrients. Acquisition of nutrients includes the uptake of phosphorus, nitrate or ammonium, iron,

magnesium Magnesium is a chemical element; it has Symbol (chemistry), symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 ...

, and other ions. Many ectomycorrhizal fungi form fungal mats in the upper layers of soils surrounding host plants. These mats have significantly higher carbon and fixed nitrogen concentrations than surrounding soils. Because these mats are nitrogen sinks, leaching of nutrients is reduced. Mycelial mats can also help maintain the structure of soils by holding organic matter in place and preventing

erosion Erosion is the action of surface processes (such as Surface runoff, water flow or wind) that removes soil, Rock (geology), rock, or dissolved material from one location on the Earth's crust#Crust, Earth's crust and then sediment transport, tran ...

. Often, these networks of mycelium provide support for smaller organisms in the soil, such as bacteria and microscopic

arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...

s. Bacteria feed on the exudates released by mycelium and colonise the soil surrounding them. Microscopic arthropods such as mites feed directly on mycelium and release valuable nutrients for the uptake of other organisms. Thus, truffle fungi and other ectomycorrhizal fungi facilitate a complex system of nutrient exchange between plants, animals, and microbes.

Importance in arid-land ecosystems

Plant community structure is often affected by the availability of compatible mycorrhizal fungi. In arid-land ecosystems, these fungi become essential for the survival of their host plants by enhancing the ability to withstand drought. A foundation species in arid-land ecosystems of the Southwest United States is '' Pinus edulis'', commonly known as pinyon pine. ''P. edulis'' associates with the subterranean fungi ''Geopora'' and ''Rhizopogon''. As global temperatures rise, so does the occurrence of severe droughts, detrimentally affecting the survival of aridland plants. This variability in climate has increased the mortality of ''P. edulis''. Therefore, the availability of compatible mycorrhizal inoculum can greatly affect the successful establishment of ''P. edulis'' seedlings. Associated ectomycorrhizal fungi will likely play a significant role in the survival of ''P. edulis'' with continuing global

climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...

Extraction

Because truffles are subterranean, they are often located with the help of an animal (sometimes called a truffler) possessing a refined sense of smell. Traditionally, pigs have been used to extract truffles. Both the female pig's natural truffle-seeking and her intent to eat the truffle were thought to be due to a compound within the truffle similar to androstenol, the sex

pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...

of boar saliva, to which the sow is keenly attracted. Studies in 1990 demonstrated that the compound actively recognised by both truffle pigs and dogs is dimethyl sulfide. In Italy, the use of pigs to hunt truffles has been prohibited since 1985 because of damage caused by animals to truffle

during the digging that dropped the production rate of the area for some years. An alternative to truffle pigs are dogs. Dogs offer an advantage because they do not have a strong desire to eat truffles, so they can be trained to locate sporocarps without digging them up. Pigs attempt to dig up truffles. Fly species of the genus '' Suillia'' can also detect the volatile compounds associated with subterranean fruiting bodies. These flies lay their eggs above truffles to provide food for their young. At ground level, ''Suilla'' flies can be seen flying above truffles.

Volatile constituents

The mycelia or fruiting bodies release the volatile constituents responsible for the natural aroma of truffles or derive from truffle-associated microbes. The chemical ecology of truffle volatiles is complex, interacting with plants, insects, and mammals, which contribute to

spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...

dispersal. Depending on the truffle species, lifecycle, or location, they include: *

Sulfur Sulfur ( American spelling and the preferred IUPAC name) or sulphur ( Commonwealth spelling) is a chemical element; it has symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms ...

volatiles, which occur in all truffle species, such as dimethyl mono- (DMS), di- (DMDS) and tri- (DMTS) sulfides, as well as 2-methyl-4,5-dihydrothiophene, characteristic of the white truffle ''T. borchii'' and 2,4-Dithiapentane occurring in all species but mostly characteristic of the white truffle ''T. magnatum''. Some very aromatic white truffles are notably pungent, even irritating the eye when cut or sliced. * Metabolites of nonsulfur

amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...

constituents (simple and branched-chain hydrocarbons) such as ethylene (produced by mycelia of white truffles affecting root architecture of host tree), as well as 2-methylbutanal, 2-methylpropanal, and 2-phenylethanol (also common in baker's yeast). *

Fatty acid In chemistry, in particular in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated and unsaturated compounds#Organic chemistry, saturated or unsaturated. Most naturally occurring fatty acids have an ...

-derived volatiles (C8-alcohols and aldehydes with a characteristic fungal odor, such as 1-octen-3-ol and 2-octenal). The former is derived from linoleic acid and produced by mature white truffle ''T. borchii''. * Thiophene derivatives appear to be produced by bacterial symbionts living in the truffle body. The most abundant of these, 3-methyl, 4-5 dihydrothiophene, contributes to the white truffle's aroma. Several truffle species and varieties are differentiated based on their relative contents or absence of

sulfide Sulfide (also sulphide in British English) is an inorganic anion of sulfur with the chemical formula S2− or a compound containing one or more S2− ions. Solutions of sulfide salts are corrosive. ''Sulfide'' also refers to large families o ...

s, ethers or

alcohol Alcohol may refer to: Common uses * Alcohol (chemistry), a class of compounds * Ethanol, one of several alcohols, commonly known as alcohol in everyday life ** Alcohol (drug), intoxicant found in alcoholic beverages ** Alcoholic beverage, an alco ...

s, respectively. The sweaty-musky aroma of truffles is similar to that of the

androstenol that also occurs in humans. , the volatile profiles of seven black and six white truffle species have been studied.

Cultivation

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

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

Truffles can be cultivated. As early as 1808, attempts to cultivate truffles, known in French as , were successful. People had long observed that truffles were growing among the roots of certain trees, and in 1808, Joseph Talon, from Apt ( of Vaucluse) in southern

France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...

, had the idea of transplanting some seedlings that he had collected at the foot of oak trees known to host truffles in their root system. For discovering how to cultivate truffles, some sources now give priority to Pierre II Mauléon (1744–1831) of Loudun (in western France), who began to cultivate truffles around 1790. Mauléon saw an "obvious symbiosis" between the oak tree, the rocky soil, and the truffle and attempted to reproduce such an environment by taking acorns from trees known to have produced truffles and sowing them in chalky soil. His experiment was successful, with truffles found in the soil around the newly grown oak trees years later. In 1847, Auguste Rousseau of Carpentras (in Vaucluse) planted of oak trees (again from acorns found on the soil around truffle-producing oak trees), and he subsequently obtained large harvests of truffles. He received a prize at the 1855 World's Fair in

Paris Paris () is the Capital city, capital and List of communes in France with over 20,000 inhabitants, largest city of France. With an estimated population of 2,048,472 residents in January 2025 in an area of more than , Paris is the List of ci ...

Others imitated these successful attempts in France and Italy. In the late 19th century, an epidemic of

phylloxera Grape phylloxera is an insect pest of grapevines worldwide, originally native to eastern North America. Grape phylloxera (''Daktulosphaira vitifoliae'' (Fitch 1855) belongs to the family Phylloxeridae, within the order Hemiptera, bugs); orig ...

destroyed many of the vineyards in southern France. Another epidemic killed most of the silkworms there, too, making the fields of mulberry trees useless. Trufficulture became an important source of income for those affected. The calcareous and exposed vineyard soils were well-suited to the cultivation of truffles. By 1890, truffières (truffle plantations) covered 750 km² of land in France, and 2,000 tonnes of truffles were produced in that year. From the 19th century to the present, truffle production fell by 97–99% to 20–50 tonnes annually. Reasons given for this decline include the

Industrial Revolution The Industrial Revolution, sometimes divided into the First Industrial Revolution and Second Industrial Revolution, was a transitional period of the global economy toward more widespread, efficient and stable manufacturing processes, succee ...

, the subsequent

rural flight Rural flight (also known as rural-to-urban migration, rural depopulation, or rural exodus) is the Human migration, migratory pattern of people from rural areas into urban areas. It is urbanization seen from the rural perspective. In Industriali ...

and the multiple European wars of the 20th century, which reduced the rural population. For example,

World War I World War I or the First World War (28 July 1914 – 11 November 1918), also known as the Great War, was a World war, global conflict between two coalitions: the Allies of World War I, Allies (or Entente) and the Central Powers. Fighting to ...

resulted in the mobilisation of 65% of the agricultural workers from the region of Lot alone. Knowledge of truffle cultivation, the soil and the seasons was lost along with the people. Another consequence was no more grazing sheep or shepherds who pruned trees for feed and fuelwood, so former truffle plantations turned into closed forests that no longer produced truffles. Truffles were once sold at weekly markets (bi-weekly in the case of a market in Martel, Lot) and in quantities of two to six tonnes in good weeks, but only Lalbenque and Limogne today have weekly truffle markets. Prices have increased so that truffles, once seen as a food of the middle class, have become a luxury. The situation changed in the late 1960s and early 1970s, with researchers in France and Italy establishing mycorrhizas with truffle spores. Beginning from the 1980s, truffle plantations are compensating for some of the decline in wild truffles, and exist in various countries including France, Italy, Spain and Australia. About 80% of the truffles now produced in France come from specially planted truffle groves. Investments in cultivated plantations are underway in many parts of the world using controlled irrigation for regular and resilient production. A critical phase of the cultivation is the quality control of the mycorrhizal plants. Between 7 and 10 years are needed for the truffles to develop their mycorrhizal network, and only after that do the host plants come into production. Complete soil analysis to avoid contamination by other dominant fungi and very strict control of the formation of mycorrhizae are necessary to ensure the success of a plantation. Total investment per hectare for an irrigated and barrier-sealed plantation (against wild boars) can cost up to €10,000. Considering the level of initial investment and the maturity delay, farmers who have not taken care of both soil conditions and seedling conditions are at high risk of failure.

New Zealand and Australia

The first black truffles ('' Tuber melanosporum'') to be produced in the Southern Hemisphere were harvested in

Gisborne, New Zealand Gisborne is a List of cities in New Zealand, city in northeastern New Zealand and the largest settlement in the Gisborne District (or Gisborne Region). It has a population of Gisborne District Council has its headquarters in the central city. ...

in 1993. New Zealand's first burgundy truffle was found in July 2012 at a Waipara truffle farm. It weighed 330 g and was found by the farm owner's beagle. In 1999, the first Australian truffles were harvested in

Tasmania Tasmania (; palawa kani: ''Lutruwita'') is an island States and territories of Australia, state of Australia. It is located to the south of the Mainland Australia, Australian mainland, and is separated from it by the Bass Strait. The sta ...

, the result of eight years of work. Trees were inoculated with the truffle fungus to create a local truffle industry. Their success and the value of the resulting truffles has encouraged a small industry to develop. Truffle production has expanded into the colder regions of Victoria,

New South Wales New South Wales (commonly abbreviated as NSW) is a States and territories of Australia, state on the Eastern states of Australia, east coast of :Australia. It borders Queensland to the north, Victoria (state), Victoria to the south, and South ...

and

Western Australia Western Australia (WA) is the westernmost state of Australia. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to the south, the Northern Territory to the north-east, and South Australia to the south-east. Western Aust ...

. In 2014, over of truffles were harvested by Truffle Hill, Manjimup, Western Australia. In June 2014, a grower harvested Australia's largest truffle from their property at Robertson, in the Southern Highlands of New South Wales. It was a French black périgord fungus weighing in at and was valued at over $2,000 per kilogram.

United States

Périgord truffles were first farmed in Tennessee in 2007. At its peak in the 2008–2009 season, one farm produced about 200 pounds of truffles, but Eastern filbert blight almost entirely wiped out the hazel trees by 2013 and production dropped, essentially ending the business. Eastern filbert blight similarly destroyed the orchards of other once-promising commercial farms in East Tennessee, while newer farms in California, North Carolina, Oregon, and Arkansas were started. , the Appalachian truffle ('' Tuber canaliculatum'') was being developed as a potential market.

Uses

Shaved black truffles with pasta - Spago Las Vegas - March 2022 - Sarah Stierch

Because of their high price and their strong aroma, truffles are used sparingly. Supplies can be found commercially as unadulterated fresh produce or preserved, typically in a light brine. Their chemical compounds infuse well with fats such as butter, cream, cheeses, avocados, and coconut cream. As the volatile aromas dissipate quicker when heated, truffles are generally served raw and shaved over warm, simple foods which highlight their flavour, such as buttered

pasta Pasta (, ; ) is a type of food typically made from an Leavening agent, unleavened dough of wheat flour mixed with water or Eggs as food, eggs, and formed into sheets or other shapes, then cooked by boiling or baking. Pasta was originally on ...

or eggs. Thin truffle slices may be inserted into

meat Meat is animal Tissue (biology), tissue, often muscle, that is eaten as food. Humans have hunted and farmed other animals for meat since prehistory. The Neolithic Revolution allowed the domestication of vertebrates, including chickens, sheep, ...

s, under the skins of roasted

fowl Fowl are birds belonging to one of two biological orders, namely the gamefowl or landfowl ( Galliformes) and the waterfowl ( Anseriformes). Anatomical and molecular similarities suggest these two groups are close evolutionary relatives; toget ...

, in ''

foie gras ; (, ) is a specialty food product made of the liver of a Domestic duck, duck or Domestic goose, goose. According to French law, ''foie gras'' is defined as the liver of a duck or goose fattened by ''gavage'' (force feeding). ''Foie gras'' i ...

'' preparations, in ''

pâté Pâté ( , , ) is a forcemeat. Originally, the dish was cooked in a pastry case; in more recent times it is more usually cooked without pastry in a terrine. Various ingredients are used, which may include meat from pork, poultry, fish or bee ...

s'', or in

stuffing Stuffing, filling, or dressing is an edible mixture, often composed of herbs and a Starch#Food, starch such as bread, used to fill a cavity in the preparation of another food item. Many foods may be stuffed, including poultry, seafood, and v ...

s. Some speciality

cheese Cheese is a type of dairy product produced in a range of flavors, textures, and forms by coagulation of the milk protein casein. It comprises proteins and fat from milk (usually the milk of cows, buffalo, goats or sheep). During prod ...

s contain truffles, as well. Truffles are also used for producing truffle salt and truffle honey. While chefs once peeled truffles, in modern times, most restaurants brush the truffle carefully and shave it or dice it with the skin on to make the most of the valuable ingredient. Some restaurants stamp out circular discs of truffle flesh and use the skins for sauces. Truffle olive oil with Tuber melanosporum

Truffle olive oil with Tuber melanosporum

Oil

Truffle oil is used as a lower-cost and convenient substitute for truffles, to provide flavouring, or to enhance the flavour and aroma of truffles in cooking. Some products called "truffle oils" contain no truffles or include pieces of inexpensive, unprized truffle varietals, which have no culinary value, simply for show. The vast majority is oil that has been artificially flavoured using a synthetic agent such as 2,4-dithiapentane. The scientific name is included on the ingredient list of truffle oils infused with natural truffles.

Vodka

Because more aromatic molecules in truffles are soluble in alcohol, they can carry a more complex and accurate truffle flavour than oil without synthetic flavourings. Many commercial producers use 2,4-dithiapentane regardless, as it has become the dominant flavour most consumers, unexposed to fresh truffles but familiar with oils, associate with them. Because most Western nations do not have ingredient labelling requirements for spirits, consumers often do not know if artificial flavourings have been used. It is used as a spirit in its own right, a

cocktail A cocktail is a mixed drink, usually alcoholic beverage, alcoholic. Most commonly, a cocktail is a combination of one or more liquor, spirits mixed with other ingredients, such as juices, flavored syrups, tonic water, Shrub (drink), shrubs, and ...

mix or a food flavouring.

Cultural history

Antiquity

The first mention of truffles appears in the inscriptions of the neo-Sumerians regarding their Amorite enemy's eating habits (

Third Dynasty of Ur The Third Dynasty of Ur or Ur III was a Sumerian dynasty based in the city of Ur in the 22nd and 21st centuries BC ( middle chronology). For a short period they were the preeminent power in Mesopotamia and their realm is sometimes referred to by ...

, 2nd century BCE) and later in writings of

Theophrastus Theophrastus (; ; c. 371 – c. 287 BC) was an ancient Greek Philosophy, philosopher and Natural history, naturalist. A native of Eresos in Lesbos, he was Aristotle's close colleague and successor as head of the Lyceum (classical), Lyceum, the ...

in the 4th century BCE. In classical times, their origins were a mystery that challenged many;

Plutarch Plutarch (; , ''Ploútarchos'', ; – 120s) was a Greek Middle Platonist philosopher, historian, biographer, essayist, and priest at the Temple of Apollo (Delphi), Temple of Apollo in Delphi. He is known primarily for his ''Parallel Lives'', ...

and others thought them to be the result of lightning, warmth, and water in the soil, while

Juvenal Decimus Junius Juvenalis (), known in English as Juvenal ( ; 55–128), was a Roman poet. He is the author of the '' Satires'', a collection of satirical poems. The details of Juvenal's life are unclear, but references in his works to people f ...

thought thunder and rain to be instrumental in their origin.

Cicero Marcus Tullius Cicero ( ; ; 3 January 106 BC – 7 December 43 BC) was a Roman statesman, lawyer, scholar, philosopher, orator, writer and Academic skeptic, who tried to uphold optimate principles during the political crises tha ...

deemed them children of the earth, while

Dioscorides Pedanius Dioscorides (, ; 40–90 AD), "the father of pharmacognosy", was a Greek physician, pharmacologist, botanist, and author of (in the original , , both meaning "On Materia medica, Medical Material") , a 5-volume Greek encyclopedic phar ...

thought they were tuberous roots.

Rome Rome (Italian language, Italian and , ) is the capital city and most populated (municipality) of Italy. It is also the administrative centre of the Lazio Regions of Italy, region and of the Metropolitan City of Rome. A special named with 2, ...

and

Thracia Thracia or Thrace () is the ancient name given to the southeastern Balkans, Balkan region, the land inhabited by the Thracians. Thrace was ruled by the Odrysian kingdom during the Classical Greece, Classical and Hellenistic period, Hellenis ...

in the Classical period identified three kinds of truffles: '' Tuber melanosporum'', '' T. magnificus'', and '' T. magnatum''. The Romans instead used a variety of fungus called terfez, also sometimes called a "desert truffle". Terfez used in Rome came from Lesbos,

Carthage Carthage was an ancient city in Northern Africa, on the eastern side of the Lake of Tunis in what is now Tunisia. Carthage was one of the most important trading hubs of the Ancient Mediterranean and one of the most affluent cities of the classic ...

, and especially

Libya Libya, officially the State of Libya, is a country in the Maghreb region of North Africa. It borders the Mediterranean Sea to the north, Egypt to Egypt–Libya border, the east, Sudan to Libya–Sudan border, the southeast, Chad to Chad–L ...

, where the coastal climate was less dry in ancient times. Their substance is pale, tinged with rose. Unlike truffles, terfez have little inherent flavour. The Romans used the terfez as a flavour carrier because the terfez tends to absorb surrounding flavours. Because Ancient Roman cuisine used many spices and flavourings, the terfez may have been appropriate in that context.

Middle Ages

Truffles were rarely used during the

Middle Ages In the history of Europe, the Middle Ages or medieval period lasted approximately from the 5th to the late 15th centuries, similarly to the post-classical period of global history. It began with the fall of the Western Roman Empire and ...

. Truffle hunting is mentioned by Bartolomeo Platina, the papal historian, in 1481, when he recorded that the sows of Notza were without equal in hunting truffles, but they should be muzzled to prevent them from eating the prize.

Renaissance and modernity

During the

Renaissance The Renaissance ( , ) is a Periodization, period of history and a European cultural movement covering the 15th and 16th centuries. It marked the transition from the Middle Ages to modernity and was characterized by an effort to revive and sur ...

, truffles regained popularity in Europe and were honoured at the court of King

Francis I of France Francis I (; ; 12 September 1494 – 31 March 1547) was King of France from 1515 until his death in 1547. He was the son of Charles, Count of Angoulême, and Louise of Savoy. He succeeded his first cousin once removed and father-in-law Louis&nbs ...

. They were popular in Parisian markets in the 1780s, imported seasonally from truffle grounds, where peasants had long enjoyed them. Brillat-Savarin (1825) noted that they were so expensive they appeared only at the dinner tables of great nobles and kept women. They were sometimes served with turkey.

References

Additional resources

* * * * {{DEFAULTSORT:Truffle (Fungus) Edible fungi French cuisine Fungi in cultivation Italian cuisine Mushroom types Fungus common names

Taxonomic details

Species

Black

Summer or burgundy

White

Whitish

Other ''Tuber''

Beyond ''Tuber''

Phylogeny

Etymology

Nutrient exchange

Nutrient cycling

Importance in arid-land ecosystems

Extraction

Volatile constituents

Cultivation

New Zealand and Australia

United States

Uses

Oil

Vodka

Cultural history

Antiquity

Middle Ages

Renaissance and modernity

See also

References

Additional resources