Black truffles
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''Tuber melanosporum'', called the black truffle, Périgord truffle or French black truffle, is a species of
truffle A truffle is the fruiting body of a subterranean ascomycete fungus, predominantly one of the many species of the genus ''Tuber''. In addition to ''Tuber'', many other genera of fungi are classified as truffles including '' Geopora'', '' Pe ...
native to Southern Europe. It is one of the most expensive edible mushrooms in the world.


Taxonomy

Italian naturalist
Carlo Vittadini Carlo Vittadini (11 June 1800, in Bertonico – 20 November 1865, in Milan) was an Italian doctor and mycologist. Life He studied in Milan and at the University of Pavia, where he attended the classes given by Giuseppe L. Moretti (1782–1 ...
described the black truffle in 1831.


Description


External characteristics

The round, dark brown fruiting bodies (
ascocarp An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mos ...
s) have a black-brown skin with small pyramidal cusps. They have a strong, aromatic smell and normally reach a size of up to . Some may be significantly larger, such as a black truffle found in 2012 in
Dordogne Dordogne ( , or ; ; oc, Dordonha ) is a large rural department in Southwestern France, with its prefecture in Périgueux. Located in the Nouvelle-Aquitaine region roughly half-way between the Loire Valley and the Pyrenees, it is named ...
with a mass of . Their flesh is initially white, then dark. It is permeated by white veins, which turn brown with age. The spores are elliptical and measure about 22–55 
µm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer ( American spelling), also commonly known as a micron, is a unit of length in the International System of Uni ...
by 20–35 µm. They are dark brown and covered with large spikes.


Aroma

The fruiting bodies of the black truffle exude a scent reminiscent of undergrowth, strawberries, wet earth, or dried fruit with a hint of
cocoa Cocoa may refer to: Chocolate * Chocolate * ''Theobroma cacao'', the cocoa tree * Cocoa bean, seed of ''Theobroma cacao'' * Chocolate liquor, or cocoa liquor, pure, liquid chocolate extracted from the cocoa bean, including both cocoa butter an ...
. Their taste, which fully develops after the truffles are heated, is slightly peppery and bitter. If stored at room temperature, the aromatic compounds dissipate, while storage around the freezing point (0°C) leads to an increased synthesis of these compounds. The volatile compounds that contribute to the aroma and are developed by the fruiting bodies include 2-methyl-1-butanol,
isoamyl alcohol Isoamyl alcohol is a colorless liquid with the formula , specifically (H3C–)2CH–CH2–CH2–OH. It is one of several isomers of amyl alcohol (pentanol). It is also known as isopentyl alcohol, isopentanol, or (in the IUPAC recommended nomencla ...
, 2-methylbutyraldehyde, and 3-methylbutyraldehyde, as well as traces of sulfur compounds. One of these,
dimethyl sulfide Dimethyl sulfide (DMS) or methylthiomethane is an organosulfur compound with the formula (CH3)2S. Dimethyl sulfide is a flammable liquid that boils at and has a characteristic disagreeable odor. It is a component of the smell produced from co ...
, is what attracts truffle dogs, truffle hogs and truffle flies to the fruiting bodies. Several species of
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constit ...
, which produce part of the aromatic compounds, have been isolated from ''Tuber melanosporum'' and '' Tuber magnatum''.


Genome

The
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
of the black truffle was published in 2010. It contains 125 million
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both D ...
s, 58% of the genome consists of
transposable element A transposable element (TE, transposon, or jumping gene) is a nucleic acid sequence in DNA that can change its position within a genome, sometimes creating or reversing mutations and altering the cell's genetic identity and genome size. Transp ...
s, and the genome contains only 7500 identified protein-encoding
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
s. During symbiosis, genes involved in the decomposition of plant
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mec ...
s and
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids in ...
s are induced. This indicates that black truffles decompose the cell walls of their host plants at the beginning of the symbiosis.


Identification

The black truffle is morphologically very similar to the commercially less valuable Chinese truffle ('' Tuber indicum''). To avoid fraud or misidentifications in commerce, a RFLP genetic test has been developed to distinguish the two species. Externally, they can be distinguished by their skin, which is smoother and dark red or dark brown in the Chinese truffle. Two other similar truffle species are the summer truffle ('' Tuber aestivum'') and the winter truffle ('' Tuber brumale''), whose flesh is of a lighter color.


Ecology


Development and phenology

Black truffles grow at a depth of to as
ectomycorrhiza An ectomycorrhiza (from Greek ἐκτός ', "outside", μύκης ', "fungus", and ῥίζα ', "root"; pl. ectomycorrhizas or ectomycorrhizae, abbreviated EcM) is a form of symbiotic relationship that occurs between a fungal symbiont, or my ...
e, preferably in loose calcareous soil, close to the roots of their plant
symbiont Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasi ...
s. These include holm oaks,
French oaks The Prix de Diane, sometimes referred to as the French Oaks, is a Group 1 flat horse race in France open to three-year-old thoroughbred fillies. It is run at Chantilly over a distance of 2,100 metre ...
,
hazel The hazel (''Corylus'') is a genus of deciduous trees and large shrubs native to the temperate Northern Hemisphere. The genus is usually placed in the birch family Betulaceae,Germplasmgobills Information Network''Corylus''Rushforth, K. (1999). ...
,
cherry A cherry is the fruit of many plants of the genus ''Prunus'', and is a fleshy drupe (stone fruit). Commercial cherries are obtained from cultivars of several species, such as the sweet ''Prunus avium'' and the sour '' Prunus cerasus''. The n ...
and other deciduous trees. The symbiosis of holm oak saplings and black truffles has been shown to improve
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
and root growth in the plant. Black truffles suppress the growth of plants around their symbiont, creating the impression of a burnt (''brûlé'') area around it. They do so by
parasitizing Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
the roots of other plants, which may lead to
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. Necrosis is caused by factors external to the cell or tissue, such as infection, or trauma which result in the unregulated diges ...
of the root bark and the death of the parasitized plant. Moreover, part of the scent emitted by the truffles may limit the growth of other plants through
oxidative stress Oxidative stress reflects an imbalance between the systemic manifestation of reactive oxygen species and a biological system's ability to readily detoxify the reactive intermediates or to repair the resulting damage. Disturbances in the normal ...
. In the Northern Hemisphere, the fruiting bodies develop from April to June and are harvested from November to March.


Reproduction

Boar The wild boar (''Sus scrofa''), also known as the wild swine, common wild pig, Eurasian wild pig, or simply wild pig, is a suid native to much of Eurasia and North Africa, and has been introduced to the Americas and Oceania. The species is ...
s and the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e of the truffle fly ('' Suillia tuberiperda''), which eat the fruiting bodies, aid in the distribution of the species by excreting the indigestible spores. Their excrement likely also serves to fertilize the spores. Black truffles are sometimes found together with winter truffles, which aid the growth of black truffles in wet soils. Until 2010, all truffle species were thought to be homothallic, that is, capable of
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
from a single organism. Subsequent research indicated that black truffles are heterothallic; that is, sexual reproduction requires contact between the
mycelia Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates in ...
of different
mating type Mating types are the microorganism equivalent to sexes in multicellular lifeforms and are thought to be the ancestor to distinct sexes. They also occur in macro-organisms such as fungi. Definition Mating types are the microorganism equivalent to ...
s. If mycelia of different mating types surround a tree, eventually, one type becomes predominant. This is relevant for the operation of truffle plantations: To improve production, planters must ensure that neighboring trees harbor mycelia of different mating types, such as by inoculating new saplings with a mycelium of a particular type.


Habitat and cultivation

The natural
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
of the black truffle includes various regions in Spain, France, Italy, and Croatia. These are presumably the areas where the host plants found refuge during the last Ice Age. In these areas, the search for black truffles and their cultivation is a tradition going back more than 200 years. Truffles are still collected manually in a traditional way in large areas of natural forests. For example, the county of Alto Maestrazgo (province of Castellón, Spain) has an ideal ground with suitable conditions for cultivating truffles. Albocàsser, Atzaneta, Culla, and Morella are just some of the villages in this region where one can find black truffles in large amounts. Nevertheless, climate change has increasingly affected this form of recollection, and since 2010, a significant drop in productivity has occurred in naturally producing forests. In exchange, cultivated areas are increasingly popular, and in central Spain, several thousands of hectares are dedicated to truffle cultivation (the Mecca of the black truffle being in Sarrión, Teruel province in the Aragon region). Some experiments have also been conducted in burnt areas, with promising results, as legally, no need to ask for a land-use change exists when planting truffles, as it can be considered (EU-28) as forest land. Black truffles are now also cultivated in Australia, New Zealand, Chile, North America, Argentina, South Africa, and Wales. Cultivation involves the planting of, for example, hazel trees whose roots are inoculated with truffle mycelium. The first fruiting bodies can be harvested about 4-10 years after planting the trees.madrimasd
"New technique to grow black truffles"
ScienceDaily, 23 Oct. 2012. Web. 5 Aug. 2013

2 September 2021 at archive.today.


Production

France accounts for around 45% of the world's production of black truffles, Spain 35%, and Italy 20%. Smaller amounts are produced in the United States, South Africa, Slovenia, Croatia, and the Australian states of Tasmania and Western Australia. In 2005, black truffles were found in
Serbia Serbia (, ; Serbian: , , ), officially the Republic of Serbia ( Serbian: , , ), is a landlocked country in Southeastern and Central Europe, situated at the crossroads of the Pannonian Basin and the Balkans. It shares land borders with Hu ...
. About 80% of the French production comes from southeast France: upper
Provence Provence (, , , , ; oc, Provença or ''Prouvènço'' , ) is a geographical region and historical province of southeastern France, which extends from the left bank of the lower Rhône to the west to the Italian border to the east; it is bo ...
('' départements'' of
Vaucluse Vaucluse (; oc, Vauclusa, label= Provençal or ) is a department in the southeastern French region of Provence-Alpes-Côte d'Azur. It had a population of 561,469 as of 2019.Alpes-de-Haute-Provence Alpes-de-Haute-Provence or sometimes abbreviated as AHP (; oc, Aups d'Auta Provença; ) is a department in the Provence-Alpes-Côte d'Azur region of France, bordering Alpes-Maritimes and Italy to the east, Var to the south, Vaucluse to the w ...
), part of
Dauphiné The Dauphiné (, ) is a former province in Southeastern France, whose area roughly corresponded to that of the present departments of Isère, Drôme and Hautes-Alpes. The Dauphiné was originally the Dauphiné of Viennois. In the 12th centu ...
(''département'' of
Drôme Drôme (; Occitan: ''Droma''; Arpitan: ''Drôma'') is the southernmost department in the Auvergne-Rhône-Alpes region of Southeastern France. Named after the river Drôme, it had a population of 516,762 as of 2019.
), and part of
Languedoc The Province of Languedoc (; , ; oc, Lengadòc ) is a former province of France. Most of its territory is now contained in the modern-day region of Occitanie in Southern France. Its capital city was Toulouse. It had an area of approximately ...
(''département'' of
Gard Gard () is a department in Southern France, located in the region of Occitanie. It had a population of 748,437 as of 2019;Quercy Quercy (; oc, Carcin , locally ) is a former province of France located in the country's southwest, bounded on the north by Limousin, on the west by Périgord and Agenais, on the south by Gascony and Languedoc, and on the east by Rouergue and ...
(''département'' of Lot) and
Périgord Périgord ( , ; ; oc, Peiregòrd / ) is a natural region and former province of France, which corresponds roughly to the current Dordogne department, now forming the northern part of the administrative region of Nouvelle-Aquitaine. It is div ...
. The largest truffle market in France (and probably also in the world) is at
Richerenches Richerenches (; oc, Richarenchas) is a commune in the Vaucluse department in the Provence-Alpes-Côte d'Azur region in southeastern France. Truffles The village is famous for its truffle market as well as for its ''messe aux truffes'' in the ch ...
in Vaucluse. The largest truffle market in southwest France is at
Lalbenque Lalbenque (; oc, L'Albenca) is a commune in the Lot department in south-western France. See also *Communes of the Lot department The following is a list of the 313 communes An intentional community is a voluntary residential community w ...
in
Quercy Quercy (; oc, Carcin , locally ) is a former province of France located in the country's southwest, bounded on the north by Limousin, on the west by Périgord and Agenais, on the south by Gascony and Languedoc, and on the east by Rouergue and ...
. These markets are busiest in January when the black truffles have their highest perfume. Production has considerably diminished during the 21st century, dropping to around 20 metric tonnes per year, with peaks of about 46 tonnes in the best years. By comparison, in 1937, France produced about 1,000 metric tonnes of black truffles. The following table shows the production in the EU of'' T. melanosporum'' in Spain, France, and Italy. Production data are in metric tonnes and country weights in percentage and come from the Groupe Européen Truffe et Trufficulture, an association of the leading European producers. As the data show, France has been the leading producer of black truffles in the last decade and a half but is rapidly challenged by Spain, where regions have made use of the EU-funded Rural Development Programme to subsidise cultivated plantations. This is particularly visible in the
Teruel Teruel () is a city in Aragon, located in eastern Spain, and is also the capital of Teruel Province. It has a population of 35,675 in 2014 making it the least populated provincial capital in the country. It is noted for its harsh climate, with ...
province of the
Aragón Aragon ( , ; Spanish and an, Aragón ; ca, Aragó ) is an autonomous communities of Spain, autonomous community in Spain, coextensive with the medieval Kingdom of Aragon. In northeastern Spain, the Aragonese autonomous community comprises th ...
region, where the black truffle represents the first and main economic activity (in GDP and employment), especially since 2010-2011 when many plantations opened under the last Rural Development Programme 2000-2006 came into production phase.


Commerce and use

With a price of about 1,000 to 2,000 euros per kilogram, black truffles are the second-most expensive truffles after
white truffle ''Tuber magnatum'', the white truffle (Italian language, Italian: ), is a species of truffle in the order Pezizales and family Tuberaceae. It is found in southern Europe. Distribution It is found mainly in the Langhe and Montferrat areas of the ...
s and one of the most sought-after edible mushrooms in the world. In cooking, black truffles are used to refine the taste of meat, fish, soups, cheeses, and
risotto Risotto (, , from meaning "rice") is a northern Italian rice dish cooked with broth until it reaches a creamy consistency. The broth can be derived from meat, fish, or vegetables. Many types of risotto contain butter, onion, white wine, and Pa ...
. Unlike white truffles, the aroma of black truffles does not diminish when they are heated but becomes more intense. They are most commonly shaved into or on top of a dish raw or infused with high-quality olive oil or butter.


Physiology

Truffles contain the
endocannabinoid Cannabinoids () are several structural classes of compounds found in the cannabis plant primarily and most animal organisms (although insects lack such receptors) or as synthetic compounds. The most notable cannabinoid is the phytocannabinoid tet ...
anandamide Anandamide (ANA), also known as ''N''-arachidonoylethanolamine (AEA), is a fatty acid neurotransmitter. Anandamide was the first endocannabinoid to be discovered: it participates in the body's endocannabinoid system by binding to cannabinoid r ...
(AEA) and the major metabolic enzymes of the
endocannabinoid system The endocannabinoid system (ECS) is a biological system composed of endocannabinoids, which are endogenous lipid-based retrograde neurotransmitters that bind to cannabinoid receptors (CBRs), and cannabinoid receptor proteins that are expressed ...
(ECS). The AEA content increases in the late stages of truffles’ development. AEA and ECS metabolic enzymes may have evolved earlier than endocannabinoid-binding receptors, and AEA might be an ancient attractant to truffle-eating animals, which are well-equipped with endocannabinoid-binding receptors.


References


External links

* ' {{Taxonbar, from=Q1093314 melanosporum Truffles (fungi) Fungi described in 1831