What kind of an organism is a truffle?
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Organism Truffle
In biology, an organism is any contiguous living system, such as an animal, plant, fungus, archaeon, or bacterium. All known types of organisms are capable of some degree of response to stimuli, reproduction, growth and development and homeostasis. An organism consists of one or more cells; when it has one cell it is known as a unicellular organism; and when it has more than one it is known as a multicellular organism. Most unicellular organisms are of microscopic size and are thus classified as microorganisms. Humans are multicellular organisms composed of many trillions of cells grouped into specialized tissues and organs.

An organism  may be either a prokaryote or a eukaryote. Prokaryotes are represented by two separate domains, the Bacteria and Archaea. Eukaryotic organisms are characterized by the presence of a membrane-bound cell nucleus and contain additional membrane-bound compartments called organelles (such as mitochondria in animals and plants and plastids in plants and algae, all generally considered to be derived from endosymbiotic bacteria). Fungi, animals and plants are examples of kingdoms of organisms within the eukaryotes.

Estimates on the number of Earth's current species range from 10 million to 14 million,  of which only about 1.2 million have been documented.  More than 99% of all species, amounting to over five billion species,  that ever lived on Earth are estimated to be extinct.   In July 2016, scientists reported identifying a set of 355 genes from the Last Universal Common Ancestor (LUCA) of all organisms living on Earth. 

Etymology 

The term "organism" (from Greek ὀργανισμός, organismos, from ὄργανον, organon, i.e. "instrument, implement, tool, organ of sense or apprehension") first appeared in the English language in 1703 and took on its current definition by 1834 (Oxford English Dictionary). It is directly related to the term "organization". There is a long tradition of defining organisms as self-organizing beings. 

There has been controversy about the best way to define the organism         and indeed about whether or not such a definition is necessary.   Several contributions  are responses to the suggestion that the category of "organism" may well not be adequate in biology. 

Semantics

The word organism may broadly be defined as an assembly of molecules functioning as a more or less stable whole that exhibits the properties of life. However, many sources propose definitions that exclude viruses and theoretically possible man-made non-organic life forms. Viruses are dependent on the biochemical machinery of a host cell for reproduction.

Chambers Online Reference provides a broad definition: "any living structure, such as a plant, animal, fungus or bacterium, capable of growth and reproduction".

In multicellular terms, "organism" usually describes the whole hierarchical assemblage of systems (for example circulatory, digestive, or reproductive) themselves collections of organs; these are, in turn, collections of tissues, which are themselves made of cells. In some plants and the nematode Caenorhabditis elegans, individual cells are totipotent.

A superorganism is an organism consisting of many individuals working together as a single functional or social unit.

Non-cellular life 

Viruses are not typically considered to be organisms because they are incapable of autonomous reproduction, growth or metabolism. This controversy is problematic because some cellular organisms are also incapable of independent survival (but are capable of independent metabolism and procreation) and live as obligatory intracellular parasites. Although viruses have a few enzymes and molecules characteristic of living organisms, they have no metabolism of their own; they cannot synthesize and organize the organic compounds from which they are formed. Naturally, this rules out autonomous reproduction: they can only be passively replicated by the machinery of the host cell. In this sense they are similar to inanimate matter. While viruses sustain no independent metabolism, and thus are usually not classified as organisms, they do have their own genes, and they do evolve by mechanisms similar to the evolutionary mechanisms of organisms.

The most common argument in support of viruses as living organisms is their ability to undergo evolution and replicate through self-assembly. Some scientists argue that viruses neither evolve, nor self- reproduce. In fact, viruses are evolved by their host cells, meaning that there was co-evolution of viruses and host cells. If host cells did not exist, viral evolution would be impossible. This is not true for cells. If viruses did not exist, the direction of cellular evolution could be different, but cells would nevertheless be able to evolve. As for the reproduction, viruses totally rely on hosts' machinery to replicate. The discovery of viral megagenomes with genes coding for energy metabolism and protein synthesis fueled the debate about whether viruses belong in the tree of life. The presence of these genes suggested that viruses were once able to metabolize. However, it was found later that the genes coding for energy and protein metabolism have cellular origin. Most likely, these genes were acquired through horizontal gene transfer from viral hosts.

Organizational terminology

All organisms are classified by the science of alpha taxonomy into either taxa or clades.

Taxa are ranked groups of organisms, which run from the general (domain) to the specific (species). A broad scheme of ranks in hierarchical order is:

# Domain
# Kingdom
# Phylum
# Class
# Order
# Family
# Genus
# Species

To give an example, Homo sapiens is the Latin binomial equating to modern "humans". All members of the species sapiens are, at least in theory, genetically able to interbreed. Several species may belong to a genus, but the members of different species within a genus are usually unable to interbreed to produce fertile offspring. (The red wolf Canis lupus rufus, however, may be a result of interbreeding between the grey wolf Canis lupus and the coyote Canis latrans.) Homo only has one surviving species (sapiens), Homo erectus, Homo neanderthalensis, etc. having become extinct thousands of years ago; some scientists argue for interbreeding between H. sapiens and H. neanderthalensis with fertile progeny. Several genera belong to the same family and so on up the hierarchy. Eventually, the relevant kingdom (Animalia, in the case of humans) is placed into one of the three domains depending upon certain genetic and structural characteristics.

All living organisms known to science are given classification by this system such that the species within a particular family are more closely related and genetically similar than the species within a particular phylum.

Since viruses are not living organisms, their classification is a challenging task. At first, viruses were classified according to their hosts: plant viruses, animal viruses, bacteriophages. Later, they were classified by the disease that they cause. For example, respiratory viruses, enterics. Now, viruses are classified based on the nucleic acid content, capsid symmetry and the presence or absence of the envelope.

Chemistry

Organisms are complex chemical systems, organized in ways that promote reproduction and some measure of sustainability or survival. The same laws that govern non-living chemistry govern the chemical processes of life. It is generally the phenomena of entire organisms that determine their fitness to an environment and therefore the survivability of their DNA-based genes.

Organisms clearly owe their origin, metabolism, and many other internal functions to chemical phenomena, especially the chemistry of large organic molecules. Organisms are complex systems of chemical compounds that, through interaction and environment, play a wide variety of roles.

Organisms are semi-closed chemical systems. Although they are individual units of life (as the definition requires), they are not closed to the environment around them. To operate they constantly take in and release energy. Autotrophs produce usable energy (in the form of organic compounds) using light from the sun or inorganic compounds while heterotrophs take in organic compounds from the environment.

The primary chemical element in these compounds is carbon. The chemical properties of this element such as its great affinity for bonding with other small atoms, including other carbon atoms, and its small size making it capable of forming multiple bonds, make it ideal as the basis of organic life. It is able to form small three-atom compounds (such as carbon dioxide), as well as large chains of many thousands of atoms that can store data (nucleic acids), hold cells together, and transmit information (protein).

Macromolecules

Compounds that make up organisms may be divided into macromolecules and other, smaller molecules. The four groups of macromolecule are nucleic acids, proteins, carbohydrates and lipids. Nucleic acids (specifically deoxyribonucleic acid, or DNA) store genetic data as a sequence of nucleotides. The particular sequence of the four different types of nucleotides (adenine, cytosine, guanine, and thymine) dictate many characteristics that constitute the organism. The sequence is divided up into codons, each of which is a particular sequence of three nucleotides and corresponds to a particular amino acid. Thus a sequence of DNA codes for a particular protein that, due to the chemical properties of the amino acids it is made from, folds in a particular manner and so performs a particular function.

These protein functions have been recognized:
# Enzymes, which catalyze all of the reactions of metabolism
# Structural proteins, such as tubulin, or collagen
# Regulatory proteins, such as transcription factors or cyclins that regulate the cell cycle
# Signaling molecules or their receptors such as some hormones and their receptors
# Defensive proteins, which can include everything from antibodies of the immune system, to toxins (e.g., dendrotoxins of snakes), to proteins that include unusual amino acids like canavanine

A bilayer of phospholipids makes up the membrane of cells that constitutes a barrier, containing everything within the cell and preventing compounds from freely passing into, and out of, the cell. Due to the selective permeability of the phospholipid membraine only specific compounds can pass through it. In some multicellular organisms they serve as a storage of energy and mediate communication between cells. Carbohydrates are more easily broken down than lipids and yield more energy to compare to lipids and proteins. In fact, carbohydrates are the number one source of energy for all living organisms.

Structure

All organisms consist of monomeric units called cells; some contain a single cell (unicellular) and others contain many units (multicellular). Multicellular organisms are able to specialize cells to perform specific functions. A group of such cells is a tissue, and in animals these occur as four basic types, namely epithelium, nervous tissue, muscle tissue, and connective tissue. Several types of tissue work together in the form of an organ to produce a particular function (such as the pumping of the blood by the heart, or as a barrier to the environment as the skin). This pattern continues to a higher level with several organs functioning as an organ system to allow for reproduction, digestion, etc. Many multicellular organisms consist of several organ systems, which coordinate to allow for life.

Cell

The cell theory, first developed in 1839 by Schleiden and Schwann, states that all organisms are composed of one or more cells; all cells come from preexisting cells; all vital functions of an organism occur within cells, and cells contain the hereditary information necessary for regulating cell functions and for transmitting information to the next generation of cells.

There are two types of cells, eukaryotic and prokaryotic. Prokaryotic cells are usually singletons, while eukaryotic cells are usually found in multicellular organisms. Prokaryotic cells lack a nuclear membrane so DNA is unbound within the cell; eukaryotic cells have nuclear membranes.

All cells, whether prokaryotic or eukaryotic, have a membrane, which envelops the cell, separates its interior from its environment, regulates what moves in and out, and maintains the electric potential of the cell. Inside the membrane, a salty cytoplasm takes up most of the cell volume. All cells possess DNA, the hereditary material of genes, and RNA, containing the information necessary to build various proteins such as enzymes, the cell's primary machinery. There are also other kinds of biomolecules in cells.

All cells share several similar characteristics of: 
* Reproduction by cell division (binary fission, mitosis or meiosis).
* Use of enzymes and other proteins coded by DNA genes and made via messenger RNA intermediates and ribosomes.
* Metabolism, including taking in raw materials, building cell components, converting energy, molecules and releasing by-products. The functioning of a cell depends upon its ability to extract and use chemical energy stored in organic molecules. This energy is derived from metabolic pathways.
* Response to external and internal stimuli such as changes in temperature, pH or nutrient levels.
* Cell contents are contained within a cell surface membrane that contains proteins and a lipid bilayer.

Life span

One of the basic parameters of an organism is its life span. Some organisms live as short as one day, while some plants and fungi can live thousands of years.

Evolution

Last universal ancestor

The last universal ancestor is the most recent organism from which all organisms now living on Earth descend.  Thus it is the most recent common ancestor (MRCA) of all current life on Earth. The LUA is estimated to have lived some 3.5 to 3.8 billion years ago (sometime in the Paleoarchean era).   The earliest evidences for life on Earth are graphite found to be biogenic in 3.7 billion-year-old metasedimentary rocks discovered in Western Greenland  and microbial mat fossils found in 3.48 billion-year-old sandstone discovered in Western Australia.   Although more than 99 percent of all species that ever lived on the planet are estimated to be extinct, there are currently 10–14 million species of life on Earth.

Information about the early development of life includes input from many different fields, including geology and planetary science. These sciences provide information about the history of the Earth and the changes produced by life. However, a great deal of information about the early Earth has been destroyed by geological processes over the course of time.

All organisms on Earth are descended from a common ancestor or ancestral gene pool. Evidence for common descent may be found in traits shared between all living organisms. In Darwin's day, the evidence of shared traits was based solely on visible observation of morphologic similarities, such as the fact that all birds have wings, even those that do not fly.

Today, there is strong evidence from genetics that all organisms have a common ancestor. For example, every living cell makes use of nucleic acids as its genetic material, and uses the same twenty amino acids as the building blocks for proteins. All organisms use the same genetic code (with some extremely rare and minor deviations) to translate nucleic acid sequences into proteins. The universality of these traits strongly suggests common ancestry, because the selection of many of these traits seems arbitrary.

In 2000, W. Ford Doolittle  discussed variability in the genetic code (see: genetic code variation) and suggested that horizontal gene transfer might make it difficult to study the last universal ancestor. In 2010, a formal mathematical test calculated from the genetic record (and in particular the universal use of the same genetic code, same nucleotides, and same amino acids), that the factor in favor of a common ancestor's existence is 10^2489. 

Reproduction

Sexual reproduction (sex) is widespread among current eukaryotes. Dacks and Rogers  proposed, on the basis of a phylogenetic analysis, that sex was present in the common ancestor of all eukaryotes.  The finding of a core set of genes necessary for meiosis in the descendants of lineages that diverged early form the eukaryotic evolutionary tree also led Ramesh et al.  and Malik et al.  to suggest that sex was present in a common ancestor of all eukaryotes. Furthermore, this view is supported by evidence that eukaryotes previously regarded as "ancient asexuals", such as amoeba, were likely sexual in the past, and that most present day asexual amoeboid lineages likely arose recently and independently. 

In prokaryotes, natural bacterial transformation involves the transfer of DNA from one bacterium to another and integration of the donor DNA into the recipient chromosome by recombination. Natural bacterial transformation is considered to be a primitive sexual process and occurs in both bacteria and archaea, although it has been studied mainly in bacteria. Transformation is clearly a bacterial adaptation and not an accidental occurrence, because it depends on numerous gene products that specifically interact with each other to enter a state of natural competence to perform this complex process.  Transformation is a common mode of DNA transfer, and over 60 prokaryotic species are known to be naturally competent for transformation. 

History of life

The chemical evolution from self-catalytic chemical reactions to life (see Origin of life) is not a part of biological evolution, but it is unclear at which point such increasingly complex sets of reactions became what we would consider, today, to be living organisms.

Little is known about the earliest developments in life. However, all existing organisms share certain traits, including cellular structure and genetic code. Most scientists interpret this to mean all existing organisms share a common ancestor, which had already developed the most fundamental cellular processes, but there is no scientific consensus on the relationship of the three domains of life (Archaea, Bacteria, Eukaryota) or the origin of life. Attempts to shed light on the earliest history of life generally focus on the behavior of macromolecules, particularly RNA, and the behavior of complex systems.

The emergence of oxygenic photosynthesis (around 3 billion years ago) and the subsequent emergence of an oxygen-rich, non-reducing atmosphere can be traced through the formation of banded iron deposits, and later red beds of iron oxides. This was a necessary prerequisite for the development of aerobic cellular respiration, believed to have emerged around 2 billion years ago.

In the last billion years, simple multicellular plants and animals began to appear in the oceans. Soon after the emergence of the first animals, the Cambrian explosion (a period of unrivaled and remarkable, but brief, organismal diversity documented in the fossils found at the Burgess Shale) saw the creation of all the major body plans, or phyla, of modern animals. This event is now believed to have been triggered by the development of the Hox genes. About 500 million years ago, plants and fungi colonized the land, and were soon followed by arthropods and other animals, leading to the development of today's land ecosystems.

The evolutionary process may be exceedingly slow. Fossil evidence indicates that the diversity and complexity of modern life has developed over much of the history of the earth. Geological evidence indicates that the Earth is approximately 4.6 billion years old. Studies on guppies by David Reznick at the University of California, Riverside, however, have shown that the rate of evolution through natural selection can proceed 10 thousand to 10 million times faster than what is indicated in the fossil record.  Such comparative studies however are invariably biased by disparities in the time scales over which evolutionary change is measured in the laboratory, field experiments, and the fossil record.

Horizontal gene transfer

The ancestry of living organisms has traditionally been reconstructed from morphology, but is increasingly supplemented with phylogenetics—the reconstruction of phylogenies by the comparison of genetic (DNA) sequence.
Sequence comparisons suggest recent horizontal transfer of many genes among diverse species including across the boundaries of phylogenetic "domains". Thus determining the phylogenetic history of a species can not be done conclusively by determining evolutionary trees for single genes. 

Biologist Gogarten suggests "the original metaphor of a tree no longer fits the data from recent genome research", therefore "biologists (should) use the metaphor of a mosaic to describe the different histories combined in individual genomes and use (the) metaphor of a net to visualize the rich exchange and cooperative effects of HGT among microbes." 

Future of life (cloning and synthetic organisms)

In modern terms, the category of organism cloning refers to the procedure of creating a new multicellular organism, genetically identical to another. However, the techniques used in cloning have the potential of creating entirely new species of organisms. Organism cloning is the subject of much ethical debate. (see Bioethics, Ethics of cloning, and Designer baby articles)

In 2008 the J. Craig Venter Institute assembled a synthetic bacterial genome, Mycoplasma genitalium, by using recombination in yeast of 25 overlapping DNA fragments in a single step. The use of yeast recombination greatly simplifies the assembly of large DNA molecules from both synthetic and natural fragments. Other companies, such as Synthetic Genomics, have already been formed to take advantage of the many commercial uses of custom designed genomes. A truffle is the fruiting body of a subterranean Ascomycete fungus, predominantly one of the many species of the genus Tuber. Truffles are ectomycorrhizal fungi and are therefore usually found in close association with the roots of trees. Spore dispersal is accomplished through fungivores, animals that eat fungi.

Some of the truffle species are highly prized as food. French gourmet Jean Anthelme Brillat-Savarin called truffles "the diamond of the kitchen".  Edible truffles are held in high esteem in Croatian, French, Georgian, Greek, Italian, Middle Eastern, and Spanish cooking, as well as in international .

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, 20th century BCE)  and later in writings of Theophrastus in the fourth century BCE. In classical times, their origins were a mystery that challenged many; Plutarch and others thought them to be the result of lightning, warmth and water in the soil, while Juvenal thought thunder and rain to be instrumental in their origin. Cicero deemed them children of the earth, while Dioscorides thought they were tuberous roots.

Italy in the Classical period produced three kinds of truffles: the , the  and the . The Romans, however, only used the terfez (Terfezia bouderi), a fungus of similar appearance, which the Romans called truffles, and which is sometimes called "desert truffle". Terfez used in Rome came from Lesbos, Carthage, and especially Libya, where the coastal climate was less dry in ancient times. Their substance is pale, tinged with rose. Unlike truffles, terfez have no taste of their own. The Romans used the terfez as a carrier of flavour, because the terfez have the property to absorb surrounding flavours. Indeed, Ancient Roman cuisine used many spices and flavours, and terfez were perfect in that context.

Middle Ages 

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

According to a hadith narrated by Saeed bin Zaid, Muhammad said that truffles are like manna and that water from truffles "heals eye diseases". 

Renaissance and modern times 

During the Renaissance, truffles regained popularity in Europe and were honoured at the court of King Francis I of France. However, it was not until the 17th century that Western (and in particular French) cuisine abandoned "heavy" oriental spices, and rediscovered the natural flavour of foodstuffs. Truffles were very popular in Parisian markets in the 1780s. They were imported seasonally from truffle grounds, where peasants had long enjoyed their secret. Brillat-Savarin (1825) noted characteristically that they were so expensive they appeared only at the dinner tables of great nobles and kept women. A great delicacy was a truffled turkey.

Cultivation 

Truffles long eluded techniques of domestication, 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.

However, truffles can be cultivated.  As early as 1808, there were successful attempts to cultivate truffles, known in French as . People had long observed that truffles were growing among the roots of certain trees, and in 1808, Joseph Talon, from Apt (département of Vaucluse) in southern France, had the idea of transplanting some seedlings that he'd 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 the Vienne department), who began to cultivate truffles ca. 1790.  See:  Thérèse Dereix de Laplane (2010) "Des truffes sauvages aux truffes cultivées en Loudunais" (From wild truffles to cultivated truffles in the area of Loudun), Mémoires de l’Académie des Sciences, Arts et Belles-Lettres de Touraine, 23 :  215–241.  Available on-line at:  [http://academie-de-touraine.com/Tome_23_files/191_215-241.pdf Academy of Touraine]  From pp. 224–225:  " … le paysan, a alors l'idée, vers 1790 — puisqu'il y a symbiose évidente entre le chêne, les galluches et la truffe — de provoquer la formation de truffières, en reproduisant leur environnement naturel par des semis de glands dans ses "terres galluches".  Avec "les glands venus sur les chênes donnant les truffes, des semis furent faits dans les terrains calcaires voisins" … " ( … the farmer [viz, Pierre II Mauléon] then had the idea, around 1790 — because there is an obvious symbiosis between the oak tree, the rocky soil, and truffles — of inducing the formation of truffle patches, by reproducing their natural environment by sowing acorns in his rocky soils.  With "the acorns [that] came from the oak trees producing truffles, sowings were made in the neighboring chalky plots" … )  See also:  Léon de la Toubette (1869) [https://books.google.com/books?id97oAAAAAYAAJ&pg
RA1-PA300#vonepage&q&f
false "Culture de la truffe à Loudun et à Richelieu,"] Annales de la Société d'Agriculture Sciences, Arts, et Belles-lettres du Départment d'Indre-et-Loire, 10th series, 48 :  300–302 ; see p. 300.

The experiment was successful: Years later, truffles were found in the soil around the newly grown oak trees. In 1847, Auguste Rousseau of Carpentras (in Vaucluse) planted 7 ha 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. 

These successful attempts were met with enthusiasm in southern France, which possessed the sweet limestone soils and dry, hot weather that truffles need to grow. In the late 19th century, an epidemic of phylloxera destroyed many of the vineyards in southern France. Another epidemic destroyed most of the silkworms there, too, making the fields of mulberry trees useless. Thus, large tracts of land were set free for the cultivation of truffles. Thousands of truffle-producing trees were planted, and production reached peaks of hundreds of tonnes at the end of the 19th century. In 1890, there were 75000 ha of truffle-producing trees.

In the 20th century, however, with the growing industrialization of France and the subsequent rural exodus, many of these truffle fields ( or ) returned to wilderness. The First World War also dealt a serious blow to the French countryside, killing 20% or more of the male working force. As a consequence, newly acquired techniques of  were lost. Also, between the two world wars, the truffle groves planted in the 19th century stopped being productive. (The average life cycle of a truffle-producing tree is 30 years.) Consequently, after 1945, the production of truffles plummeted, and the prices have risen dramatically. In 1900, truffles were used by most people, and on many occasions. Today, they are a rare delicacy reserved for the rich, or used on very special occasions.

In the last 30 years, new attempts for mass production of truffles have been started. Eighty percent of the truffles now produced in France come from specially planted truffle groves. Nonetheless, production has yet to recover its 1900s peaks. Local farmers are sometimes opposed to a return of mass production, which would possibly decrease the price of truffles (though it is commonly stated that demand is 10 times higher than offer). In exchange there are heavy investments in cultivated plantations under way in many parts of the world. Thanks to controlled irrigation, regular and resilient production is indeed possible.   There are now truffle-growing areas in the United Kingdom, United States, Spain, Sweden, New Zealand, Australia, Chile and South Africa.

A critical phase of the cultivation is the quality control of the michorized plants. It takes indeed between 7 and 10 years for the truffles to develop their micchorizes network, and only after that the host-plants come into production. Both a complete soil analysis to avoid contamination by other dominant fungus and a very strict control of the michorization process 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.  Regarding the level of initial investment and the maturity delay, farmers that have not taken care of both soil conditions and seedlings conditions are at high risk of failure.

In New Zealand and Australia 

The first black truffles () to be produced in the Southern Hemisphere were harvested in Gisborne, New Zealand, in 1993. 

In 1999, the first Australian truffles were harvested in Tasmania,  the result of eight years of work. Trees were inoculated with the truffle fungus in the hope of creating a local truffle industry. Their success and the value of the resulting truffles has encouraged a small industry to develop. A Western Australian venture, The Wine and Truffle Company, had its first harvest in 2004, and in 2005 they unearthed a 1-kg (2.2-lb) truffle. In 2008, an estimated 600 kg of truffles were removed from the rich ground of Manjimup. Each year, The Wine and Truffle Company has expanded its production, moving into the colder regions of Victoria and New South Wales.

In June 2010, Tasmanian growers Michael and Gwynneth Williams harvested Australia's largest truffle from their property at Myrtle Bank, near Launceston. It weighed in at .  Mrs. Williams told ABC Radio in Australia  that it is valued at about A$1,500 per kg.

New Zealand's first burgundy truffle was found in July 2012 at a Waipara truffle farm. It weighed 330 g, and was found by Rosie, the farm owner's beagle. 

Etymology 

The origin of the word truffle appears to be the Latin term , meaning "swelling" or "lump", which became tufer- and gave rise to the various European terms: Croatian , Danish ', Dutch , English truffle, French , German , Italian , Polish , Spanish , Swedish and .

The German word  ("potato") is derived from the Italian term for truffle because of superficial similarities.  In Portuguese, the words  and  are synonyms, the latter closer to the Latin term.

Biology 

The mycelia of truffles form symbiotic, mycorrhizal relationships with the roots of several tree species including beech, birch, hazel, hornbeam, oak, pine, and poplar.   They prefer argillaceous or calcareous soils that are well drained and neutral or alkaline.   Truffles fruit throughout the year, depending on the species, and can be found buried between the leaf litter and the soil.

Phylogeny 

The phylogeny and biogeography of truffles was investigated in 2008  using internal transcribed spacers (ITS) of nuclear DNA with five major clades (Aestivum, Excavatum, Rufum, Melanosporum and Puberulum); this was later improved and expanded in 2010 using large subunits (LSU) of mitochondrial DNA to nine major clades.  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. 

Species 

Black truffle 

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

The following table shows the production in the EU of Tuber melanosporum in Spain, France and Italy. Production data is in metric tons, and country weights in percentage, and comes from the Groupe Européen Truffe et Trufficulture, an association of the main producers in Europe.

As the data shows, France has been the main producer of black truffles in the last century and a half, but is rapidly challenged by Spain, where regions have made use of the EU-funded Rural Development Programme (EARFD funding) to subsidise cultivated plantations. This is particularly visible in the Teruel province of the Aragón region where the black truffle represents the first and main economic activity (in GDP and employment). Especially since 2010-2011 when many plantations done under the last Rural Development Programme 2000-2006 came into production phase.

Summer or burgundy truffle 

The black summer truffle () is found across Europe and is prized for its culinary value. Burgundy truffles (Tuber uncinatum) are harvested in autumn until December and have aromatic flesh of a darker colour.

White truffle 

The "white truffle" or "trifola d'Alba Madonna" ("Truffle of the White Mother" in Italian) (Tuber magnatum) is found mainly in the Langhe and Montferrat areas  of the Piedmont region in northern Italy and, most famously, in the countryside around the cities of Alba and Asti;  in Italy it can also be found in Molise, Abruzzo, and in the hills around San Miniato, in Tuscany. It is also found on the Istria peninsula, in Croatia in the Motovun forest along the Mirna river,   and in Slovenia along the Dragonja and Rizana river,  as well as in the Drome area in France. Growing symbiotically with oak, hazel, poplar and beech and fruiting in autumn, they can reach 12 cm diameter and 500 g, though are usually much smaller. The flesh is pale cream or brown with white marbling. Italian white truffles are very highly esteemed (illustration, left) and are the most valuable on the market: The white truffle market in Alba is busiest in the months of October and November when the Fiera del Tartufo (truffle fair) takes place. In 2001, the Tuber magnatum truffles sold for between $1000–$2200 per pound ($2000–$4500 per kg);  as of December 2009 they were being sold at $14,203.50 per kilogram.

In 1999, Giancarlo Zigante and his dog Diana found one of the largest truffles in the world near Buje, Croatia. The truffle weighed  and has entered the Guinness Book of Records. 

The record price paid for a single white truffle was set in December 2007, when Macau casino owner Stanley Ho paid $330,000 (£165,000) for a specimen weighing , discovered by Luciano Savini and his dog Rocco. One of the largest truffles found in decades, it was unearthed near Pisa, Italy, and sold at an auction held simultaneously in Macau, Hong Kong, and Florence.  This record was then matched on November 27, 2010 when Ho again paid $330,000 for a pair of white truffles, including one weighing nearly a kilogram.

In December 2014, a White Alba’s Truffle weighing in at 4.16 pounds or 1.89 kilos was unearthed in the Umbrian region of Italy by the Balestra Family of Sabatino. It was auctioned at Sotheby’s in New York.  While some had expected it to sell for $1 million,  it was sold for $61,000 to a Taiwanese buyer. 

A variety of white truffle (Tuber magnatum pico) is found in other parts of northern and central Italy, but is as aromatic as those from Piedmont.

Whitish truffle

The "whitish truffle" (Tuber borchii) is a similar species found in Tuscany, Abruzzo, Romagna, Umbria, the Marche and Molise. It is not as aromatic as those from Piedmont, although those from Città di Castello come quite close.

Other species 

A less common truffle is "garlic truffle" (Tuber macrosporum).

In the U.S. Pacific Northwest, several species of truffle are harvested both recreationally and commercially, most notably, the Leucangium carthusianum, "Oregon white truffles", Tuber gibbosum, and Tuber oregonense.

The "pecan truffle" (Tuber lyonii)  syn. texense  is found in the Southern United States, usually associated with pecan trees. Chefs who have experimented with them agree "they are very good and have potential as a food commodity".  Although pecan farmers used to find them along with pecans and discard them, considering them a nuisance, they sell for about $100 a pound and have been used in some gourmet restaurants.

Truffle-like species 

The term "truffle" has been applied to several other genera of similar underground fungi. The genera Terfezia and Tirmania of the family Terfeziaceae are known as the "desert truffles" of Africa and the Middle East. "Hart's truffle" is a name for Elaphomycetaceae. Pisolithus tinctorius, which was historically eaten in parts of Germany, is sometimes called "Bohemian truffle".

Extraction 

See also truffle hog.

The female pig's natural truffle-seeking, as well as her usual intent to eat the truffle, is due to a compound within the truffle similar to androstenol, the sex pheromone of boar saliva, to which the sow is keenly attracted.

In Italy, the use of the pig to hunt truffles has been prohibited since 1985 because of damage caused by animals to truffle's mycelia during the digging that dropped the production rate of the area for some years.

Volatile constituents 

The volatile constituents responsible for the natural aroma of truffles are released by the mycelia, fruiting body or derive from truffle-associated microbes.  The chemical ecology of truffle volatiles is complex, interacting with plants, insects and mammals, which contribute to spore dispersal. Depending on the truffle species, life cycle or location, these are most notably:
 
* Sulfur 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 bis(methylthio)methane occurring in all species but mostly characteristic of the white truffle T. magnatum. Some of the very aromatic white truffles are notably pungent, even irritating the eye when cut or sliced. 
* Metabolites of non-sulfur amino acid constituents (simple and branched chain hydrocarbons) such as ethylene (produced by mycelia of white truffles affecting root architecture of host tree), as well as 2-methylbutanal,  2-methylpropanal and 2-phenylethanol (also common in baker’s yeast).
* Fatty acid-derived volatiles (C8-alcohols and 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.
A number of truffle species and varieties are differentiated based on their relative contents or absence of sulfides, ethers or alcohols, respectively.  
The sweaty-musky aroma of truffles is similar to that of the pheromone androstenol that also occurs in humans. 
, the volatile profiles of seven black and six white truffle species have been studied. 

Culinary use 

Because of their high price  and their pungent aroma, truffles are used sparingly. Supplies can be found commercially as unadulterated fresh produce or preserved, typically in a light brine.

White truffles are generally served raw, and shaved over steaming buttered pasta, salads, or fried eggs. White or black paper-thin truffle slices may be inserted into meats, under the skins of roasted fowl, in foie gras preparations, in pâtés, or in stuffings. Some speciality cheeses contain truffles, as well.

The flavor of black truffles is far less pungent and more refined than that of white truffles. Their strong flavor is often described as syrupy sweet. Black truffles also are used for producing truffle salt and truffle honey.

While in the past chefs used to peel truffles, in modern times, most restaurants brush the truffle carefully and shave it or dice it with the skin on so as to make the most of this valuable ingredient. A few restaurants in Switzerland still stamp out circular discs of truffle flesh and use the skins for sauces.

Truffle oil 

Truffle oil is used as a lower-cost and convenient substitute for truffles, to provide flavoring, or to enhance the flavor and aroma of truffles in cooking. Most "truffle oil", however, does not contain any truffles. The vast majority is olive oil which has been artificially flavoured using a synthetic agent such as 2,4-dithiapentane.

Truffle vodka 

The bulk of truffle oil on the market is made with a synthetic ingredient, as are many other truffle products. However, alcohol is now being used to carry the truffle flavour without the need for synthetic flavourings. The first truffle vodka, Black Moth Vodka, is a natural vodka infused with black Périgord truffles (Tuber melanosporum). Although primarily used as a spirit in its own right and mixed in a range of cocktails, truffle vodka is also used by various chefs to flavour dishes by evaporating the alcohol through cooking whilst retaining the truffle aroma.
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