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'''''Theobroma cacao''''' also '''cacao tree''' and '''cocoa tree''', is a small ({{convert|4|-|8|m|ft|abbr=on}} tall) [[evergreen]] [[tree]] in the family [[Sterculiaceae]] (alternatively [[Malvaceae]]), native to the deep [[tropical]] region of the Americas. Its seeds are used to make [[cocoa powder]] and [[chocolate]].
'''''Theobroma cacao''''' also '''cacao tree''' and '''cocoa tree''', is a small ({{convert|4|-|8|m|ft|abbr=on}} tall) [[evergreen]] [[tree]] in the family [[Malvaceae]], native to the deep [[tropical]] region of the Americas. Its seeds are used to make [[cocoa powder]] and [[chocolate]].


==Description==
==Description==
Butterflies [[leaf|leaves]] are alternate, entire, unlobed, {{convert|10|-|40|cm|in|abbr=on}} long and {{convert|5|-|20|cm|in|abbr=on}} broad.
[[leaf|Leaves]] are alternate, entire, unlobed, {{convert|10|-|40|cm|in|abbr=on}} long and {{convert|5|-|20|cm|in|abbr=on}} broad.


The [[flower]]s are produced in clusters directly on the [[Trunk (botany)|trunk]] and older branches; they are small, {{convert|1|-|2|cm|in|abbr=on}} diameter, with pink calyx. While many of the world's flowers are pollinated by [[bee]]s ([[Hymenoptera]]) or [[Butterfly|butterflies]]/[[moth]]s ([[Lepidoptera]]), cacao flowers are pollinated by tiny flies, [[Forcipomyiinae|''Forcipomyia'']] [[midge]]s in the order [[Diptera]].<ref>{{cite journal | author = Hernández B, J. | title = Insect pollination of cacao (Theobroma cacao L.) in Costa Rica | publisher = University of Wisconsin | year = 1965 | url = http://orton.catie.ac.cr/cgi-bin/wxis.exe/?IsisScript=orton.xis&expresion=mfn=032019}}</ref> The [[fruit]], called a cacao pod, is ovoid, {{convert|15|-|30|cm|in|abbr=on}} long and {{convert|8|-|10|cm|in|abbr=on}} wide, ripening yellow to orange, and weighs about {{convert|500|g|lb|abbr=on}} when ripe. The pod contains 20 to 60 [[seed]]s, usually called "beans", embedded in a white pulp. The seeds are the main ingredient of [[chocolate]], while the pulp is used in some countries to prepare a refreshing [[juice]]. Each seed contains a significant amount of fat (40–50%) as [[cocoa butter]]. Their most noted active constituent is [[theobromine]], a compound similar to [[caffeine]].
The [[flower]]s are produced in clusters directly on the [[Trunk (botany)|trunk]] and older branches; they are small, {{convert|1|-|2|cm|in|abbr=on}} diameter, with pink calyx. While many of the world's flowers are pollinated by [[bee]]s ([[Hymenoptera]]) or [[Butterfly|butterflies]]/[[moth]]s ([[Lepidoptera]]), cacao flowers are pollinated by tiny flies, [[Forcipomyiinae|''Forcipomyia'']] [[midge]]s in the order [[Diptera]].<ref>{{cite journal | author = Hernández B, J. | title = Insect pollination of cacao (Theobroma cacao L.) in Costa Rica | publisher = University of Wisconsin | year = 1965 | url = http://orton.catie.ac.cr/cgi-bin/wxis.exe/?IsisScript=orton.xis&expresion=mfn=032019}}</ref> The [[fruit]], called a cacao pod, is ovoid, {{convert|15|-|30|cm|in|abbr=on}} long and {{convert|8|-|10|cm|in|abbr=on}} wide, ripening yellow to orange, and weighs about {{convert|500|g|lb|abbr=on}} when ripe. The pod contains 20 to 60 [[seed]]s, usually called "beans", embedded in a white pulp. The seeds are the main ingredient of [[chocolate]], while the pulp is used in some countries to prepare a refreshing [[juice]]. Each seed contains a significant amount of fat (40–50%) as [[cocoa butter]]. Their most noted active constituent is [[theobromine]], a compound similar to [[caffeine]].


==Taxonomy and nomenclature==
==Taxonomy and nomenclature==
Cacao (''Theobroma cacao'') belongs to the [[genus]] ''[[Theobroma]]'' classified under the subfamily [[Malvaceae|Sterculioidea]] of the mallow [[Family (biology)|family]] [[Malvaceae]]. Cacao, together with the [[kola nut]], was once classified under the now obsolete family [[Sterculiaceae]].
Cacao (''Theobroma cacao'') belongs to the [[genus]] ''[[Theobroma]]'' classified under the subfamily [[Malvaceae|Sterculioidea]] of the mallow [[Family (biology)|family]] [[Malvaceae]]. Cacao is one of 22 species of ''Theobroma''.


<!--NOTE: Study below deals with cacao germplasm, not taxonomy, not sure if it's even relevant:
<!--NOTE: Study below deals with cacao germplasm, not taxonomy, not sure if it's even relevant:
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The [[cupuaçu]], ''Theobroma grandiflorum'', is a closely related species also grown in Brazil. Like the cacao, it is also the source for a kind of chocolate known as ''cupulate'' or ''cupuaçu'' chocolate.<ref>http://www.fruitipedia.com/cupuacu_theobroma_grandiflorum.htm</ref> The cupuaçu is considered of high potential by the food and cosmetics industries.
The [[cupuaçu]], ''Theobroma grandiflorum'', is a closely related species also grown in Brazil. Like the cacao, it is also the source for a kind of chocolate known as ''cupulate'' or ''cupuaçu'' chocolate.<ref>http://www.fruitipedia.com/cupuacu_theobroma_grandiflorum.htm</ref> The cupuaçu is considered of high potential by the food and cosmetics industries.


==Distribution and habitat==
==Distribution and domestication==
''T. cacao'' is widely distributed from southeastern [[Mexico]] to the [[Amazon basin]]. There were originally two hypotheses about its domestication; one said that there were two foci for domestication, one in the Lacandon area of Mexico and another in lowland [[South America]]. More recent studies of patterns of DNA diversity, however, suggest that this is not the case. Motomayor ''et al.'' <ref>{{cite journal|author=Juan C. Motamayor, Philippe Lachenaud, Jay Wallace da Silva e Mota, Rey Loor, David N. Kuhn, J. Steven Brown, Raymond J. Schnell |title=Geographic and Genetic Population Differentiation of the Amazonian Chocolate Tree (''Theobroma cacao'' L.) |journal=[[PLoS ONE]] |volume=3 |issue=10 |doi=10.1371/journal.pone.0003311 |year=2008 |url=http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0003311}}</ref> sampled 1241 trees and classified them into 10 distinct genetic clusters. This study also identified areas, for example around [[Iquitos]] in modern [[Peru]], where representatives of several genetic clusters originated. This result suggests that this is where ''T. cacao'' was originally domesticated, probably for the pulp that surrounds the beans, which is eaten as a snack and fermented into a mildly alcoholic beverage. <ref>{{cite journal|last=Clement|first=Charles R.|coauthors=de Cristo-Araújo, Michelly; d’Eeckenbrugge, Geo Coppens; Alves Pereira, Alessandro; Picanço-Rodrigues, Doriane|title=Origin and Domestication of Native Amazonian Crops|journal=Diversity|date=6 January 2010|year=2010|volume=2|issue=1|pages=72–106|doi=10.3390/d2010072|accessdate=9 November 2012}}</ref> Using the DNA sequences obtained by Motomayor ''et al.'' and comparing them with data derived from climate models and the known conditions suitable for cacao, Thomas ''et al.'' have further refined the view of domestication, linking the area of greatest cacao genetic diversity to a bean-shaped area that encompasses the border between Brazil and Peru and the southern part of the Colombian-Brazilian border. <ref>{{cite journal|last=Thomas|first=Evert|coauthors=van Zonneveld, Maarten; Loo, Judy; Hodgkin, Toby; Galluzzi, Gea; van Etten, Jacob; Fuller, Dorian Q.|title=Present Spatial Diversity Patterns of Theobroma cacao L. in the Neotropics Reflect Genetic Differentiation in Pleistocene Refugia Followed by Human-Influenced Dispersal|journal=PLoS ONE|date=24 October 2012|volume=7|issue=10|pages=e47676|doi=10.1371/journal.pone.0047676}}</ref> Climate models indicate that at the peak of the previous [[ice age]] 21,000 years ago, when habitat suitable for cacao was at its most reduced, this area was still suitable, and so provided a [[Refugium_(population_biology)|refugium]] for the species. Thomas ''et al.'' speculate that from there people took cacao to Mexico, where selection for the beans took place.
Two prominent hypotheses conflict about the origins of the domestication of the originally wild ''Theobroma cacao'' tree. One is that wild examples were originally distributed from southeastern [[Mexico]] to the [[Amazon basin]], with domestication taking place both in the Lacandon area of Mexico and in lowland [[South America]]. But recent studies of ''T. cacao'' genetics seem to show the plant originated in the Amazon and was distributed by humans throughout [[Central America]] and [[Mesoamerica]].


Cacao trees grow well as [[understory]] plants in humid forest ecosystems. This is equally true of abandoned cultivated trees, making it difficult to distinguish truly wild trees from those whose parents may originally have been cultivated.
The tree is today found only growing wild in the low foothills of the [[Andes]] at elevations of around {{convert|200|-|400|m|ft|abbr=on}} in the [[Amazon River|Amazon]] and [[Orinoco]] [[river basin]]s. It requires a humid climate with regular [[rainfall]] and good soil. It is an [[understory]] tree, growing best with some overhead shade.


==History of cultivation==
==History of cultivation==
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The initial domestication was probably related to the making of a fermented, thus alcoholic beverage.<ref>{{cite journal | author=J.S. Henderson 'et al.' |title= Chemical and archaeological evidence for the earliest cacao beverages |journal=[[Proceedings of the National Academy of Sciences]] |year= 2007 |volume= 104 |pages= 18937–18940}}</ref>
The initial domestication was probably related to the making of a fermented, thus alcoholic beverage.<ref>{{cite journal | author=J.S. Henderson 'et al.' |title= Chemical and archaeological evidence for the earliest cacao beverages |journal=[[Proceedings of the National Academy of Sciences]] |year= 2007 |volume= 104 |pages= 18937–18940}}</ref>


⚫
Several mixtures of cacao are described in ancient texts, for ceremonial or medicinal, as well as culinary, purposes. Some mixtures included [[maize]], [[Capsicum|chili]], [[vanilla]] (''Vanilla planifolia''), and honey. [[Archaeological]] evidence for use of cacao, while relatively sparse, has come from the recovery of whole cacao beans at [[Uaxactun]], [[Guatemala]] <ref>(Kidder 1947)</ref> and from the preservation of wood fragments of the cacao tree at [[Belize]] sites including [[Cuello]] and [[Pulltrouser Swamp]].<ref>(Hammond and Miksicek 1981; Turner and Miksicek 1984)</ref> In addition, analysis of residues from ceramic vessels has found traces of [[theobromine]] and [[caffeine]] in early formative vessels from Puerto Escondido, Honduras (1100-900 BC) and in middle formative vessels from [[Colha, Belize]] (600-400 BC) using similar techniques to those used to extract chocolate residues from four classic period (''circa'' 400 AD) vessels from a tomb at the archaeological site of [[Rio Azul]]. As cacao is the only known commodity from Mesoamerica containing both of these [[alkaloid]] compounds, it seems likely these vessels were used as containers for cacao drinks. In addition, cacao is named in a [[hieroglyph]]ic text on one of the Rio Azul vessels. Cacao was also believed to be ground by the Aztecs and mixed with tobacco for smoking purposes{{citation needed|date=September 2012}}.
In a study of ''Theobroma cacao'' tree genetics,<ref>{{cite journal | author= Motamayor J.C., Risterucci A.-M., Lopez P.A., Ortiz C.F., Moreno A., Lanaud C. |title=Cacao domestication I: the origin of the cacao cultivated by the Mayas. |journal= [[Heredity (journal)|Heredity]] |volume= 89 |issue=5 |pages=380–386 |year=2002 |pmid= 12399997 |doi= 10.1038/sj.hdy.6800156}}</ref> Motamayor ''et al.'' suggest a domestication and spread from lowland Amazonia, contesting an earlier hypothesis that the tree was domesticated independently in both the [[Lacandon]] area of Mexico, and in Amazonia. The cacao tree belongs to the ''Theobroma'' genus, in the Sterculiaceae family, that contains 22 species. Today, the most common of the cultivated species is ''Theobroma cacao'', with two subspecies and three forms. Wild cacaos fall into two groups. The South American subspecies ''T. c. spaerocarpum'' has a fairly smooth, melon-like fruit. In contrast, the Mesoamerican cacao subspecies has ridged, elongated fruits. At some unknown early date, this subspecies ''T. cacao'' reached the southern lowlands of Mesoamerica and came into wide usage.


⚫
===Currency system===
⚫
Cacao beans constituted both a ritual beverage and a major [[currency]] system in [[pre-Columbian]] [[Mesoamerica]]n civilizations. At one point, the Aztec empire received a yearly tribute of 980 loads (''xiquipil'' in Nahuatl) of cacao, in addition to other goods. Each load represented exactly 8,000 beans.<ref>J. Bergmann (1969).</ref> The buying power of quality beans was such that 80-100 beans could buy a new cloth mantle. The use of cacao beans as currency is also known to have spawned counterfeiters during the Aztec empire.<ref>S. Coe (1994).</ref>

==Mythology==
[[File:Kakaw (Mayan word).png|left|thumb|150px|''Kakaw'' (cacao) written in the [[Maya script]]: The word was also written in several other ways in old Maya texts.]]
[[File:Kakaw (Mayan word).png|left|thumb|150px|''Kakaw'' (cacao) written in the [[Maya script]]: The word was also written in several other ways in old Maya texts.]]
The [[Maya civilization|Maya]] believed the ''kakaw'' (cacao) was discovered by the gods in a mountain that also contained other delectable foods to be used by them. According to [[Maya mythology]], the [[Plumed Serpent]] gave cacao to the Maya after humans were created from maize by divine grandmother goddess [[Xmucane]].<ref>(Bogin 1997, Coe 1996, Montejo 1999, Tedlock 1985)</ref> The Maya celebrated an annual festival in April to honor their cacao god, [[Ek Chuah]], an event that included the [[Dogs in Mesoamerica|sacrifice of a dog]] with cacao-colored markings, additional animal sacrifices, offerings of cacao, feathers and incense, and an exchange of gifts. In a similar creation story, the [[Mexica]] ([[Aztec]]) god [[Quetzalcoatl]] discovered cacao (''cacahuatl'': "bitter water"), in a mountain filled with other plant foods.<ref>(Coe 1996, Townsend 1992)</ref> Cacao was offered regularly to a pantheon of Mexica deities and the [[Madrid Codex (Maya)|Madrid Codex]] depicts priests lancing their ear lobes (autosacrifice) and covering the cacao with blood as a suitable sacrifice to the gods. The cacao beverage as ritual was used only by men, as it was believed to be [[toxic]] for women and children.{{Citation needed|date=February 2011}}
The [[Maya civilization|Maya]] believed the ''kakaw'' (cacao) was discovered by the gods in a mountain that also contained other delectable foods to be used by them. According to [[Maya mythology]], the [[Plumed Serpent]] gave cacao to the Maya after humans were created from maize by divine grandmother goddess [[Xmucane]].<ref>(Bogin 1997, Coe 1996, Montejo 1999, Tedlock 1985)</ref> The Maya celebrated an annual festival in April to honor their cacao god, [[Ek Chuah]], an event that included the [[Dogs in Mesoamerica|sacrifice of a dog]] with cacao-colored markings, additional animal sacrifices, offerings of cacao, feathers and incense, and an exchange of gifts. In a similar creation story, the [[Mexica]] ([[Aztec]]) god [[Quetzalcoatl]] discovered cacao (''cacahuatl'': "bitter water"), in a mountain filled with other plant foods.<ref>(Coe 1996, Townsend 1992)</ref> Cacao was offered regularly to a pantheon of Mexica deities and the [[Madrid Codex (Maya)|Madrid Codex]] depicts priests lancing their ear lobes (autosacrifice) and covering the cacao with blood as a suitable sacrifice to the gods. The cacao beverage as ritual was used only by men, as it was believed to be [[toxic]] for women and children.{{Citation needed|date=February 2011}}


==Modern History==
⚫
Several mixtures of cacao are described in ancient texts, for ceremonial or medicinal, as well as culinary, purposes. Some mixtures included [[maize]], [[Capsicum|chili]], [[vanilla]] (''Vanilla planifolia''), and honey. [[Archaeological]] evidence for use of cacao, while relatively sparse, has come from the recovery of whole cacao beans at [[Uaxactun]], [[Guatemala]] <ref>(Kidder 1947)</ref> and from the preservation of wood fragments of the cacao tree at [[Belize]] sites including [[Cuello]] and [[Pulltrouser Swamp]].<ref>(Hammond and Miksicek 1981; Turner and Miksicek 1984)</ref> In addition, analysis of residues from ceramic vessels has found traces of [[theobromine]] and [[caffeine]] in early formative vessels from Puerto Escondido, Honduras (1100-900 BC) and in middle formative vessels from [[Colha, Belize]] (600-400 BC) using similar techniques to those used to extract chocolate residues from four classic period (''circa'' 400 AD) vessels from a tomb at the archaeological site of [[Rio Azul]]. As cacao is the only known commodity from Mesoamerica containing both of these [[alkaloid]] compounds, it seems likely these vessels were used as containers for cacao drinks. In addition, cacao is named in a [[hieroglyph]]ic text on one of the Rio Azul vessels. Cacao was also believed to be ground by the Aztecs and mixed with tobacco for smoking purposes{{citation needed|date=September 2012}}.

The first Europeans to encounter cacao were [[Christopher Columbus]] and his crew [[Voyages of Christopher Columbus#Fourth voyage|in 1502]],{{Citation needed|date=February 2011}} when they captured a canoe at [[Guanaja]] that contained a quantity of mysterious-looking "almonds". The first real European knowledge about chocolate came in the form of a beverage which was first introduced to the Spanish at their meeting with [[Moctezuma II|Moctezuma]] in the [[Aztec]] capital of [[Tenochtitlan]] in 1519. Cortez and others noted the vast quantities of this beverage the Aztec emperor consumed, and how it was carefully whipped by his attendants beforehand. Examples of cacao beans, along with other agricultural products, were brought back to Spain at that time, but it seems the beverage made from cacao was introduced to the Spanish court in 1544 by Kekchi Maya nobles brought from the New World to Spain by [[Dominican Order|Dominican]] [[friar]]s to meet [[Philip II of Spain|Prince Philip]].<ref>(Coe and Coe 1996)</ref> Within a century, the culinary and [[medical]] uses of chocolate had spread to France, England and elsewhere in [[Western Europe]]. Demand for this beverage led the French to establish cacao [[plantation]]s in the [[Caribbean]], while Spain subsequently developed their cacao plantations in their [[History of the Philippines (1521-1898)|Philippine colony]] (Bloom 1998, Coe 1996). The Nahuatl-derived Spanish word ''cacao'' entered scientific nomenclature in 1753 after the [[Sweden|Swedish]] naturalist [[Linnaeus]] published his taxonomic binomial system and coined the genus and species ''Theobroma cacao''.
The first Europeans to encounter cacao were [[Christopher Columbus]] and his crew [[Voyages of Christopher Columbus#Fourth voyage|in 1502]],{{Citation needed|date=February 2011}} when they captured a canoe at [[Guanaja]] that contained a quantity of mysterious-looking "almonds". The first real European knowledge about chocolate came in the form of a beverage which was first introduced to the Spanish at their meeting with [[Moctezuma II|Moctezuma]] in the [[Aztec]] capital of [[Tenochtitlan]] in 1519. Cortez and others noted the vast quantities of this beverage the Aztec emperor consumed, and how it was carefully whipped by his attendants beforehand. Examples of cacao beans, along with other agricultural products, were brought back to Spain at that time, but it seems the beverage made from cacao was introduced to the Spanish court in 1544 by Kekchi Maya nobles brought from the New World to Spain by [[Dominican Order|Dominican]] [[friar]]s to meet [[Philip II of Spain|Prince Philip]].<ref>(Coe and Coe 1996)</ref> Within a century, the culinary and [[medical]] uses of chocolate had spread to France, England and elsewhere in [[Western Europe]]. Demand for this beverage led the French to establish cacao [[plantation]]s in the [[Caribbean]], while Spain subsequently developed their cacao plantations in their [[History of the Philippines (1521-1898)|Philippine colony]] (Bloom 1998, Coe 1996). The Nahuatl-derived Spanish word ''cacao'' entered scientific nomenclature in 1753 after the [[Sweden|Swedish]] naturalist [[Linnaeus]] published his taxonomic binomial system and coined the genus and species ''Theobroma cacao''.


Traditional [[pre-Hispanic]] beverages made with cacao are still consumed in [[Mesoamerica]]. These include the [[Oaxaca]]n beverage known as ''[[tejate]]''.
Traditional [[pre-Hispanic]] beverages made with cacao are still consumed in [[Mesoamerica]]. These include the [[Oaxaca]]n beverage known as ''[[tejate]]''.

⚫
===Currency system===
⚫
Cacao beans constituted both a ritual beverage and a major [[currency]] system in [[pre-Columbian]] [[Mesoamerica]]n civilizations. At one point, the Aztec empire received a yearly tribute of 980 loads (''xiquipil'' in Nahuatl) of cacao, in addition to other goods. Each load represented exactly 8,000 beans.<ref>J. Bergmann (1969).</ref> The buying power of quality beans was such that 80-100 beans could buy a new cloth mantle. The use of cacao beans as currency is also known to have spawned counterfeiters during the Aztec empire.<ref>S. Coe (1994).</ref>


==Cultivation==
==Cultivation==
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<nowiki>*</nowiki>Based on 1999–2001 international prices
<nowiki>*</nowiki>Based on 1999–2001 international prices


Cacao production has increased from 1.5 million tons in 1983-1984 to 3.5 million tons in 2003-2004, almost entirely due to the expansion of the production area rather than to yield increases. Cacao is grown both by large agroindustrial plantations and small producers.
Cacao production has increased from 1.5 million tons in 1983-1984 to 3.5 million tons in 2003-2004, almost entirely due to the expansion of the production area rather than to yield increases. Cacao is grown both by large agroindustrial plantations and small producers, the bulk of production coming from millions of farmers who have a few trees each. <ref>{{cite web|last=Anon|title=A strategy to safeguard the future of chocolate|url=http://www.bioversityinternational.org/index.php?id=6817|publisher=Bioversity International|accessdate=9 November 2012}}</ref>


A tree begins to bear when it is four or five years old. A mature tree may have 6,000 flowers in a year, yet only about 20 pods. About 300-600 seeds (10 pods) are required to produce 1&nbsp;kg (2.2&nbsp;lb) of [[chocolate liquor|cocoa paste]].
A tree begins to bear when it is four or five years old. A mature tree may have 6,000 flowers in a year, yet only about 20 pods. About 300-600 seeds (10 pods) are required to produce 1&nbsp;kg (2.2&nbsp;lb) of [[chocolate liquor|cocoa paste]].


There are three main [[cultivar|cultivar groups]] of cacao beans used to make cocoa and chocolate.<ref>[http://www.xocoatl.org/variety.htm All about Chocolate -- Varieties]</ref> The most prized, rare, and expensive is the Criollo group, the cocoa bean used by the [[Maya civilization|Maya]]. Only 10% of chocolate is made from Criollo, which is less bitter and more aromatic than any other bean. The cacao bean in 80% of chocolate is made using beans of the Forastero group. Forastero trees are significantly hardier than Criollo trees, resulting in cheaper cacao beans. Trinitario, a hybrid of Criollo and Forastero, is used in about 10% of chocolate.
Historically, chocolate makers have recognized three main [[cultivar|cultivar groups]] of cacao beans used to make cocoa and chocolate.<ref>[http://www.xocoatl.org/variety.htm All about Chocolate -- Varieties]</ref> The most prized, rare, and expensive is the Criollo group, the cocoa bean used by the [[Maya civilization|Maya]]. Only 10% of chocolate is made from Criollo, which is less bitter and more aromatic than any other bean. The cacao bean in 80% of chocolate is made using beans of the Forastero group. Forastero trees are significantly hardier than Criollo trees, resulting in cheaper cacao beans. Trinitario, a hybrid of Criollo and Forastero, is used in about 10% of chocolate. The new, genetically-based classification into 10 groups <ref>{{cite journal|author=Juan C. Motamayor, Philippe Lachenaud, Jay Wallace da Silva e Mota, Rey Loor, David N. Kuhn, J. Steven Brown, Raymond J. Schnell |title=Geographic and Genetic Population Differentiation of the Amazonian Chocolate Tree (''Theobroma cacao'' L.) |journal=[[PLoS ONE]] |volume=3 |issue=10 |doi=10.1371/journal.pone.0003311 |year=2008 |url=http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0003311}}</ref> may well help breeders to create new varieties that are both pest- and disease-resistant and contain valued flavours.


Major [[cocoa bean]] processors include [[The Hershey Company|Hershey's]], [[Nestlé]] and [[Mars, Incorporated|Mars]], all of which purchase cocoa beans via various sources.
Major [[cocoa bean]] processors include [[The Hershey Company|Hershey's]], [[Nestlé]] and [[Mars, Incorporated|Mars]], all of which purchase cocoa beans via various sources.
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* [[Mistletoe]]
* [[Mistletoe]]
*[[Rat]]s and other [[vertebrate]] pests ([[squirrel]]s, [[woodpecker]]s, ''etc.'')
*[[Rat]]s and other [[vertebrate]] pests ([[squirrel]]s, [[woodpecker]]s, ''etc.'')

==Conservation==
The pests and diseases to which cacao is subject, along with climate change, mean that new varieties will be needed to respond to these challenges. Breeders rely on the genetic diversity conserved in field [[genebank]]s to create new varieties, because cacao has [[recalcitrant seed]]s that cannot be stored in a conventional genebank. <ref>{{cite web|title=Cacao Collections|url=https://sites.google.com/a/cgxchange.org/cacaonet/cacao-genebanks|publisher=CacaoNet|accessdate=9 November 2012}}</ref> In an effort to improve the diversity available to breeders, and ensure the future of the field genebanks, experts have drawn up a [https://sites.google.com/a/cgxchange.org/cacaonet/cacaonet-global-strategy A Global Strategy for the Conservation and Use of Cacao Genetic Resources, as the Foundation for a Sustainable Cocoa Economy.] The strategy has been adopted by the cacao producers and their clients, and seeks to improve the characterization of cacao diversity, the sustainability and diversity of the cacao collections, the usefulness of the collections, and to ease access to better information about the conserved material. Some natural areas of cacao diversity are protected by various forms of conservation, for example national parks. However, a recent study of genetic diversity and predicted climates <ref>{{cite journal|last=Thomas|first=Evert|coauthors=van Zonneveld, Maarten; Loo, Judy; Hodgkin, Toby; Galluzzi, Gea; van Etten, Jacob; Fuller, Dorian Q.|title=Present Spatial Diversity Patterns of Theobroma cacao L. in the Neotropics Reflect Genetic Differentiation in Pleistocene Refugia Followed by Human-Influenced Dispersal|journal=PLoS ONE|date=24 October 2012|volume=7|issue=10|pages=e47676|doi=10.1371/journal.pone.0047676}}</ref> suggests that many of those protected areas will no longer be suitable for cacao by 2050. It also identifies an area around [[Iquitos]] in [[Peru]] that will remain suitable for cacao and that is home to considerable genetic diversity, and recommends that this area be considered for protection.


==Cacao genome==
==Cacao genome==

Revision as of 10:30, 9 November 2012

For other uses, see Cacao (disambiguation).

Theobroma cacao
Cacao fruits on the tree
Scientific classification
Kingdom:
Plantae
(unranked):
Angiosperms
(unranked):
Eudicots
(unranked):
Rosids
Order:
Malvales
Family:
Malvaceae
Genus:
Theobroma
Species:
T. cacao
Binomial name
Theobroma cacao

Theobroma cacao also cacao tree and cocoa tree, is a small (4–8 m (13–26 ft) tall) evergreen tree in the family Malvaceae, native to the deep tropical region of the Americas. Its seeds are used to make cocoa powder and chocolate.

Description

Leaves are alternate, entire, unlobed, 10–40 cm (3.9–15.7 in) long and 5–20 cm (2.0–7.9 in) broad.

The flowers are produced in clusters directly on the trunk and older branches; they are small, 1–2 cm (0.39–0.79 in) diameter, with pink calyx. While many of the world's flowers are pollinated by bees (Hymenoptera) or butterflies/moths (Lepidoptera), cacao flowers are pollinated by tiny flies, Forcipomyia midges in the order Diptera.[1] The fruit, called a cacao pod, is ovoid, 15–30 cm (5.9–11.8 in) long and 8–10 cm (3.1–3.9 in) wide, ripening yellow to orange, and weighs about 500 g (1.1 lb) when ripe. The pod contains 20 to 60 seeds, usually called "beans", embedded in a white pulp. The seeds are the main ingredient of chocolate, while the pulp is used in some countries to prepare a refreshing juice. Each seed contains a significant amount of fat (40–50%) as cocoa butter. Their most noted active constituent is theobromine, a compound similar to caffeine.

Taxonomy and nomenclature

Cacao (Theobroma cacao) belongs to the genus Theobroma classified under the subfamily Sterculioidea of the mallow family Malvaceae. Cacao is one of 22 species of Theobroma.

The generic name is derived from the Greek for "food of the gods"; from θεος (theos), meaning "god," and βρῶμα (broma), meaning "food".

The specific name cacao is derived from the native name of the plant in indigenous Mesoamerican languages. The cacao was known as kakaw in Tzeltal, K’iche’ and Classic Maya; kagaw in Sayula Popoluca; and cacahuatl[dubious – discuss] in Nahuatl.

Cacao flowers.

The cupuaçu, Theobroma grandiflorum, is a closely related species also grown in Brazil. Like the cacao, it is also the source for a kind of chocolate known as cupulate or cupuaçu chocolate.[2] The cupuaçu is considered of high potential by the food and cosmetics industries.

Distribution and domestication

T. cacao is widely distributed from southeastern Mexico to the Amazon basin. There were originally two hypotheses about its domestication; one said that there were two foci for domestication, one in the Lacandon area of Mexico and another in lowland South America. More recent studies of patterns of DNA diversity, however, suggest that this is not the case. Motomayor et al. [3] sampled 1241 trees and classified them into 10 distinct genetic clusters. This study also identified areas, for example around Iquitos in modern Peru, where representatives of several genetic clusters originated. This result suggests that this is where T. cacao was originally domesticated, probably for the pulp that surrounds the beans, which is eaten as a snack and fermented into a mildly alcoholic beverage. [4] Using the DNA sequences obtained by Motomayor et al. and comparing them with data derived from climate models and the known conditions suitable for cacao, Thomas et al. have further refined the view of domestication, linking the area of greatest cacao genetic diversity to a bean-shaped area that encompasses the border between Brazil and Peru and the southern part of the Colombian-Brazilian border. [5] Climate models indicate that at the peak of the previous ice age 21,000 years ago, when habitat suitable for cacao was at its most reduced, this area was still suitable, and so provided a refugium for the species. Thomas et al. speculate that from there people took cacao to Mexico, where selection for the beans took place.

Cacao trees grow well as understory plants in humid forest ecosystems. This is equally true of abandoned cultivated trees, making it difficult to distinguish truly wild trees from those whose parents may originally have been cultivated.

History of cultivation

Toasted cacao beans grown at the La Chonita Hacienda in Tabasco, Mexico

Cultivation, use, and cultural elaboration of cacao were early and extensive in Mesoamerica. Ceramic vessels with residues from the preparation of cacao beverages have been found at archaeological sites dating back to the Early Formative (1900-900 BC) period. For example, one such vessel found at an Olmec archaeological site on the Gulf Coast of Veracruz, Mexico dates cacao's preparation by pre-Olmec peoples as early as 1750 BC.[6] On the Pacific coast of Chiapas, Mexico, a Mokaya archaeological site provides evidence of cacao beverages dating even earlier, to 1900 BC.[6] The initial domestication was probably related to the making of a fermented, thus alcoholic beverage.[7]

Several mixtures of cacao are described in ancient texts, for ceremonial or medicinal, as well as culinary, purposes. Some mixtures included maize, chili, vanilla (Vanilla planifolia), and honey. Archaeological evidence for use of cacao, while relatively sparse, has come from the recovery of whole cacao beans at Uaxactun, Guatemala [8] and from the preservation of wood fragments of the cacao tree at Belize sites including Cuello and Pulltrouser Swamp.[9] In addition, analysis of residues from ceramic vessels has found traces of theobromine and caffeine in early formative vessels from Puerto Escondido, Honduras (1100-900 BC) and in middle formative vessels from Colha, Belize (600-400 BC) using similar techniques to those used to extract chocolate residues from four classic period (circa 400 AD) vessels from a tomb at the archaeological site of Rio Azul. As cacao is the only known commodity from Mesoamerica containing both of these alkaloid compounds, it seems likely these vessels were used as containers for cacao drinks. In addition, cacao is named in a hieroglyphic text on one of the Rio Azul vessels. Cacao was also believed to be ground by the Aztecs and mixed with tobacco for smoking purposes[citation needed].

Currency system

Cacao beans constituted both a ritual beverage and a major currency system in pre-Columbian Mesoamerican civilizations. At one point, the Aztec empire received a yearly tribute of 980 loads (xiquipil in Nahuatl) of cacao, in addition to other goods. Each load represented exactly 8,000 beans.[10] The buying power of quality beans was such that 80-100 beans could buy a new cloth mantle. The use of cacao beans as currency is also known to have spawned counterfeiters during the Aztec empire.[11]

Mythology

Kakaw (cacao) written in the Maya script: The word was also written in several other ways in old Maya texts.

The Maya believed the kakaw (cacao) was discovered by the gods in a mountain that also contained other delectable foods to be used by them. According to Maya mythology, the Plumed Serpent gave cacao to the Maya after humans were created from maize by divine grandmother goddess Xmucane.[12] The Maya celebrated an annual festival in April to honor their cacao god, Ek Chuah, an event that included the sacrifice of a dog with cacao-colored markings, additional animal sacrifices, offerings of cacao, feathers and incense, and an exchange of gifts. In a similar creation story, the Mexica (Aztec) god Quetzalcoatl discovered cacao (cacahuatl: "bitter water"), in a mountain filled with other plant foods.[13] Cacao was offered regularly to a pantheon of Mexica deities and the Madrid Codex depicts priests lancing their ear lobes (autosacrifice) and covering the cacao with blood as a suitable sacrifice to the gods. The cacao beverage as ritual was used only by men, as it was believed to be toxic for women and children.[citation needed]

Modern History

The first Europeans to encounter cacao were Christopher Columbus and his crew in 1502,[citation needed] when they captured a canoe at Guanaja that contained a quantity of mysterious-looking "almonds". The first real European knowledge about chocolate came in the form of a beverage which was first introduced to the Spanish at their meeting with Moctezuma in the Aztec capital of Tenochtitlan in 1519. Cortez and others noted the vast quantities of this beverage the Aztec emperor consumed, and how it was carefully whipped by his attendants beforehand. Examples of cacao beans, along with other agricultural products, were brought back to Spain at that time, but it seems the beverage made from cacao was introduced to the Spanish court in 1544 by Kekchi Maya nobles brought from the New World to Spain by Dominican friars to meet Prince Philip.[14] Within a century, the culinary and medical uses of chocolate had spread to France, England and elsewhere in Western Europe. Demand for this beverage led the French to establish cacao plantations in the Caribbean, while Spain subsequently developed their cacao plantations in their Philippine colony (Bloom 1998, Coe 1996). The Nahuatl-derived Spanish word cacao entered scientific nomenclature in 1753 after the Swedish naturalist Linnaeus published his taxonomic binomial system and coined the genus and species Theobroma cacao.

Traditional pre-Hispanic beverages made with cacao are still consumed in Mesoamerica. These include the Oaxacan beverage known as tejate.

Cultivation

See also: Cocoa bean
Cacao seed in the fruit or pocha
Young cacao plantation
Macrophotography of theobroma cacao flower (closed) (University of Vienna)
Macrophotography of theobroma cacao flower (open) (University of Vienna)

Cacao is cultivated on roughly 17,000,000 acres (27,000 sq mi; 69,000 km2) worldwide.[15] According to the Food and Agriculture Organization (FAO), the top 20 cacao-producing countries in 2005 were as follows:[16]

Rank, Country Value
(Int'l $1,000*) 		Production
(Metric tons)
1  Côte d'Ivoire (Ivory Coast) 1,024,339 1,330,000
2  Ghana 566,852 736,000
3  Indonesia 469,810 610,000
4  Nigeria 281,886 366,000
5  Brazil 164,644 213,774
6  Cameroon 138,632 180,000
7  Ecuador 105,652 137,178
8  Colombia 42,589 55,298
9  Mexico 37,281 48,405
10  Papua New Guinea 32,733 42,500
11  Malaysia 25,742 33,423
12  Dominican Republic 24,646 32,000
13  Peru 21,950 28,500
14  Venezuela 13,093 17,000
15  Sierra Leone 8,472 11,000
16  Togo 6,547 8,500
17  India 6,161 8,000
18  Philippines 4,352 5,650
19 Republic of the Congo Congo, Rep. 4,336 5,630
20  Solomon Islands 3,851 5,000

*Based on 1999–2001 international prices

Cacao production has increased from 1.5 million tons in 1983-1984 to 3.5 million tons in 2003-2004, almost entirely due to the expansion of the production area rather than to yield increases. Cacao is grown both by large agroindustrial plantations and small producers, the bulk of production coming from millions of farmers who have a few trees each. [17]

A tree begins to bear when it is four or five years old. A mature tree may have 6,000 flowers in a year, yet only about 20 pods. About 300-600 seeds (10 pods) are required to produce 1 kg (2.2 lb) of cocoa paste.

Historically, chocolate makers have recognized three main cultivar groups of cacao beans used to make cocoa and chocolate.[18] The most prized, rare, and expensive is the Criollo group, the cocoa bean used by the Maya. Only 10% of chocolate is made from Criollo, which is less bitter and more aromatic than any other bean. The cacao bean in 80% of chocolate is made using beans of the Forastero group. Forastero trees are significantly hardier than Criollo trees, resulting in cheaper cacao beans. Trinitario, a hybrid of Criollo and Forastero, is used in about 10% of chocolate. The new, genetically-based classification into 10 groups [19] may well help breeders to create new varieties that are both pest- and disease-resistant and contain valued flavours.

Major cocoa bean processors include Hershey's, Nestlé and Mars, all of which purchase cocoa beans via various sources.

In June 2009, Mars Botanicals, a division of Mars, launched Cirku, a cocoa extract product that provides cocoa flavanols made with a patented process that contains a high level of phytonutrients.[20]

Pests

See also: List of cacao diseases

Various plant pests and diseases can cause serious problems for cacao production; see: Illustrated guide to pests and their management.

Conservation

The pests and diseases to which cacao is subject, along with climate change, mean that new varieties will be needed to respond to these challenges. Breeders rely on the genetic diversity conserved in field genebanks to create new varieties, because cacao has recalcitrant seeds that cannot be stored in a conventional genebank. [21] In an effort to improve the diversity available to breeders, and ensure the future of the field genebanks, experts have drawn up a A Global Strategy for the Conservation and Use of Cacao Genetic Resources, as the Foundation for a Sustainable Cocoa Economy. The strategy has been adopted by the cacao producers and their clients, and seeks to improve the characterization of cacao diversity, the sustainability and diversity of the cacao collections, the usefulness of the collections, and to ease access to better information about the conserved material. Some natural areas of cacao diversity are protected by various forms of conservation, for example national parks. However, a recent study of genetic diversity and predicted climates [22] suggests that many of those protected areas will no longer be suitable for cacao by 2050. It also identifies an area around Iquitos in Peru that will remain suitable for cacao and that is home to considerable genetic diversity, and recommends that this area be considered for protection.

Cacao genome

The genome of T. cacao is diploid, its size is 430 Mbp, and it comprises 10 chromosome pairs (2n=2x=20). In September 2010, a team of scientists announced a draft sequence of the cacao genome (Matina1-6 genotype).[23] In a second, unrelated project, the International Cocoa Genome Sequencing Consortium-ICGS, co-ordinated by CIRAD,[24] first published[25] in December 2010 (online, paper publication in January 2011), the sequence of the cacao genome, of the Criollo cacao (of a landrace from Belize, B97-61/B2). In their publication, they reported a detailed analysis of the genomic and genetic data.

The sequence of the cacao genome identified 28,798 protein-coding genes, compared to the roughly 23,000 protein-coding genes of the human genome. About 20% of the cacao genome consists of transposable elements, a low proportion compared to other plant species. Many genes were identified as coding for flavonoids, aromatic terpenes, theobromine and many other metabolites involved in cocoa flavor and quality traits, among which a relatively high proportion code for polyphenols, which constitute up to 8% of cacao pods dry weight. The cacao genome appears close to the hypothetical hexaploid ancestor of all dicotyledonous plants,[26] and it is proposed as an evolutionary mechanism by which the 21 chromosomes of the dicots' hypothetical hexaploid ancestor underwent major fusions leading to cacao's 10 chromosome pairs.

The genome sequence will help accelerate research on cacao molecular biology and breeding for elite varieties through marker-assisted selection, in particular for genetic resistance to fungal, oomycete and viral diseases responsible for huge yield losses each year.

Notes

  1. ^ Hernández B, J. (1965). "Insect pollination of cacao (Theobroma cacao L.) in Costa Rica". University of Wisconsin. {{cite journal}}: Cite journal requires |journal= (help)
  2. ^ http://www.fruitipedia.com/cupuacu_theobroma_grandiflorum.htm
  3. ^ Juan C. Motamayor, Philippe Lachenaud, Jay Wallace da Silva e Mota, Rey Loor, David N. Kuhn, J. Steven Brown, Raymond J. Schnell (2008). "Geographic and Genetic Population Differentiation of the Amazonian Chocolate Tree (Theobroma cacao L.)". PLoS ONE. 3 (10). doi:10.1371/journal.pone.0003311.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  4. ^ Clement, Charles R. (6 January 2010). "Origin and Domestication of Native Amazonian Crops". Diversity. 2 (1): 72–106. doi:10.3390/d2010072. {{cite journal}}: |access-date= requires |url= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: date and year (link) CS1 maint: unflagged free DOI (link)
  5. ^ Thomas, Evert (24 October 2012). "Present Spatial Diversity Patterns of Theobroma cacao L. in the Neotropics Reflect Genetic Differentiation in Pleistocene Refugia Followed by Human-Influenced Dispersal". PLoS ONE. 7 (10): e47676. doi:10.1371/journal.pone.0047676. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: unflagged free DOI (link)
  6. ^ a b Terry G. Powis, W. Jeffrey Hurst, María del Carmen Rodríguez, Ponciano Ortíz C., Michael Blake, David Cheetham, Michael D. Coe & John G. Hodgson (2007). "Oldest chocolate in the New World". Antiquity. 81 (314). ISSN 0003-598X. Retrieved 2011-02-15. {{cite journal}}: Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link)
  7. ^ J.S. Henderson; et al. (2007). "Chemical and archaeological evidence for the earliest cacao beverages". Proceedings of the National Academy of Sciences. 104: 18937–18940. {{cite journal}}: Explicit use of et al. in: |author= (help)
  8. ^ (Kidder 1947)
  9. ^ (Hammond and Miksicek 1981; Turner and Miksicek 1984)
  10. ^ J. Bergmann (1969).
  11. ^ S. Coe (1994).
  12. ^ (Bogin 1997, Coe 1996, Montejo 1999, Tedlock 1985)
  13. ^ (Coe 1996, Townsend 1992)
  14. ^ (Coe and Coe 1996)
  15. ^ http://findarticles.com/p/articles/mi_m1134/is_6_112/ai_105371465/pg_3/ CBS Interactive Business Network
  16. ^ FAO.org
  17. ^ Anon. "A strategy to safeguard the future of chocolate". Bioversity International. Retrieved 9 November 2012.
  18. ^ All about Chocolate -- Varieties
  19. ^ Juan C. Motamayor, Philippe Lachenaud, Jay Wallace da Silva e Mota, Rey Loor, David N. Kuhn, J. Steven Brown, Raymond J. Schnell (2008). "Geographic and Genetic Population Differentiation of the Amazonian Chocolate Tree (Theobroma cacao L.)". PLoS ONE. 3 (10). doi:10.1371/journal.pone.0003311.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
  20. ^ Cocoa-derived Cirku launched for better circulation
  21. ^ "Cacao Collections". CacaoNet. Retrieved 9 November 2012.
  22. ^ Thomas, Evert (24 October 2012). "Present Spatial Diversity Patterns of Theobroma cacao L. in the Neotropics Reflect Genetic Differentiation in Pleistocene Refugia Followed by Human-Influenced Dispersal". PLoS ONE. 7 (10): e47676. doi:10.1371/journal.pone.0047676. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: unflagged free DOI (link)
  23. ^ The Cacao Genome Database Project. Retrieved September 24, 2010
  24. ^ The International Cocoa Genome Sequencing Consortium federates efforts from circa 20 different institutions from six countries (France, USA, Côte d’Ivoire, Brazil, Venezuela and Trinidad et Tobago). Financing comes from several public and private sources from France, USA and Venezuela, among which the chocolate brands VALRHONA (France) and HERSHEY’S (USA). See : http://www.cirad.fr/actualites/toutes-les-actualites/communiques-de-presse/2010/decryptage-du-genome-du-cacaoyer
  25. ^ Xavier Argout, Jerome Salse, Jean-Marc Aury, Mark J Guiltinan, Gaetan Droc, Jerome Gouzy, Mathilde Allegre, Cristian Chaparro, .. & Claire Lanaud (2011). "The genome of Theobroma cacao". Nature Genetics. 43 (2): 101–108. doi:10.1038/ng.736. PMID 21186351.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  26. ^ Olivier Jaillon, Jean-Marc Aury, Benjamin Noel, Alberto Policriti, Christian Clepet, Alberto Casagrande .. & Patrick Wincker (2007). "The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla". Nature. 449 (7161): 463–467. doi:10.1038/nature06148. PMID 17721507.{{cite journal}}: CS1 maint: multiple names: authors list (link)

References

  • Coe, Sophie D. (1994). America's First Cuisines. Austin: University of Texas Press. ISBN 0-292-71155-7. {{cite book}}: templatestyles stripmarker in |author= at position 1 (help)CS1 maint: numeric names: authors list (link)
  • Coe, Sophie D. (1996). The True History of Chocolate. London: Thames & Hudson. ISBN 0-500-01693-3. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); templatestyles stripmarker in |author= at position 1 (help); templatestyles stripmarker in |coauthors= at position 5 (help)CS1 maint: numeric names: authors list (link)
  • Dienhart, John M. (1997). "The Mayan Languages- A Comparative Vocabulary" (electronic version (PDF)). Odense University. Retrieved 2007-02-14. {{cite web}}: templatestyles stripmarker in |author= at position 1 (help)CS1 maint: numeric names: authors list (link)
  • McNeil, Cameron (editor) (2006). Chocolate in Mesoamerica: A Cultural History of Cacao. Gainesville: University of Florida Press. ISBN 0-8130-2953-8. {{cite book}}: |author= has generic name (help); templatestyles stripmarker in |author= at position 1 (help)CS1 maint: numeric names: authors list (link)

External links

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