View source for Theobroma cacao

{{Short description|Species of tree grown for its seeds}}
{{Other uses|Cacao (disambiguation){{!}}Cacao|Cocoa (disambiguation){{!}}Cocoa}}
{{Use dmy dates|date=June 2018}}
{{Speciesbox
|image = Matadecacao.jpg
|image_caption = Cacao fruits on the tree
|genus = Theobroma
|species = cacao
|authority = [[Carl Linnaeus|L.]]<ref name="WFO">{{cite web |title=''Theobroma cacao'' L. Sp. Pl. : 782 (1753) |url=http://www.worldfloraonline.org/taxon/wfo-0000458440 |website=World Flora Online  |date=2022 |publisher=World flora Consortium |access-date=14 December 2022}}</ref>
|synonyms = *''Cacao mina'' {{small|Gaertn.}} 
*''Cacao minus '' <small>Gaertn.</small>
*''Cacao sativa'' <small> Aubl.</small> 
*''Cacao theobroma'' <small> Tussac </small> 
*''Theobroma cacao f. leiocarpum '' <small>(Bernoulli) Ducke</small> 
*''Theobroma cacao subsp. leiocarpum '' <small>(Bernoulli) Cuatrec.</small> 
*''Theobroma cacao var. leiocarpum '' <small>(Bernoulli) Cif.</small>
*''Theobroma cacao subsp. sativa '' <small>(Aubl.) León</small>
*''Theobroma cacao var. typica '' <small>Cif.</small> 
*''Theobroma caribaea '' <small>Sweet</small> 
*''Theobroma integerrima '' <small>Stokes</small> 
*''Theobroma kalagua'' <small> De Wild.</small>
*''Theobroma leiocarpum'' <small> Bernoulli</small> 
*''Theobroma pentagonum '' <small>Bernoulli</small>
*''Theobroma saltzmanniana '' <small>Bernoulli</small>
*''Theobroma sapidum '' <small>Pittier</small> 
*''Theobroma sativa'' <small> (Aubl.) Lign. & Le Bey</small> 
*''Theobroma sativa var. leucosperma '' <small>A. Chev.</small> 
*''Theobroma sativa var. melanosperma '' <small>A. Chev.</small> 
*''Theobroma sativum'' <small>(Aubl.) Lign. & Bey</small>
| synonyms_ref = <ref name="WFO"/><ref name="POWO">{{cite web |url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:320783-2#synonyms |title=''Mentha canadensis'' L. |date=2022 |work=Plants of the World Online |publisher=Royal Botanic Gardens, Kew|access-date=14 December 2022 }}</ref>
}}

'''''Theobroma cacao''''' ('''cacao tree''' or '''cocoa tree''') is a small ({{convert|6|-|12|m|ft|abbr=on|0}} tall) [[evergreen]] [[tree]] in the [[Malvaceae]] family.<ref name="WFO" /><ref>{{cite encyclopedia |title=Theobroma cacao |url=http://eol.org/pages/484592/overview |encyclopedia=Encyclopedia of Life |access-date=9 November 2012 |archive-date=4 November 2012 |archive-url=https://web.archive.org/web/20121104063223/http://eol.org/pages/484592/overview |url-status=live}}</ref> Its seeds{{Mdash}}[[cocoa bean]]s{{Mdash}}are used to make [[chocolate liquor]], [[cocoa solids]], [[cocoa butter]] and [[chocolate]].<ref name="drugs">{{cite web |url=https://www.drugs.com/npp/cocoa.html |title=Cocoa |publisher=Drugs.com |date=6 January 2020 |access-date=3 September 2020 |archive-date=20 January 2021|archive-url=https://web.archive.org/web/20210120103652/https://www.drugs.com/npp/cocoa.html|url-status=live}}</ref>  Although the tree is native to the [[tropics]] of the [[Americas]], the largest producer of cocoa beans in 2022 was [[Ivory Coast]]. The plants [[leaf|leaves]] are alternate, entire, unlobed, {{convert|10|-|50|cm|in|abbr=on|0}} long and {{convert|5|-|10|cm|in|abbr=on|0}} broad.

== Description ==

=== Flowers ===

The [[flower]]s are produced in clusters directly on the [[Trunk (botany)|trunk]] and older branches; this is known as [[cauliflory]]. The flowers are small, {{convert |1 |- |2 |cm |in |abbr=on |frac=16}} diameter, with pink [[Calyx (botany)|calyx]]. The [[floral formula]], used to represent the structure of a [[flower]] using numbers, is ✶ K5 C5 A(5°+5<sup>2</sup>) {{underline |G}}(5).<ref name="craene">{{cite book |last=Ronse De Craene |first=Louis P. |title=Floral Diagrams: An Aid to Understanding Flower Morphology and Evolution |publisher=[[Cambridge University Press]] |location=Cambridge |date=4 February 2010 |url=https://books.google.com/books?id=24p-LgWPA50C |page=224 |isbn=978-0-521-49346-8 |access-date=16 September 2017 |archive-date=28 April 2021 |archive-url=https://web.archive.org/web/20210428023554/https://books.google.com/books?id=24p-LgWPA50C |url-status=live}}</ref>

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, ''[[Forcipomyia]]'' biting [[midge]]s.<ref>{{cite web |last=Hernández B. |first=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&method=post&formato=2&cantidad=1&expresion=mfn=032010 |access-date=27 November 2012 |archive-date=4 October 2013 |archive-url=https://web.archive.org/web/20131004220619/http://orton.catie.ac.cr/cgi-bin/wxis.exe/?IsisScript=ORTON.xis&method=post&formato=2&cantidad=1&expresion=mfn=032010 |url-status=dead}}</ref><ref name=":2">{{cite journal |last1=Forbes |first1=Samantha J. |last2=Northfield |first2=Tobin D. |date=26 December 2016 |title=Increased pollinator habitat enhances cacao fruit set and predator conservation |journal=[[Ecological Applications]] |doi=10.1002/eap.1491 |pmid=28019052 |volume=27 |issue=3 |pages=887–899 |s2cid=6858627}}</ref> Using the natural pollinator ''Forcipomyia'' midges produced more fruit than using artificial pollinators.<ref name=":2" />

[[File:Theobroma cacao2.jpg|thumb|center|Cacao flowers]]

=== Fruit ===

The [[fruit]], called a cacao pod, is ovoid, {{convert|15|-|30|cm|in|abbr=on|0}} long and {{convert|8|-|10|cm|in|abbr=on|0}} wide, ripening yellow to orange, and weighs about {{convert|500|g|lb|abbr=on|0}} 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 [[juice]], [[smoothie]]s, [[Jelly (fruit preserves)|jelly]], and cream. Usually discarded until practices changed in the 21st century, the fermented pulp may be distilled into an alcoholic beverage.<ref>{{cite web |last=Bell |first=Katie K. |url=https://www.forbes.com/sites/katiebell/2014/09/10/cacao-cocktails-a-new-tequila-like-spirit-distilled-from-cacao-fruit/ |title=Cacao Cocktails: A New Tequila-Like Spirit Distilled from Cacao Fruit  |website=[[Forbes]] |access-date=17 March 2015 |archive-date=28 April 2015 |archive-url=https://web.archive.org/web/20150428064621/http://www.forbes.com/sites/katiebell/2014/09/10/cacao-cocktails-a-new-tequila-like-spirit-distilled-from-cacao-fruit/ |url-status=live}}</ref> The pulp tastes like [[lychee]] or [[Melon|tropical melon]].<ref>{{Cite web |last=Martin |first=Carla D |author-link=Carla Martin (anthropologist) |date=March 9, 2020 |title=What Makes Chocolate "Good"? |url=https://www.youtube.com/watch?v=PFGKUZdGap4 |url-status= |access-date=February 9, 2025 |via=[[YouTube]] |publisher=[[Peabody Museum of Archaeology and Ethnology]] |at=19:35}}</ref> Each seed contains a significant amount of fat (40–50%) as [[cocoa butter]].

The fruit's active constituent is the stimulant [[theobromine]], a compound similar to [[caffeine]].<ref>{{cite book |last=Wilson |first=Philip K. |editor-last1=Wilson |editor-first1=P. |editor-last2=Hurst |editor-first2=W. |year=2015 |chapter=Chocolate in Science, Nutrition and Therapy: A Historical Perspective |title=Chocolate and Health: Chemistry, Nutrition and Therapy |url=https://pubs.rsc.org/en/content/ebook/978-1-84973-912-2 |location=Cambridge, UK |publisher=Royal Society of Chemistry |page=18 |isbn=978-1-84973-912-2}}</ref>

[[File:Chocolate in its Rawest Form (27583224425).jpg|thumb|upright|Fruit]]

== Nomenclature==

The generic name ''[[Theobroma]]'' is derived from the [[Greek language|Greek]] for "food of the gods"; from {{wikt-lang|grc|θεός}} ({{lang|grc-Latn|theós}}), meaning 'god' or 'divine', and {{wikt-lang|grc|βρῶμα}} ({{lang|grc|brôma}}), meaning 'food'. The specific name ''cacao'' is the [[Spanish language|Hispanization]] of the name given to the plant in indigenous [[Mesoamerican languages]] such as {{lang|emy-Latn|kakaw}} in [[Tzeltal language|Tzeltal]], [[Kʼicheʼ language|Kʼicheʼ]] and [[Classic Maya language|Classic Maya]]; {{lang|pos|kagaw}} in [[Sayula Popoluca]]; and {{lang|nah|cacahuatl}} in [[Nahuatl]] meaning "bean of the cocoa-tree".<ref>{{cite web |title=Cacao |url=https://www.etymonline.com/word/cacao |publisher=Etymology Online Dictionary, Douglas Harper |access-date=19 August 2018 |date=2018 |archive-date=19 August 2018 |archive-url=https://web.archive.org/web/20180819182509/https://www.etymonline.com/word/cacao |url-status=live }}</ref> While the tree and seeds are generally called cacao, they are sometimes referred to as 'cocoa'.<ref>{{Cite book |last=Coe |first=Sophie |author-link=Sophie Coe |title=The Oxford Companion to Food |last2=Dalby |first2=Andrew |publisher=[[Oxford University Press]] |year=2014 |isbn=9780199677337 |editor-last=Davidson |editor-first=Alan |edition=3rd |chapter=Chocolate: Botany and early history |editor-last2=Jaine |editor-first2=Tom}}</ref>

=== Taxonomy ===

Cacao (''Theobroma cacao'') is one of 26 species belonging to the [[genus]] ''[[Theobroma]]'' classified under the subfamily [[Byttnerioideae]] of the mallow [[family (biology)|family]] [[Malvaceae]].<ref name=WFO/>

In 2008, researchers proposed a new classification based upon [[morphology (biology)|morphological]], geographic, and [[genomic]] criteria: 10 groups have been named according to their geographic origin or the traditional [[cultivar]] name. These groups are: Amelonado, Criollo, Nacional, Contamana, Curaray, Cacao guiana, Iquitos, Marañon, Nanay, and Purús.<ref name="Juan C. Motamayor">{{cite journal |first1=Juan C. |last1=Motamayor |first2=Philippe |last2=Lachenaud |first3=Jay Wallace |last3=da Silva e Mota |first4=Rey |last4=Loor |first5=David N. |last5=Kuhn |first6=J. Steven |last6=Brown |first7=Raymond J. |last7=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 |article-number=e3311 |pmid=18827930 |pmc=2551746 |bibcode=2008PLoSO...3.3311M |doi-access=free }}</ref>

==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 Jungle]] area of Mexico and another in lowland [[South America]].{{citation needed|date=January 2024}} More recent studies of patterns of DNA diversity, however, suggest that this is not the case. One study sampled 1241 trees and classified them into 10 distinct genetic clusters.<ref name="Juan C. Motamayor" /> This study also identified areas, for example around [[Iquitos]] in modern [[Peru]] and [[Ecuador]], where representatives of several genetic clusters originated more than 5000 years ago, leading to development of the variety, [[Nacional (cocoa bean)|Nacional cocoa bean]].<ref>{{cite web |url=http://www.anecacao.com/es/quienes-somos/historia-del-cacao.html |title=History of cocoa (translated) |publisher=National Association of Exporters and Industrialists of Cacao of Ecuador |date=2015 |access-date=20 February 2018 |archive-date=21 February 2018 |archive-url=https://web.archive.org/web/20180221035529/http://www.anecacao.com/es/quienes-somos/historia-del-cacao.html|url-status=live}}</ref> 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 |last1=Clement |first1=Charles R. |last2=de Cristo-Araújo |first2=Michelly |last3=d'Eeckenbrugge |first3=Geo Coppens |last4=Alves Pereira |first4=Alessandro |last5=Picanço-Rodrigues |first5=Doriane |title=Origin and Domestication of Native Amazonian Crops |journal=Diversity |date=6 January 2010 |volume=2 |issue=1 |pages=72–106 |doi=10.3390/d2010072 |url=http://agritrop.cirad.fr/553502/ |doi-access=free |access-date=5 September 2019 |archive-date=5 September 2019 |archive-url=https://web.archive.org/web/20190905095250/http://agritrop.cirad.fr/553502/ |url-status=live}}</ref> Using the DNA sequences and comparing them with data derived from climate models and the known conditions suitable for cacao, one study refined the view of domestication, linking the area of greatest cacao genetic diversity to a bean-shaped area that encompasses [[Ecuador]], the border between Brazil and Peru and the southern part of the Colombian–Brazilian border.<ref name="Thomas e47676">{{cite journal|last1=Thomas|first1=Evert|last2=van Zonneveld|first2=Maarten|last3=Loo|first3=Judy |last4=Hodgkin|first4=Toby|last5=Galluzzi|first5=Gea|last6=van Etten|first6=Jacob|last7=Fuller|first7=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 |article-number=e47676 |doi=10.1371/journal.pone.0047676 |pmid=23112832 |pmc=3480400 |bibcode=2012PLoSO...747676T |doi-access=free}}</ref> Climate models indicate that at the peak of the [[Last glacial period|last 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 species.

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.{{citation needed|date=January 2024}}

== Cultivation ==

{{See also |Cocoa bean}}

[[File:CacaoChonita14.JPG|thumb|Toasted cacao beans, ready for use]]

In 2023, cocoa beans were cultivated on roughly {{convert |11650000 |ha |acres}} worldwide.<ref name="faostat2023">{{cite web |url=https://www.fao.org/faostat/en/#data/QCL/visualize |title=Crops and livestock products; Production quantities of Cocoa beans by country |publisher=United Nations Food and Agriculture Organization, Statistics Division (FAOSTAT) |date=2023 |access-date=22 May 2025}}</ref> Cocoa beans are grown by large agroindustrial plantations and small producers, the bulk of production coming from millions of farmers with small plots.<ref>{{cite web |title=A strategy to safeguard the future of chocolate |url=http://www.bioversityinternational.org/index.php?id=6817 |archive-url=https://archive.today/20130414081745/http://www.bioversityinternational.org/index.php?id=6817 |url-status=dead |archive-date=14 April 2013 |publisher=Bioversity International |access-date=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 1,200 seeds (40 pods) are required to produce {{convert |1 |kg |lb |abbr=on}} of [[chocolate liquor|cocoa paste]].
Historically, chocolate makers have recognized three main [[cultivar |cultivar groups]] of cacao beans used to make cocoa and chocolate: Forastero, Criollo and Trinitario.<ref>{{cite web |url=http://www.xocoatl.org/variety.htm |website=All about Chocolate |title=Varieties |access-date=3 June 2017 |archive-date=2 July 2017 |archive-url=https://web.archive.org/web/20170702090148/http://xocoatl.org/variety.htm |url-status=live}}</ref> The most prized, rare, and expensive is the Criollo group, the cocoa bean used by the [[Maya civilization|Maya]].<ref>{{Cite journal |last1=Lanaud |first1=Claire |last2=Vignes |first2=Hélène |last3=Utge |first3=José |last4=Valette |first4=Gilles |last5=Rhoné |first5=Bénédicte |last6=Garcia Caputi |first6=Mariella |last7=Angarita Nieto |first7=Natalia Sofía |last8=Fouet |first8=Olivier |last9=Gaikwad |first9=Nilesh |last10=Zarrillo |first10=Sonia |last11=Powis |first11=Terry G. |last12=Cyphers |first12=Ann |last13=Valdez |first13=Francisco |last14=Olivera Nunez |first14=S. Quirino |last15=Speller |first15=Camilla |display-authors=5 |date=2024-03-07 |title=A revisited history of cacao domestication in pre-Columbian times revealed by archaeogenomic approaches |url= |journal=[[Scientific Reports]] |volume=14 |issue=1 |page=2972 |doi=10.1038/s41598-024-53010-6 |issn=2045-2322 |doi-access=free |pmid=38453955 |pmc=10920634 |bibcode=2024NatSR..14.2972L }}</ref> Only 10% of chocolate is made from Criollo, which is arguably less bitter and more aromatic than any other bean. In November 2000, the cacao beans coming from [[Chuao]] were awarded an appellation of origin under the title {{lang |es |Cacao de Chuao}} (from [[Spanish language|Spanish]]: 'cacao of Chuao').<ref>{{cite web |url=http://www.wipo.int/ipadvantage/en/details.jsp?id=2618 |title=Branding Matters: The Success of Chuao Cocoa Bean |access-date=3 June 2017 |archive-date=3 March 2017 |archive-url=https://web.archive.org/web/20170303044314/http://www.wipo.int/ipadvantage/en/details.jsp?id=2618 |url-status=live}}</ref>

The cacao bean in 80% of chocolate is made using beans of the Forastero group, the main and most ubiquitous variety being the Amenolado variety, while the Arriba variety (such as the Nacional variety) are less commonly found in Forastero produce. Forastero trees are significantly hardier and more disease-resistant than Criollo trees, resulting in cheaper cacao beans.<ref>{{Cite journal |last1=Kongor |first1=John Edem |last2=Hinneh |first2=Michael |last3=de Walle |first3=Davy Van |last4=Afoakwa |first4=Emmanuel Ohene |last5=Boeckx |first5=Pascal |last6=Dewettinck |first6=Koen |date=2016-04-01 |title=Factors influencing quality variation in cocoa (Theobroma cacao) bean flavour profile — A review |url=https://www.sciencedirect.com/science/article/pii/S0963996916300163 |journal=Food Research International |language=en |volume=82 |pages=44–52 |doi=10.1016/j.foodres.2016.01.012 |issn=0963-9969|url-access=subscription }}</ref>

Major cocoa bean processors include [[The Hershey Company|Hershey's]], [[Nestlé]] and [[Mars, Incorporated|Mars]]. Chocolate can be made from ''T. cacao'' through a process of steps that involve harvesting, fermenting of ''T. cacao'' pulp, drying, harvesting, and then extraction.<ref name="Zzaman">{{Cite journal |last1=Zzaman |first1=Wahidu |last2=Bhat |first2=Rajeev |last3=Yang |first3=Tajul Aris |last4=Mat Easa |first4=Azhar |date=2 March 2017 |title=Influences of superheated steam roasting on changes in sugar, amino acid and flavor active components of cocoa bean (Theobroma cacao) |journal=Journal of the Science of Food and Agriculture |volume=97 |issue=13 |pages=4429–4437 |doi=10.1002/jsfa.8302 |issn=1097-0010 |pmid=28251656 |bibcode=2017JSFA...97.4429Z }}</ref> Roasting ''T. cacao'' by using superheated steam was found to be better than conventional oven-roasting because it resulted in the same quality of cocoa beans in a shorter time.<ref name="Zzaman"/>

== Production ==
{{Table alignment}}
{| class="wikitable floatright sortable col2right"
|+ Cocoa bean production (2023)<ref name="faostat2023"/>
|-
! Country
! Production<br/>(metric tonnes)
|-
| {{CIV}}
| 2,377,442
|-
| {{GHA}}
| 653,700
|-
| {{IDN}}
| 641,741
|-
| {{ECU}}
| 375,719
|-
| {{BRA}}
|296,145
|-
| {{CMR}}
| 295,819
|-
| {{NGA}}
| 284,232
|- class="sortbottom"
! World
! 5,600,100
|}

In 2023, world production of cocoa beans was 5.6 million [[tonne]]s, led by [[Ivory Coast]] with 42.5% of the total (table). Other major producers were [[Ghana]] (11.7%) and [[Indonesia]] (11.5%).

== 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|access-date=9 November 2012|archive-date=14 May 2015|archive-url=https://web.archive.org/web/20150514215705/https://sites.google.com/a/cgxchange.org/cacaonet/cacao-genebanks|url-status=live}}</ref> In an effort to improve the diversity available to breeders, and ensure the future of the field genebanks, experts have drawn up A Global Strategy for the Conservation and Use of Cacao Genetic Resources, as the Foundation for a Sustainable Cocoa Economy.<ref>{{cite web|url=https://sites.google.com/a/cgxchange.org/cacaonet/cacaonet-global-strategy|title=Global Strategy – cacaonet|website=sites.google.com|access-date=19 July 2018|archive-date=8 October 2018|archive-url=https://web.archive.org/web/20181008150644/https://sites.google.com/a/cgxchange.org/cacaonet/cacaonet-global-strategy|url-status=live}}</ref> 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 name="Thomas e47676"/> 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.  Other projects, such as the [[International Cocoa Quarantine Centre]], aim to combat cacao diseases and preserve genetic diversity.

Phytopathogens (parasitic organisms) cause much damage to ''Theobroma cacao'' plantations around the world. Many of those phytopathogens, which include many of the pests named below, were analyzed using mass spectrometry and allow for guiding on the correct approaches to get rid of the specific phytopathogens. This method was found to be quick, reproducible, and accurate showing promising results in the future to prevent damage to ''Theobroma cacao'' by various phytopathogens.<ref>{{Cite journal|last1=Dos Santos|first1=Fábio Neves|last2=Tata |first2=Alessandra|last3=Belaz|first3=Kátia Roberta Anacleto|last4=Magalhães|first4=Dilze Maria Argôlo|last5=Luz |first5=Edna Dora Martins Newman|last6=Eberlin|first6=Marcos Nogueira|date=1 March 2017 |title=Major phytopathogens and strains from cocoa (Theobroma cacao L.) are differentiated by MALDI-MS lipid and/or peptide/protein profiles |journal=Analytical and Bioanalytical Chemistry |volume=409 |issue=7 |pages=1765–1777 |doi=10.1007/s00216-016-0133-5 |pmid=28028594 |s2cid=3764608}}</ref>

A specific bacterium ''Streptomyces camerooniansis'' was found to be beneficial for ''T. cacao'' by helping plant growth by accelerating seed germination of ''T. cacao,'' inhibiting growth of various types of microorganisms (such as different oomycetes, fungi, and bacteria), and preventing rotting by ''[[Phytophthora megakarya]]''.<ref>{{cite journal |last1=Boudjeko|first1=Thaddée|last2=Tchinda|first2=Romaric Armel Mouafo|last3=Zitouni|first3=Mina|last4=Nana|first4=Joëlle Aimée Vera Tchatchou |last5=Lerat|first5=Sylvain |last6=Beaulieu|first6=Carole |date=4 March 2017|title=Streptomyces cameroonensis sp. nov., a Geldanamycin Producer That Promotes Theobroma cacao Growth|journal=Microbes and Environments|doi=10.1264/jsme2.ME16095|issn=1347-4405|pmid=28260703|pmc=5371071|volume=32|issue=1|pages=24–31}}</ref>

=== Pests ===

{{See also|List of cacao diseases}}

Various plant pests and diseases can cause serious problems for cacao production.<ref>{{cite web|title=Cocoa Crop Protection|url=http://www.dropdata.org/cocoa/|access-date=9 November 2012|archive-date=9 February 2013|archive-url=https://web.archive.org/web/20130209060931/http://www.dropdata.org/cocoa/|url-status=live}}</ref>

*[[Insect]]s
**Cocoa [[Miridae|mirids or capsids]] worldwide (but especially ''Sahlbergella singularis'' and ''Distantiella theobroma'' in [[West Africa]] and ''[[Helopeltis]]'' spp. in [[Southeast Asia]])
**''[[Bathycoelia thalassina]]'' - West Africa
**''[[Conopomorpha cramerella]]'' (cocoa pod borer – in [[Southeast Asia]])
**''[[Carmenta theobromae]]'' - C. & S. America
*[[Fungus|Fungi]]
**''[[Moniliophthora roreri]]'' (frosty pod rot)
**''[[Moniliophthora perniciosa]]'' (witches' broom)
**''[[Ceratocystis cacaofunesta]]'' (''mal de machete'') or (''Ceratocystis'' wilt)
**''[[Verticillium dahliae]]''
**''[[Oncobasidium theobromae]]'' (vascular streak dieback)
*[[Oomycete]]s
**''[[Phytophthora]]'' spp. ([[black pod disease|black pod]]) especially ''[[Phytophthora megakarya]]'' in West Africa
*[[Virus]]es
**[[Cacao swollen shoot virus]]
* [[Mistletoe]]
*[[Rat]]s and other [[vertebrate]] pests ([[squirrel]]s, [[woodpecker]]s, ''etc.'')

== Genome ==

{{Infobox genome 
| image = Cacao Genetic Clusters.png
| caption = Map showing genetic clusters of ''Theobroma cacao''
| taxId = 572
| ploidy = diploid
| chromosomes = 10 pairs
| size = 345.99 Mb
| year = 2010
}}

The [[genome]] of ''T. cacao'' is diploid, its size is 430 M[[Base pair|bp]], 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).<ref>{{cite web|url=http://www.cacaogenomedb.org/|title=Welcome to the Cacao Genome Project – Cacao Genome Database|website=Cacaogenomedb.org|access-date=19 July 2018|archive-date=20 July 2018|archive-url=https://web.archive.org/web/20180720051951/https://www.cacaogenomedb.org/|url-status=usurped}}</ref> In a second, unrelated project, the International Cocoa Genome Sequencing Consortium-ICGS, coordinated by CIRAD,<ref>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 {{Webarchive|url=https://web.archive.org/web/20110116220553/http://www.cirad.fr/actualites/toutes-les-actualites/communiques-de-presse/2010/decryptage-du-genome-du-cacaoyer |date=16 January 2011 }}</ref> first published<ref name="Argout">{{cite journal|first1=Xavier |last1=Argout |first2=Jerome  |last2=Salse|first3=Jean-Marc |last3=Aury|first4=Mark J. |last4=Guiltinan|first5=Gaetan |last5=Droc |first6=Jerome |last6=Gouzy|first7=Mathilde |last7=Allegre|first8=Cristian |last8=Chaparro|display-authors=etal|year=2011|title=The genome of Theobroma cacao|journal=[[Nature Genetics]]|volume=43|issue=2|pmid=21186351|pages=101–108|doi=10.1038/ng.736|s2cid=4685532 |url=http://agritrop.cirad.fr/558337/1/document_558337.pdf }}</ref> 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 [[flavonoid]]s, [[Aromaticity|aromatic]] [[terpene]]s, [[theobromine]] and many other [[metabolite]]s involved in cocoa flavor and quality traits, among which a relatively high proportion code for [[polyphenol]]s, which constitute up to 8% of cacao pods dry weight. The cacao genome appears close to the hypothetical [[Polyploid|hexaploid]] ancestor of all [[dicotyledonous]] plants,<ref>{{cite journal |first1=Olivier |last1=Jaillon |first2=Jean-Marc |last2=Aury |first3=Benjamin |last3=Noel |first4=Alberto |last4=Policriti |first5=Christian |last5=Clepet |first6=Alberto |last6=Casagrande |display-authors=etal|year=2007|title=The grapevine genome sequence suggests ancestral hexaploidization in major angiosperm phyla|journal=[[Nature (journal)|Nature]]|volume=449|issue=7161|pmid=17721507|pages=463–467|doi=10.1038/nature06148|bibcode=2007Natur.449..463J |doi-access=free |hdl=11577/2430527 |hdl-access=free }}</ref> 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 enables cacao molecular biology and breeding for elite varieties through marker-assisted selection, in particular for genetic resistance to [[Fungus|fungal]], oomycete and viral diseases responsible for huge yield losses each year. In 2017–18, due to concerns about survivability of cacao plants in an era of [[global warming]] in which climates become more extreme in the narrow band of latitudes where cacao is grown (20 degrees north and south of the [[equator]]), the commercial company, [[Mars, Incorporated]] and the [[University of California, Berkeley]] are using [[CRISPR]] to adjust [[DNA]] for improved [[hardiness (plants)|hardiness]] of cacao in hot climates.<ref name="mars">{{cite web |last1=Brodwin|first1=Erin |title=Chocolate is on track to go extinct in 40 years |url=http://www.businessinsider.com/when-chocolate-extinct-2017-12 |work=Business Insider |access-date=31 January 2018 |date=31 December 2017|archive-date=31 January 2018 |archive-url=https://web.archive.org/web/20180131040813/http://www.businessinsider.com/when-chocolate-extinct-2017-12|url-status=live}}</ref>

== History of cultivation ==

=== Domestication ===

The cacao tree, native of the Amazon rainforest, was first domesticated at least 5,300 years ago, in equatorial South America from the [[Santa Ana (La Florida)|Santa Ana-La Florida]] (SALF) site in what is present-day southeast [[Ecuador]] ([[Zamora-Chinchipe Province]]) by the [[Mayo-Chinchipe]] culture before being introduced in Mesoamerica.<ref name=":02">{{cite journal |last1=Zarrillo |first1=Sonia |last2=Gaikwad |first2=Nilesh |last3=Lanaud |first3=Claire |last4=Powis |first4=Terry |last5=Viot |first5=Christopher |last6=Lesur |first6=Isabelle |last7=Fouet |first7=Olivier |last8=Argout |first8=Xavier |last9=Guichoux |first9=Erwan |last10=Salin |first10=Franck |last11=Solorzano |first11=Rey Loor |last12=Bouchez |first12=Olivier |last13=Vignes |first13=Hélène |last14=Severts |first14=Patrick |last15=Hurtado |first15=Julio |display-authors=6 |date=2018-10-29 |title=The use and domestication of Theobroma cacao during the mid-Holocene in the upper Amazon |url=https://www.nature.com/articles/s41559-018-0697-x |journal=[[Nature Ecology & Evolution]] |language=en |volume=2 |issue=12 |pages=1879–1888 |bibcode=2018NatEE...2.1879Z |doi=10.1038/s41559-018-0697-x |pmid=30374172 |s2cid=53099825|url-access=subscription }}</ref>

In Mesoamerica, ceramic vessels with residues from the preparation of cacao beverages have been found from the [[Mesoamerican chronology|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.<ref name="aj200712">{{cite journal |url=http://antiquity.ac.uk/projgall/powis/index.html |title=Oldest chocolate in the New World |first1=Terry G. |last1=Powis |first2=W. Jeffrey |last2=Hurst |first3=María |last3=del Carmen Rodríguez |first4=Ponciano |last4=Ortíz C. |first5=Michael |last5=Blake |first6=David |last6=Cheetham |first7=Michael D. |last7=Coe |first8=John G. |last8=Hodgson |journal=[[Antiquity (journal)|Antiquity]] |volume=81 |issue=314 |date=December 2007 |access-date=15 February 2011 |archive-date=15 September 2018 |archive-url=https://web.archive.org/web/20180915102140/http://antiquity.ac.uk/projgall/powis/index.html |url-status=live }}</ref> On the Pacific coast of [[Chiapas]], Mexico, a [[Mokaya]] archaeological site provides evidence of even earlier cacao beverages, to 1900 BC.<ref name="aj200712"/> The initial domestication was probably related to the making of a fermented alcoholic beverage.<ref>{{cite journal |first=J. S. |last=Henderson |title=Chemical and archaeological evidence for the earliest cacao beverages |journal=[[Proceedings of the National Academy of Sciences]] |year=2007 |volume= 104 |issue=48 |pages=18937–18940 |doi=10.1073/pnas.0708815104 |display-authors=etal |pmid=18024588 |pmc=2141886 |doi-access=free}}</ref> In 2018, researchers who analysed the genome of cultivated cacao trees concluded that the domesticated cacao trees all originated from a single domestication event that occurred about 3,600 years ago somewhere in Central America.<ref>{{Cite web |url=https://www.eurekalert.org/pub_releases/2018-10/wsu-cad102218.php |title="Cacao analysis dates the dawn of domesticated chocolate trees to 3,600 years ago", Eurekalert, October 24, 2018 |access-date=24 October 2018 |archive-date=24 October 2018 |archive-url=https://web.archive.org/web/20181024203409/https://www.eurekalert.org/pub_releases/2018-10/wsu-cad102218.php |url-status=live }}</ref>

=== Ancient uses ===

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 (Maya site)|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&nbsp;BC) using similar techniques to those used to extract chocolate residues from four classic period (around 400&nbsp;AD) vessels from a tomb at the Maya 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 is also believed to have been ground by the Aztecs and mixed with tobacco for smoking purposes.{{citation needed|date=September 2012}} Cocoa was being domesticated by the Mayo Chinchipe of the upper Amazon around 3,000 BC.<ref>{{cite news |last1=Davis |first1=Nicola |title=Origin of chocolate shifts 1,400 miles and 1,500 years |url=https://www.theguardian.com/science/2018/oct/29/origin-of-chocolate-shifts-1400-miles-and-1500-years-cacao-ecuador |access-date=31 October 2018 |work=[[The Guardian]] |date=29 October 2018 |archive-date=30 October 2018 |archive-url=https://web.archive.org/web/20181030234709/https://www.theguardian.com/science/2018/oct/29/origin-of-chocolate-shifts-1400-miles-and-1500-years-cacao-ecuador |url-status=live }}</ref>

The [[Maya civilization|Maya]] believed the {{lang|yua|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 ({{lang|nci|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 was used as a [[ritual]] only by men, as it was believed to be an intoxicating food unsuitable for women and children.<ref>{{Cite journal|last1=Grivetti|first1=Louis E. |last2=Lowe|first2=Diana Salazar |last3=Jimenez|first3=Martha |last4=Escárcega|first4=Sylvia |last5=Barriga |first5=Patricia |last6=Dillinger|first6=Teresa L. |date=2000-08-01 |title=Food of the Gods: Cure for Humanity? A Cultural History of the Medicinal and Ritual Use of Chocolate |journal=The Journal of Nutrition |volume=130 |issue=8 |pages=2057S–72S |doi=10.1093/jn/130.8.2057S |pmid=10917925 |doi-access=free}}</ref>

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 ({{langx|nci|xiquipilli}}) 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>

<gallery mode=packed heights=180>
File:Cacao written as 2ka-wa Kettunen-midres.png|''kakaw'' (cacao) written in the [[Maya script]]
File:Aztec. Man Carrying a Cacao Pod, 1440-1521.jpg|Sculpture of a man carrying a cacao pod. Aztec, 1440-1521 AD
File:Codex Fejéváry-Mayer 01 cacao tree.JPG|A cacao tree in the Aztec [[Codex Fejérváry-Mayer]]
</gallery>

=== Modern history ===

[[File:East Indian Immigrants in Cacao Estate Trinidad 1903-2.jpg|thumb|alt=Immigrant workers from India relaxing on a cacao estate in Trinidad, 1903.|Immigrant workers from India relaxing on a cacao estate in [[Trinidad]], 1903{{Sfn|Head|1903|p=ii|ps=&nbsp;(frontispiece)}} ]]

The first 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.{{citation needed|date=January 2018}} [[Hernán Cortés|Cortés]] 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, 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 [[Venezuela]]n and [[History of the Philippines (1521-1898)|Philippine colonies]] (Bloom 1998, Coe 1996).<ref>Alexander Walker (1822). ''[https://books.google.com/books?id=jXBOAQAAIAAJ Colombia, relación geográfica, topográfica, agrícola, comercial y política de este país: adaptada para todo lector en general y para el comerciante y colono en particular]''. Tomo II. Londres: Banco de la República, pp. 284.</ref> A painting by Dutch Golden Age artist [[Albert Eckhout]] shows a wild cacao tree in mid-seventeenth century Dutch Brazil. The Nahuatl-derived Spanish word ''cacao'' entered scientific nomenclature in 1753 after the [[Sweden|Swedish]] naturalist [[Carl Linnaeus|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]]''.

== Gallery ==

<gallery mode="packed" heights="180px">
File:Floral diagram -- theobroma cacao.svg|[[Floral diagram]] showing partial inflorescence
File:Naturalis Biodiversity Center - L.0939563 - Bernecker, A. - Theobroma cacao - Artwork.jpeg|Leaves, fruits and seed. A. Bernecker, 1864.
File:Cacao fruit in Côte d'Ivoire (7).JPG|Young trees, Côte d'Ivoire
Image:Theobroma cacao - flower1.JPG|Macrophotography of flower (closed)
Image:Theobroma cacao - flower2.JPG|Macrophotography of flower (open)
Image:Theobroma cacao flowers at Kunnathurpadi (5).jpg|Flowers
Image:Cacao-pod-k4636-14.jpg|Cacao seed in the fruit or ''pocha''
File:Theobroma cacao MHNT.BOT.2004.0.204.jpg|Fruit, dried
</gallery>

==See also==

* ''[[Ceratonia siliqua]]'', the carob tree
* [[Kola nut]]

==References==

{{Reflist|30em}}

== Sources ==

* {{cite book |last=Head |first=Brandon |title=The Food of the Gods: A Popular Account of Cocoa |url=http://archive.org/stream/foodgodsapopula00headgoog#page/n9/mode/2up |year=1903 |publisher=R. B. Johnson |location=London |page=ii}}

== Further reading ==

* {{cite book |last1=Coe |first1=Sophie D. |last2=Coe |first2=Michael D. |author-link2=Michael D. Coe |year=1996 |title=The True History of Chocolate |location=London |publisher=Thames & Hudson |isbn=0-500-01693-3 |url=https://archive.org/details/truehistoryofcho00coes |ref=none}}
* {{cite web |last=Dienhart |first=John M. |year=1997 |title=The Mayan Languages – A Comparative Vocabulary |url=http://maya.hum.sdu.dk/proto-forms/cacao.pdf |publisher=[[Odense University]] |access-date=14 February 2007 |archive-date=5 July 2007 |archive-url=https://web.archive.org/web/20070705170059/http://maya.hum.sdu.dk/proto-forms/cacao.pdf |url-status=dead |ref=none}}
* {{cite book |editor-last=McNeil |editor-first=Cameron |year=2006 |title=Chocolate in Mesoamerica: A Cultural History of Cacao  |location=Gainesville |publisher=University of Florida Press |isbn=0-8130-2953-8 |ref=none}}

==External links==

{{Commons|Theobroma cacao}}
{{Wikispecies|Theobroma cacao}}
* [http://www.icco.org International Cocoa Organization (ICCO)] – includes cacao daily market prices and charts

{{Chocolate}}
{{Taxonbar|from=Q42385}}
{{Authority control}}

[[Category:Theobroma|cacao]]
[[Category:Caffeine|Cacao]]
[[Category:Agriculture in Mesoamerica]]
[[Category:Agriculture in Ecuador]]
[[Category:Chocolate]]
[[Category:Cocoa production]]
[[Category:Components of chocolate]]
[[Category:Crops]]
[[Category:Crops originating from Ecuador]]
[[Category:Crops originating from Peru]]
[[Category:Plants described in 1753]]
[[Category:Botanical taxa named by Carl Linnaeus]]
[[Category:Crops originating from indigenous Americans]]