Cacao God Sculpture
Habitat & Distribution
Life History and Behavior
Although the full story of cacao pollination is not yet known, there seems little doubt that the flower is not self-pollinating, as flowers bagged to exclude insects invariably shed (Gnanaratnam 1954). Also, some plants are self-incompatible but set fruit well if pollinated with pollen from compatible trees (Chats 1953, Cope 1958, Knight and Rogers 1955). The method of the transfer of the pollen in nature is the somewhat questionable factor. The sticky pollen is not carried by the wind. Furthermore, it is produced and released in the petal pouches where wind is unlikely to disturb it (Cobley 1966*, Gnanaratnam 1954). Glendenning (1962) noted that pollen found on a stigma was usually from more than one flower, but the amount of foreign pollen depended on proximity to other plants. Little pollen seemed to move more than a couple of trees' distance.
There is general belief that small insects are the primary pollinating agents of cacao, but no general agreement as to which insects are responsible. Numerous authorities credit midges, especially Forcipomyia quasiingrami Macfie and Lasiohela nana Macfie (Barroga 1964, Chatt 1953, Fontanilla-Barroga 1965, Macfie 1944, Saunders 1959, Toxopeus 1969). Others credit ants (Crematogaster spp.), aphids (Aphis gossypii Glover and Toxoptera spp.), thrips (Frankliniella parvula Hood), and unidentified wild bees (Billes 1941; Cope 1940; Harland 1925a, b; Hernandez 1966; Jones 1912; Muntzing 1947; Posnette 1942a, b, 1944, 1950; Posnette and Entwistle 1957; Urquhart 1961; Voelcker 1940).
Thrips and aphids move about but slightly from tree to tree, yet Glendenning (1958) reported, after a study of albino trees, that a considerable proportion of pollination takes place across two intervening trees, though less than over shorter distances. This would indicate an agent with considerable movement between trees.
The ants Wasmannia suropunctata (Roger) and Solenopsis geminata (F.) and the wild bee Trigona jaty Smith were occasional visitors. Glendenning (1958) concluded that the midges (Forcipomyia spp.) were the main pollinators, accounting for twice the pollination service performed by all of the other species combined. This was verified in various experiments with different numbers of insects per cage over cacao flowers. Hernandez (1965) reported pollination percentages ranging from 1 to 52 percent when he used midges, bees, thrips, and ants. However, Hernandez did not, indicate how pollination was accomplished.
Although midges seem to get the most credit as pollinators of cacao, there is clearly a lack of knowledge as to which insects are responsible in the different areas for the commercial set of fruit of this important crop.
Harland (1925a) found that of 5 percent of the flowers on trees not infested by ants and aphids, only 0.3 percent set fruit; whereas, on trees heavily infested by these insects, 35 percent of the flowers were pollinated and 2 percent set. At the same time, 5 percent of the hand pollinated flowers set fruit.
Little has been said about the adequacy of pollination of the individual flower or the minimum number of seed in relation to fruit set or shedding. However, at least as many pollen grains must fall upon the stigma as there are subsequently developed seeds. Thus, a minimum of 60 pollen grains is necessary to set the highest number of seed.
Many of the flowers are never pollinated (Harland 1925b), at least under Trinidad conditions. Apparently, wherever the crop is grown the lack of adequate pollination is a strongly limiting factor in production of the beans. Sumner (1962) stated that most of the pollination occurs 2 to 3 hours after dawn with a second much smaller peak in the afternoon, but only 2 to 5 percent of the flowers ever get pollinated, and these may not set if pollinated too late or with incompatible pollen. Urquhart (1961) stated that only about 5 percent of the stigmas ever get pollinated; Harland (1925b) found only 9 percent to be pollinated. Because some plants are self-incompatible - some are male sterile or sterile (Gnanaratnam 1954) - many of the flowers would appear to be doomed to shed. Knoke and Saunders (1966) tried a mist blower for mechanical transfer of pollen but achieved uneconomical success.
The use of honey bees under saturated pollination conditions has never been tried, probably because the blossom has no aroma and produces no nectar. Quite conceivably, however, honey bee colonies could be concentrated in numbers sufficient to exhaust the supply of pollen and nectar on competing plants and the bees induced to visit the flowers of this important crop for pollen and increase the percentage of cross- pollination and fruit set. A search for a selection of cacao pollen-loving honey bees might produce an acceptable and controllable pollinating agent. One or more of the various species of pollen-foraging wild bees might be found that could be controlled and used as a profitable pollinating agent of cacao.
Pollination Recommendations and Practices:
There are no recommendations on the use or manipulation of insect pollinators of cacao. According to Faegri and van der Pijl (1966*), the Forcipomyia spp in Africa breed mainly in decaying pods. If the pods are removed by too scrupulous cleaning of the plantations, these midges might also be removed. This would result in deficient pollination of the flowers. Otherwise, the presence or numbers of insect pollinators are left entirely to chance on this billion-dollar crop.
- McGregor, S.E. 1976. Insect Pollination Of Cultivated Crop Plants. USDA (online publication)
- BARROGA S. F. 1964. PROGRESS REPORT ON THE STUDY OF INSECTS, PARTICULARLY MIDGES ASSOCIATED WITH POLLINATION OF Theobroma cacao, APRIL 1963. Philippine Jour. Plant Indus. 29(3/4): 123 - 133.
- BILLES, D. J. 1941. POLLINATION OF Theobroma cacao L. IN TRINIDAD, B.W.L Trop. Agr. [Trinidad] 18: 151-156.
- CHATT, E. M.1953. COCOA. 302 pp. Interscience Publishers Inc., New York.
- CHEESEMAN, E. E. 1932. THE ECONOMIC BOTANY OF CACAO. A CRITICAL SURVEY OF THE LITERATURE TO THE END OF 1930. Trop. Agr. [Trinidad] Sup., v. 9, June, 16 pp.
- COPE, F. W. 1940. AGENTS OF POLLINATION IN CACAO. St. Augustine, Trinidad, Imperial College of Tropical Agr. [Trinidad], Ninth Ann. Rpt. on Cacao Res. 1939: 13-19.
- ______ 1958. INCOMPATIBILITY IN Theobroma cacao. Nature 181: 279.
- FONTANILLA-BARROGA, S. 1965. A PROGRESS REPORT ON THE STUDY OF INSECTS ASSOCIATED WITH POLLINATION OF Theobroma cacao WITH SPECIAL EMPHASIS ON MIDGES. Philippine Jour. Agr. 27(3/4): 147-159.
- GLENDINNING, D. R. 1958. PLANT BREEDING AND SELECTION. Cocoa Res. Inst. Rpt. of West Africa, 1957-58, pp. 50-54.
- ______ 1962. NATURAL POLLINATION OF COCOA. Nature 193(4822): 1305.
- GNANARATNAM, J. K. 1954. POLLINATION MECHANISM OF THE CACAO FLOWER. Trop. Agr. [Ceylon] 110: 98 - 104.
- HALL, C. J. J. VAN. 1932. CACAO Ed. 2, 514 pp. Macmillan, London.
- HARLAND, S. C. 1925a. STUDIES IN CACAO. THE METHOD OF POLLINATION. Ninth West Indian Agr. Conf. Proc. Kingston, Jamaica, 1924: 61 - 69.
- ______ 1925b. STUDIES IN CACAO. PART I. THE METHOD OF POLLINATION. Ann. Appl. Biol. 12: 403-409.
- HERNANDEZ, B. J. 1965. INSECT POLLINATION OF Theobroma cacao L.) IN COSTA RICA. 173 pp. Ph.D. thesis and Diss. Abs. 28(1): 2B-3B, 1967, AA-257/71, Wis, Univ., Madison.
- JONES, G. A. 1912. THE STRUCTURE AND POLLINATION OF THE CACAO FLOWER. West Indian Bull 12: 347 - 350.
- KNIGHT, R., and ROGERS, H. H. 1955. INCOMPATIBILITY IN Theobroma cacao. Heredity 9: 69 - 77.
- KNOKE J. K., and SAUNDERS, J. L. 1966. INDUCED FRUIT SET OF Theobroma cacao BY MISTBLOWER APPLICATIONS OF INSECTICIDES. Jour. Econ. Ent. 59: 1427-1430.
- MACFIE, J. W. S. 1944. Ceratopogonidae COLLECTED IN TRINIDAD FROM CACAO FLOWERS. Bul.:Ent. Res. [England] 35: 297 - 300.
- MUNTZING, A. 1947. SOME OBSERVATIONS ON POLLINATION AND FRUIT-SETTING IN ECUADORIAN CACAO. Hereditas 33: 397 - 404.
- POSNETTE, A. F. 1924a. NATURAL POLLINATION OF COCOA, Theobroma leiocarpa, ON THE GOLD COAST. Trop. Agr. [Trinidad] 19: 12-16.
- ______ 1942b. NATURAL POLLINATION OF COCOA. Theobroma leiocarpa, BERN., ON THE GOLD COAST II. Trop. Agr. [Trinidad] 19(10): 188-191.
- ______ 1944. POLLINATION OF CACAO IN TRINIDAD. Trop. Agr. [Trinidad] 21(6): 115-118.
- ______ 1950. THE POLLINATION OF CACAO IN THE GOLD COAST. Jour. Hort. Sci. 25: 155 - 168.
- ______and ENTWISTLE, H. M. 1957. THE POLLINATION OF COCOA FLOWERS. Rpt. Cocoa Conf. Grosvenor House, London, Sept. 10-12, pp. 66-69. (Abs.) Plant Breeding 28(4): 4550. Oct. 1958.
- SAUNDERS, L. G. 1959. METHODS FOR STUDYING Forcipomyia MIDGES, WITH SPECIAL REFERENCE TO CACAO-POLLINATING SPECIES (Diptera, Ceratopogonidae). Canad. Jour. Zool. 37: 33-51.
- STEJSKAL, M. 1969. [NECTAR AND AROMA OF THE CACAO FLOWER.] Oriente Agropecuario 1(2): 75-92. [In Spanish, English summary.]
- SUMNER, H. M. 1962. [COCOA] POLLINATION. In Wills, J. B., ed., Agriculture and Land Use in Ghana, pp. 260 - 261. Oxford University Press, London, Accra, New York.
- TOXOPEUS, H.1969. CACAO. In Ferwerda, F. P., and Wit, F., eds., Outlines of Perennial Crop Breeding in the Tropics, pp. 79-109. H. Veenman and Zonen, N. V. Wageningen, The Netherlands.
- URQUHART. D. H. 1961. COCOA. Ed. 2, 293 pp. Longmans, Green & Co., Ltd., London.
- VOELCKER, O. J. 1940. THE DEGREE OF CROSS-POLLINATION IN CACAO IN NIGERIA. Trop. Agr. [Trinidad] 17: 184-186.
Molecular Biology and Genetics
Cocoa butter contains glycerides, and its sterols include ergosterol, sitosterol and stigmasterol. Seed alkaloids include caffeine and the diuretic theobromine. Seed phenolic compounds include coumarin, esculetine and catechol. Leaf contains genistic acid, an antirheumatic and analgesic.
Statistics of barcoding coverage: Theobroma cacao
Public Records: 0
Specimens with Barcodes: 1
Species With Barcodes: 1
- Phylota TaxBrowser: Sequence Diversity and Cluster Set Summaries
- Kuhn, D.N., Narasimhan, G., Nakamura, K., Brown, J.S., Schnell II, R.J., Meerow, A.W. 2006. Identification of cacao tir-nbs-lrr resistance gene analogs and their use as genetic markers. Journal of the American Society for Horticultural Science. 131(6):806-813.
- Schnell II, R.J. 2004. Can modern plant science drive the agro-ecology of an ancient crop? Proceedings of the National Academy of Sciences.
- USDA Research Project: Analysis of Genetic Diversity and Biochemistry of Cacao
How to Grow
Frost Tolerance: Needs uniformly high temperatures, green house only in Phoenix, for short duration cold, foliage damaged at 33° F (0.5° C), serious damage at 28° F (-2° C)
Sun Exposure: Light shade
Origin: Central America, Mexico
Growth Habits: Tropical evergreen tree to 25 feet (7.5 m)
Watering Needs: Abundant water and humidity
Propagation: Cuttings, budding, grafting, seeds
Propagation and Management
Cocoa seeds readily germinate when sown and do not pass through a dormancy period. They lose viability within 5-7 days of extraction from the pod unless specially treated, and germinate within 7-10 days. The plant can easily be propagated vegetatively by leaf-bud cutting, multiple-bud cutting, marcotting, budding, grafting and layering.
Weeding and temporary shade are essential within the 1st 3-4 years of establishment before the canopy closes. Plantain appears to meet most of cocoa’s requirements in this respect, whereas bananas compete heavily for moisture during the dry season. The young trees should be mulched before the onset of the 1st dry season to conserve soil moisture. Light pruning is recommended to remove low-hanging, broken and dead branches, as well as for the regeneration of fallen or damaged trees. Farmers plant cocoa at high densities of 3000-4000 trees/ha because the resulting tall trees develop fewer lateral branches and more vertical suckers. This encourages flowering on the main stem at the expense of branches, particularly suitable for some lower Amazon Forastero varieties.
Seed storage behaviour is recalcitrant. Storage temperature between 4 and 15 deg. C is damaging to seed viability and germination. Optimum storage temperature appears to be 17 deg. C. Seeds tolerate desiccation to 25% mc when dried at 20 deg. C, while only about 40-60% survive when dried at 10 deg. C; seeds stored in pods at 5 or 10 deg. C are killed within 2 days, and there is 100% survival when stored in pods at temperatures of 15-30 deg. C for 3 weeks. Viability is reduced from 92% to 18% on desiccation from 45% to 36.7% mc; no seeds survive desiccation to 26% mc; 24% germination after 8 months subimbibed storage (41-42% mc) at 98% rh and 20 deg. C with Thiram fungicide. Similarly, no seeds survive desiccation to below 20% mc, and no fresh seeds survive in storage at 4 deg. C or 15 deg. C.
History of Cultivation
Movies & Media
- “Like Water for Chocolate” (1992)
- “Chacolat” (2000)
- “Diseases Imperil Chocolate Production”. Randy Ploetz. Talk of the Nation. NPR, 9 June 2006.
- Buford, Bill. “Notes of a Gastronome: Extreme Chocolate.” The New Yorker. (29 Oct 2007) p 68.
- “Chocolate lorry goes to Timbuktu”. BBC News, (23 Nov 2007)
- Dahl, Roald. 1964. Charlie and the Chocolate Factory, Knopf.
- “Montezuma’s Request” Raymond Sokolov, From the column “A Matter of Taste” Natural History Magazine.
- Reid, TR. “Caffeine.” National Geographic.
- Sacha, Bob. “Mouth-Watering Art.” National Geographic.
- The Worldwide Gourmet. "Cocoa — The Saga of Chocolate".
- Young, Gordon. “Chocolate: Food for the Gods.” National Geographic, (Nov 1984) 664-87.
Cacao is cultivated on over 70,000 km² (27,000 mi²) worldwide. Hershey's produces 40% of world cacao, Nestle and Mars each produce about 15%.
A tree begins to bear when it is four or five years old. In one year, when mature, it may have 6,000 flowers, but only about 20 pods. About 300-600 seeds (10 pods) are required to produce around 1 kg (2.2 lb) of cocoa paste.
There are three main cultivar groups of cacao beans used to make cocoa and chocolate. 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. For details of processing, see cocoa.
All about Chocolate -- Varieties
History of Cultivation
Cultivation, cultural elaboration and use of cacao were extensive and early in Mesoamerica. Studies of the Theobroma cacao tree genetics suggests 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 spaerocarpum has a fairly smooth melon-like fruit. In contrast, the Mesoamerican cacao subspecies has ridged, elongated fruits. At some unknown early date, the subspecies T. cacao cacao reached the southern lowlands of Mesoamerica and came into wide usage.
The Maya believed that the kakaw (cacao) was discovered by the gods in a mountain that also contained other delectable foods to be used by the Maya. According to Maya mythology, the Plumed Serpent gave cacao to the Maya after humans were created from maize by divine grandmother goddess Xmucane (Bogin 1997, Coe 1996, Montejo 1999, Tedlock 1985). 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 (Coe 1996, Townsend 1992). 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 were used only by men, as it was believed to be toxic for women and children.
There are several mixtures of cacao described in ancient texts, for ceremonial, medicinal uses as well as culinary purposes. Some mixtures included maize, chili, vanilla (Vanilla planifolia), peanut butter and honey. Archaeological evidence for use of cacao, while relatively sparse, has come from the recovery of whole cacao beans at Uaxactun, Guatemala (Kidder 1947) and from the preservation of wood fragments of the cacao tree at Belize sites including Cuello and Pulltrouser Swamp (Hammond and Miksicek 1981; Turner and Miksicek 1984). 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 B.C.) and in middle formative vessels from Colha, Belize (600-400 B.C.) using similar techniques to those used to extract chocolate residues from four classic period (ca. 400 A.D.) 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 that 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.
The first Europeans to encounter cacao were Christopher Columbus and his crew in 1502, 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 Montezuma in the Aztec capital of Tenochtitlan in 1519. Cortez and others noted the vast quantities of this beverage that 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 that 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 (Coe and Coe 1996). 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 ("food of the gods") cacao.
Relevance to Humans and Ecosystems
Seed: Used as an emollient by the French Guiana Palikur in a remedy to extract splinters or prickles embedded in the skin, when mixed with stems of Chromolaena odorata and wood of Cecropia obtusa. Leaf-bud: Infusion used with incense to treat diarrhoea in French Guiana. Fruit: Infusion of dry pods used to decrease leprosy spots. Seed: Infusion of baked seed-membranes is drunk for remedying anemia.
There are at least 63 plant species known to produce caffeine(1). The most important harvested plants are coffee, tea, cacao, maté, kola (the original caffeine ingredient in carbonated drinks) and guarana (common in health supplements). In tea and maté, the leaves are harvested; for coffee, cacao, kola and guarana, the fruit and seeds. The highest concentrations of caffeine tend to occur in tea and guarana, but all species can vary widely in caffeine content, depending on variety, climate, and cultivation (Graham, 2009), and whether the plant is male or female. Some reported concentrations:
Plant_______ Scientific Name_____ Caffeine content___Source
Tea________Camellia sinensis____ 2.7-4.1%__________Kaplan et al, 1974 Guarana____Paullinia cupana______1.6-4.3%__________Baumann et al, 1995
Kola_______ Cola acuminate ______1-2.2%___________ Somorin, 1973
& Cola nitida
Coffee______Coffea arabica_______ 0.8-1.8%__________Kaplan et al, 1974
Maté_______ Ilex paraguayensis ___ 0.8-1.7%__________Dellacassa et al, 2007 Cacao______Theobroma cacao____ 0.07-1.7%_________Asamoa and Wurziger, 1976
On top of this natural variation, preparation of a serving can greatly vary the strength of the caffeine dose. A prepared cup of regular coffee (not counting decaf) can vary five-fold in caffeine concentration among brands and methods (McCusker et al, 2003). The same study found, after purchasing the same beverage from the same coffee shop for six days in a row, that the dose among those six cups varied up to a factor of two, from 259-564mg per cup.
- Grassi, D., Necozione, S., Lippi, C., Croce, G., Valeri, L., Pasqualetti, P., Desideri, G., Blumberg, J.B., Ferri, C. 2005. Cocoa reduces blood pressure and insulin resistance and improves endothelium-dependent vasodilation in hypertensives. Hypertension. 46(2):398-405.
- Engler, M.B., Engler, M.M., Chen, C.Y., Malloy, M.J., Bowne, A., Chiu, E.Y., Kwak, H., Milbury, P., Blumberg, J., Mietus-Snyder, M.L. 2004. Flavanoid-rich dark chocolate improves endothelial function by increasing plasma epicatechin concentration in health adults. Journal of American College of Nutrition. 23(3):197-204.
- Kelishadi, Roya. 2005. Cacao to Cocoa to Chocolate: Healthy Food? ARYA Journal. 1(1):29-35.
- Tarkan, Laurie. “Chocolate a Health Food? Maybe, but Keep the Aspirin.” New York Times. 30 Oct 2000.
- Raloff, Janet. 2000. Chocolate Hearts: Yummy and good medicine? Science News. 157(12): 188.
- The Science of Chocolate
- McNeil, Donald. “This is Your Brain on Chocolate.” New York Times. 23 Aug 2005.
- Death By Chocolate (Chocolate Toxicity & Pets)
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 8000 beans. 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.
In some areas, such as Yucatán, cacao beans were still used in place of small coins as late as the 1840s.
- Bergmann, John (1969). "The Distribution of Cacao Cultivation in Pre-Columbian America". Annals of the Association of American Geographers 59: 85-96.
- Coe, Sophie D. (1994). America's First Cuisines. Austin: University of Texas Press. ISBN 0-292-71155-7.
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 regions of Central and South America. Its seeds, cocoa beans, are used to make cocoa mass, cocoa powder, and chocolate.
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; this is known as cauliflory. The flowers are small, 1–2 cm (0.39–0.79 in) diameter, with pink calyx. The floral formula is ✶ K5 C5 A(5°+5²) G(5). 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. 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 refreshing juice, smoothies, jelly, and nata. The fermented pulp, until recently discarded in Ecuador, Dominican Republic and Peru, is now being distilled there into a popular alcoholic beverage sold in the United States. 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
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 ] in Nahuatl.
Cupuaçu, Theobroma grandiflorum, is a closely related species found in Colombia, Peru, Bolivia and Brazil. Like cacao, it is also the source for a kind of chocolate known as cupulate or cupuaçu chocolate. Cupuaçu is considered as having 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. 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. 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. Climate models indicate that at the peak of the 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 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
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. On the Pacific coast of Chiapas, Mexico, a Mokaya archaeological site provides evidence of cacao beverages dating even earlier, to 1900 BC. The initial domestication was probably related to the making of a fermented, thus alcoholic beverage.
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 and from the preservation of wood fragments of the cacao tree at Belize sites including Cuello and Pulltrouser Swamp. 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 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 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.
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. 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.
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. 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. 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.
The first Europeans to encounter cacao were Christopher Columbus and his crew in 1502, 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. 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 Venezuelan and Philippine colonies (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.
Cacao is cultivated on roughly 17,000,000 acres (27,000 sq mi; 69,000 km2) worldwide. According to the Food and Agriculture Organization (FAO), the top 20 cacao-producing countries in 2005 were as follows:
|1 Côte d'Ivoire (Ivory Coast)||1,024,339||1,330,000|
|10 Papua New Guinea||32,733||42,500|
|12 Dominican Republic||24,646||32,000|
|15 Sierra Leone||8,472||11,000|
|19 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.
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 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. 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 criollo cacao beans from Chuao in Aragua, Venezuela are widely regarded as some of the finest in the world. In November 2000, the cacao beans coming from said region were awarded an appellation of origin under the title "Cacao de Chuao" (from Spanish-cacao of Chuao) effectively making this one of the most expensive and sought after types of cacao.
A new, genetically based classification of 10 groups may well help breeders to create new varieties that are both pest- and disease-resistant and contain valued flavours.
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.
Various plant pests and diseases can cause serious problems for cacao production.
- Phytophthora spp. (black pod) especially Phytophthora megakarya in West Africa
- Rats and other vertebrate pests (squirrels, woodpeckers, etc.)
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. 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. 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 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.
Map showing genetic clusters of Theobroma cacao
|NCBI genome ID|
|Genome size||345.99 Mb|
|Number of chromosomes||10 pairs|
|Year of completion||2010|
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). In a second, unrelated project, the International Cocoa Genome Sequencing Consortium-ICGS, co-ordinated by CIRAD, first published 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, 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.
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