Theobroma cacao, also called the cacao tree and the 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, confectionery, ganache and chocolate.
Floral diagram showing partial inflorescence
2 Taxonomy and nomenclature
3 Distribution and domestication
4 History of cultivation
6 Modern history
12 See also
14 Further reading
15 External links
Leaves are alternate, entire, unlobed, 10–40 cm
(3.9–15.7 in) long and 5–20 cm (2.0–7.9 in)
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
Forcipomyia midges in the subfamily Forcipomyiinae.
Having the natural pollinator
Forcipomyia midges for
was shown to have more fruit production than using artificial
pollinators. 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. Usually
discarded until practices changed in the 21st century, the fermented
pulp may be distilled into an alcoholic beverage sold in the United
States. 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.
Taxonomy and nomenclature
Theobroma cacao) belongs to the genus
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 θεός (theos), meaning "god", and βρῶμα (broma), meaning
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 in Nahuatl.
Theobroma grandiflorum, is a closely related species found
in Brazil, Colombia,
Peru and Bolivia. Like cacao, it is also the
source for a kind of chocolate known as cupulate or cupuaçu
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 Jungle 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. One study sampled 1241 trees and
classified them into 10 distinct genetic clusters. This study also
identified areas, for example around
Iquitos in modern
Ecuador, where representatives of several genetic clusters originated
more than 5000 years ago, leading to development of the variety,
Nacional cocoa bean. 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 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
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.
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
La Chonita Hacienda in Tabasco,
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,
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
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
Belize (600–400 BC) using similar techniques to
those used to extract chocolate residues from four classic period
(around 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
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. 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 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.[citation
needed] Cortés and others noted the vast quantities of this beverage
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,
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). 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 Swedish naturalist
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.
See also: Cocoa bean
Cacao seed in the fruit or pocha
In 2016, cocoa beans were cultivated on roughly 10,196,725 hectares
(25,196,660 acres) worldwide. Cocoa beans are grown by large
agroindustrial plantations and small producers, the bulk of production
coming from millions of farmers with small plots. 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: Forastero,
Criollo and Trinitario. 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 arguably less bitter and more
aromatic than any other bean. In November 2000, the cacao beans coming
Chuao were awarded an appellation of origin under the title
"Cacao de Chuao" (from Spanish-cacao of Chuao).
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.
Major cocoa bean processors include Hershey's,
Nestlé and Mars, all
of which purchase cocoa beans via various sources.
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. Roasting T. cacao by using superheated steam was found
to be better than conventional roasting (use of ovens) because it
resulted in same quality of cocoa beans in a shorter amount of
Cocoa bean production – 2016
FAOSTAT of the United Nations
In 2016, world production was 4.5 million tonnes, led by Ivory Coast
with 33% of global production and
Ghana with 19% of the total
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
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 above, 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.
A specific type of bacteria 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
See also: List of cacao diseases
Various plant pests and diseases can cause serious problems for cacao
Cocoa mirids or capsids worldwide (but especially Sahlbergella
singularis and Distantiella theobroma in
West Africa and Helopeltis
spp. in Southeast Asia)
Conopomorpha cramerella (cocoa pod borer – in Southeast Asia)
Moniliophthora roreri (frosty pod rot)
Moniliophthora perniciosa (witches' broom)
Ceratocystis cacaofunesta (mal de machete) or (Ceratocystis wilt)
Oncobasidium theobromae (vascular streak dieback)
Phytophthora spp. (black pod) especially
Phytophthora megakarya in
Rats and other vertebrate pests (squirrels, woodpeckers, etc.)
Map showing genetic clusters of
NCBI genome ID
Number of chromosomes
Year of completion
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
The genome sequence enables 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. 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
Mars, Incorporated and the University of
California, Berkeley are using
CRISPR to adjust
DNA for improved
hardiness of cacao in hot climates.
Young cacao plantation
Immigrant workers from India relaxing on a cacao estate in Trinidad,
Theobroma cacao flower (closed) (University of
Macrophotography of T. cacao flower (open) (University of Vienna)
T. cacao fruit
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