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''Pentadiplandra brazzeana'' is an evergreen shrub or liana that is the only species assigned to the genus ''Pentadiplandra'', and has been placed in a family of its own called Pentadiplandraceae. It produces large red berries, sometimes mottled with grey. It is known from West-Central Tropical Africa, between northern Angola, eastern Nigeria and western Democratic Republic of Congo. The berry is sweet in taste due to the protein,
brazzein Brazzein is a sweet-tasting protein extracted from the West African fruit of the climbing plant Oubli (''Pentadiplandra brazzeana'' Henri Ernest Baillon, Baillon). It was first isolated by the University of Wisconsin–Madison in 1994. Brazzein ...
, which is substantially sweeter than
saccharose Sucrose, a disaccharide, is a sugar composed of glucose and fructose subunits. It is produced naturally in plants and is the main constituent of white sugar. It has the molecular formula . For human consumption, sucrose is extracted and refine ...
. Brazzein may be useful as a low-calorie
sweetener {{Wiktionary, sweetener A sweetener is a substance added to food or drink to impart the flavor of sweetness, either because it contains a type of sugar, or because it contains a sweet-tasting sugar substitute. Many artificial sweeteners have been ...
, but is not yet allowed as a food additive in the United States and the European Union.


Description

''Pentadiplandra brazzeana'' is a
monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system alongside gynomonoecy, andromonoecy and trimonoecy. Monoecy is conne ...
shrub of maximally , but can also develop into a
liana A liana is a long- stemmed, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the canopy in search of direct sunlight. The word ''liana'' does not refer to a ta ...
, climbing up to high in the trees. The shrub
morph Morph may refer to: Biology * Morph (zoology), a visual or behavioral difference between organisms of distinct populations in a species * Muller's morphs, a classification scheme for genetic mutations * "-morph", a suffix commonly used in tax ...
usually has a mass of branched bulging roots, while the liana morph has a large, fleshy tuber. The branches are without hair and carry alternately set, simple and entire leaves, without
stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...
s at the base of the ½–1 cm (0.2–0.4 in) long
leaf stalk In botany, the petiole () is the stalk that attaches the leaf blade to the stem, and is able to twist the leaf to face the sun. This gives a characteristic foliage arrangement to the plant. Outgrowths appearing on each side of the petiole in so ...
. The hairless leaf blade is
elliptical Elliptical may mean: * having the shape of an ellipse, or more broadly, any oval shape ** in botany, having an elliptic leaf shape ** of aircraft wings, having an elliptical planform * characterised by ellipsis (the omission of words), or by conc ...
to
oblanceolate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
, long and 1½–5 cm (0.6–2 in) wide, with a wedge-shaped base, a pointed tip, a dull or shining dark green upper surface and a dull dark green lower surface, and a central
vein Veins are blood vessels in humans and most other animals that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are the pulmonary and umbilical veins, both of which carry oxygenated b ...
that branches feather-like into five to eleven pairs of side veins. The flowers are in
raceme A raceme ( or ) or racemoid is an unbranched, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are produced as the s ...
s in the axils of the leaves or at the tip of the branches, with the common inflorescence stalk much longer in terminal racemes, up to long. The individual flowers can be functionally only male, only female or
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...
, all on the same plant. They sit on a long stalk, and carry five elliptical to
lanceolate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
, ½–1 cm (0.2-0.4 in) long, green
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coined b ...
s with a violet margin, which are slightly buiging and fused at their base. Inside are five free white to yellowish
petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
s of between 2 and 2½ cm (0.8–1.0 in) long, set between the neighboring sepals. They consist of a separate, wide, pouch-like base or alternatively described as having a conspicuous scale, fringed by hairs that form a "roof" over a chamber and also make the base of the sepals cling together. The petal further consists of a lanceolate to oblanceolate plate, with irregular burgundy-colored splashing, and a pointy tip. Stamens and pistol are separated from the sepals and petals by a firm stalk (or androgynophore), which carries at its top ten to thirteen
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
s with about long thin filaments connected only at the base forming two or three worls. The
anther The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
s on top are attached at the base, have two cavities that release pollen by lengthwise slits. The ovary, which may be clearly distanced from the base of the stamens by a
gynophore A gynophore is the stalk of certain flowers which supports the gynoecium (the ovule-producing part of a flower), elevating it above the branching points of other floral parts. Plant genera that have flowers with gynophores include '' Telopea'', ''P ...
, has (four or) five cavities and carries a short
style Style is a manner of doing or presenting things and may refer to: * Architectural style, the features that make a building or structure historically identifiable * Design, the process of creating something * Fashion, a prevailing mode of clothing ...
topped by a (4– or) 5-lobed stigma. In male flowers the
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
is rudimentary, and in female flowers the stamens are rudimentary. The fruit is a globe-shaped
berry A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit, although many pips or seeds may be present. Common examples are strawberries, raspb ...
of 3½–5 cm (1.4–2 in) in diameter, entirely red or mottled with grey, containing many seeds attached to the axis surrounded by pink pulp. The seeds are kidney-shaped, and have an outer layer of wooly, white, one-celled hairs.


Phytochemistry

''Pentadiplandra'' contains
thiocarbamate In organic chemistry, thiocarbamates (thiourethanes) are a family of organosulfur compounds. As the prefix ''thio-'' suggests, they are sulfur analogues of carbamates. There are two isomeric forms of thiocarbamates: ''O''-thiocarbamates, (ester ...
s such as methyl N-benzylthiocarbamate, methyl and ethyl N-methoxybenzylthiocarbamate, and
glucosinolate Glucosinolates are natural components of many pungent plants such as mustard, cabbage, and horseradish. The pungency of those plants is due to mustard oils produced from glucosinolates when the plant material is chewed, cut, or otherwise damaged. T ...
s such as benzyl- and 4-methoxybenzyl glucosinolates. It has cells containing
myrosinase Myrosinase (, ''thioglucoside glucohydrolase'', ''sinigrinase'', and ''sinigrase'') is a family of enzymes involved in plant defense against herbivores, specifically the mustard oil bomb. The three-dimensional structure has been elucidated and is ...
.


Taxonomy

''Pentadiplandra brazzeana'' was first described by French botanist and physician
Henri Ernest Baillon Henri Ernest Baillon was a French botanist and physician. He was born in Calais on 30 November 1827 and died in Paris on 19 July 1895. Baillon spent his professional life as a professor of natural history, and he published numerous works on bo ...
in 1886, who assigned it to the family Capparaceae, based on a specimen from
Osika Osika is a Slavic surname. Notable people with the surname include: * Craig Osika (born 1979), American football player * Ron Osika (born 1939), Canadian politician See also * Osaka (surname) Osaka or Ōsaka is a Japanese surname that may refer t ...
in Congo by Jacques de Brazza. In 1897,
Ernest Friedrich Gilg Ernest (or Ernst) Friedrich Gilg (12 January 1867 in Baden-Württemberg, Germany – 11 October 1933 in Berlin) was a German botanist. Life Gilg was curator of the Botanical Museum in Berlin. With fellow botanist Adolf Engler, he co-authored ...
described ''Cercopetalum dasyanthum'' in the Capparaceae. Otto Stapf described ''Cotylonychia chevalieri'' in 1908 as part of the
Sterculiaceae Sterculiaceae was a family of flowering plant based on the genus ''Sterculia''. Genera formerly included in Sterculiaceae are now placed in the family Malvaceae, in the subfamilies: Byttnerioideae, Dombeyoideae, Helicteroideae and Sterculioideae. A ...
. In 2000,
Clemens Bayer Clemens is both a Late Latin masculine given name and a surname meaning "merciful". Notable people with the name include: Surname * Adelaide Clemens (born 1989), Australian actress. * Andrew Clemens (b. 1852 or 1857–1894), American folk artist * ...
showed ''Cotylonychia'' to be
synonymous A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
to ''Pentadiplandra''.
Arthur Wallis Exell Arthur Wallis Exell OBE (21 May 1901, in Birmingham – 15 January 1993, in Cheltenham) was initially an assistant and later Deputy Keeper of Botany at the British Museum during the years 1924–1939 and 1950–1962. A noted cryptographer, taxo ...
introduced the name ''Pentadiplandra gossweileri'' in 1927. The family Pentadiplandraceae was proposed by John Hutchinson and British botanist, physician and scientific explorer
John McEwan Dalziel John is a common English name and surname: * John (given name) * John (surname) John may also refer to: New Testament Works * Gospel of John, a title often shortened to John * First Epistle of John, often shortened to 1 John * Second ...
in the ''Flora of West Tropical Africa'' in 1928.


Phylogeny

''P. brazzeana'' is the only known species in the genus ''Pentadiplandra'', and has been assigned its own family named Pentadiplandraceae. Analyses of the development of the flower and anatomic features suggest that ''Pentadiplandra'' is a
relict A relict is a surviving remnant of a natural phenomenon. Biology A relict (or relic) is an organism that at an earlier time was abundant in a large area but now occurs at only one or a few small areas. Geology and geomorphology In geology, a r ...
genus branching off near the base of the core
Brassicales The Brassicales (or Cruciales) are an order (biology), order of flowering plants, belonging to the eurosids II group of dicotyledons under the APG II system. One character common to many members of the order is the production of glucosinolate (mu ...
. It has many characters in common with the American genus ''
Tovaria ''Tovaria'' is a genus of herbs native to Jamaica and South America. There are two species, ''Tovaria pendula'' and ''Tovaria diffusa''. The genus is the only one in the family Tovariaceae. References

* http://www.mobot.org/MOBOT/Research/A ...
''. Current insights in the relationships of the Brassicaceae, based on a 2012 DNA-analysis, are summarized in the following tree.


Etymology

''Pentadiplandra'' is the contraction of the Greek words πέντε (pente), meaning "five", διπλόος (diploos), "double", and ἀνδρὸς (andros), "male" or "stamen", a reference to the usually two worls of five stamens each. ''brazzeana'' is probably derived from the name of the collector of the type specimen, J. de Brazza


Distribution and habitat

The plant grows in
Angola , national_anthem = " Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordina ...
, the
Democratic Republic of the Congo The Democratic Republic of the Congo (french: République démocratique du Congo (RDC), colloquially "La RDC" ), informally Congo-Kinshasa, DR Congo, the DRC, the DROC, or the Congo, and formerly and also colloquially Zaire, is a country in ...
, the
Central African Republic The Central African Republic (CAR; ; , RCA; , or , ) is a landlocked country in Central Africa. It is bordered by Chad to the north, Sudan to the northeast, South Sudan to the southeast, the DR Congo to the south, the Republic of th ...
, the
Republic of the Congo The Republic of the Congo (french: République du Congo, ln, Republíki ya Kongó), also known as Congo-Brazzaville, the Congo Republic or simply either Congo or the Congo, is a country located in the western coast of Central Africa to the w ...
,
Cameroon Cameroon (; french: Cameroun, ff, Kamerun), officially the Republic of Cameroon (french: République du Cameroun, links=no), is a country in west-central Africa. It is bordered by Nigeria to the west and north; Chad to the northeast; the C ...
,
Gabon Gabon (; ; snq, Ngabu), officially the Gabonese Republic (french: République gabonaise), is a country on the west coast of Central Africa. Located on the equator, it is bordered by Equatorial Guinea to the northwest, Cameroon to the north ...
and
Nigeria Nigeria ( ), , ig, Naìjíríyà, yo, Nàìjíríà, pcm, Naijá , ff, Naajeeriya, kcg, Naijeriya officially the Federal Republic of Nigeria, is a country in West Africa. It is situated between the Sahel to the north and the Gulf o ...
. It is not uncommon in upland primary forest dominated by '' Scorodophloeus zenkeri'', but also occurs regularly on the banks of rivers and in secondary forest. The species is found in particular in forest margins bordering savanna's in Cameroon. It does not appear in clusters anywhere.


Sweetness and calories

Most primates have a
genotype The genotype of an organism is its complete set of genetic material. Genotype can also be used to refer to the alleles or variants an individual carries in a particular gene or genetic location. The number of alleles an individual can have in a ...
of the
taste receptor A taste receptor or tastant is a type of cellular receptor which facilitates the sensation of taste. When food or other substances enter the mouth, molecules interact with saliva and are bound to taste receptors in the oral cavity and other locat ...
protein, taste receptor type 1 member 3 (TAS1R3), that enables them to taste the protein,
brazzein Brazzein is a sweet-tasting protein extracted from the West African fruit of the climbing plant Oubli (''Pentadiplandra brazzeana'' Henri Ernest Baillon, Baillon). It was first isolated by the University of Wisconsin–Madison in 1994. Brazzein ...
. To humans, the fruit is intensely sweet, but provides few
calories The calorie is a unit of energy. For historical reasons, two main definitions of "calorie" are in wide use. The large calorie, food calorie, or kilogram calorie was originally defined as the amount of heat needed to raise the temperature of on ...
. Such proteins may imitate sweetness to lure wild animals to eat the berries and disperse the seeds.
Western lowland gorilla The western lowland gorilla (''Gorilla gorilla gorilla'') is one of two Critically Endangered subspecies of the western gorilla (''Gorilla gorilla'') that lives in Montane ecosystems#Montane forests, montane, Old-growth forest, primary and sec ...
s (''Gorilla gorilla''), however, have two
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mi ...
s in the TAS1R3
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
, and although its diet contains a high proportion of fruit, scientists have not witnessed gorillas consuming ''P. brazzeana'' berries. If factual, this avoidance behavior and the taste gene mutations may indicate a counter-adaptation to deter gorillas from foraging for low-calorie foods.


Uses


Traditional uses

The berries, leaves, roots, and tubers of these plants have been used in local traditional culture. Roots hung in the house are thought to repel snakes. Powdered root bark is an ingredient of "African whiskey in sachets", which is said to be cheap but dangerous. The root is said to be eaten occasionally as a vegetable. A syrup made from the root is marketed throughout the
Congo Basin The Congo Basin (french: Bassin du Congo) is the sedimentary basin of the Congo River. The Congo Basin is located in Central Africa, in a region known as west equatorial Africa. The Congo Basin region is sometimes known simply as the Congo. It con ...
. The pulp of the berries is eaten by children, and is used as a sweetener in maize porridge.


Modern uses

Brazzein has been isolated from the plant and a company was formed to bring it to market in 2008, which initially said it would start selling the product by 2010 once it obtained agreement from the FDA that its brazzein was generally recognized as safe (GRAS). In 2012 the company said that regulatory approval might take an additional one or two years and in 2014 it still had not obtained a GRAS waiver from the FDA and was seeking partners, and the product was still not on the market as of 2016.


References

{{Taxonbar, from=Q134544 Monotypic Brassicales genera Brassicales Flora of West-Central Tropical Africa Taxa named by Henri Ernest Baillon