Kiwi (Actinidia chinensis) 1 Luc Viatour

botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...

, a berry is a fleshy

fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in partic ...

without a stone (pit) produced from a single

flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...

containing one

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. ...

. Berries so defined include

grape A grape is a fruit, botanically a berry, of the deciduous woody vines of the flowering plant genus '' Vitis''. Grapes are a non- climacteric type of fruit, generally occurring in clusters. The cultivation of grapes began perhaps 8,000 years a ...

s, currants, and

tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...

es, as well as

cucumber Cucumber (''Cucumis sativus'') is a widely-cultivated creeping vine plant in the Cucurbitaceae family that bears usually cylindrical fruits, which are used as culinary vegetables.eggplant Eggplant ( US, Canada), aubergine ( UK, Ireland) or brinjal (Indian subcontinent, Singapore, Malaysia, South Africa) is a plant species in the nightshade family Solanaceae. ''Solanum melongena'' is grown worldwide for its edible fruit. Mo ...

s (aubergines) and

banana A banana is an elongated, edible fruit – botanically a berry – produced by several kinds of large herbaceous flowering plants in the genus ''Musa''. In some countries, bananas used for cooking may be called "plantains", disting ...

s, but exclude certain fruits that meet the culinary definition of berries, such as

strawberries The garden strawberry (or simply strawberry; ''Fragaria × ananassa'') is a widely grown hybrid species of the genus '' Fragaria'', collectively known as the strawberries, which are cultivated worldwide for their fruit. The fruit is widely ap ...

and raspberries. The berry is the most common type of fleshy fruit in which the entire outer layer of the ovary wall ripens into a potentially edible " pericarp". Berries may be formed from one or more carpels from the same flower (i.e. from a simple or a compound ovary). The

seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...

s are usually embedded in the fleshy interior of the ovary, but there are some non-fleshy exceptions, such as peppers, with air rather than pulp around their seeds. Many berries are edible, but others, such as the fruits of the

potato The potato is a starchy food, a tuber of the plant ''Solanum tuberosum'' and is a root vegetable native to the Americas. The plant is a perennial in the nightshade family Solanaceae. Wild potato species can be found from the southern Uni ...

and the deadly nightshade, are

poison Poison is a chemical substance that has a detrimental effect to life. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figuratively, with a broa ...

ous to humans. A plant that bears berries is said to be bacciferous or baccate (a fruit that resembles a berry, whether it actually is a berry or not, can also be called "baccate"). In everyday English, a "

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, rasp ...

" is any small edible fruit. Berries are usually juicy, round, brightly coloured, sweet or sour, and do not have a stone or pit, although many small seeds may be present.

Botanical berries

botanical Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...

language, a berry is a simple fruit having

s and fleshy pulp (the pericarp) produced from the

of a single flower. The ovary can be inferior or

superior Superior may refer to: *Superior (hierarchy), something which is higher in a hierarchical structure of any kind Places *Superior (proposed U.S. state), an unsuccessful proposal for the Upper Peninsula of Michigan to form a separate state *Lake ...

. It is indehiscent, i.e. it does not have a special "line of weakness" along which it splits to release the seeds when ripe. The pericarp is divided into three layers. The outer layer is called the "exocarp" or " epicarp"; the middle layer, the " mesocarp" or "sarcocarp"; the inner layer, the " endocarp". Botanists have not applied these terms consistently. Exocarp and endocarp may be restricted to more-or-less single-layered "skins", or may include tissues adjacent to them; thus on one view, the exocarp extends inwards to the layer of vascular bundles ("veins"). The inconsistency in usage has been described as "a source of confusion". The nature of the endocarp distinguishes a berry from a drupe, which has a hardened or stony endocarp (see also below). The two kinds of fruit intergrade, depending on the state of the endocarp. Some sources have attempted to quantify the difference, e.g. requiring the endocarp to be less than 2 mm thick in a berry. Examples of botanical berries include: *

Avocado The avocado (''Persea americana'') is a medium-sized, evergreen tree in the laurel family ( Lauraceae). It is native to the Americas and was first domesticated by Mesoamerican tribes more than 5,000 years ago. Then as now it was prized for ...

contains a single large seed surrounded by an imperceptible endocarp. Avocados are however also sometimes classified as drupes. *

Banana A banana is an elongated, edible fruit – botanically a berry – produced by several kinds of large herbaceous flowering plants in the genus ''Musa''. In some countries, bananas used for cooking may be called "plantains", disting ...

Barberry ''Berberis'' (), commonly known as barberry, is a large genus of deciduous and evergreen shrubs from tall, found throughout temperate and subtropical regions of the world (apart from Australia). Species diversity is greatest in South America an ...

(''Berberis''),

Oregon-grape ''Mahonia aquifolium'', the Oregon grape or holly-leaved barberry, is a species of flowering plant in the family Berberidaceae, native to western North America. It is an evergreen shrub growing tall and wide, with pinnate leaves consisting of s ...

(''Mahonia aquifolium'') and mayapple (''Podophyllum'' spp.) ( Berberidaceae) * Strawberry tree (''Arbutus unedo'') (not to be confused with the

strawberry The garden strawberry (or simply strawberry; ''Fragaria × ananassa'') is a widely grown hybrid species of the genus '' Fragaria'', collectively known as the strawberries, which are cultivated worldwide for their fruit. The fruit is widely ap ...

(''Fragaria''), which is an accessory fruit), bearberry (''Arctostaphylos'' spp.),

bilberry Bilberries (), or sometimes European blueberries, are a primarily Eurasian species of low-growing shrubs in the genus '' Vaccinium'' (family Ericaceae), bearing edible, dark blue berries. The species most often referred to is ''Vaccinium myrti ...

, blueberry, cranberry, lingonberry/cowberry ('' Vaccinium vitis-idaea''),

crowberry ''Empetrum nigrum'', crowberry, black crowberry, or, in western Alaska, blackberry, is a flowering plant species in the heather family Ericaceae with a near circumboreal distribution in the Northern Hemisphere. It is usually dioecious, but ther ...

(''Empetrum'' spp.) (family Ericaceae) *

Coffee Coffee is a drink prepared from roasted coffee beans. Darkly colored, bitter, and slightly acidic, coffee has a stimulating effect on humans, primarily due to its caffeine content. It is the most popular hot drink in the world. Seeds of ...

berries ( Rubiaceae) (also described as drupes) "fruit a berry containing two (rarely one) seeds" *

Gooseberry Gooseberry ( or (American and northern British) or (southern British)) is a common name for many species of '' Ribes'' (which also includes currants), as well as a large number of plants of similar appearance. The berries of those in the gen ...

and currant (''Ribes'' spp.;

Grossulariaceae ''Ribes'' is a genus of about 200 known species of flowering plants, most of them native to the temperate regions of the Northern Hemisphere. The various species are known as currants or gooseberries, and some are cultivated for their edi ...

), red, black, and white types * Aubergine/

Eggplant Eggplant ( US, Canada), aubergine ( UK, Ireland) or brinjal (Indian subcontinent, Singapore, Malaysia, South Africa) is a plant species in the nightshade family Solanaceae. ''Solanum melongena'' is grown worldwide for its edible fruit. Mo ...

goji Goji, goji berry, or wolfberry () is the fruit of either '' Lycium barbarum'' or ''Lycium chinense'', two closely related species of boxthorn in the nightshade family, Solanaceae. ''L. barbarum'' and ''L. chinense'' fruits are simil ...

berries (wolfberry) and other species of the family Solanaceae * Elderberry (''Sambucus niger''; Adoxaceae) * Indian gooseberry ''(Phyllanthus emblica)'' ( Phyllanthaceae) * '' Garcinia gummi-gutta'', '' Garcinia mangostana'' (mangosteen) and ''

Garcinia indica ''Garcinia indica'', a plant in the mangosteen family (Clusiaceae), commonly known as ''kokum'', is a fruit-bearing tree that has culinary, pharmaceutical, and industrial uses. It primarily grows in the Western Ghats, especially the Goa and Konka ...

'' in the family Clusiaceae *

Sapodilla ''Manilkara zapota'', commonly known as sapodilla (), sapote, naseberry, nispero or chicle, is a long-lived, evergreen tree native to southern Mexico, Central America and the Caribbean. An example natural occurrence is in coastal Yucatán in the ...

(''Manilkara zapota''), Sapotaceae *

Grape A grape is a fruit, botanically a berry, of the deciduous woody vines of the flowering plant genus '' Vitis''. Grapes are a non- climacteric type of fruit, generally occurring in clusters. The cultivation of grapes began perhaps 8,000 years a ...

, ''

Vitis vinifera ''Vitis vinifera'', the common grape vine, is a species of flowering plant, native to the Mediterranean region, Central Europe, and southwestern Asia, from Morocco and Portugal north to southern Germany and east to northern Iran. There are cur ...

'' in the family Vitaceae *

Honeysuckle Honeysuckles are arching shrubs or twining vines in the genus ''Lonicera'' () of the family Caprifoliaceae, native to northern latitudes in North America and Eurasia. Approximately 180 species of honeysuckle have been identified in both con ...

: the berries of ''some'' species are edible and are called honeyberries, but others are poisonous (''Lonicera'' spp.;

Caprifoliaceae The Caprifoliaceae or honeysuckle family is a clade of dicotyledonous flowering plants consisting of about 860 species, in 33, to 42 genera, with a nearly cosmopolitan distribution. Centres of diversity are found in eastern North America and ...

) * Persimmon (

Ebenaceae The Ebenaceae are a family of flowering plants belonging to order Ericales. The family includes ebony and persimmon among about 768 species of trees and shrubs. It is distributed across the tropical and warmer temperate regions of the world. I ...

) * Pumpkin,

Modified berries

"True berries", or "baccae", may also be required to have a thin outer skin, not self-supporting when removed from the berry. This distinguishes, for example, a '' Vaccinium'' or '' Solanum'' berry from an ''

Adansonia ''Adansonia'' is a genus made up of eight species of medium-to-large deciduous trees known as baobabs ( or ). They are placed in the Malvaceae family, subfamily Bombacoideae. They are native to Madagascar, mainland Africa, and Australia.Trop ...

'' (baobab) amphisarca, which has a dry, more rigid and self-supporting skin. The fruit of

citrus ''Citrus'' is a genus of flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, lemons, grapefruits, pomelos, and limes. The genus ''Citrus'' is native to ...

, such as the orange, kumquat and lemon, is a berry with a thick rind and a very juicy interior divided into segments by

septa The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five c ...

, that is given the special name " hesperidium". A specialized term,

pepo PEPO Lappeenranta (abbreviated PEPO) is a football club from Lappeenranta in Finland. The club was formed in 1958 and their home ground is at the Kimpinen Sports Centre. The men's first team currently plays in the Ykkönen (Second Division). ...

, is also used for fruits of the gourd family, Cucurbitaceae, which are modified to have a hard outer rind, but are not internally divided by septae. The fruits of ''

Passiflora ''Passiflora'', known also as the passion flowers or passion vines, is a genus of about 550 species of flowering plants, the type genus of the family Passifloraceae. They are mostly tendril-bearing vines, with some being shrubs or trees. The ...

'' ( passion fruit) and '' Carica'' (papaya) are sometimes also considered pepos. Berries that develop from an inferior ovary are sometimes termed epigynous berries or false berries, as opposed to true berries, which develop from a superior ovary. In epigynous berries, the berry includes tissue derived from parts of the flower besides the ovary. The floral tube, formed from the basal part of the sepals, petals and stamens can become fleshy at maturity and is united with the ovary to form the fruit. Common fruits that are sometimes classified as epigynous berries include

coffee Coffee is a drink prepared from roasted coffee beans. Darkly colored, bitter, and slightly acidic, coffee has a stimulating effect on humans, primarily due to its caffeine content. It is the most popular hot drink in the world. Seeds of ...

, members of the genus '' Vaccinium'' (e.g., cranberries and blueberries), and members of the family Cucurbitaceae (gourds,

cucumber Cucumber (''Cucumis sativus'') is a widely-cultivated creeping vine plant in the Cucurbitaceae family that bears usually cylindrical fruits, which are used as culinary vegetables.melon A melon is any of various plants of the family Cucurbitaceae with sweet, edible, and fleshy fruit. The word "melon" can refer to either the plant or specifically to the fruit. Botanically, a melon is a kind of berry, specifically a "pepo". T ...

s and squash).

Berry-like fruits

Many fruits commonly referred to as berries are not actual berries by the scientific definition, but fall into one of the following categories:

Drupes

Drupes are varyingly distinguished from botanical berries. Drupes are fleshy fruits produced from a (usually) single-seeded ovary with a hard woody layer (called the endocarp) surrounding the seed. Familiar examples include the stonefruits of the genus ''

Prunus ''Prunus'' is a genus of trees and shrubs, which includes (among many others) the fruits plums, cherries, peaches, nectarines, apricots, and almonds. Native to the North American temperate regions, the neotropics of South America, and the ...

'' (

peach The peach (''Prunus persica'') is a deciduous tree first domesticated and cultivated in Zhejiang province of Eastern China. It bears edible juicy fruits with various characteristics, most called peaches and others (the glossy-skinned, n ...

es,

plum A plum is a fruit of some species in ''Prunus'' subg. ''Prunus'.'' Dried plums are called prunes. History Plums may have been one of the first fruits domesticated by humans. Three of the most abundantly cultivated species are not found ...

s and

cherries A cherry is the fruit of many plants of the genus '' Prunus'', and is a fleshy drupe (stone fruit). Commercial cherries are obtained from cultivars of several species, such as the sweet ''Prunus avium'' and the sour ''Prunus cerasus''. The ...

olive The olive, botanical name ''Olea europaea'', meaning 'European olive' in Latin, is a species of small tree or shrub in the family Oleaceae, found traditionally in the Mediterranean Basin. When in shrub form, it is known as ''Olea europaea'' ' ...

coconut The coconut tree (''Cocos nucifera'') is a member of the palm tree family (Arecaceae) and the only living species of the genus ''Cocos''. The term "coconut" (or the archaic "cocoanut") can refer to the whole coconut palm, the seed, or the f ...

, dates,

bayberry ''Myrica'' is a genus of about 35–50 species of small trees and shrubs in the family Myricaceae, order Fagales. The genus has a wide distribution, including Africa, Asia, Europe, North America and South America, and missing only from Aust ...

and ''

Persea ''Persea'' is a genus of about 150 species of evergreen trees belonging to the laurel family, Lauraceae. The best-known member of the genus is the avocado, ''P. americana'', widely cultivated in subtropical regions for its large, edible fruit. ...

'' species. Some definitions make the mere presence of an internally differentiated endocarp the defining feature of a drupe; others qualify the nature of the endocarp required in a drupe, e.g. defining berries to have endocarp less than 2 mm thick. The term "drupaceous" is used of fruits that have the general structure and texture of a drupe, without necessarily meeting the full definition. Other drupe-like fruits with a single seed that lack the stony endocarp include sea-buckthorn (''Hippophae rhamnoides'', Elaeagnaceae), which is an

achene An achene (; ), also sometimes called akene and occasionally achenium or achenocarp, is a type of simple dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and indehiscent (they do not o ...

, surrounded by a swollen

hypanthium In angiosperms, a hypanthium or floral cup is a structure where basal portions of the calyx, the corolla, and the stamens form a cup-shaped tube. It is sometimes called a floral tube, a term that is also used for corolla tube and calyx tube. It ...

that provides the fleshy layer. Fruits of ''Coffea'' species are described as either drupes or berries.

Pomes

The pome fruits produced by plants in subtribe Pyrinae of family

Rosaceae Rosaceae (), the rose family, is a medium-sized family of flowering plants that includes 4,828 known species in 91 genera. The name is derived from the type genus ''Rosa''. Among the most species-rich genera are '' Alchemilla'' (270), ''Sorb ...

, such as apples and pears, have a structure (the core) in which tough tissue clearly separates the seeds from the outer softer pericarp. Pomes are not berries. However, some of the smaller pomes are sometimes referred to as berries. ''

Amelanchier ''Amelanchier'' ( ), also known as shadbush, shadwood or shadblow, serviceberry or sarvisberry (or just sarvis), juneberry, saskatoon, sugarplum, wild-plum or chuckley pear,A Digital Flora of Newfoundland and Labrador Vascular Plants/ref> is a g ...

'' pomes become so soft at maturity that they resemble a blueberry and are known as Juneberries, serviceberries or Saskatoon berries.

Aggregate fruits

Aggregate or compound fruits contain seeds from different ovaries of a single flower, with the individual "fruitlets" joined at maturity to form the complete fruit. Examples of aggregate fruits commonly called "berries" include members of the genus ''Rubus'', such as

blackberry The blackberry is an edible fruit produced by many species in the genus ''Rubus'' in the family Rosaceae, hybrids among these species within the subgenus ''Rubus'', and hybrids between the subgenera ''Rubus'' and ''Idaeobatus''. The taxonomy ...

and

raspberry The raspberry is the edible fruit of a multitude of plant species in the genus ''Rubus'' of the rose family, most of which are in the subgenus '' Idaeobatus''. The name also applies to these plants themselves. Raspberries are perennial with ...

. Botanically, these are not berries. Other large aggregate fruits, such as soursop (''

Annona muricata Soursop (also called ''graviola, guyabano'', and in Hispanic America, ''guanábana'') is the fruit of ''Annona muricata'', a broadleaf, flowering, evergreen tree. It is native to the tropical regions of the Americas and the Caribbean and is wi ...

''), are not usually called "berries", although some sources do use this term.

Multiple fruits

Multiple fruits are not botanical berries. Multiple fruits are the fruits of two or more multiple flowers that are merged or packed closely together. The

mulberry ''Morus'', a genus of flowering plants in the family Moraceae, consists of diverse species of deciduous trees commonly known as mulberries, growing wild and under cultivation in many temperate world regions. Generally, the genus has 64 ident ...

is a berry-like example of a multiple fruit; it develops from a cluster of tiny separate flowers that become compressed as they develop into fruit.

Accessory fruits

Accessory fruits are not botanical berries. In accessory fruits, the edible part is not generated by the ovary. Berry-like examples include: *

Strawberry The garden strawberry (or simply strawberry; ''Fragaria × ananassa'') is a widely grown hybrid species of the genus '' Fragaria'', collectively known as the strawberries, which are cultivated worldwide for their fruit. The fruit is widely ap ...

– the non-fleshy aggregate of seed-like

s on its exterior is actually the "fruit", derived from an aggregate of ovaries; the fleshy part develops instead from the receptacle. *

Mock strawberry ''Potentilla indica'', known commonly as mock strawberry, Indian-strawberry, or false strawberry, often referred to as a backyard strawberry, mainly in North America, is a flowering plant in the family Rosaceae. It has foliage and an aggregate ac ...

(''Duchesnea indica)'' – structured just like a strawberry. * Sea grape ('' Coccoloba uvifera'';

Polygonaceae The Polygonaceae are a family of flowering plants known informally as the knotweed family or smartweed—buckwheat family in the United States. The name is based on the genus ''Polygonum'', and was first used by Antoine Laurent de Jussieu in ...

) – the fruit is a dry capsule surrounded by fleshy calyx.

Berry-like conifer seed cones

The female seed cones of some

conifer Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...

s have fleshy and merged scales, giving them a berry-like appearance. Juniper "berries" (family

Cupressaceae Cupressaceae is a conifer family, the cypress family, with worldwide distribution. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 species in total. They are monoecious, subdioeci ...

), in particular those of '' Juniperus communis'', are used to flavour gin. The seed cones of species in the families

Podocarpaceae Podocarpaceae is a large family of mainly Southern Hemisphere conifers, known in English as podocarps, comprising about 156 species of evergreen trees and shrubs.James E. Eckenwalder. 2009. ''Conifers of the World''. Portland, Oregon: Timber P ...

and

Taxaceae Taxaceae (), commonly called the yew family, is a coniferous family which includes six extant and two extinct genera, and about 30 species of plants, or in older interpretations three genera and 7 to 12 species. Description They are many-bran ...

have a bright colour when fully developed, increasing the resemblance to true berries. The "berries" of yews (''

Taxus ''Taxus'' is a genus of coniferous trees or shrubs known as yews in the family Taxaceae. They are relatively slow-growing and can be very long-lived, and reach heights of , with trunk girth averaging . They have reddish bark, lanceolate, flat ...

'' species) consist of a female seed cone with which develops a fleshy red

aril An aril (pronounced ), also called an arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode or false aril is sometimes distinguished: whereas an aril grows from the attachment point of the see ...

partially enclosing the poisonous seed.

History of terminology

The Latin word ' or ' (plural ') was originally used for "any small round fruit".

Andrea Caesalpinus Andrea Cesalpino ( Latinized as Andreas Cæsalpinus) (6 June 1524 – 23 February 1603) was a Florentine physician, philosopher and botanist. In his works he classified plants according to their fruits and seeds, rather than alphabetically o ...

(1519–1603) classified plants into trees and herbs, further dividing them by properties of their flowers and fruit. He did not make the modern distinction between "fruits" and "seeds", calling hard structures like nuts ' or seeds. A fleshy fruit was called a '. For Caesalpinus, a true ' or berry was a ' derived from a flower with a superior ovary; one derived from a flower with an inferior ovary was called a '. In 1751,

Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...

wrote '' Philosophia Botanica'', considered to be the first textbook of descriptive systematic botany. He used eight different terms for fruits, one of which was ' or berry, distinguished from other types of fruit such as ' (drupe) and ' (pome). A ' was defined as "'", meaning "unvalved solid pericarp, containing otherwise naked seeds". The adjective "'" here has the sense of "solid with tissue softer than the outside; stuffed". A berry or ' was distinguished from a drupe and a pome, both of which also had an unvalved solid pericarp; a drupe also contained a nut (') and a pome a capsule ('), rather than the berry's naked seeds. Linnaeus' use of ' and ' was thus significantly different from that of Caesalpinus. Botanists continue to differ on how fruit should be classified.

Joseph Gaertner Joseph Gaertner (12 March 1732 – 14 July 1791) was a German botanist, best known for his work on seeds, ''De Fructibus et Seminibus Plantarum'' (1788-1792). Biography He was born in Calw, and studied in Göttingen under Albrecht von Haller. ...

published a two-volume work, ''

De Fructibus et Seminibus Plantarum ''De Fructibus et Seminibus Plantarum'', also known by its standard botanical abbreviation ''Fruct. Sem. Pl.'', is a three-volume botanic treatise by Joseph Gaertner. The first volume was published in December 1788. The second volume was published ...

'' (on the fruits and seeds of plants) between 1788 and 1792. In addition to Linnaeus' eight terms, he introduced seven more, including ' for the berry-like fruits of cucurbits. A pepo was distinguished by being a fleshy berry with the seeds distant from the axis, and so nearer the fruit wall (i.e. by having " parietal placentation" in modern terminology). Nicaise Auguste Desvaux in 1813 used the terms ' and ' as further subdivisions of berries. A hesperidium, called by others ' (berry with a cortex), had separate internal compartments (''""'' in the original French) and a separable membraneous epicarp or skin. An amphisarca was described as woody on the outside and fleshy on the inside. "Hesperidium" remains in general use, but "amphisarca" is rarely used. There remains no universally agreed system of classification for fruits, and there continues to be "confusion over classification of fruit types and the definitions given to fruit terms".

Evolution and phylogenetic significance

By definition, berries have a fleshy, indehiscent pericarp, as opposed to a dry, dehiscent pericarp.

Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

s show that early

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...

s had dry fruits; fleshy fruits, such as berries or drupes, appeared only towards the end of the Cretaceous Period or the beginning of the Paleogene Period, about . The increasing importance of seed dispersal by fruit-eating vertebrates, both mammals and birds, may have driven the evolution of fleshy fruits. Alternatively, the causal direction may be the other way round. Large fleshy fruits are associated with moist habitats with closed tree canopies, where wind dispersal of dry fruits is less effective. Such habitats were increasingly common in the Paleogene and the associated change in fruit type may have led to the evolution of fruit eating in mammals and birds. Fruit type has been considered to be a useful character in classification and in understanding the

phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological s ...

of plants. The evolution of fruits with a berry-like pericarp has been studied in a wide range of flowering plant families. Repeated transitions between fleshy and dry pericarps have been demonstrated regularly. One well-studied family is the Solanaceae, because of the commercial importance of fruit such as tomatoes, bell peppers, and eggplants or aubergines. Capsules, which are dry dehiscent fruits, appear to be the original form of the fruit in the earliest diverging members of the family. Berries have then evolved at least three times: in '' Cestrum'', '' Duboisia'', and in the subfamily Solanoideae. Detailed anatomical and developmental studies have shown that the berries of ''Cestrum'' and those of the Solanoideae are significantly different; for example, expansion of the fruit during development involves cell divisions in the mesocarp in Solanoideae berries, but not in ''Cestrum'' berries. When fruits described as berries were studied in the family

Melastomaceae Melastomataceae is a family of dicotyledonous flowering plants found mostly in the tropics (two-thirds of the genera are from the New World tropics) comprising c. 175 genera and c. 5115 known species. Melastomes are annual or perennial herbs, sh ...

, they were found to be highly variable in structure, some being soft with an endocarp that soon broke down, others having a hard, persistent endocarp, even woody in some species. Fruits classified as berries are thus not necessarily homologous, with the fleshy part being derived from different parts of the ovary, and with other structural and developmental differences. The presence or absence of berries is not a reliable guide to phylogeny. Indeed, fruit type in general has proved to be an unreliable guide to flowering plant relationships.

Uses

Culinary

Berries, defined loosely, have been valuable as a food source to humans since before the start of agriculture, and remain among the primary food sources of other primates. Botanically defined berries with culinary uses include: * Berries in the strictest sense: including bananas and plantains, blueberries, cranberries, coffee berries, gooseberries, red-, black- and white currants, tomatoes, grapes and peppers (''

Capsicum ''Capsicum'' () is a genus of flowering plants in the nightshade family Solanaceae, native to the Americas, cultivated worldwide for their chili pepper or bell pepper fruit. Etymology and names The generic name may come from Latin , me ...

'' fruits) * Hesperidia: citrus fruits, including oranges, lemons and limes * Pepos: cucurbits, including squashes, cucumbers, melons and watermelons Some berries are brightly coloured, due to

plant pigment Biological pigments, also known simply as pigments or biochromes, are substances produced by living organisms that have a color resulting from selective color absorption. Biological pigments include plant pigments and flower pigments. Many biol ...

s such as

anthocyanin Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical com ...

s and other flavonoids. These pigments are localized mainly in the outer surface and the

s. Such pigments have

antioxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubrica ...

properties ''

in vitro ''In vitro'' (meaning in glass, or ''in the glass'') studies are performed with microorganisms, cells, or biological molecules outside their normal biological context. Colloquially called " test-tube experiments", these studies in biology a ...

'', but there is no reliable evidence that they have antioxidant or any other useful functions within the human body. Consequently, it is not permitted to claim that foods containing plant pigments have antioxidant health value on product labels in the United States or Europe. Some

spice A spice is a seed, fruit, root, bark, or other plant substance primarily used for flavoring or coloring food. Spices are distinguished from herbs, which are the leaves, flowers, or stems of plants used for flavoring or as a garnish. Spices a ...

s are prepared from berries. Allspice is made from the dried berries of ''Pimenta dioica''. The fruits (berries) of different

cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...

s of '' Capsicum annuum'' are used to make paprika (mildly hot), chili pepper (hot) and cayenne pepper (very hot).

Others

Pepos, characterized by a hard outer rind, have also been used as containers by removing the inner flesh and seeds and then drying the remaining exocarp. The English name of '' Lagenaria siceraria'', "bottle gourd", reflects its use as a liquid container. Some true berries have also been used as a source of

dye A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution and ...

s. In

Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...

, these included berries from a species of '' Dianella'', used to produce blue, and berries from black nightshade ('' Solanum americanum''), used to produce green.

History

Cucurbit berries or pepos, particularly from ''Cucurbita'' and ''Lagenaria'', are the earliest plants known to be domesticated – before 9,000–10,000 BP in the Americas, and probably by 12,000–13,000 BP in Asia. Peppers were domesticated in Mesoamerica by 8,000 BP. Many other early cultivated plants were also berries by the strict botanical definition, including grapes, domesticated by 8,000 BP and known to have been used in wine production by 6,000 BP. Bananas were first domesticated in

Papua New Guinea Papua New Guinea (abbreviated PNG; , ; tpi, Papua Niugini; ho, Papua Niu Gini), officially the Independent State of Papua New Guinea ( tpi, Independen Stet bilong Papua Niugini; ho, Independen Stet bilong Papua Niu Gini), is a country i ...

and

Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainland ...

Archaeological Archaeology or archeology is the scientific study of human activity through the recovery and analysis of material culture. The archaeological record consists of artifacts, architecture, biofacts or ecofacts, sites, and cultural landscap ...

and palaeoenvironmental evidence at Kuk Swamp in the Western Highlands Province of Papua New Guinea suggests that banana cultivation there goes back to at least 7,000 BP, and possibly to 10,000 BP. The history of cultivated

citrus fruit ''Citrus'' is a genus of flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, lemons, grapefruits, pomelos, and limes. The genus ''Citrus'' is native to ...

remains unclear, although some recent research suggests a possible origin in

Papuasia Papuasia is a Level 2 botanical region defined in the World Geographical Scheme for Recording Plant Distributions (WGSRPD). It lies in the southwestern Pacific Ocean, in the Melanesia ecoregion of Oceania and Tropical Asia. It comprises th ...

rather than continental southeast Asia. Chinese documents show that mandarins and pomelos were established in cultivation there by around 4,200 BP.

Commercial production

According to

FAOSTAT The Food and Agriculture Organization Corporate Statistical Database (FAOSTAT) website disseminates statistical data collected and maintained by the Food and Agriculture Organization (FAO). FAOSTAT data are provided as a time-series from 1961 in ...

data, in 2013 four of the five top fruit crops in terms of world production by weight were botanical berries. The other was a pome (apples). According to FAOSTAT, in 2001, bananas (including plantains) and citrus comprised over 25% by value of the world's exported fruits and vegetables, citrus fruits being more valuable than bananas. Export quantities of fruit are not entirely comparable with production quantities, since slightly different categories are used. The top five fruit exports by weight in 2012 are shown in the table below. The top two places are again occupied by bananas and citrus.

Notes

References

External links

{{Authority control Fruit morphology