Redcurrants, a type of berry derived from a simple (one-locule) inferior ovaryKiwifruit, a berry derived from a compound (many carpellate) superior ovary
Inbotany, aberry is a fleshyfruit produced from a singleflower containing oneovary.[1] Berries so defined includegrapes,currants, andtomatoes, as well ascucumbers,eggplants (aubergines),persimmons andbananas, but exclude certain fruits that meet theculinary definition of berries, such asstrawberries andraspberries. 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 morecarpels from the same flower (i.e. from a simple or a compound ovary).[2]: 291 Theseeds are usually embedded in the fleshy interior of the ovary, but there are some non-fleshy exceptions, such asCapsicum species, with air rather than pulp around their seeds.
A plant that bears berries is said to bebacciferous orbaccate[a] (from Latinbacca).
In everyday English, a "berry" is any small edible fruit. Berries are usually juicy, round, brightly coloured,sweet orsour, and do not have astone or pit, although many small seeds may be present.[3]
Diagram of a grape berry, showing the pericarp and its layersCoffee cherries (Coffea arabica) – described as drupes or berries
Inbotanical language, a berry is a simple fruit havingseeds and fleshy pulp (thepericarp) produced from theovary of a single flower. The ovary can beinferior orsuperior. It isindehiscent, i.e. it does not have a special "line of weakness" along which it splits to release the seeds when ripe.[4] 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 ofvascular bundles ("veins"). The inconsistency in usage has been described as "a source of confusion".[5]
The nature of the endocarp distinguishes a berry from adrupe, 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.[6]
Examples of botanical berries include:
Avocado contains a single large seed surrounded by an imperceptible endocarp.[7] Avocados are, however, also sometimes classified as drupes.[8]
Cross-section of a cucumber pepo (Cucumis sativus)
"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, aVaccinium orSolanum berry from anAdansonia (baobab) amphisarca, which has a dry, more rigid and self-supporting skin.[13] The fruit ofcitrus, such as theorange,kumquat andlemon, is a berry with a thick rind and a very juicy interior divided into segments bysepta, which is given the special name "hesperidium".[13] A special term,pepo, is also used for fruits of the gourd family,Cucurbitaceae, which have a hard outer rind but are not internally divided by septa.[13] The fruits ofPassiflora (passion fruit) andCarica (papaya) are sometimes also considered pepos.[13]
Berries that develop from an inferior ovary are sometimes termedepigynous 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 other than 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 includebananas,coffee, members of the genusVaccinium (e.g., cranberries and blueberries), and members of the family Cucurbitaceae (gourds,cucumbers,melons andsquash).[14]
In the first image, only theblueberry is botanically a berry:blackberries are aggregate fruit composed of many drupelets, andstrawberries are aggregate accessory fruits. Sloes, the fruits ofPrunus spinosa, are drupes.Avocado fruit are described as either drupes or berries. Serviceberries, fruits ofAmelanchier species, arepomes. Mulberries, the fruits ofMorus nigra, are multiple fruits.
Many fruits which are berries in the culinary definition are not berries in the botanic sense, but fall into one of the following categories:
Drupes are varyingly distinguished from botanical berries. Drupes are fleshy fruits produced from a (usually) single-seeded ovary with a hard woody layer (called theendocarp) surrounding the seed. Familiar examples include the stonefruits of the genusPrunus (peaches,plums andcherries),olives,coconut,dates,bayberry andPersea species. Some definitions make the mere presence of an internally differentiated endocarp the defining feature of a drupe;[13] others qualify the nature of the endocarp required in a drupe, e.g. defining berries to have endocarp less than 2 mm thick.[6] The term "drupaceous" is used to describe fruits that have the general structure and texture of a drupe,[15] without necessarily meeting the full definition. Other drupe-like fruits with a single seed that lack the stony endocarp includesea-buckthorn (Hippophae rhamnoides,Elaeagnaceae), which is anachene, surrounded by a swollenhypanthium that provides the fleshy layer.[16] Fruits ofCoffea species are described as either drupes or berries.[11]
The pome fruits produced by plants in subtribe Pyrinae of familyRosaceae, such as apples and pears, have a structure (the core) in which tough tissue separates the seeds from the outer softer pericarp.[17] Although pomes are not botanical berries,Amelanchier pomes become soft at maturity, resembling ablueberry, and are commonly called Juneberries, serviceberries orSaskatoon berries.[18]
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.[19] Examples of aggregate fruits commonly called "berries" include members of the genusRubus, such asblackberry andraspberry.[20] Botanically, these are not berries. Other large aggregate fruits, such as soursop (Annona muricata),[21] are not usually called "berries", although some sources do use this term.[22]
Multiple fruits are not botanical berries. Multiple fruits are the fruits of two or more multiple flowers that are merged or packed closely together.[23] Themulberry 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.[24]
Accessory fruits are not botanical berries. In accessory fruits, the edible part is not generated by the ovary. Berry-like examples include:
Strawberry – the non-fleshy aggregate of seed-likeachenes on its exterior is actually the "fruit", derived from an aggregate of ovaries; the fleshy part develops instead from thereceptacle.
Mock strawberry (Duchesnea indica) – structured just like a strawberry.
The female seedcones of someconifers have fleshy and merged scales, giving them a berry-like appearance.Juniper "berries" (familyCupressaceae), in particular those ofJuniperus communis, are used to flavourgin. The seed cones of species in the familiesPodocarpaceae andTaxaceae have a bright colour when fully developed, increasing the resemblance to true berries. The "berries" of yews (Taxus species) consist of a female seed cone with which develops a fleshy redaril partially enclosing the poisonous seed.[citation needed]
Some fruits classified asbacca (berries) by Gaertner (De Fructibus et Seminibus Plantarum, Tab. 28)
The Latin wordbaca orbacca (pluralbaccae) was originally used for "any small round fruit".[25]Andrea Caesalpinus (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 nutssemina or seeds. A fleshy fruit was called apericarpium. For Caesalpinus, a truebacca or berry was apericarpium derived from a flower with a superior ovary; one derived from a flower with an inferior ovary was called apomum.[26]
In 1751,Carl Linnaeus wrotePhilosophia Botanica, considered to be the first textbook of descriptive systematic botany.[27] He used eight different terms for fruits, one of which wasbacca or berry, distinguished from other types of fruit such asdrupa (drupe) andpomum (pome).[28] Abacca was defined as "pericarpium farctum evalve, semina ceteroquin nuda continens", meaning "unvalved solid pericarp, containing otherwise naked seeds".[29] The adjective "farctus" here has the sense of "solid with tissue softer than the outside; stuffed".[30] A berry orbacca was distinguished from a drupe and a pome, both of which also had an unvalved solid pericarp; a drupe also contained a nut (nux) and a pome a capsule (capsula), rather than the berry's naked seeds.[29] Linnaeus' use ofbacca andpomum was thus significantly different from that of Caesalpinus. Botanists continue to differ on how fruit should be classified.[28]
Joseph Gaertner published a two-volume work,De Fructibus et Seminibus Plantarum (on the fruits and seeds of plants) between 1788 and 1792. In addition to Linnaeus' eight terms, he introduced seven more, includingpepo for the berry-like fruits of cucurbits.[28] A pepo was distinguished by being a fleshy berry with the seeds distant from the axis, and so nearer the fruit wall[31] (i.e. by having "parietal placentation" in modern terminology).Nicaise Auguste Desvaux in 1813 used the termshesperidium andamphisarca as further subdivisions of berries.[28] A hesperidium, called by othersbacca corticata (berry with a cortex), had separate internal compartments ("loges" 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.[32] "Hesperidium" remains in general use, but "amphisarca" is rarely used.[28]
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".[28]
By definition, berries have a fleshy, indehiscent pericarp, as opposed to a dry, dehiscent pericarp.Fossils show that earlyflowering plants had dry fruits; fleshy fruits, such as berries or drupes, appeared only towards the end of theCretaceous Period or the beginning of thePaleogene Period, about66 million years ago. 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 around. 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.[33]
Fruit type has been considered to be a useful character in classification and in understanding thephylogeny of plants.[34] The evolution of fruits with a berry-like pericarp has been studied in various flowering plant families. Repeated transitions between fleshy and dry pericarps have been demonstrated regularly. One well-studied family is theSolanaceae, 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: inCestrum,Duboisia, and in the subfamilySolanoideae. Detailed anatomical and developmental studies have shown that the berries ofCestrum 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 inCestrum berries.[35]
When fruits described as berries were studied in the familyMelastomaceae, 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.[34] 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.[33] Indeed, fruit type in general has proved to be an unreliable guide to flowering plant relationships.[34]
Berries, defined loosely, have been valuable as a food source to humans since before the start of agriculture,[citation needed] 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 fruits)
Hesperidia: citrus fruits, including oranges, lemons and limes
Pepos: cucurbits, including squashes, cucumbers, melons and watermelons
Some berries are brightly coloured, due toplant pigments such asanthocyanins and otherflavonoids. These pigments are localized mainly in the outer surface and theseeds.[36] Such pigments haveantioxidant propertiesin vitro,[37] 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.[38][39]
Somespices are prepared from berries.Allspice is made from the dried berries ofPimenta dioica.[40] The fruits (berries) of differentcultivars ofCapsicum annuum are used to make paprika (mildly hot), chili pepper (hot) and cayenne pepper (very hot).[41]
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 ofLagenaria siceraria, "bottle gourd", reflects its use as a liquid container.[42]
Some true berries have also been used as a source ofdyes. InHawaii, these included berries from a species ofDianella, used to produce blue, and berries from black nightshade (Solanum americanum), used to produce green.[43]
Cucurbit berries or pepos, particularly fromCucurbita andLagenaria, are the earliest plants known to bedomesticated – before 9,000–10,000 BP in the Americas, and probably by 12,000–13,000 BP in Asia.[42] Peppers were domesticated in Mesoamerica by 8,000 BP.[44][45] Many other early cultivated plants were also berries by the strict botanical definition, including grapes, domesticated by 8,000 BP[46][47] and known to have been used in wine production by 6,000 BP.[48]
The history of cultivatedcitrus fruit remains unclear, although some recent research suggests a possible origin inPapuasia rather than continental southeast Asia. Chinese documents show thatmandarins andpomelos were established in cultivation there by around 4,200 BP.[51]
Four banana and plantain cultivarsWatermelonGrapes
According toFAOSTAT 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).[Note 1]
Worldwide fruit production in 2013
Name
Thousands of tonnes
Fruit type
Bananas & plantains
144,592
Berry
Citrus fruit†
135,761
Berry (hesperidium)
Watermelons
109,279
Berry (pepo)
Apples
80,823
Pome
Grapes
77,181
Berry
†Citrus fruit includes but is not limited to, oranges, lemons, limes, grapefruit (including pomelos), tangerines, mandarins, clementines, and satsumas. Oranges make up 53% of the total.
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.[53] 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. Bananas and citrus occupy the top two places.[Note 2]
Worldwide fruit export in 2012
Name
Thousands of tonnes
Fruit type
Bananas & plantains
19,725
Berry
Citrus fruit†
15,262
Berry (hesperidium)
Apples
8,271
Pome
Prepared fruit‡
7,120
–
Grapes
4,051
Berry
†Citrus fruit includes oranges, lemons, limes, grapefruit (including pomelos), tangerines, mandarins, clementines, and satsumas. Oranges make up 43% of the total. ‡Prepared fruit here is "fruit, nuts and peel, including frozen, prepared or preserved, jam, paste, marmalade, purée and cooked fruits, other than those listed separately".[54]
^A fruit that resembles a berry, whether it is one or not, can also be called "baccate".
^Production Quantity data for 2013 for the World total for all items was downloaded to a spreadsheet from FAOSTAT.[52] Items coded 486 to 626, corresponding to fruit, were retained. Data for bananas and plantains were combined, as these are not distinguished by all countries. Data for all citrus fruits were also combined since some countries provide data under a general heading of "citrus fruit". The resulting table was then sorted and the top five entries used.
^Export Quantity data for 2012 for the World total for all items was downloaded to a spreadsheet from FAOSTAT.[52] As for production, items coded 486 to 626 (but now excluding 564 wine, not present in the production data) were retained. Data for bananas and plantains were combined, as was data for all citrus fruits. The resulting table was then sorted and the top five entries were used.
^Kiger, Robert W. & Porter, Duncan M. (2001)."Find term 'berry'".Categorical Glossary for the Flora of North America Project. Archived fromthe original on 20 May 2019. Retrieved14 August 2015.
^abcHeywood, V.H.; Brummitt, R.K.; Culham, A.; Seberg, O. (2007).Flowering plant families of the world. Firefly Books.ISBN978-1-55407-206-4.
^abcdeSpjut, Richard W."Classification of Fruit Types".A Systematic Treatment of Fruit Types. The World Botanical Associates. Archived fromthe original on 29 March 2015. Retrieved16 August 2015.
^Kiger, Robert W. & Porter, Duncan M. (2001)."Find term 'drupaceous'".Categorical Glossary for the Flora of North America Project. Archived fromthe original on 20 May 2019. Retrieved14 August 2015.
^Kiger, Robert W. & Porter, Duncan M. (2001)."Find term 'pome'".Categorical Glossary for the Flora of North America Project. Archived fromthe original on 20 May 2019. Retrieved21 August 2015.
^Campbell, Christopher S.; Burgess, Michael B. & Cushman, Kevin R."Amelanchier". In Flora of North America Editorial Committee (ed.).Flora of North America(online). eFloras.org. Retrieved21 August 2015.
^Kiger, Robert W. & Porter, Duncan M. (2001)."Find term 'aggregate fruit'".Categorical Glossary for the Flora of North America Project. Archived fromthe original on 20 May 2019. Retrieved14 August 2015.
^Lu, Lingdi & Boufford, David E."Rubus". In Wu, Zhengyi; Raven, Peter H. & Hong, Deyuan (eds.).Flora of China(online). eFloras.org. Retrieved21 August 2015.
^Mabberley, D.J. (1978). "Annonaceae". In Heywood, V.H.; Moore, D.M.; Richardson, I.B.K. & Stearn, W.T. (eds.).Flowering Plants of the World. Oxford University Press.ISBN978-0-19-217674-5.
^"Annonaceae".Encyclopædia Britannica. Retrieved22 August 2015.
^Kiger, Robert W. & Porter, Duncan M. (2001)."Find term 'multiple fruit'".Categorical Glossary for the Flora of North America Project. Archived fromthe original on 20 May 2019. Retrieved14 August 2015.
^Stearn, W.T. (2004).Botanical Latin (4th (p/b) ed.). Portland, Oregon: Timber Press.ISBN978-0-7153-1643-6. p. 376.
^Vines, S.H. (1913)."Robert Morison and John Ray". In Oliver, F.W. (ed.).Makers of British Botany (Project Gutenberg EBook ed.). Cambridge University Press. Retrieved18 August 2015.
^Gaertner, Joseph (1788–1792).De Fructibus et Seminibus Plantarum (in Latin). p. XCVII.Pepo, qui vulgo majorem Cucurbitae fructum denotat, nobis generatim dicitur bacca carnosa, cuius loculamenta ab axi remota, et prope fructus peripheriam ita posita sunt, ut etiam semina, ejus parietibus affigantur. (a fleshy berry, whose locules are remote from the axis, and are so positioned near the periphery of the fruit, that as well as the seeds, they are affixed to its walls, is generally called by us 'pepo', which usually denotes the larger fruit of cucurbits)
^Desvaux, N.A. (1813). "Essai sur les différens genres de fruits des plantes phanérogames".Journal de Botanique, appliqué a l'Agriculture, a la Pharmacie, a la Médecine, et aux Arts (in French).2:161–183.
^abKnapp, Sandra (2002), "Tobacco to tomatoes: a phylogenetic perspective on fruit diversity in the Solanaceae",Journal of Experimental Botany,53 (377):2001–2022,doi:10.1093/jxb/erf068,PMID12324525
^abcClausing, Gudrun; Meyer, Karsten & Renner, Susanne S. (2000), "Correlations among fruit traits and evolution of different fruits within Melastomataceae",Botanical Journal of the Linnean Society,133 (3):303–326,doi:10.1006/bojl.1999.0340
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