Berry (botany)
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In
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 wo ...

botany
, a berry is a fleshy
fruit In 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 ...

fruit
without a stone (pit) produced from a single
flower A flower, sometimes known as a bloom or blossom Cherry blossoms in Paris in full bloom. In botany, blossoms are the flowers of stone fruit fruit tree, trees (genus ''Prunus'') and of some other plants with a similar appearance that flower prof ...

flower
containing one
ovary The ovary is an organ found in the female reproductive system 400px, 1. Labia_majora.html"_;"title="Vulva: 2. Labia_majora">Vulva: 2. Labia_majora; 3. Labia_minora; 4. Vulval_vestibule.html" ;"title="Labia_minora.html" ...
. Berries so defined include
grape A grape is a fruit In botany, a fruit is the seed-bearing structure in flowering plants (also known as angiosperms) formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate seeds. Edible fruits, in ...

grape
s,
currants
currants
, and
tomato The tomato is the edible berry A berry is a small, pulpy, and often edible fruit In botany, a fruit is the seed-bearing structure in flowering plants (also known as angiosperms) formed from the ovary after flowering. Fruits are ...

tomato
es, as well as
cucumber Cucumber (''Cucumis sativus'') is a widely-cultivated creeping vine plant in the '' Cucurbitaceae'' gourd family that bears cucumiform fruits In botany Botany, also called , plant biology or phytology, is the science Scien ...

cucumber
s,
eggplant Eggplant ( US, Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous ...

eggplant
s (aubergines) and
banana A banana is an elongated, edible fruit – botanically a berry (botany), berry – produced by several kinds of large herbaceous flowering plants in the genus ''Musa (genus), Musa''. In some countries, Cooking banana, bananas used for c ...

banana
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 (biology), hybrid species of the genus ''Fragaria'', collectively known as the strawberries, which are cultivated worldwide for their fruit. The fr ...

strawberries
and
raspberries The raspberry is the edible fruit of a multitude of plant species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is ...

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 Fruit anatomy is the plant anatomy of the internal structure of fruit In botany, a fruit is the seed-bearing structure in flowering plants (also known as angiosperms) formed from the ovary after flowering. Fruits are the means by which ...
". Berries may be formed from one or more
carpels '' stigmas and style Gynoecium (, from Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided in ...

carpels
from the same flower (i.e. from a simple or a compound ovary). The
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seed
s are usually embedded in the fleshy interior of the ovary, but there are some non-fleshy exceptions, such as
peppers Pepper or peppers may refer to: Food and spice * Piperaceae or the pepper family, a large family of flowering plant ** Black pepper * ''Capsicum'' or pepper, a genus of flowering plants in the nightshade family Solanaceae ** Bell pepper ** Chili p ...

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 starch Starch or amylum is a polymeric A polymer (; Greek '' poly-'', "many" + '' -mer'', "part") is a substance Substance may refer to: * Substance (Jainism), a term in Jain ontology to denote the base or owner of attr ...

potato
and the
deadly nightshade ''Atropa belladonna'', commonly known as belladonna or deadly nightshade, is a poisonous perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is ...
, are
poison In , poisons are that can cause , injury or harm to , , , and usually by chemical reactions or other on the scales, when an organism is exposed to a sufficient quantity. In a metaphorical broader use of term it may refer to any thing deemed ...

poison
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 In botany, a fruit is the seed-bearing structure in flowering plants (also known as angiosperms) formed from the ovary after flowering. Fruits are the means by which angiosperms dissem ...

berry
" is any small edible fruit. Berries are usually juicy, round, brightly coloured,
sweet Sweetness is a Taste#Basic tastes, basic taste most commonly Perception, perceived when eating foods rich in sugars. Sweet tastes are generally regarded as pleasure, pleasurable, except when in excess. In addition to sugars like sucrose, many ot ...

sweet
or
sour The gustatory system or sense of taste is the sensory system The sensory nervous system is a part of the nervous system responsible for processing sensory information. A sensory system consists of sensory neurons (including the sensory re ...
, and do not have a stone or pit, although many small seeds may be present.


Botanical berries

In botany, botanical language, a berry is a simple fruit having
seed A seed is an embryonic ''Embryonic'' is the twelfth studio album by experimental rock band the Flaming Lips released on October 13, 2009, on Warner Bros. Records, Warner Bros. The band's first double album, it was released to generally positiv ...

seed
s and fleshy pulp (the
pericarp Fruit anatomy is the plant anatomy of the internal structure of fruit In botany, a fruit is the seed-bearing structure in flowering plants (also known as angiosperms) formed from the ovary after flowering. Fruits are the means by which ...
) produced from the ovary (plants), ovary of a single flower. The ovary can be inferior ovary, inferior or superior ovary, superior. It is Dehiscence (botany), 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 contains a single large seed surrounded by an imperceptible endocarp. Avocados are however also sometimes classified as drupes. * Banana * Berberis, Barberry (''Berberis''), Oregon-grape (''Mahonia aquifolium'') and Podophyllum peltatum, mayapple (''Podophyllum'' spp.) (Berberidaceae) * Arbutus unedo, Strawberry tree (''Arbutus unedo'') (not to be confused with the strawberry (''Fragaria''), which is an accessory fruit), bearberry (''Arctostaphylos'' spp.), bilberry, blueberry, cranberry, lingonberry/cowberry (''Vaccinium vitis-idaea''), crowberry (''Empetrum'' spp.) (family Ericaceae) * Coffea, coffee berries (Rubiaceae) (also described as drupes) "fruit a berry containing two (rarely one) seeds" * Gooseberry and Ribes, currant (''Ribes'' spp.; Grossulariaceae), red, black, and white types * Aubergine/Eggplant,
tomato The tomato is the edible berry A berry is a small, pulpy, and often edible fruit In botany, a fruit is the seed-bearing structure in flowering plants (also known as angiosperms) formed from the ovary after flowering. Fruits are ...

tomato
, goji berries (wolfberry) and other species of the family Solanaceae * Sambucus, Elderberry (''Sambucus niger''; Adoxaceae) * Indian gooseberry ''(Phyllanthus emblica)'' (Phyllanthaceae) * ''Garcinia gummi-gutta'', ''Garcinia mangostana'' (mangosteen) and ''Garcinia indica'' in the family Clusiaceae * Sapodilla (''Manilkara zapota''), Sapotaceae * Grape, ''Vitis vinifera'' in the family Vitaceae * Honeysuckle: the berries of ''some'' species are edible and are called Lonicera caerulea, honeyberries, but others are poisonous (''Lonicera'' spp.; Caprifoliaceae) * Persimmon (Ebenaceae) * Pumpkin,
cucumber Cucumber (''Cucumis sativus'') is a widely-cultivated creeping vine plant in the '' Cucurbitaceae'' gourd family that bears cucumiform fruits In botany Botany, also called , plant biology or phytology, is the science Scien ...

cucumber
and watermelon in the family Cucurbitaceae


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'' (baobab) amphisarca, which has a dry, more rigid and self-supporting skin. The fruit of citrus, such as the orange (fruit), orange, kumquat and lemon, is a berry with a thick rind and a very juicy interior divided into segments by Septum, septa, that is given the special name "hesperidium". A specialized term, Glossary of botanical terms#pepo, pepo, 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'' (passion fruit) and ''Carica'' (papaya) are sometimes also considered pepos. Berries that develop from an inferior ovary are sometimes termed Glossary of botanical terms#epigynous, 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
banana A banana is an elongated, edible fruit – botanically a berry (botany), berry – produced by several kinds of large herbaceous flowering plants in the genus ''Musa (genus), Musa''. In some countries, Cooking banana, bananas used for c ...

banana
s, Coffea arabica, coffee, 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'' gourd family that bears cucumiform fruits In botany Botany, also called , plant biology or phytology, is the science Scien ...

cucumber
s, melons and Squash (plant), squash).


Fruits not botanical berries

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'' (peaches, plums and Cherry, cherries), olives, coconut, bayberry and ''Persea'' 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, surrounded by a swollen hypanthium 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, 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'' 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 together at maturity to form the complete fruit. Examples of aggregate fruits commonly called "berries" include members of the genus ''Rubus'', such as blackberry and raspberry. Botanically, these are not berries. Other large aggregate fruits, such as soursop (''Annona muricata''), 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 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 aggregate of seed-like achenes is actually the "fruit", derived from an aggregate of ovaries, and the fleshy part develops from the Receptacle (botany), receptacle. * Mock strawberry, ''Duchesnea indica'' – structured just like a strawberry * Sea grape (''Coccoloba uvifera''; Polygonaceae) – the fruit is a dry capsule surrounded by fleshy calyx * Eastern teaberry (''Gaultheria procumbens'') – the fruit is a dry capsule surrounded by fleshy calyx


Berry-like conifer seed cones

The female seed Conifer cone, cones of some conifers have fleshy and merged scales, giving them a berry-like appearance. Juniper berry, Juniper "berries" (family Cupressaceae), in particular those of ''Juniperus communis'', are used to flavour gin. The seed cones of species in the families Podocarpaceae and Taxaceae 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 red aril partially enclosing the poisonous seed.


History of terminology

The Latin word ' or ' (plural ') was originally used for "any small round fruit". 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 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 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 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, 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 "Gynoecium#Placentation, 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. Fossils show that early flowering plants 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 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. Capsule (botany), 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, 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 prior to 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'' 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 pigments such as anthocyanins and other flavonoids. These pigments are localized mainly in the outer surface and the
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seed
s. Such pigments have antioxidant properties ''in vitro'', 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 spices are prepared from berries. Allspice is made from the dried berries of ''Pimenta dioica''. The fruits (berries) of different cultivars 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 ''Calabash, Lagenaria siceraria'', "bottle gourd", reflects its use as a liquid container. Some true berries have also been used as a source of dyes. In Hawaii, these included berries from a species of ''Dianella (plant), 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 domestication, 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 and Southeast Asia. Archaeology, Archaeological 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, citrus fruit remains unclear, although some recent research suggests a possible origin in Papuasia rather than continental southeast Asia. Chinese documents show that Mandarin orange, mandarins and pomelos were established in cultivation there by around 4,200 BP.


Commercial production

According to Food and Agriculture Organization Corporate Statistical Database, FAOSTAT 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.


See also

*List of culinary fruits *List of inedible fruits


Notes


References


External links

{{Authority control Berries, Fruit morphology