In botany, a berry is a fleshy fruit without a stone (pit) produced from a single flower containing one ovary. Berries so defined include grapes, currants, and tomatoes, as well as cucumbers, eggplants (aubergines), persimmons and bananas, but exclude certain fruits that meet the culinary definition of berries, such as strawberries 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 seeds are usually embedded in the fleshy interior of the ovary, but there are some non-fleshy exceptions, such as Capsicum species, with air rather than pulp around their seeds.
Many berries are edible, but others, such as the fruits of the potato and the deadly nightshade, are poisonous to humans.
A plant that bears berries is said to be bacciferous or baccate [lower-alpha 1] (from Latin bacca ).
In everyday English, a "berry" 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. [1]
In botanical language, a berry is a simple fruit having seeds and fleshy pulp (the pericarp) produced from the ovary of a single flower. The ovary can be inferior or superior. It is indehiscent, i.e. it does not have a special "line of weakness" along which it splits to release the seeds when ripe. [2] 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". [3]
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. [4]
Examples of botanical berries include:
"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. [11] The fruit of citrus, such as the orange, kumquat and lemon, is a berry with a thick rind and a very juicy interior divided into segments by septa, that is given the special name "hesperidium". [11] A specialized term, 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 septa. [11] The fruits of Passiflora (passion fruit) and Carica (papaya) are sometimes also considered pepos. [11]
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 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 include bananas, coffee, members of the genus Vaccinium (e.g., cranberries and blueberries), and members of the family Cucurbitaceae (gourds, cucumbers, melons and squash). [12]
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 the endocarp) surrounding the seed. Familiar examples include the stonefruits of the genus Prunus (peaches, plums and cherries), olives, coconut, dates, bayberry and Persea species. Some definitions make the mere presence of an internally differentiated endocarp the defining feature of a drupe; [11] others qualify the nature of the endocarp required in a drupe, e.g. defining berries to have endocarp less than 2 mm thick. [4] The term "drupaceous" is used of fruits that have the general structure and texture of a drupe, [13] 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. [14] Fruits of Coffea species are described as either drupes or berries. [9]
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. [15] Although pomes are not botanical berries, Amelanchier pomes become soft at maturity, resembling a blueberry, and are commonly called Juneberries, serviceberries or Saskatoon berries. [16]
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. [17] Examples of aggregate fruits commonly called "berries" include members of the genus Rubus, such as blackberry and raspberry. [18] Botanically, these are not berries. Other large aggregate fruits, such as soursop ( Annona muricata ), [19] are not usually called "berries", although some sources do use this term. [20]
Multiple fruits are not botanical berries. Multiple fruits are the fruits of two or more multiple flowers that are merged or packed closely together. [21] 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. [22]
Accessory fruits are not botanical berries. In accessory fruits, the edible part is not generated by the ovary. Berry-like examples include:
The female seed cones of some conifers have fleshy and merged scales, giving them a berry-like appearance. 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.[ citation needed ]
The Latin word baca or bacca (plural baccae) was originally used for "any small round fruit". [23] 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 semina or seeds. A fleshy fruit was called a pericarpium. For Caesalpinus, a true bacca or berry was a pericarpium derived from a flower with a superior ovary; one derived from a flower with an inferior ovary was called a pomum. [24]
In 1751, Carl Linnaeus wrote Philosophia Botanica , considered to be the first textbook of descriptive systematic botany. [25] He used eight different terms for fruits, one of which was bacca or berry, distinguished from other types of fruit such as drupa (drupe) and pomum (pome). [26] A bacca was defined as "pericarpium farctum evalve, semina ceteroquin nuda continens", meaning "unvalved solid pericarp, containing otherwise naked seeds". [27] The adjective "farctus" here has the sense of "solid with tissue softer than the outside; stuffed". [28] A berry or bacca 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. [27] Linnaeus' use of bacca and pomum was thus significantly different from that of Caesalpinus. Botanists continue to differ on how fruit should be classified. [26]
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 pepo for the berry-like fruits of cucurbits. [26] A pepo was distinguished by being a fleshy berry with the seeds distant from the axis, and so nearer the fruit wall [29] (i.e. by having "parietal placentation" in modern terminology). Nicaise Auguste Desvaux in 1813 used the terms hesperidium and amphisarca as further subdivisions of berries. [26] A hesperidium, called by others bacca 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. [30] "Hesperidium" remains in general use, but "amphisarca" is rarely used. [26]
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". [26]
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 66 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 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. [31]
Fruit type has been considered to be a useful character in classification and in understanding the phylogeny of plants. [32] 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. [33]
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. [32] 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. [31] Indeed, fruit type in general has proved to be an unreliable guide to flowering plant relationships. [32]
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:
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 seeds. [34] Such pigments have antioxidant properties in vitro , [35] 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. [36] [37]
Some spices are prepared from berries. Allspice is made from the dried berries of Pimenta dioica. [38] The fruits (berries) of different cultivars of Capsicum annuum are used to make paprika (mildly hot), chili pepper (hot) and cayenne pepper (very hot). [39]
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. [40]
Some true berries have also been used as a source of dyes. In Hawaii, these included berries from a species of Dianella , used to produce blue, and berries from black nightshade ( Solanum americanum ), used to produce green. [41]
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. [40] Peppers were domesticated in Mesoamerica by 8,000 BP. [42] [43] Many other early cultivated plants were also berries by the strict botanical definition, including grapes, domesticated by 8,000 BP [44] [45] and known to have been used in wine production by 6,000 BP. [46]
Bananas were first domesticated in Papua New Guinea and Southeast Asia. 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. [47] [48]
The history of cultivated citrus fruit remains unclear, although some recent research suggests a possible origin in Papuasia rather than continental southeast Asia. Chinese documents show that mandarins and pomelos were established in cultivation there by around 4,200 BP. [49]
According to 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). [Note 1]
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. [51] 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. [Note 2]
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". [52]
In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering.
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 of berries in the culinary sense are strawberries, raspberries, blueberries, blackberries, white currants, blackcurrants, and redcurrants. In Britain, soft fruit is a horticultural term for such fruits.
In botany, a drupe is a type of fruit in which an outer fleshy part surrounds a single shell of hardened endocarp with a seed (kernel) inside. Drupes do not split open to release the seed, i.e. they are indehiscent. These fruits usually develop from a single carpel, and mostly from flowers with superior ovaries.
Prunus is a genus of trees and shrubs in the flowering plant family Rosaceae that includes plums, cherries, peaches, nectarines, apricots, and almonds. The genus has a cosmopolitan distribution, being native to the North American temperate regions, the neotropics of South America, and temperate and tropical regions of Eurasia and Africa, There are about 340 accepted species as of March 2024. Many members of the genus are widely cultivated for their fruit and for decorative purposes. Prunus fruit are drupes, or stone fruits. The fleshy mesocarp surrounding the endocarp is edible while the endocarp itself forms a hard, inedible shell called the pyrena. This shell encloses the seed, which is edible in some species, but poisonous in many others. Besides being eaten off the hand, most Prunus fruit are also commonly used in processing, such as jam production, canning, drying, and the seeds for roasting.
Cucurbita is a genus of herbaceous fruits in the gourd family, Cucurbitaceae, native to the Andes and Mesoamerica. Five edible species are grown and consumed for their flesh and seeds. They are variously known as squash, pumpkin, or gourd, depending on species, variety, and local parlance. Other kinds of gourd, also called bottle-gourds, are native to Africa and belong to the genus Lagenaria, which is in the same family and subfamily as Cucurbita, but in a different tribe. These other gourds are used as utensils or vessels, and their young fruits are eaten much like those of the Cucurbita species.
A nut is a fruit consisting of a hard or tough nutshell protecting a kernel which is usually edible. In general usage and in a culinary sense, many dry seeds are called nuts. In a botanical context, "nut" implies that the shell does not open to release the seed (indehiscent).
In botany, a pome is a type of fruit produced by flowering plants in the subtribe Malinae of the family Rosaceae. Pome fruits consist of a central "core" containing multiple small seeds, which is enveloped by a tough membrane and surrounded by an edible layer of flesh. Pome fruit trees are deciduous, and undergo a dormant winter period that requires cold temperatures to break dormancy in spring. Well-known pomes include the apple, pear, and quince.
An aril, 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 seed to the ovary, an arillode forms from a different point on the seed coat. The term "aril" is sometimes applied to any fleshy appendage of the seed in flowering plants, such as the mace of the nutmeg seed. Arils and arillodes are often edible enticements that encourage animals to transport the seed, thereby assisting in seed dispersal. Pseudarils are aril-like structures commonly found on the pyrenes of Burseraceae species that develop from the mesocarp of the ovary. The fleshy, edible pericarp splits neatly in two halves, then falling away or being eaten to reveal a brightly coloured pseudaril around the black seed.
Sclerocarya birrea, commonly known as the marula, is a medium-sized deciduous fruit-bearing tree, indigenous to the miombo woodlands of Southern Africa, the Sudano-Sahelian range of West Africa, the savanna woodlands of East Africa and Madagascar.
Arctostaphylos uva-ursi is a plant species of the genus Arctostaphylos widely distributed across circumboreal regions of the subarctic Northern Hemisphere. Kinnikinnick is a common name in Canada and the United States. Growing up to 30 centimetres in height, the leaves are evergreen. The flowers are white to pink and the fruit is a red berry.
A pyrena or pyrene is the fruitstone within a drupe or drupelet produced by the ossification of the endocarp or lining of the fruit. It consists of a hard endocarp tissue surrounding one or more seeds. The hardened endocarp which constitutes the pyrene provides a protective physical barrier around the seed, shielding it from pathogens and herbivory.
Ferrocalamus, or iron bamboo, is a genus of Chinese bamboo in the grass family. endemic to China. The plant is known only from southern Yunnan, at elevations of 900 to 1,200 m above sea level.
In the flowering plants, an ovary is a part of the female reproductive organ of the flower or gynoecium. Specifically, it is the part of the pistil which holds the ovule(s) and is located above or below or at the point of connection with the base of the petals and sepals. The pistil may be made up of one carpel or of several fused carpels, and therefore the ovary can contain part of one carpel or parts of several fused carpels. Above the ovary is the style and the stigma, which is where the pollen lands and germinates to grow down through the style to the ovary, and, for each individual pollen grain, to fertilize one individual ovule. Some wind pollinated flowers have much reduced and modified ovaries.
A seedless fruit is a fruit developed to possess no mature seeds. Since eating seedless fruits is generally easier and more convenient, they are considered commercially valuable.
Peel, also known as rind or skin, is the outer protective layer of a fruit or vegetable which can be peeled off. The rind is usually the botanical exocarp, but the term exocarp also includes the hard cases of nuts, which are not named peels since they are not peeled off by hand or peeler, but rather shells because of their hardness.
Fruits are the mature ovary or ovaries of one or more flowers. They are found in three main anatomical categories: aggregate fruits, multiple fruits, and simple fruits.
Simple fruits are the result of the ripening-to-fruit of a simple or compound ovary in a single flower with a single pistil. In contrast, a single flower with numerous pistils typically produces an aggregate fruit; and the merging of several flowers, or a 'multiple' of flowers, results in a 'multiple' fruit. A simple fruit is further classified as either dry or fleshy.
Prunus rivularis, known variously by the common names creek plum, hog plum, or wild-goose plum is a thicket-forming shrub. It prefers calcareous clay soil or limestone-based woodland soils. This deciduous plant belongs to the rose family, Rosaceae, and is found mainly in the central United States. It is a shrub consisting of slender stems with umbel clusters of white blossoms. The fruit is a drupe that resembles a large berry; though it has a bitter taste, it serves as a source of food for birds and other wildlife. "Prunus" is Latin for plum, whereas "rivularis" means being near a stream.
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)