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Fruit anatomy is the
plant anatomy Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. Originally it included plant morphology, the description of the physical form and external structure of plants, but since the mid-20th century plan ...
of the internal structure of
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 ...
. Fruits are the mature
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. ...
or ovaries of one or more
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 ...
s. They are found in three main anatomical categories:
aggregate fruit A raspberry fruit (shown with a raspberry beetle larva) is an aggregate fruit, an aggregate of drupe">raspberry_beetle.html" ;"title="raspberry fruit (shown with a raspberry beetle">raspberry fruit (shown with a raspberry beetle larva) is an aggre ...
s,
multiple fruit Multi-fruits, also called collective fruits, are fruiting bodies formed from a cluster of flowers, the '' inflorescence''. Each flower in the inflorescence produces a fruit, but these mature into a single mass. After flowering the mass is calle ...
s, and simple fruits. Aggregate fruits are formed from a single compound flower and contain many ovaries or fruitlets. Examples include raspberries and blackberries. Multiple fruits are formed from the fused ovaries of multiple flowers or inflorescence. Examples include
fig The fig is the edible fruit of ''Ficus carica'', a species of small tree in the flowering plant family Moraceae. Native to the Mediterranean and western Asia, it has been cultivated since ancient times and is now widely grown throughout the world ...
,
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 ...
, and
pineapple The pineapple (''Ananas comosus'') is a tropical plant with an edible fruit; it is the most economically significant plant in the family Bromeliaceae. The pineapple is indigenous to South America, where it has been cultivated for many centuri ...
. Simple fruits are formed from a single ovary and may contain one or many
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. They can be either fleshy or dry. In fleshy fruit, during development, the pericarp (ovary wall) and other accessory structures become the fleshy portion of the fruit. The types of fleshy fruits are
berries 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, ras ...
,
pome In botany, a pome is a type of fruit produced by flowering plants in the subtribe Malinae of the family Rosaceae. Well-known pomes include the apple, pear, and quince. Etymology The word ''pome'' entered English in the late 14th century, and re ...
s, and
drupe In botany, a drupe (or stone fruit) is an indehiscent fruit in which an outer fleshy part ( exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pit'', ''stone'', or ''pyrena'') of hardened endocarp with a seed (''kernel ...
s. In some fruits, the edible portion is not derived from the ovary, but rather from the
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 ...
, such as the
mangosteen Mangosteen (''Garcinia mangostana''), also known as the purple mangosteen, is a tropical evergreen tree with edible fruit native to tropical lands surrounding the Indian Ocean. Its origin is uncertain due to widespread prehistoric cultivation. ...
or pomegranate, and the
pineapple The pineapple (''Ananas comosus'') is a tropical plant with an edible fruit; it is the most economically significant plant in the family Bromeliaceae. The pineapple is indigenous to South America, where it has been cultivated for many centuri ...
from which tissues of the flower and
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
provide food. The
grain A grain is a small, hard, dry fruit ( caryopsis) – with or without an attached hull layer – harvested for human or animal consumption. A grain crop is a grain-producing plant. The two main types of commercial grain crops are cereals and legum ...
s of grasses are single-seed simple fruits wherein the pericarp and seed coat are fused into one layer. This type of fruit is called a caryopsis. Examples include
cereal grain A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more food ...
s, such as
wheat Wheat is a grass widely cultivated for its seed, a cereal grain that is a worldwide staple food. The many species of wheat together make up the genus ''Triticum'' ; the most widely grown is common wheat (''T. aestivum''). The archaeologi ...
,
barley Barley (''Hordeum vulgare''), a member of the grass family, is a major cereal grain grown in temperate climates globally. It was one of the first cultivated grains, particularly in Eurasia as early as 10,000 years ago. Globally 70% of barley p ...
, oats and
rice Rice is the seed of the grass species '' Oryza sativa'' (Asian rice) or less commonly ''Oryza glaberrima'' (African rice). The name wild rice is usually used for species of the genera '' Zizania'' and '' Porteresia'', both wild and domesticat ...
.


Categories of fruits

Fruits are found in three main anatomical categories:
aggregate fruit A raspberry fruit (shown with a raspberry beetle larva) is an aggregate fruit, an aggregate of drupe">raspberry_beetle.html" ;"title="raspberry fruit (shown with a raspberry beetle">raspberry fruit (shown with a raspberry beetle larva) is an aggre ...
s,
multiple fruit Multi-fruits, also called collective fruits, are fruiting bodies formed from a cluster of flowers, the '' inflorescence''. Each flower in the inflorescence produces a fruit, but these mature into a single mass. After flowering the mass is calle ...
s, and simple fruits. Aggregate fruits are formed from a single compound flower and contain many ovaries or fruitlets. Examples include raspberries and blackberries. Multiple fruits are formed from the fused ovaries of multiple flowers or inflorescence. An example of multiple fruits are the fig, mulberry, and the
pineapple The pineapple (''Ananas comosus'') is a tropical plant with an edible fruit; it is the most economically significant plant in the family Bromeliaceae. The pineapple is indigenous to South America, where it has been cultivated for many centuri ...
. Simple fruits are formed from a single ovary and may contain one or many seeds. They can be either fleshy or dry. In fleshy fruit, during development, the pericarp and other accessory structures become the fleshy portion of the fruit. The types of fleshy fruits are berries, pomes, and drupes. In berries, the entire pericarp is fleshy but this excludes the exocarp which acts as more as a skin. There are berries that are known as pepo, a type of berry with an inseparable rind, or hesperidium, which has a separable rind. An example of a pepo is the
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.lemon The lemon (''Citrus limon'') is a species of small evergreen trees in the flowering plant family Rutaceae, native to Asia, primarily Northeast India (Assam), Northern Myanmar or China. The tree's ellipsoidal yellow fruit is used for culin ...
would be an example of a hesperidium. The fleshy portion of the pomes is developed from the floral tube and like the berry most of the pericarp is fleshy but the endocarp is cartilaginous; an
apple An apple is an edible fruit produced by an apple tree (''Malus domestica''). Apple trees are cultivated worldwide and are the most widely grown species in the genus '' Malus''. The tree originated in Central Asia, where its wild ancest ...
is an example of a pome. Lastly, drupes are known for being one-seeded with a fleshy mesocarp; an example of this would be 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 ...
. However, there are fruits where the fleshy portion is developed from tissues that are not the ovary, such as in 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 ...
. The edible part of the strawberry is formed from the receptacle of the flower. Due to this difference the strawberry is known as a false fruit or an accessory fruit. There is a shared method of seed dispersal within fleshy fruits. These fruits depend on animals to eat the fruits and disperse the seeds in order for their populations to survive. Dry fruits also develop from the ovary, but unlike the fleshy fruits they do not depend on the mesocarp but the endocarp for
seed dispersal In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vector ...
. Dry fruits depend more on physical forces, like wind and water. Dry fruits' seeds can also perform pod shattering, which involve the seed being ejected from the seed coat by shattering it. Some dry fruits are able to perform seed pod explosions, such as wisteria, resulting the seed to be dispersed over long distances. Like fleshy fruits, dry fruits can also depend on animals to spread their seeds by adhering to animal's fur and skin, this is known as epizoochory. Types of dry fruits include
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 ...
s, capsules, follicles or nuts. Dry fruits can also be separated into dehiscent and indehiscent fruits. Dry dehiscent fruits are described as a fruit where the pod has an increase in internal tension to allow seeds to be released. These include the sweet pea,
soybean The soybean, soy bean, or soya bean (''Glycine max'') is a species of legume native to East Asia, widely grown for its edible bean, which has numerous uses. Traditional unfermented food uses of soybeans include soy milk, from which tofu ...
,
alfalfa Alfalfa () (''Medicago sativa''), also called lucerne, is a perennial flowering plant in the legume family Fabaceae. It is cultivated as an important forage crop in many countries around the world. It is used for grazing, hay, and silage, as ...
,
milkweed ''Asclepias'' is a genus of herbaceous, perennial, flowering plants known as milkweeds, named for their latex, a milky substance containing cardiac glycosides termed cardenolides, exuded where cells are damaged. Most species are toxic to hum ...
, mustard,
cabbage Cabbage, comprising several cultivars of ''Brassica oleracea'', is a leafy green, red (purple), or white (pale green) biennial plant grown as an annual vegetable crop for its dense-leaved heads. It is descended from the wild cabbage ( ''B.&n ...
and poppy. Dry indehiscent fruit differ in that they do not have this mechanism and simply depend on physical forces. Examples of species indehiscent fruit are
sunflower The common sunflower (''Helianthus annuus'') is a large annual forb of the genus ''Helianthus'' grown as a crop for its edible oily seeds. Apart from cooking oil production, it is also used as livestock forage (as a meal or a silage plant), ...
seeds, nuts, and dandelions.


Evolutionary history

There is a wide variety in the structures of fruit across the different species of plants. Evolution has selected for certain traits in plants that would increase their fitness. This diversity arose through the selection of advantageous methods for seed protection and dispersal in different environments. It is known that dry fruits were present before fleshy fruits and fleshy fruits diverged from them. A study looking at the family
Rubiaceae The Rubiaceae are a family of flowering plants, commonly known as the coffee, madder, or bedstraw family. It consists of terrestrial trees, shrubs, lianas, or herbs that are recognizable by simple, opposite leaves with interpetiolar stipules a ...
found that within the family, fleshy fruits had evolved independently at least 12 times. This means that fleshy fruits were not passed on to following generations but that this form of fruit was selected for in different species. This may imply that fleshy fruit is a favorable and beneficial trait because not only does it disperse the seeds, but it also protects them. There is also a variety of dispersal methods that are used by different plants. The origins of these modes of dispersal have been found to be a more recent evolutionary change. Of the methods of dispersal, the plants that use animals have not changed in many ways from the original trait. Due to this, it may be assumed that animal dispersal is an efficient form of dispersal, however there has been no evidence that it increases dispersal distances. Therefore, the question remains of what evolutionary mechanism causes such dramatic diversity. It has been found, however, that simple changes within developmental regulatory genes can cause large alterations within the anatomical structure of the fruit. Even without knowing the mechanism involved in the biodiversity of fruit, it is clear that this diversity is important to the continuation of plant populations.


Anatomy of simple fruits

In berries and
drupe In botany, a drupe (or stone fruit) is an indehiscent fruit in which an outer fleshy part ( exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pit'', ''stone'', or ''pyrena'') of hardened endocarp with a seed (''kernel ...
s, the pericarp forms the edible tissue around the seeds. In other fruits such as
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 ...
and stone fruits (
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 ...
) only some layers of the pericarp are eaten. In accessory fruits, other tissues develop into the edible portion of the fruit instead, for example the receptacle of the flower in
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 ...
.


Pericarp layers

In fleshy fruits, the pericarp is typically made up of three distinct layers: the epicarp (also known as exocarp), which is the outermost layer; the mesocarp, which is the middle layer; and the endocarp, which is the inner layer surrounding the ovary or the seeds. In a citrus fruit, the epicarp and mesocarp make up the peel. In dry fruits, the layers of the pericarp are not clearly distinguishable


Epicarp

Epicarp (from el, epi-, "on" or "upon" + ''-carp'', "fruit") is a
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 ...
term for the outermost layer of the pericarp (or fruit). The epicarp forms the tough outer skin of the fruit, if there is one. The epicarp is sometimes called the exocarp, or, especially in
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 ...
, the flavedo.


Flavedo

Flavedo is mostly composed of cellulosic material but also contains other components, such as
essential oil An essential oil is a concentrated hydrophobic liquid containing volatile (easily evaporated at normal temperatures) chemical compounds from plants. Essential oils are also known as volatile oils, ethereal oils, aetheroleum, or simply as the o ...
s,
paraffin wax Paraffin wax (or petroleum wax) is a soft colorless solid derived from petroleum, coal, or oil shale that consists of a mixture of hydrocarbon molecules containing between 20 and 40 carbon atoms. It is solid at room temperature and begins t ...
es,
steroid A steroid is a biologically active organic compound with four rings arranged in a specific molecular configuration. Steroids have two principal biological functions: as important components of cell membranes that alter membrane fluidity; and ...
s and
triterpenoid Triterpenes are a class of chemical compounds composed of three terpene units with the molecular formula C30H48; they may also be thought of as consisting of six isoprene units. Animals, plants and fungi all produce triterpenes, including squal ...
s,
fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, f ...
s,
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...
s (
carotenoid Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, ...
s,
chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to ...
s,
flavonoid Flavonoids (or bioflavonoids; from the Latin word ''flavus'', meaning yellow, their color in nature) are a class of polyphenolic secondary metabolites found in plants, and thus commonly consumed in the diets of humans. Chemically, flavonoids ...
s), bitter principles ( limonin), and
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s. In citrus fruits, the flavedo constitutes the peripheral surface of the pericarp. It is composed of several cell layers that become progressively thicker in the internal part; the epidermic layer is covered with wax and contains few
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bo ...
ta, which in many cases are closed when the fruit is ripe. When ripe, the flavedo cells contain carotenoids (mostly xanthophyll) inside chromoplasts, which, in a previous developmental stage, contained
chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to ...
. This hormonally controlled progression in development is responsible for the fruit's change of color from green to yellow upon ripening. Citrus flavedo may be scraped off the fruit to create zest. The internal region of the flavedo is rich in multicellular bodies with spherical or
pyriform Piriform, sometimes ''pyriform'', means pear-shaped (from Latin ''pirum'' "pear" and ''forma'' "shape"). It may also refer to: Going pear-shaped * Going wrong or going pear-shaped Anatomy * Piriform aperture, more commonly known as anterior ...
shapes, which are full of
essential oil An essential oil is a concentrated hydrophobic liquid containing volatile (easily evaporated at normal temperatures) chemical compounds from plants. Essential oils are also known as volatile oils, ethereal oils, aetheroleum, or simply as the o ...
s.


Mesocarp

The mesocarp (from Greek: ''meso-'', "middle" + ''-carp'', "fruit") is the fleshy middle layer of the pericarp of a fruit; it is found between the epicarp and the endocarp. It is usually the part of the fruit that is eaten. For example, the mesocarp makes up most of the edible part of a peach, and a considerable part of a tomato. "Mesocarp" may also refer to any fruit that is fleshy throughout. In a hesperidium, as is found in
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 ...
, the mesocarp is also referred to as ''albedo'' or pith. It is the inner part of the peel and is commonly removed before eating. In citron fruit, where the mesocarp is the most prominent part, it is used to produce succade.


Endocarp

Endocarp (from Greek: ''endo-'', "inside" + ''-carp'', "fruit") is a
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 ...
term for the inside layer of the pericarp (or fruit), which directly surrounds the
seeds 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 angiosperm ...
. It may be membranous as in
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 ...
where it is the only part consumed, or thick and hard as in the
pyrena A pyrena or pyrene (commonly called a "pit" or "stone") 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 (al ...
s of
drupe In botany, a drupe (or stone fruit) is an indehiscent fruit in which an outer fleshy part ( exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pit'', ''stone'', or ''pyrena'') of hardened endocarp with a seed (''kernel ...
fruits of the 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
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,
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 ...
,
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
apricot An apricot (, ) is a fruit, or the tree that bears the fruit, of several species in the genus '' Prunus''. Usually, an apricot is from the species '' P. armeniaca'', but the fruits of the other species in ''Prunus'' sect. ''Armeniaca'' are al ...
s. In nuts, it is the stony layer that surrounds the kernel of pecans, walnuts, etc., and that is removed before consumption. In
citrus fruits ''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 nativ ...
, the endocarp is separated into sections, which are called ''segments''. These segments are filled with juice vesicles, which contain the juice of the fruit.


Anatomy of grass fruits

The grains of
grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns a ...
es are single-seed simple fruits wherein the pericarp (ovary wall) and seed coat are fused into one layer. This type of fruit is called a caryopsis. Examples include cereal grains, such as wheat, barley, and rice. The dead pericarp of dry fruits represents an elaborated layer that is capable of storing active proteins and other substances for increasing survival rate of
germinating Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fern ...
seeds.


See also

*
Gynoecium Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) '' pist ...


References


External links

* {{Fruits Fruit morphology