Endocarp
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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 flowers. They are found in three main anatomical categories: aggregate fruits,
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 called a ...
s, and simple fruits. 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 or pomegranate, and the pineapple 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 grains, such as wheat, barley, oats and rice.


Categories of fruits

Fruits are found in three main anatomical categories: aggregate fruits,
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 called a ...
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. 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 Vine#Horticultural climbing plants, creeping vine plant in the Cucurbitaceae family that bears usually cylindrical Fruit, fruits, which are used as culinary vegetables.
and a lemon 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 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. 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. 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 ope ...
s, capsules, follicles or
nuts Nut often refers to: * Nut (fruit), fruit composed of a hard shell and a seed, or a collective noun for dry and edible fruits or seeds * Nut (hardware), fastener used with a bolt Nut or Nuts may also refer to: Arts, entertainment, and media Com ...
. 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,
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 w ...
, milkweed, 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.&nb ...
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), as ...
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 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

Fruit anatomy is the plant anatomy of the internal structure of fruit. 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 and stone fruits ('' Prunus'') 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 Receptacle may refer to: Biology * Receptacle (botany), a plant anatomical part * Seminal receptacle, a sperm storage site in some insects Electrical engineering * Automobile auxiliary power outlet, formerly known as ''cigarette lighter recep ...
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 Peel or Peeling may refer to: Places Australia * Peel (Western Australia) * Peel Island, Queensland *Peel, New South Wales * Peel River (New South Wales) Canada * Peel Parish, New Brunswick * Peel, New Brunswick, an unincorporated communi ...
. 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 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, the flavedo.


Flavedo

Flavedo is mostly composed of
cellulosic material Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
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 waxes,
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 a ...
s and triterpenoids, fatty acids, pigments (
carotenoid Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic compound, organic pigments that are produced by plants and algae, as well as several bacteria, and Fungus, fungi. Carotenoids give the characteristic color to pumpki ...
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 a ...
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 enzymes. 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 bor ...
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 a ...
. 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 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, the mesocarp is also referred to as ''albedo'' or pith. It is the inner part of the
peel Peel or Peeling may refer to: Places Australia * Peel (Western Australia) * Peel Island, Queensland *Peel, New South Wales * Peel River (New South Wales) Canada * Peel Parish, New Brunswick * Peel, New Brunswick, an unincorporated communi ...
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 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 pl ...
. It may be membranous as in citrus 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 (also ...
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), ''Sorbus ...
such as peaches,
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 nam ...
,
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 i ...
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 also ...
s. In
nuts Nut often refers to: * Nut (fruit), fruit composed of a hard shell and a seed, or a collective noun for dry and edible fruits or seeds * Nut (hardware), fastener used with a bolt Nut or Nuts may also refer to: Arts, entertainment, and media Com ...
, it is the stony layer that surrounds the kernel of pecans, walnuts, etc., and that is removed before consumption. In citrus fruits, 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 grasses 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


References


External links

* {{Fruits Fruit morphology