
Fruits are the mature
ovary
The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...
or ovaries of one or more
flower
Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s. They are found in three main anatomical categories:
aggregate fruits,
multiple fruit
Multiple 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 call ...
s, and
simple fruits.
Fruitlike structures may develop directly from the seed itself rather than the ovary, such as a fleshy
aril
An aril (), also called 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 ova ...
or
sarcotesta
The sarcotesta is a fleshy seedcoat, a type of testa. Examples of seeds with a sarcotesta are pomegranate, ginkgo
''Ginkgo'' is a genus of non-flowering seed plants, assigned to the gymnosperms. The scientific name is also used as the Eng ...
.
The
grain
A grain is a small, hard, dry fruit (caryopsis) – with or without an attached husk, 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 ...
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 a grass cultivated for its edible grain. Cereals are the world's largest crops, and are therefore staple foods. They include rice, wheat, rye, oats, barley, millet, and maize (Corn). Edible grains from other plant families, suc ...
s, such as
wheat
Wheat is a group of wild and crop domestication, domesticated Poaceae, grasses of the genus ''Triticum'' (). They are Agriculture, cultivated for their cereal grains, which are staple foods around the world. Well-known Taxonomy of wheat, whe ...
,
barley
Barley (), a member of the grass family, is a major cereal grain grown in temperate climates globally. It was one of the first cultivated grains; it was domesticated in the Fertile Crescent around 9000 BC, giving it nonshattering spikele ...
,
oats and
rice
Rice is a cereal grain and in its Domestication, domesticated form is the staple food of over half of the world's population, particularly in Asia and Africa. Rice is the seed of the grass species ''Oryza sativa'' (Asian rice)—or, much l ...
.
Categories of fruits
Fruits are found in three main anatomical categories:
aggregate fruits,
multiple fruit
Multiple 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 call ...
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 vegetation, 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 culti ...
.
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
A hesperidium (: hesperidia) is a modified berry with a tough, leathery rind, as in the oranges and lemons of the genus ''Citrus''.
Etymology
Carl Linnaeus gave the name Hesperideæ to an order containing the genus ''Citrus'', in allusion to ...
, which has a separable rind.
A
cucumber is an example of a pepo, while a
lemon
The lemon (''Citrus'' × ''limon'') is a species of small evergreen tree in the ''Citrus'' genus of the flowering plant family Rutaceae. A true lemon is a hybrid of the citron and the bitter orange. Its origins are uncertain, but some ...
is 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 a round, edible fruit produced by an apple tree (''Malus'' spp.). Fruit trees of the orchard or domestic apple (''Malus domestica''), the most widely grown in the genus, are agriculture, cultivated worldwide. The tree originated ...
is an example of a pome.
Lastly, drupes are known for being one-seeded with a fleshy mesocarp; an example of this is the
peach
The peach (''Prunus persica'') is a deciduous tree first domesticated and Agriculture, cultivated in China. It bears edible juicy fruits with various characteristics, most called peaches and the glossy-skinned, non-fuzzy varieties called necta ...
.
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 (biology), hybrid plant cultivated worldwide for its fruit. The genus ''Fragaria'', the strawberries, is in the rose family, Rosaceae. The fruit ...
. 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
An accessory fruit is a fruit that contains tissue derived from plant parts other than the Ovary (botany), ovary. In other words, the flesh of the fruit develops not from the floral ovary, but from some adjacent tissue exterior to the carpel (f ...
.
There is a shared method of seed dispersal within fleshy fruits. These fruits depend on animals to eat the fruits and disperse the seeds (
endozoochory) 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 vectors, ...
.
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
''Wisteria'' is a genus of flowering plants in the legume family, Fabaceae (Leguminosae). The genus includes four species of woody twining vines that are native to China, Japan, Korea, Vietnam, southern Canada, the Eastern United States, and nor ...
, 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 fruit, simple dry fruits, dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and Dehi ...
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
The sweet pea, ''Lathyrus odoratus'', is a flowering plant in the genus '' Lathyrus'' in the family Fabaceae ( legumes), native to Sicily, southern Italy and the Aegean Islands.
It is an annual climbing plant, growing to a height of , where ...
,
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. Soy is a staple crop, the world's most grown legume, and an important animal feed.
Soy is a key source o ...
,
alfalfa
Alfalfa () (''Medicago sativa''), also called lucerne, is a perennial plant, 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, ...
,
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.& ...
and
poppy
A poppy is a flowering plant in the subfamily Papaveroideae of the family Papaveraceae. Poppies are herbaceous plants, often grown for their colourful flowers. One species of poppy, '' Papaver somniferum'', is the source of the narcotic drug ...
.
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 species of large annual forb of the daisy family Asteraceae. The common sunflower is harvested for its edible oily seeds, which are often eaten as a snack food. They are also used in the pr ...
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
Rubiaceae () is a family (biology), 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 Petiole ( ...
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
Plant anatomy or phytotomy is the general term for the study of the internal Anatomy, structure of plants. Originally, it included plant morphology, the description of the physical form and external structure of plants, but since the mid-20th centu ...
of the internal structure of
fruit
In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering.
Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...
.
In berries and
drupe
In botany, a drupe (or stone fruit) is a type of fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pip'' (UK), ''pit'' (US), ''stone'', or ''pyrena'') of hardened endocarp with a seed ...
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 family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes.
''Citrus'' is nativ ...
and stone fruits (''
Prunus
''Prunus'' is a genus of flowering plant, flowering trees and shrubs from the family (biology), family Rosaceae. The genus includes plums, cherries, peaches, nectarines, apricots and almonds (collectively Drupe, stonefruit). The genus has a cosm ...
'') only some layers of the pericarp are eaten. In
accessory fruit
An accessory fruit is a fruit that contains tissue derived from plant parts other than the Ovary (botany), ovary. In other words, the flesh of the fruit develops not from the floral ovary, but from some adjacent tissue exterior to the carpel (f ...
s, other tissues develop into the edible portion of the fruit instead, for example the
receptacle of the flower in
strawberries.
Pericarp layers
In fleshy fruits, the pericarp is typically made up of three distinct layers: the outer epicarp, the middle mesocarp and the inner endocarp. These layers vary in thickness and texture, and may blend into each other. In a
hesperidium
A hesperidium (: hesperidia) is a modified berry with a tough, leathery rind, as in the oranges and lemons of the genus ''Citrus''.
Etymology
Carl Linnaeus gave the name Hesperideæ to an order containing the genus ''Citrus'', in allusion to ...
like
lemon
The lemon (''Citrus'' × ''limon'') is a species of small evergreen tree in the ''Citrus'' genus of the flowering plant family Rutaceae. A true lemon is a hybrid of the citron and the bitter orange. Its origins are uncertain, but some ...
, the epicarp and mesocarp make up the
peel; in many
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 fruit, stone or pit (fruit), pit although many wikt:pip#Etymology 2, pips or seeds may be p ...
like
melons or cucumbers (pepo), the mesocarp and endocarp make up the flesh.
In dry fruits, the layers of the pericarp are usually hard, dry and not clearly distinguishable.
Epicarp

Epicarp (from , "on" or "upon" + ''-carp'', "fruit") is a
botanical
Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
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 family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes.
''Citrus'' is nativ ...
, the flavedo (zest).
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 ...
s,
pigment
A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), chemically unreactive in water or another medium; in contrast, dyes are colored sub ...
s (
carotenoid
Carotenoids () are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, archaea, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, corn, tomatoes, cana ...
s,
chlorophyll
Chlorophyll 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 allows plants to absorb energy ...
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),
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 melting poi ...
es,
steroid
A steroid is an organic compound with four fused compound, fused rings (designated A, B, C, and D) arranged in a specific molecular configuration.
Steroids have two principal biological functions: as important components of cell membranes t ...
s and
triterpenoids,
fatty acid
In chemistry, in particular in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated and unsaturated compounds#Organic chemistry, saturated or unsaturated. Most naturally occurring fatty acids have an ...
s, bitter principles (
limonin), and
enzyme
An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
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 (: stomata, from Greek language, Greek ''στόμα'', "mouth"), also called a stomate (: stomates), is a pore found in the Epidermis (botany), epidermis of leaves, stems, and other organs, that controls the rate of gas exc ...
ta, which in many cases are closed when the fruit is ripe.
When ripe, the flavedo cells contain carotenoids (mostly
xanthophyll) inside
chromoplast
Chromoplasts are plastids, heterogeneous organelles responsible for pigment synthesis and storage in specific Photosynthesis, photosynthetic eukaryotes. It is thought (according to symbiogenesis) that like all other plastids including chloroplast ...
s, which, in a previous developmental stage, contained
chlorophyll
Chlorophyll 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 allows plants to absorb energy ...
. 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 ...
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
A hesperidium (: hesperidia) is a modified berry with a tough, leathery rind, as in the oranges and lemons of the genus ''Citrus''.
Etymology
Carl Linnaeus gave the name Hesperideæ to an order containing the genus ''Citrus'', in allusion to ...
, the mesocarp is the inner part of the
peel and is commonly removed before eating, as is found in
citrus fruit
''Citrus'' is a genus of flowering plant, flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as Orange (fruit), oranges, Mandarin orange, mandarins, lemons, grapefruits, ...
.
It is also referred to as ''albedo'' or
pith
Pith, or medulla, is a tissue in the stems of vascular plants. Pith is composed of soft, spongy parenchyma cells, which in some cases can store starch. In eudicotyledons, pith is located in the center of the stem. In monocotyledons, it ex ...
. 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 science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
term for the inside layer of the pericarp (or fruit), which directly surrounds the
seeds. It may be membranous as in
citrus
''Citrus'' is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes.
''Citrus'' is nativ ...
where it is the only part consumed, or thick and hard as in the
pyrenas of
drupe
In botany, a drupe (or stone fruit) is a type of fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pip'' (UK), ''pit'' (US), ''stone'', or ''pyrena'') of hardened endocarp with a seed ...
fruits of the family
Rosaceae
Rosaceae (), the rose family, is a family of flowering plants that includes 4,828 known species in 91 genera.
The name is derived from the type genus '' Rosa''. The family includes herbs, shrubs, and trees. Most species are deciduous, but som ...
such as
peach
The peach (''Prunus persica'') is a deciduous tree first domesticated and Agriculture, cultivated in China. It bears edible juicy fruits with various characteristics, most called peaches and the glossy-skinned, non-fuzzy varieties called necta ...
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 name ...
,
plum
A plum is a fruit of some species in Prunus subg. Prunus, ''Prunus'' subg. ''Prunus'.'' Dried plums are often called prunes, though in the United States they may be labeled as 'dried plums', especially during the 21st century.
Plums are ...
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, it is the stony layer that surrounds the kernel of
pecans,
walnut
A walnut is the edible seed of any tree of the genus '' Juglans'' (family Juglandaceae), particularly the Persian or English walnut, '' Juglans regia''. They are accessory fruit because the outer covering of the fruit is technically an i ...
s, 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
Juice is a drink made from the extraction or Cold-pressed juice, pressing of the natural liquid contained in fruit and vegetables. It can also refer to liquids that are flavored with concentrate or other biological food sources, such as meat ...
of the fruit.
Anatomy of grass fruits
The grains of
grass
Poaceae ( ), also called Gramineae ( ), is a large and nearly ubiquitous family (biology), family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos, the grasses of natural grassland and spe ...
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 seeds.
See also
*
Gynoecium
Gynoecium (; ; : gynoecia) 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 (botany), whorl of a flower; it consists ...
References
External links
*
{{Fruits
Fruit morphology