botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...

a capsule is a type of simple, dry, though rarely fleshy dehiscent fruit produced by many species of angiosperms (

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s).

Origins and structure

The capsule (

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

: ''capsula'', small box) is derived from a compound (multicarpeled) ovary. A capsule is a structure composed of two or more

carpel 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) '' pistils' ...

s. In (flowering plants), the term locule (or cell) is used to refer to a chamber within the fruit. Depending on the number of locules in the ovary, fruit can be classified as uni-locular (unilocular), bi-locular, tri-locular or multi-locular. The number of locules present in a gynoecium may be equal to or less than the number of carpels. The locules contain the ovules or

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 angiospe ...

s and are separated by septa.

Dehiscence

In most cases the capsule is dehiscent, i.e. at maturity, it splits apart (dehisces) to release the

s within. A few capsules are indehiscent, for example those of '' Adansonia digitata'', '' Alphitonia'', and '' Merciera''. Capsules are often classified into four types, depending on the type and location of dehiscence (see Simpson Fig 9.41 and Hickey & King ). Lilium candidum MHNT

Loculicidal capsules possess longitudinal lines of dehiscence radially aligned with the

locules A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usu ...

, ''i.e.'' not at the septa, along the midrib or dorsal suture (seam) of the locules. If septa are absent, the dehiscence lines lie between the placentae. This type is common among many members of the Liliaceae such as

Lilium ''Lilium'' () is a genus of herbaceous flowering plants growing from bulbs, all with large prominent flowers. They are the true lilies. Lilies are a group of flowering plants which are important in culture and literature in much of the world. M ...

(see illustration). Septicidal capsules have dehiscence lines aligned with the sutures of the ovary septa or placentae, that is between the carpels. Both loculocidal and septicidal capsules split into distinguishable segments called valves. The valves are a part of the

pericarp Fruit anatomy is the plant anatomy of the internal structure of fruit. 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. Aggr ...

(fruit wall) that has split away, without enclosing the seed or seeds. The borders of the valves may or may not coincide with the borders of carpels. These valves may remain attached to the fruit or fall off. In septicidal capsules the valves remain in place. In some capsules, the split occurs between carpels, and in others each carpel splits open. Circumscissile capsules (pyxide, pyxis, pyxidium or lid capsule) have a transverse, rather than longitudinal, dehiscence line, so that the upper part of the capsule dehisces, usually forming a terminal lid ( operculum) that opens (see illustration). An example is ''

Plantago ''Plantago'' is a genus of about 200 species of flowering plants in the family Plantaginaceae, commonly called plantains or fleaworts. The common name plantain is shared with the unrelated cooking plantain. Most are herbaceous plants, though ...

''. A variant is the Septifragal capsule (valvular capsule) in which the outer walls break away from the septa and (usually axile) placentae as valves. Poricidal capsules dehisce through pores (openings) in the capsule, as in ''

Papaver ''Papaver'' is a genus of 70–100 species of frost-tolerant annuals, biennials, and perennials native to temperate and cold regions of Eurasia, Africa and North America. It is the type genus of the poppy family, Papaveraceae. Description The ...

'', the seeds escaping through these pores. Spathoglottis flwrs

Examples of other plants that produce capsules include

nigella ''Nigella'' is a genus of 18 species of annual plants in the family Ranunculaceae, native to Southern Europe, North Africa, South Asia, Southwest Asia and Middle East. Common names applied to members of this genus are nigella, devil-in-a-bush o ...

, orchid, willow,

cotton Cotton is a soft, fluffy staple fiber that grows in a boll, or protective case, around the seeds of the cotton plants of the genus '' Gossypium'' in the mallow family Malvaceae. The fiber is almost pure cellulose, and can contain minor pe ...

, and jimson weed.

Specialised capsules

Some dry dehiscent fruits form specialised capsule-like structures. A follicle is derived from a single carpel that splits along a suture, as in '' Magnolia'', while a legume splits along two sutures, and are a defining feature of the Fabaceae. Some variants of legumes that have retained vestigial sutures include loments that split transversely into segments, each with a single seed, and indehiscent legumes, such as ''

Arachis hypogaea The peanut (''Arachis hypogaea''), also known as the groundnut, goober (US), pindar (US) or monkey nut (UK), is a legume crop grown mainly for its edible seeds. It is widely grown in the tropics and subtropics, important to both small and la ...

'' (peanuts). Capsules derived from two carpels include silicles and siliques that dehisce along two suture lines but retain a partition called the replum, which is a septum with attached seeds. While both are characteristic of

Brassicaceae Brassicaceae () or (the older) Cruciferae () is a medium-sized and economically important family of flowering plants commonly known as the mustards, the crucifers, or the cabbage family. Most are herbaceous plants, while some are shrubs. The leav ...

, silicles have a ratio of length to width of no more than 3:1. A

schizocarp A schizocarp is a dry fruit that, when mature, splits up into mericarps. There are different definitions: * Any dry fruit composed of multiple carpels that separate. : Under this definition the mericarps can contain one or more seeds (the m ...

is derived from a compound ovary with two or more locules which then separate radially as one of the above types, such as a schizocarp of follicles, as in '' Asclepias'' (

Asclepiadoideae The Asclepiadoideae are a subfamily of plants in the family Apocynaceae. Formerly, they were treated as a separate family under the name Asclepiadaceae, e.g. by APG II, and known as the milkweed family. They form a group of perennial herbs, twi ...

) (see illustration). A mericarp is a portion of the fruit that separates from the ovary to form a distinct locule unit which encloses the seed, usually-nut-like, as in

Apiaceae Apiaceae or Umbelliferae is a family of mostly aromatic flowering plants named after the type genus '' Apium'' and commonly known as the celery, carrot or parsley family, or simply as umbellifers. It is the 16th-largest family of flowering plan ...

in which the mericarps are joined by a stalk (carpophore). Thus a schizocarp of mericarps is a structure in which the carpels of a single ovary split to form mericarps. A schizocarp of nutlets is derived from a carpel that becomes lobed, and the lobes become nutlets that split apart. Examples include Boraginaceae and most Lamiaceae, where the styles are attached between the ovary lobes.

Nuts

Capsules are sometimes mislabeled as nuts, as in the example of the Brazil nut or the Horse-chestnut. A capsule is not a nut because it releases its seeds and it splits apart. Nuts, on the other hand, do not release seeds as they are a compound ovary containing both a single seed and the fruit. Nuts also do not split. In the Brazil nut, a lid on the capsule opens, but is too small to release the dozen or so seeds (the actual "Brazil nut" of commerce) within. These germinate inside the capsule after it falls to the ground.

References

Bibliography