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Amaryllis style and stigmas

The gynoecium (from Ancient Greek γυνή, gyne, meaning woman, and οἶκος, oikos, meaning house) is a term with several meanings in botanical usage. In reference to mosses; liverworts; and hornworts, gynoecium refers to a cluster of archegonia and any associated modified leaves or stems present on a gametophyte shoot. The more common use of gynoecium, however, is to refer to the ovule-producing part of a flower. The gynoecium in angiosperms is typically the innermost whorl of structures in a flower and is surrounded (in perfect flowers) by the androecium (stamens) and (in complete flowers) by the perianth (petals and sepals). The gynoecium is often inaccurately referred to as female because it gives rise to female (egg-producing) gametophytes.[1]

The structure from which the gynoecium is fashioned is the carpel. A gynoecium may consist of a single carpel, multiple distinct (unfused) carpels or multiple connate (fused) carpels. Flowers that bear a gynoecium but no androecium are called carpellate. Flowers lacking a gynoecium are called staminate flowers.

Gynoecia give rise to and protect ovules (within one or more ovaries). When mature, gynoecia may function to attract pollinators (through nectaries or visual cues). Gynoecia receive pollen on their surface (usually on a stigma), in some cases actively selecting genetically different pollen so as to promote outcrossing. Gynoecia may facilitate pollen tube growth to the ovule and delivery of the sperm to the egg. The gynoecium forms the pericarp of fruit.

Contents

Types of gynoecia

Center of a Ranunculus repens (Creeping Buttercup) showing multiple unfused carpels surrounded by longer stamens.
Center of a Tulipa aucheriana (Tulip) showing multiple connate carpels (a compound pistil) surrounded by stamens.

Carpels are the building blocks of the gynoecium. If a gynoecium has a single carpel, it is called monocarpous. If a gynoecium has multiple, distinct (free, unfused) carpels, it is apocarpous. If a gynoecium has multiple carpels fused into a single structure, it is syncarpous. A syncarpous gynoecium can sometimes appear very much like a monocarpous gynoecium. Because of this difficulty in interpreting carpel number and arrangement, some people use another term to describe the visible structure(s) in a gynoecium. This term is pistil (from Latin pistillum meaning pestle). The existence of two different ways of describing gynoecia has led to much confusion, even among botanists. The pistil terminology is less exact and specific, and is more cumbersome, but can be useful in keys and field descriptions where detailed studies are not possible.

Comparison of gynoecium terminology using carpel and pistil.
Gynoecium composition Carpel terminology Pistil terminology Examples
Single carpel Monocarpous, or unicarpellate gynoecium A pistil (simple) Avocado (Persea sp.), most legumes (Fabaceae)
Multiple distinct (unfused) carpels Apocarpous gynoecium Pistils (simple) Strawberry (Fragaria sp.), buttercup (Ranunculus sp.)
Multiple connate

(fused) carpels

Syncarpous gynoecium A pistil (compound) Tulip (Tulipa sp.), most flowers

Gynoecium development and arrangement is important in systematic research and identification of angiosperms, but can be the most challenging of the floral parts to interpret.

Carpel morphology

A monocarpous (single carpel) gynoecium in context. The gynoecium/carpel is made up of the ovary, style, and stigma in the center of the depicted flower.

Each structure in a gynoecium, whether it consists of a lone carpel or multiple fused carpels, usually consists of:

  • An enlarged basal portion called the ovary (from Latin ovum meaning egg), which contains placentas bearing one or more ovules (integumented megasporangia). A chamber inside an ovary, in which the ovules develop, is called a locule (or sometimes cell). No matter how many carpels fuse together to form a syncarpous ovary, there may be a single locule or more than one locule. If there are multiple locules the walls dividing them are called septa.
  • The style (from Latin stilus meaning stake, or stylus), a stalk-like conduit allowing the pollen (male gametophytes) to grow to the egg cell and deliver the sperm.
  • The stigma (from Ancient Greek στίγμα, stigma meaning mark, or puncture), usually found at the tip of the style, the portion of the structure that receives the pollen (male gametophytes). It is commonly sticky or feathery to capture pollen.

Carpels begin as small primorida on a floral apical meristem, forming later than and closer to the shoot apex than sepal, petal and stamen primordia. Morphological[2] and molecular studies of carpel ontogeny reveal that carpels are most likely homologous to leaves. In some basal angiosperm lineages, for example Degeneria, native to Fiji, carpels begin as a shallow cup and eventually form a folded, leaf-like structure, sealed at its margins by interlocking hairs. No stigma or style exists, but a broad stigmatic crest allows pollen tubes access to two rows of ovules enclosed in the carpel. Carpels, like leaves, generally bear three vascular traces

In a fully syncarpous gynoecium, the challenge is to determine how many carpels fused to form the gynoecium. Sometimes, the number of stigmas or stigma lobes is useful, as can be ridges on the ovary or locules within the ovary.

The degree of connation (fusion) in a syncarpous gynoecium can vary. Fusion can occur very early in carpel development, or later. In some gynoecia carpels are fused only at their base, but retain separate styles and stigmas. In other gynoecia, the ovaries and styles are fused but the stigmas remain separate. Sometimes (e.g., Apocynaceae) carpels are fused by their styles or stigmas but possess distinct ovaries.

In flowers with multiple distinct carpels, the carpels are nearly always single (apocarpous gynoecium).

Carpel position

Flowers and fruit (capsules) of the ground orchid, Spathoglottis plicata, illustrating an inferior ovary.

Basal angiosperm groups tend to have carpels arranged spirally around a conical or dome-shaped receptacle. In later diverging lineages, carpels tend to be in whorls.

The relationship of the other flower parts to the gynoecium can be an important systematic and taxonomic character. The ovary of the gynoecium can be above the attachment of petals and sepals to the receptacle. In this case the ovary is called superior. This corresponds to a the term hypogynous, used to describe a flower with a fully superior ovary.

In some flowers with superior ovaries, (examples include many members of the rose family, Rosaceae) the base of the stamens, petals and sepals are fused into a floral tube or hypanthium. In these flowers, the ovary can be free of or partially adnate (fused) to the hypanthium, but remains at least partially superior. These flower are called perigynous.

In some flowers, the ovary is completely fused to the hypanthium. Only the style and stigma project above the sepals, petals, and stamens, which appear to attach to the top of the ovary. This sort of ovary is called inferior ovary, and the flower is said to be epigynous. Examples of plant families with inferior ovaries include the orchids Orchidaceae, sunflower Asteraceae and the evening primrose Onagraceae.

The ovary

Ovaries are attached directly to the receptacle, but can be sunken into it (inferior, as described above), superior with no stalk, or on stalk-like extensions of the receptacle, as in Isomeris arborea.

The ovary surrounds one or more locules in which ovules develop from placenta. In monocarpous or apocarpous gynoecia, placenta are typically arranged in longitudinal strips near the margin (before fusion) of the carpel (called marginal placentation). In syncarpous gynoecia where the fused carpels form a central column and several locules from their fused margins, placenta can occur in the center of the ovary, on the column (axile placentation). In some ovaries, the placenta are found in clumps along the wall (parietal placentation) or on a central column with no septa defining locules (free central placentation). In some cases a single ovule is attached to the bottom or top of the locule (basal or apical placentation, respectively).

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The ovule

Longitudinal section of carpellate flower of squash showing ovary, ovules, pistil, and petals

The ovule (from Latin ovulum meaning small egg) is a complex structure, born inside ovaries of carpels in angiosperms. The ovule initially consists of a stalked, integumented megasporangium. Typically one cell in the megasporangium undergoes meiosis resulting in one to four megaspores. These develop into reduced megagametophytes (often called embryo sacs) within the ovule. Before fertilization, the ovule consists of one or two layers of integuments surrounding the remains of the megasporangium, called the nucellus and an embryo sac, with a small number of cells and nuclei, including one egg cell and two polar nuclei (which will form, together with a sperm cell, the primary endosperm nucleus). The gap in the integuments through which the pollen tube enters to deliver sperm to the egg is called the micropyle. The stalk attaching the ovule to the placenta is called the funiculus. Ovules are typically positioned so that the micropyle is facing the point of funiculus attachment, but other positions are found in a variety of plant groups.

The stigma and style

Stigma of a crocus flower.

The stigma is the receptive tip of the carpel, which receives pollen at pollination and on which the pollen grain germinates. The stigma is adapted to catch and trap pollen, either by combining pollen of visiting insects or by various hairs, flaps, or sculpturings.[3]

Stigmas can vary from long and slender to globe shaped to feathery. Their role in the life cycle of angiosperms is to receive and adhere to pollen from whatever vector has carried it from it flower of origin. Stigmas must distinguish and reject the pollen of other species, and in some cases the pollen from the same or genetically similar flowers. This self incompatibility function ensures outcrossing.

References

  1. ^ Judd, Campbell, Kellogg, Stevens, Donoghue, 2007 Plant Systematics: A Phylogenetic Approach, 3rd Edition Sinauer Associates, Inc. Sunderland, MA
  2. ^ Gifford, E. and Foster A. 1989Morphology and Evolution of Flowering Plants 3rd EditionW. H. Freeman and Company, New York
  3. ^ The Penguin Dictionary of Botany, edited by Elizabeth Toothill, Penguin Books 1984 ISBN 0-14-051126-1

Simple English

style and stigmas]] A gynoecium (from Ancient Greek gyne, "woman") is the female reproductive part of a flower. The male counterpart is called an androecium. A gynoecium is composed of one or more pistils. A pistil may consist of a single free carpel, or be formed from a number of carpels that are fused. The pistil itself is formed from the stigma, style, and ovary.

A plant ovary (much like an animal ovary) is the part of the pistil which contains ovules. The style is generally referred to as stalklike, without ovules located between the ovary (at the bottom of the pistil) and the stigma (located at the top portion of the pistil). In some plant species styles are not found in the pistils. Stigma is the pollen receptor within the pistil at the top of the pistil. Stigmas may be discretely defined structures or they may be within a region referred to as the stigmatic region. [1]

Pistils or ovaries can be either simple meaning only one carpel or compound meaning two or more carpels.[1]

Contents

Carpel anatomy

A carpel is the basic unit of the female reproductive organ of a flower, the gynoecium. A flower may have zero, one, or more carpels. Multiple carpels may combine into a single pistil, or into multiple pistils.

The parts of the carpel are:

  • the stigma (from Ancient Greek stigma "mark, puncture"), usually the terminal (end) portion that receives the pollen (male gametophytes); it is commonly somewhat glutinous or viscid;
  • the style (from Latin stilus "stake, stylus"), a stalk connecting the stigma with the ovary below containing the transmitting tract, which facilitates the growth of the pollen tube and hence the movement of the male gamete to the ovule; and
  • the ovary (from Latin ovum "egg") or megasporophyll (see sporophyll) containing the female reproductive cell or ovule.

The pistil

plicata, illustrating an inferior ovary.]]

A pistil (from Latin pistillum "pestle") is made up of a carpel (if single) or carpels (if fused). A gynoecium that consists of a single free carpel is termed monocarpous. That with two or more fused carpels (called a compound ovary or compound pistil) is termed syncarpous. However, if the gynoecium consists of one or more free, simple, and distinct carpels, each carpel makes an individual pistil and the gynoecium is termed apocarpous. Fertilization of the ovule or ovules results in development of the carpel(s) into a fruit.

When two or more carpels are fused or joined together its called syncarpy. In a compound pistil, the carpels are fused together in one of two basic ways:

  • the carpels are fused at or near their margins (parietal placentation), usually forming a single large cavity — an example would be the violet.
  • the folded carpels extend in towards the center, being fused along their outer faces (laterally concrescent), with the placentae arranged around a central column of tissue (axile placentation). There may be as many locules as there are carpels; and tissue of the receptacle may be involved in forming the axillary column. An example of axile placentation would be the lily.

A complicating factor in all of this is the fact that in some species syncarpy is present only at the base of the carpels, the pistil being apocarpous in the upper part. The manner of fusing of the carpels can also vary from one part of the pistil to another.

Inferior vs. superior ovaries

The gynoecium, the collective term for all the carpels, is the innermost whorl of the parts of a flower, and in many flowers the other parts (sepals, petals, and stamens) are attached to the receptacle beneath the gynoecium. In such cases, where the ovary lies above the attachments of the other distinct floral parts, the flower is described as hypogynous or as having a superior ovary. In some species (examples include plum, cherry, and blackberry), the other (noncarpellary) floral parts are fused to form a cup called a floral tube or hypanthium. In these flowers, the ovary lies physically lower than the lobes of the sepals and petals and below the point of attachment of the stamen filaments — the ovary is still considered to be superior but the flower is termed perigynous.

In those flowers in which the floral tube is fused with the ovary, the sepals, petals, and stamens appear to grow out from the top of the ovary, and the flower is said to be epigynous and have an inferior ovary. Examples of plant families with inferior ovaries include orchid, sunflower, and cactus. The position of the ovary is an important consideration in the identification and classification of plant species, as well as the kind of fruit that develops after fertilization.

The ovule

The ovule (from Latin ovulum "small egg"), which represents the megasporangium, when mature, consists of one or two coats surrounding the central nucellus, except at the apex where an opening, the micropyle, is left. The nucellus is a cellular tissue enveloping one large cell, the embryo-sac or megaspore. The germination of the megaspore consists in the repeated division of its nucleus to form two groups of four, one group at each end of the embryo-sac. One nucleus from each group, the polar nucleus, passes to the centre of the sac, where the two fuse to form the so-called definitive nucleus. Of the three cells at the micropylar end of the sac, all naked cells (the so-called egg-apparatus), one is the egg-cell or oosphere, the other two, which may be regarded as representing abortive egg-cells (in rare cases capable of fertilization), are known as synergidae. The three cells at the opposite end are known as antipodal cells and become invested with a cell-wall.

The carpel of a simple apocarpous gynoecium appears as a folded structure, differentiated into a basal fertile part (ovary) and an upper sterile part (style). Various interpretations of the origin from a leaf-like structure have been made (Esau, 1965), but the important anatomical description is that of a variously folded tissue surrounding a cavity (called a locule) within which projects one or more ovules, attached by or along a placenta. Typically, a carpel has two placentae. An example of a simple carpel is that of a pea, bean or Arabidopsis: the fruit develops from the single carpel consisting of two rows of ovules aligned beside one another along the placental margin.

References

  • Esau, K. 1965. Plant Anatomy, 2nd Edition. John Wiley & Sons. 767 pp.
  • This article includes text from the public domain 1911 Encyclopaedia Britannica. Please add to the article as needed.
  1. 1.0 1.1 Simpson, M.G.: "Plant Systematics", pp. 374-375. Elsevier Academic Press, 2006


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