Plant reproductive morphology

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Close-up of a flower of Schlumbergera (Christmas or Holiday Cactus), showing part of the gynoecium (the stigma and part of the style is visible) and the stamens that surround it Schlumbergera 04 ies.jpg
Close-up of a flower of Schlumbergera (Christmas or Holiday Cactus), showing part of the gynoecium (the stigma and part of the style is visible) and the stamens that surround it

Plant reproductive morphology is the study of the physical form and structure (the morphology) of those parts of plants directly or indirectly concerned with sexual reproduction.

Contents

Among all living organisms, flowers, which are the reproductive structures of angiosperms, are the most varied physically and show a correspondingly great diversity in methods of reproduction. [1] Plants that are not flowering plants (green algae, mosses, liverworts, hornworts, ferns and gymnosperms such as conifers) also have complex interplays between morphological adaptation and environmental factors in their sexual reproduction. The breeding system, or how the sperm from one plant fertilizes the ovum of another, depends on the reproductive morphology, and is the single most important determinant of the genetic structure of nonclonal plant populations. Christian Konrad Sprengel (1793) studied the reproduction of flowering plants and for the first time it was understood that the pollination process involved both biotic and abiotic interactions. Charles Darwin's theories of natural selection utilized this work to build his theory of evolution, which includes analysis of the coevolution of flowers and their insect pollinators.

Use of sexual terminology

Dioicous gametophytes of the liverwort Marchantia polymorpha. In this species, gametes are produced on different plants on umbrella-shaped gametophores with different morphologies. The radiating arms of female gametophores (left) protect archegonia that produce eggs. Male gametophores (right) are topped with antheridia that produce sperm. Marchantia polymorpha gametophytes.jpg
Dioicous gametophytes of the liverwort Marchantia polymorpha . In this species, gametes are produced on different plants on umbrella-shaped gametophores with different morphologies. The radiating arms of female gametophores (left) protect archegonia that produce eggs. Male gametophores (right) are topped with antheridia that produce sperm.

Plants have complex lifecycles involving alternation of generations. One generation, the sporophyte, gives rise to the next generation, the gametophyte asexually via spores. Spores may be identical isospores or come in different sizes (microspores and megaspores), but strictly speaking, spores and sporophytes are neither male nor female because they do not produce gametes. The alternate generation, the gametophyte, produces gametes, eggs and/or sperm. A gametophyte can be monoicous (bisexual), producing both eggs and sperm, or dioicous (unisexual), either female (producing eggs) or male (producing sperm).

In the bryophytes (liverworts, mosses, and hornworts), the sexual gametophyte is the dominant generation. In ferns and seed plants (including cycads, conifers, flowering plants, etc.) the sporophyte is the dominant generation; the obvious visible plant, whether a small herb or a large tree, is the sporophyte, and the gametophyte is very small. In bryophytes and ferns, the gametophytes are independent, free-living plants, while in seed plants, each female megagametophyte, and the megaspore that gives rise to it, is hidden within the sporophyte and is entirely dependent on it for nutrition. Each male gametophyte typically consists of two to four cells enclosed within the protective wall of a pollen grain.

The sporophyte of a flowering plant is often described using sexual terms (e.g. "female" or "male") based on the sexuality of the gametophyte it gives rise to. For example, a sporophyte that produces spores that give rise only to male gametophytes may be described as "male", even though the sporophyte itself is asexual, producing only spores. Similarly, flowers produced by the sporophyte may be described as "unisexual" or "bisexual", meaning that they give rise to either one sex of gametophyte or both sexes of the gametophyte. [2] [ page needed ]

Flowering plants

Basic flower morphology

Flower of Ranunculus glaberrimus Ranunculus glaberrimus labelled.jpg
Flower of Ranunculus glaberrimus

The flower is the characteristic structure concerned with sexual reproduction in flowering plants (angiosperms). Flowers vary enormously in their structure (morphology). A perfect flower, like that of Ranunculus glaberrimus shown in the figure, has a calyx of outer sepals and a corolla of inner petals and both male and female sex organs. The sepals and petals together form the perianth. Next inwards there are numerous stamens, which produce pollen grains, each containing a microscopic male gametophyte. Stamens may be called the "male" parts of a flower and collectively form the androecium. Finally in the middle there are carpels, which at maturity contain one or more ovules, and within each ovule is a tiny female gametophyte. [3] Carpels may be called the "female" parts of a flower and collectively form the gynoecium.

Each carpel in Ranunculus species is an achene that produces one ovule, [4] which when fertilized becomes a seed. If the carpel contains more than one seed, as in Eranthis hyemalis , it is called a follicle. Two or more carpels may be fused together to varying degrees and the entire structure, including the fused styles and stigmas may be called a pistil. The lower part of the pistil, where the ovules are produced, is called the ovary. It may be divided into chambers (locules) corresponding to the separate carpels. [5]

Variations

Alnus serrulata has unisexual flowers and is monoecious. A pair of maturing male-flower catkins are on the right; female catkins (of the previous year) on the left. Tagalder8139.jpg
Alnus serrulata has unisexual flowers and is monoecious. A pair of maturing male-flower catkins are on the right; female catkins (of the previous year) on the left.
Ilex aquifolium has unisexual flowers and is dioecious. (Above and top right): A 'shoot' with flowers from a male plant; and a male flower enlarged, showing robust stamens with pollen; and showing a female-flower stigma, reduced and sterile. + (Below and bottom right): A shoot with flowers from a female plant; and a female flower enlarged, showing a robust stigma; and showing male-flower stamens (staminodes), reduced, sterile, and with no pollen. Hollyflowers.jpg
Ilex aquifolium has unisexual flowers and is dioecious. (Above and top right): A 'shoot' with flowers from a male plant; and a male flower enlarged, showing robust stamens with pollen; and showing a female-flower stigma, reduced and sterile. + (Below and bottom right): A shoot with flowers from a female plant; and a female flower enlarged, showing a robust stigma; and showing male-flower stamens (staminodes), reduced, sterile, and with no pollen.

A perfect flower has both stamens and carpels, and is described as "bisexual" or "hermaphroditic". A unisexual flower is one in which either the stamens or the carpels are missing, vestigial or otherwise non-functional. Each flower is either staminate (having only functional stamens and thus male), or carpellate or pistillate (having only functional carpels and thus female). If separate staminate and carpellate flowers are always found on the same plant, the species is described as monoecious . If separate staminate and carpellate flowers are always found on different plants, the species is described as dioecious . [6] A 1995 study found that about 6% of angiosperm species are dioecious, and that 7% of genera contain some dioecious species. [7]

Members of the birch family (Betulaceae) are examples of monoecious plants with unisexual flowers. A mature alder tree ( Alnus species) produces long catkins containing only male flowers, each with four stamens and a minute perianth, and separate stalked groups of female flowers, each without a perianth. [8] (See the illustration of Alnus serrulata .)

Most hollies (members of the genus Ilex ) are dioecious. Each plant produces either functionally male flowers or functionally female flowers. In Ilex aquifolium (see the illustration), the common European holly, both kinds of flower have four sepals and four white petals; male flowers have four stamens, female flowers usually have four non-functional reduced stamens and a four-celled ovary. [9] Since only female plants are able to set fruit and produce berries, this has consequences for gardeners. Amborella represents the first known group of flowering plants to separate from their common ancestor. It too is dioecious; at any one time, each plant produces either flowers with functional stamens but no carpels, or flowers with a few non-functional stamens and a number of fully functional carpels. However, Amborella plants may change their "sex" over time. In one study, five cuttings from a male plant produced only male flowers when they first flowered, but at their second flowering three switched to producing female flowers. [10]

In extreme cases, almost all of the parts present in a complete flower may be missing, so long as at least one carpel or one stamen is present. This situation is reached in the female flowers of duckweeds ( Lemna ), which consist of a single carpel, and in the male flowers of spurges ( Euphorbia ) which consist of a single stamen. [11]

The basic cases of sexuality of flowering plants. Monoecy dioecy en.svg
The basic cases of sexuality of flowering plants.

A species such as Fraxinus excelsior , the common ash of Europe, demonstrates one possible kind of variation. Ash flowers are wind-pollinated and lack petals and sepals. Structurally, the flowers may be bisexual, consisting of two stamens and an ovary, or may be male (staminate), lacking a functional ovary, or female (carpellate), lacking functional stamens. Different forms may occur on the same tree, or on different trees. [8] The Asteraceae (sunflower family), with close to 22,000 species worldwide, have highly modified inflorescences made up of flowers (florets) collected together into tightly packed heads. Heads may have florets of one sexual morphology – all bisexual, all carpellate or all staminate (when they are called homogamous), or may have mixtures of two or more sexual forms (heterogamous). [12] Thus goatsbeards ( Tragopogon species) have heads of bisexual florets, like other members of the tribe Cichorieae, [13] whereas marigolds ( Calendula species) generally have heads with the outer florets bisexual and the inner florets staminate (male). [14]

Like Amborella, some plants undergo sex-switching. For example, Arisaema triphyllum (Jack-in-the-pulpit) expresses sexual differences at different stages of growth: smaller plants produce all or mostly male flowers; as plants grow larger over the years the male flowers are replaced by more female flowers on the same plant. Arisaema triphyllum thus covers a multitude of sexual conditions in its lifetime: nonsexual juvenile plants, young plants that are all male, larger plants with a mix of both male and female flowers, and large plants that have mostly female flowers. [15] Other plant populations have plants that produce more male flowers early in the year and as plants bloom later in the growing season they produce more female flowers.[ citation needed ]

Terminology

The complexity of the morphology of flowers and its variation within populations has led to a rich terminology.

Outcrossing

Outcrossing, cross-fertilization or allogamy, in which offspring are formed by the fusion of the gametes of two different plants, is the most common mode of reproduction among higher plants. About 55% of higher plant species reproduce in this way. An additional 7% are partially cross-fertilizing and partially self-fertilizing (autogamy). About 15% produce gametes but are principally self-fertilizing with significant out-crossing lacking. Only about 8% of higher plant species reproduce exclusively by non-sexual means. These include plants that reproduce vegetatively by runners or bulbils, or which produce seeds without embryo fertilization (apomixis). The selective advantage of outcrossing appears to be the masking of deleterious recessive mutations. [28]

The primary mechanism used by flowering plants to ensure outcrossing involves a genetic mechanism known as self-incompatibility. Various aspects of floral morphology promote allogamy. In plants with bisexual flowers, the anthers and carpels may mature at different times, plants being protandrous (with the anthers maturing first) or protogynous (with the carpels mature first).[ citation needed ] Monoecious species, with unisexual flowers on the same plant, may produce male and female flowers at different times.[ citation needed ]

Dioecy, the condition of having unisexual flowers on different plants, necessarily results in outcrossing, and probably evolved for this purpose. However, "dioecy has proven difficult to explain simply as an outbreeding mechanism in plants that lack self-incompatibility". [7] Resource-allocation constraints may be important in the evolution of dioecy, for example, with wind-pollination, separate male flowers arranged in a catkin that vibrates in the wind may provide better pollen dispersal. [7] In climbing plants, rapid upward growth may be essential, and resource allocation to fruit production may be incompatible with rapid growth, thus giving an advantage to delayed production of female flowers. [7] Dioecy has evolved separately in many different lineages, and monoecy in the plant lineage correlates with the evolution of dioecy, suggesting that dioecy can evolve more readily from plants that already produce separate male and female flowers. [7]

See also

Related Research Articles

<span class="mw-page-title-main">Gametophyte</span> Haploid stage in the life cycle of plants and algae

A gametophyte is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.

<span class="mw-page-title-main">Sex</span> Trait that determines an organisms sexually reproductive function

Sex is the trait that determines whether a sexually reproducing organism produces male or female gametes. A male organism produces small mobile gametes, while a female organism produces larger, non-mobile gametes. An organism that produces both types of gamete is called a hermaphrodite. During sexual reproduction, a male and a female gamete fuse to form a zygote, which develops into an offspring that inherits traits from each parent.

<span class="mw-page-title-main">Sex organ</span> Biological part involved in sexual reproduction

A sex organ, also known as a reproductive organ, is a part of an organism that is involved in sexual reproduction. Sex organs constitute the primary sex characteristics of an organism. Sex organs are responsible for producing and transporting gametes, as well as facilitating fertilization and supporting the development and birth of offspring. Sex organs are found in many species of animals and plants, with their features varying depending on the species.

<span class="mw-page-title-main">Alternation of generations</span> Reproductive cycle of plants and algae

Alternation of generations is the predominant type of life cycle in plants and algae. In plants both phases are multicellular: the haploid sexual phase – the gametophyte – alternates with a diploid asexual phase – the sporophyte.

<span class="mw-page-title-main">Stamen</span> Male organ of a flower

The stamen is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium.

<span class="mw-page-title-main">Buxales</span> Order of eudicot flowering plants

The Buxales are a small order of eudicot flowering plants, recognized by the APG IV system of 2016. The order includes the family Buxaceae; the families Didymelaceae and Haptanthaceae may also be recognized or may be included in the Buxaceae. Many members of the order are evergreen shrubs or trees, although some are herbaceous perennials. They have separate "male" (staminate) and "female" (carpellate) flowers, mostly on the same plant. Some species are of economic importance either for the wood they produce or as ornamental plants.

<i>Amborella</i> Species of shrub

Amborella is a monotypic genus of understory shrubs or small trees endemic to the main island, Grande Terre, of New Caledonia in the southwest Pacific Ocean. The genus is the only member of the family Amborellaceae and the order Amborellales and contains a single species, Amborella trichopoda. Amborella is of great interest to plant systematists because molecular phylogenetic analyses consistently place it as the sister group to all other flowering plants.

<span class="mw-page-title-main">Sporophyte</span> Diploid multicellular stage in the life cycle of a plant or alga

A sporophyte is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase.

In biology, gonochorism is a sexual system where there are only two sexes and each individual organism is either male or female. The term gonochorism is usually applied in animal species, the vast majority of which are gonochoric.

Dioecy is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly or indirectly. Dioecious reproduction is biparental reproduction. Dioecy has costs, since only the female part of the population directly produces offspring. It is one method for excluding self-fertilization and promoting allogamy (outcrossing), and thus tends to reduce the expression of recessive deleterious mutations present in a population. Plants have several other methods of preventing self-fertilization including, for example, dichogamy, herkogamy, and self-incompatibility.

<span class="mw-page-title-main">Gynoecium</span> Female organs of a flower

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 pistils and is typically surrounded by the pollen-producing reproductive organs, the stamens, collectively called the androecium. The gynoecium is often referred to as the "female" portion of the flower, although rather than directly producing female gametes, the gynoecium produces megaspores, each of which develops into a female gametophyte which then produces egg cells.

<i>Didymeles</i> Genus of trees

Didymeles is a genus of flowering plants. It is variously treated as the only genus of the family Didymelaceae — or in the family Buxaceae, as in the APG IV system.

<span class="mw-page-title-main">Flower</span> Reproductive structure in flowering plants

A flower, also known as a bloom or blossom, is the reproductive structure found in flowering plants. Flowers consist of a combination of vegetative organs – sepals that enclose and protect the developing flower, petals that attract pollinators, and reproductive organs that produce gametophytes, which in flowering plants produce gametes. The male gametophytes, which produce sperm, are enclosed within pollen grains produced in the anthers. The female gametophytes are contained within the ovules produced in the carpels.

Plant reproduction is the production of new offspring in plants, which can be accomplished by sexual or asexual reproduction. Sexual reproduction produces offspring by the fusion of gametes, resulting in offspring genetically different from either parent. Asexual reproduction produces new individuals without the fusion of gametes, resulting in clonal plants that are genetically identical to the parent plant and each other, unless mutations occur.

<span class="mw-page-title-main">Environmental sex determination</span> Method of sex-determination

Environmental sex determination is the establishment of sex by a non-genetic cue, such as nutrient availability, experienced within a discrete period after fertilization. Environmental factors which often influence sex determination during development or sexual maturation include light intensity and photoperiod, temperature, nutrient availability, and pheromones emitted by surrounding plants or animals. This is in contrast to genotypic sex determination, which establishes sex at fertilization by genetic factors such as sex chromosomes. Under true environmental sex determination, once sex is determined, it is fixed and cannot be switched again. Environmental sex determination is different from some forms of sequential hermaphroditism in which the sex is determined flexibly after fertilization throughout the organism’s life.

This page provides a glossary of plant morphology. Botanists and other biologists who study plant morphology use a number of different terms to classify and identify plant organs and parts that can be observed using no more than a handheld magnifying lens. This page provides help in understanding the numerous other pages describing plants by their various taxa. The accompanying page—Plant morphology—provides an overview of the science of the external form of plants. There is also an alphabetical list: Glossary of botanical terms. In contrast, this page deals with botanical terms in a systematic manner, with some illustrations, and organized by plant anatomy and function in plant physiology.

This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary of leaf morphology. For other related terms, see Glossary of phytopathology, Glossary of lichen terms, and List of Latin and Greek words commonly used in systematic names.

<span class="mw-page-title-main">Hermaphrodite</span> Sexually reproducing organism that produces both male and female gametes

A hermaphrodite is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are of different sexes, either male or female but not both, are gonochoric, which is the opposite of hermaphroditic.

<span class="mw-page-title-main">Sexual reproduction</span> Reproduction process that creates a new organism by combining the genetic material of two organisms

Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete with a single set of chromosomes combines with another gamete to produce a zygote that develops into an organism composed of cells with two sets of chromosomes (diploid). This is typical in animals, though the number of chromosome sets and how that number changes in sexual reproduction varies, especially among plants, fungi, and other eukaryotes.

A sexual system is a pattern of sex allocation or a distribution of male and female function across organisms in a species. Terms like reproductive system and mating system have also been used as synonyms.

References

Citations

  1. ↑ Barrett, S.C.H. (2002). "The evolution of plant sexual diversity" (PDF). Nature Reviews Genetics. 3 (4): 274–284. doi:10.1038/nrg776. PMID   11967552. S2CID   7424193.
  2. 1 2 3 4 5 Hickey, M. & King, C. (2001). The Cambridge Illustrated Glossary of Botanical Terms. Cambridge University Press.
  3. ↑ Sporne 1974, pp. 14–15.
  4. ↑ Whittemore, Alan T. "Ranunculus". Flora of North America. Retrieved 2013-03-04– via www.eFloras.org.
  5. ↑ Sporne 1974, pp. 125–127.
  6. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 Beentje, Henk (2010). The Kew Plant Glossary. Richmond, Surrey: Royal Botanic Gardens, Kew. ISBN   978-1-84246-422-9.
  7. 1 2 3 4 5 Renner, S.S. & Ricklefs, R.E. (1995). "Dioecy and its correlates in the flowering plants". American Journal of Botany. 82 (5): 596–606. doi:10.2307/2445418. JSTOR   2445418.
  8. 1 2 Stace 2010, pp. 292–296.
  9. ↑ Stace 2010, p. 669.
  10. ↑ Buzgo, Matyas; Soltis, Pamela S. & Soltis, Douglas E. (2004). "Floral Developmental Morphology of Amborella trichopoda (Amborellaceae)". International Journal of Plant Sciences. 165 (6): 925–947. doi:10.1086/424024. S2CID   84793812.
  11. ↑ Sporne 1974, pp. 15–16.
  12. ↑ Barkley, Theodore M.; Brouillet, Luc & Strother, John L. "Asteraceae". Flora of North America. Retrieved 2013-03-04– via www.eFloras.org.
  13. ↑ Barkley, Theodore M.; Brouillet, Luc & Strother, John L. "Chichorieae". Flora of North America. Retrieved 2013-03-04– via www.eFloras.org.
  14. ↑ Strother, John L. "Calendula". Flora of North America. Retrieved 2013-03-04– via www.eFloras.org.
  15. ↑ Ewing, J.W. & Klein, R.M. (1982). "Sex Expression in Jack-in-the-Pulpit". Bulletin of the Torrey Botanical Club. 109 (1): 47–50. doi:10.2307/2484467. JSTOR   2484467.
  16. 1 2 3 Janick, J. (2010). Plant Breeding Reviews. Wiley. ISBN   9780470650028.
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  18. ↑ Baskauf, Steve (2002). "Sexual systems in angiosperms". Archived from the original on 2018-07-03. Retrieved 2013-02-27.
  19. ↑ "Gynodioecious". Dictionary of Botany. Retrieved 2013-04-10.
  20. ↑ G. J. H. Grubben (2004). Vegetables. PROTA. pp.  255–. ISBN   978-90-5782-147-9.
  21. ↑ Cook 1968, p. 131.
  22. ↑ Dinesh Kumar (20 August 2008). Definitional Glossary of Agricultural Terms. I. K. International Pvt Ltd. pp. 115–. ISBN   978-81-906757-4-1.
  23. ↑ Geber, Monica A. (1999). Gender and sexual dimorphism in flowering plants. Berlin: Springer. ISBN   3-540-64597-7. p. 4
  24. ↑ Testolin, R; Cipriani, G; Costa, G (May 1995). "Sex segregation ratio and gender expression in the genus Actinidia". Sexual Plant Reproduction. 8 (3). doi:10.1007/BF00242255. S2CID   25414438 . Retrieved 30 December 2020.
  25. ↑ Olson, Matthew S. & Antonovics, Janis (2000). "Correlation between male and female reproduction in the subdioecious herb Astilbe biternata (Saxifragaceae)". American Journal of Botany. 87 (6): 837–44. doi:10.2307/2656891. JSTOR   2656891. PMID   10860914.
  26. ↑ Strittmatter, L.I.; Negrón-Ortiz, V. & Hickey, R.J. (2002). "Subdioecy in Consolea spinosissima (Cactaceae): breeding system and embryological studies". American Journal of Botany. 89 (9): 1373–1387. doi:10.3732/ajb.89.9.1373. PMID   21665739.
  27. ↑ Beentje, Henk (2016). The Kew Plant Glossary (second ed.). Richmond, Surrey: Royal Botanic Gardens, Kew. ISBN   978-1-84246-604-9.
  28. ↑ Bernstein, C. & Bernstein, H. (1991). Aging, Sex, and DNA Repair. San Diego: Academic Press. ISBN   978-0-12-092860-6.

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