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That plants employ many different strategies to engage in sexual reproduction was used, from just a structural perspective, by Carolus Linnaeus (1735) to propose a system of classification of flowering plants, and later this subject received attention from Charles Darwin (1877). Flowers , the reproductive structures of Angiosperms , are more varied than the equivalent structures of any other group of organisms, and flowering plants also have an unrivalled diversity of sexual systems (Barrett, 2002). But sexuality and the significance of sexual reproductive strategies is no less important in all of the other plant groups. The breeding system is the single most important determinant of the mating structure of nonclonal plant populations. The mating structure in turn controls the amount and distribution of genetic variation, a central element in the evolutionary process (Costich, 1995).


TERMINOLOGY


The complexity of the systems and devices used by plants to achieve sexual reproduction has resulted in botanists and evolutionary biologists proposing numerous terms to describe structures and strategies. Dellaporta and Calderon-Urrea (1993) list and define a variety of terms used to describe the modes of sexuality at different levels in flowering plants. This list is reproduced here (taken from Molner, 2004), generalized to fit more than just plants that have Flower s, and expanded to include other terms and better definitions.

is monoecious. Shown here: maturing male flower catkins on right, last year's female catkins on left]]
  • Individual reproductive unit (a flower in angiosperms):

  • --- Bisexual - Reproductive structure with both male and female equivalent parts (stamens and pistil in angiosperms; also called a '''perfect''' flower); another term widely used is ''' Hermaphrodite '''.

  • --- Unisexual - Reproductive structure that is either functionally male or functionally female. In angiosperms this condition is also called '''imperfect'''.


  • Individual plant:

  • --- Hermaphrodite - A plant that has only hermaphrodite reproductive units (flowers, conifer cones, or functionally equivalent structures). In angiosperm terminology a synonym is '''monoclinous''' from the Greek "one bed".

  • --- Monoecious - having '''unisexual''' reproductive units (flowers, conifer cones, or functionally equivalent structures) of both sexes appearing on the same plant; from Greek for "one household".

  • --- Dioecious - having '''unisexual''' reproductive units (flowers, conifer cones, or functionally equivalent structures) occurring on different individuals; from Greek for "two households".

  • --- Because many dioecious Conifer s show a tendency towards monoecy (that is, a female plant may sometimes produce small numbers of male cones or vice versa), these species are termed subdioecious (McCormick & Andresen, 1963).

  • --- In angiosperm terminology, diclinous ("two beds") includes all species with unisexual flowers, although particularly those with ''only'' unisexual flowers, i.e. the monoecious and dioecious species.

  • --- Gynoecious - has only female reproductive structures; the "female" plant.

  • --- Androecious - has only male reproductive structures; the "male" plant.

  • --- Gynomonoecious - has both hermaphrodite and female structures.

  • --- Andromonoecious - has both hermaphrodite and male structures.

  • --- Trimonoecious (polygamous) - male, female, and hermaphrodite structures all appear on the same plant.


(''Ilex aquifolium'') is dioecious: (above) shoot with flowers from male plant; (top right) male flower enlarged, showing Stamen s with Pollen and reduced, sterile Stigma ; (below) shoot with flowers from female plant; (lower right) female flower enlarged, showing stigma
and reduced, sterile stamens with no pollen]]
  • Plant population

  • --- Hermaphrodite - only hermaphrodite plants.

  • --- Monoecious - only monoecious plants.

  • --- Dioecious - only dioecious plants.

  • --- Gynodioecious - both female and hermaphrodite plants present.

  • --- Androdioecious - both male and hermaphrodite plants present.

  • --- Trioecious (or '''subdioecious''') - male, female, and hermaphrodite plants are all in the same population.


Some plants use a method known as Self-incompatibility to ensure genetic diversity within the species. In these plants, the male organs cannot fertilize the female parts of the same plant.


FLOWER MORPHOLOGY

A species, such as the Ash Tree (''Fraxinus excelsior'' L.), demonstrates the possible range of variation in morphology and functionality exhibited by flowers with respect to gender. Flowers of the ash are wind-pollinated and lack Petal s and Sepal s. Structurally, the flowers may be either male, female, or hermaphrodite, the latter consisting of two anthers and an ovary ('c' below). A male flower can be morphologically male ('a' below) or a hermaphrodite flower with anthers and a rudimentary gynoecium ('b' below; functionally 'male'). Ash flowers can also be morphologically female ('e' below) or hermaphrodite and functionally female ('d' below; with vestigial anthers).



(Illustration from Binggeli and Power, 1999)


EVOLUTION


Angiosperms

It is thought that flowering plants evolved from a common hermaphrodite ancestor, and that dioecy evolved from hermaphroditism. Hermaphroditism is very common in flowering plants—about 70% are hermaphroditic, while only about 5% are dioecious and 7% are monoecious. About 7% of species exhibit gynodioecy or androdioecy, while 10% contain both unisexual and bisexual flowers (Molner, 2004).

A fair degree of correlation (though far from complete) exists between dioecy/sub-dioecy and plants that have seeds dispersed by Bird s (both Nuts and Berries ). It is hypothesized that the concentration of fruit in half of the plants increases dispersal efficiency; female plants can produce a higher density of fruit as they do not expend resources on pollen production, and the dispersal agents (birds) need not waste time looking for fruit on male plants.


EXTERNAL LINKS



REFERENCES

  • Barrett, S.C.H. 2002. The evolution of plant sexual diversity. ''Nature Reviews Genetics'' 3(4): 274-284.

  • Binggeli, P. and J. Power. 1999. Gender variation in ash (''Fraxinus excelsior'' L.)

  • Costich, D. E. 1995. Gender specialization across a climatic gradient: experimental comparison of monoecious and dioecious ''Ecballium''. ''Ecology'', June 1995.

  • Darwin, C. 1877. ''The Different Forms of Flowers on Plants of the Same Species''.

  • Dellaporta, S.L. and A. Calderon-Urrea. 1993. Sex determination in flowering plants. ''The Plant Cell'', 5: 1241-1251

  • Linnaeus, C. 1735. '' Systema Naturae ''.

  • McCormick, J. & J. W. Andresen. 1963. A subdioecious population of ''Pinus cembroides'' in southeast Arizona. ''Ohio J. Science'', 63: 159-163.

  • Molnar, Sebastian. 2004. Plant Reproductive Systems , internet version posted February 17, 2004.