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Dichogamy

 

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Dichogamy



 
 
Dichogamy is the separation in time of gender expression in a hermaphroditic organism, a characteristic of some fishes, gastropods, and most flowering plants.

he context of the flowering plants (angiosperms), there are two forms of dichogamy: protogyny—female function precedes male function—and protandry—male function precedes female function.

Historically, dichogamy has been regarded as a mechanism for reducing inbreeding
Inbreeding

Inbreeding is biological reproduction between close Kinships, whether plant or animal. If practiced repeatedly, it leads to an increase in homozygosity of a population....
 (e.g., Darwin, 1862).






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Dichogamy is the separation in time of gender expression in a hermaphroditic organism, a characteristic of some fishes, gastropods, and most flowering plants.

In flowering plants

In the context of the flowering plants (angiosperms), there are two forms of dichogamy: protogyny—female function precedes male function—and protandry—male function precedes female function.

Historically, dichogamy has been regarded as a mechanism for reducing inbreeding
Inbreeding

Inbreeding is biological reproduction between close Kinships, whether plant or animal. If practiced repeatedly, it leads to an increase in homozygosity of a population....
 (e.g., Darwin, 1862). However, a survey of the angiosperms found that self-incompatible
Self-incompatibility in plants

Self-incompatibility is a general name for several genetic mechanisms in angiosperms, which prevent self-fertilization and thus encourage outcrossing....
 (SI) plants, which are incapable of inbreeding, were as likely to be dichogamous as were self-compatible (SC) plants (Bertin, 1993). This finding led to a reinterpretation of dichogamy as a more general mechanism for reducing the impact of pollen
Pollen

Pollen is a fine to coarse powder consisting of Gametophyte , which produce the male gametes of spermatophyta. A hard coat covering the pollen grain protects the sperm cells during the process of their movement between the stamens of the flower to the pistil of the next flower....
-pistil interference on pollen import and export (reviewed in Lloyd & Webb, 1986; Barrett, 2002). Unlike the inbreeding-avoidance hypothesis, which focused on female function, this interference-avoidance hypothesis considers both gender functions.

In many hermaphroditic species, the close physical proximity of anthers and stigma makes interference unavoidable, either within a flower
Flower

A flower, sometimes known as a bloom or blossom, is the reproduction structure found in flowering plants . The biological function of a flower is to mediate the union of male sperm with female ovum in order to produce seeds....
 or between flowers on an inflorescence
Inflorescence

An inflorescence is a group or cluster of flowers arranged on a Plant stem that is composed of a main branch or a complicated arrangement of branches....
. Within-flower interference, which occurs when either the pistil interrupts pollen removal or the anthers prevent pollen deposition, can result in autonomous or facilitated self-pollination (Lloyd & Webb, 1986; Lloyd & Schoen, 1992). Between-flower interference results from similar mechanisms, except that the interfering structures occur on different flowers within the same inflorescence and it requires pollinator
Pollinator

A pollinator is the biotic agent that moves pollen from the male anthers of a flower to the female carpel of a flower to accomplish fertilization or syngamy of the female gamete in the ovule of the flower by the male gamete from the pollen grain....
 activity. This results in geitonogamous
Geitonogamy

Geitonogamy is the pollination of a flower with the pollen from another flower on the same flowering plant. In animal pollinated systems this is accomplished by a pollinator visiting multiple flowers on the same plant....
 pollination, the transfer of pollen between flowers of the same individual (Lloyd & Schoen, 1992; de Jong et al., 1993). In contrast to within-flower interference, geitonogamy necessarily involves the same processes as outcrossing: pollinator attraction, reward provisioning, and pollen removal. Therefore, between-flower interference not only carries the cost of self-fertilization (inbreeding depression
Inbreeding depression

Inbreeding depression is reduced fitness in a given population as a result of breeding of related individuals. Breeding between closely related individuals, called inbreeding, results in more recessive deleterious traits manifesting themselves....
; Charlesworth & Charlesworth, 1987; Husband & Schemske, 1996), but also reduces the amount of pollen available for export (so-called "pollen discounting"; Harder & Wilson, 1998]). Because pollen discounting diminishes outcross siring success, interference avoidance may be an important evolutionary force in floral biology (Harder & Barrett, 1995, 1996; Harder & Wilson, 1998; Barrett, 2002).

Dichogamy may reduce between-flower interference by minimizing the temporal overlap between stigma and anthers within an inflorescence. Large inflorescences attract more pollinators, potentially enhancing reproductive success by increasing pollen import and export (Schemske, 1980; Queller, 1983; Bell, 1985; Geber, 1985; Schmid-Hempel & Speiser, 1988; Klinkhamer & de Jong, 1990). However, large inflorescences also increase the opportunities for both geitonogamy and pollen discounting, so that the opportunity for between-flower interference increases with inflorescence size (Harder & Barrett, 1996). Consequently, the evolution of floral display size may represent a compromise between maximizing pollinator visitation and minimizing geitonogamy and pollen discounting (Klinkhamer & de Jong, 1993; Barrett et al, 1994; Holsinger, 1996; Snow et al., 1996).

Protandry may be particularly relevant to this compromise, because it often results in an inflorescence structure with female phase flowers positioned below male phase flowers (Bertin & Newman, 1993). Given the tendency of many insect pollinators to forage upwards through inflorescences (Galen & Plowright, 1988), protandry may enhance pollen export by reducing between-flower interference (Darwin, 1862; Harder et al, 2000). Furthermore, this enhanced pollen export should increase as floral display size increases, because between-flower interference should increase with floral display size. These effects of protandry on between-flower interference may decouple the benefits of large inflorescences from the consequences of geitonogamy and pollen discounting. Such a decoupling would provide a significant reproductive advantage through increased pollinator visitation and siring success.

Harder et al. (2000) demonstrated experimentally that dichogamy both reduced rates of self-fertilization and enhanced outcross siring success through reductions in geitonogamy and pollen discounting, respectively. Routley & Husband (2003)examined the influence of inflorescence size on this siring advantage and found a bimodal distribution with increased siring success with both small and large display sizes.

See also

  • Plant sexuality
    Plant sexuality

    Plant sexuality covers the wide variety of sexual reproduction systems found across the plant kingdom. This article describes Morphology aspects of sexual reproduction of plants....
  • for an experimental test of the adaptive significance of protogyny.