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      Divergence in Defence against Herbivores between Males and Females of Dioecious Plant Species

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          Abstract

          Defensive traits may evolve differently between sexes in dioecious plant species. Our current understanding of this process hinges on a partial view of the evolution of resistance traits that may result in male-biased herbivory in dioecious populations. Here, we present a critical summary of the current state of the knowledge of herbivory in dioecious species and propose alternative evolutionary scenarios that have been neglected. These scenarios consider the potential evolutionary and functional determinants of sexual dimorphism in patterns of resource allocation to reproduction, growth, and defence. We review the evidence upon which two previous reviews of sex-biased herbivory have concluded that male-biased herbivory is a rule for dioecious species, and we caution readers about a series of shortcomings of many of these studies. Lastly, we propose a minimal standard protocol that should be followed in any studies that intend to elucidate the (co)evolution of interactions between dioecious plants and their herbivores.

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          Most cited references165

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          The ecology and evolution of plant tolerance to herbivory.

          The tolerance of plants to herbivory reflects the degree to which a plant can regrow and reproduce after damage from herbivores. Autoecological factors, as well as the influence of competitors and mutualists, affect the level of plant tolerance. Recent work indicates that there is a heritable basis for tolerance and that it can evolve in natural plant populations. Although tolerance is probably not a strict alternative to plant resistance, there could be inter- and intraspecific tradeoffs between these defensive strategies.
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            Sex chromosomes in land plants.

            Sex chromosomes in land plants can evolve as a consequence of close linkage between the two sex determination genes with complementary dominance required to establish stable dioecious populations, and they are found in at least 48 species across 20 families. The sex chromosomes in hepatics, mosses, and gymnosperms are morphologically heteromorphic. In angiosperms, heteromorphic sex chromosomes are found in at least 19 species from 4 families, while homomorphic sex chromosomes occur in 20 species from 13 families. The prevalence of the XY system found in 44 out of 48 species may reflect the predominance of the evolutionary pathway from gynodioecy towards dioecy. All dioecious species have the potential to evolve sex chromosomes, and reversions back from dioecy to various forms of monoecy, gynodioecy, or androdioecy have also occurred. Such reversals may occur especially during the early stages of sex chromosome evolution before the lethality of the YY (or WW) genotype is established.
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              Plant sex determination and sex chromosomes.

              Sex determination systems in plants have evolved many times from hermaphroditic ancestors (including monoecious plants with separate male and female flowers on the same individual), and sex chromosome systems have arisen several times in flowering plant evolution. Consistent with theoretical models for the evolutionary transition from hermaphroditism to monoecy, multiple sex determining genes are involved, including male-sterility and female-sterility factors. The requirement that recombination should be rare between these different loci is probably the chief reason for the genetic degeneration of Y chromosomes. Theories for Y chromosome degeneration are reviewed in the light of recent results from genes on plant sex chromosomes.
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                Author and article information

                Journal
                Int J Evol Biol
                Int J Evol Biol
                IJEB
                International Journal of Evolutionary Biology
                Hindawi Publishing Corporation
                2090-8032
                2090-052X
                2012
                23 December 2012
                : 2012
                : 897157
                Affiliations
                Department of Biology, The University of Winnipeg, Winnipeg, MB, Canada R3B 2G3
                Author notes
                *Germán Avila-Sakar: gasakar@ 123456gmail.com

                Academic Editor: Jeremy L. Marshall

                Article
                10.1155/2012/897157
                3540699
                23320247
                7dc094b5-429b-429b-8e51-2a64ae75c1bb
                Copyright © 2012 G. Avila-Sakar and C. A. Romanow.

                This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                : 29 June 2012
                : 21 September 2012
                : 17 October 2012
                Categories
                Review Article

                Evolutionary Biology
                Evolutionary Biology

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