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Variation and Evolution in Plants and Microorganisms: Toward a New Synthesis 50 Years after Stebbins (2000)
National Academy of Sciences (NAS)

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. "17 The Role of Genetic and Genomic Attributes in the Success of Polyploids." Variation and Evolution in Plants and Microorganisms: Toward a New Synthesis 50 Years after Stebbins. Washington, DC: The National Academies Press, 2000.

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Variation and Evolution in Plants and Microorganisms: TOWARD A NEW SYNTHESIS 50 YEARS AFTER STEBBINS

polyploid. Perhaps most important to Stebbins (1950) was the availability of new ecological niches. Additional hypotheses for the success of polyploids include broader ecological amplitude of the polyploid relative to its diploid parents, better colonizing ability, higher selfing rates, and increased heterozygosity.

In fact, many aspects of the genetic systems of polyploids may contribute to the success of polyploid plants. These characteristics range from the molecular level to the population level and include increased heterozygosity, reduced inbreeding depression and an associated increase in selfing rates, increased genetic diversity through multiple formations of a polyploid species, genome rearrangements, and ancient polyploidy and gene silencing. But what role, if any, do these factors really play in the success of polyploids? In this paper, we will explore the evidence for the role of these genetic attributes in the evolutionary success of polyploid plant species.

ALLO- VERSUS AUTOPOLYPLOIDY

We will distinguish among types of polyploids by using Stebbins' (1947, 1950) classification: Allopolyploids are those polyploids that have arisen through the processes of interspecific hybridization and chromosome doubling (not necessarily in that order), autopolyploids are those polyploids that have arisen from conspecific parents, and segmental allopolyploids are those that have arisen from parents with partially divergent chromosome arrangements such that some chromosomal regions are homologous between the parents and others are homoeologous; segmental allopolyploids will not be considered further in this paper. Allopolyploids are characterized by fixed (i.e., nonsegregating) heterozygosity, resulting from the combination of divergent parental genomes; bivalent formation occurs at meiosis, and disomic inheritance operates at each locus. Autopolyploids may exhibit multivalent formation at meiosis and are characterized by polysomic inheritance. Allopolyploids are considered much more prevalent in nature than are autopolyploids, but even a cursory glance at any flora (for example, see Hickman, 1993) or list of plant chromosome numbers (for example, see Federov, 1969) will reveal multiple cytotypes within many species, even though these additional ploidal levels are not typically accorded species status. Thus, autopolyploids in nature likely are much more common than typically is recognized.

INCREASED HETEROZYGOSITY

Roose and Gottlieb (1976) showed that allotetraploids in Tragopogon had fixed heterozygosity at isozyme loci, representing the combination of

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312
Front Matter (R1-R12)
Part I: Early Evolution and the Origin of Cells (1-2)
1 G. Ledyard Stebbins (1906-2000) -- An Appreciation (3-5)
2 Solution to Darwin's Dilemma: Discovery of the Missing Precambrian Record of Life (6-20)
3 The Chimeric Eukaryote: Origin of the Nucleus from the Karyomastigont in Amitochondriate Protists (21-34)
4 Dynamic Evolution of Plant Mitochondrial Genomes: Mobile Genes and Introns and Highly Variable Mutation Rates (35-58)
Part II: Viral and Bacterial Models (59-60)
5 The Evolution of RNA Viruses: A Population Genetics View (61-82)
6 Effects of Passage History and Sampling Bias on Phylogenetic Reconstruction of Human Influenza A Evolution (83-98)
7 Bacteria are Different: Observations, Interpretations, Speculations, and Opinions About the Mechanisms of Adaptive Evolution in Prokaryotes (99-114)
Part III: Protoctist Models (115-116)
8 Evolution of RNA Editing in Trypanosome Mitochondria (117-142)
9 Population Structure and Recent Evolution of Plasmodium falciparum (143-164)
Part IV: Population Variation (165-166)
10 Transposons and Genome Evolution in Plants (167-186)
11 Maize as a Model for the Evolution of Plant Nuclear Genomes (187-210)
12 Flower Color Variation: A Model for the Experimental Study of Evolution (211-234)
13 Gene Genealogies and Population Variation in Plants (235-252)
Part V: Trends and Patterns in Plant Evolution (253-254)
14 Toward a New Synthesis: Major Evolutionary Trends in the Angiosperm Fossil Record (255-270)
15 Reproductive Systems and Evolution in Vascular Plants (271-288)
16 Hybridization as a Stimulus for the Evolution of Invasiveness in Plants? (289-309)
17 The Role of Genetic and Genomic Attributes in the Success of Polyploids (310-330)
Index (331-340)