Understanding intraspecific variation in genome size in plants.

Intraspecific variation in genome size makes it possible to study ongoing processes of genome size evolution. Although there are over 200 papers on intraspecific variation in genome size, there is still limited understanding of this phenomenon, especially as many of these papers are based on weak methodology and therefore report biased or false evidence of the extent of intraspecific variation. In this paper the recent progress in understanding the spatio-temporal dynamics of intraspecific variation in genome size caused by the gradual accumulation of mutations is reviewed. The results of the case studies on Microseris douglasii, Zea mays, Silene latifolia, Hordeum spontaneum and Lolium hybrids, and in particular that on Festuca pallens, are discussed. The variation in genome size that occurs within species is caused mainly by differences in the content of repetitive DNA, in particular it is a consequence of the dynamics of transposable elements. Variation may be induced and maintained polytopically. We assume that it is probably more frequent in groups of young radiating species. Even in the initial stages, the variation in genome size generated within a population seems to be restricted by selection, which is also important in stabilizing genome size within species. The long-term persistence of the variation within a population and its further accumulation may be enhanced by gametes with different genome sizes, produced by the segregation of unequally sized homeologous chromosomes. Over large geographical scales and across contrasting environmental gradients, the distribution of genome sizes within species may be influenced by the nucleotype effect, with smaller genomes being more successful at higher latitudes and altitudes and under stressful conditions. However, the small differences in genome size within species seem generally to be of minor importance relative to other components of plant fitness that may be selectively favourable under particular environmental or habitat conditions. The processes generating variation in genome size may be associated with phenotypic variation. While the shift in the genome size of a population through selection enables adaptive evolution of genome size in a newly arising species, the spatio-temporal variation in genome size within an ancestral species allows for a rapid multiple genome size divergence of related species through random drift in genome size (founder effect, bottleneck effect) during range fragmentation, hybridization and/or polyploidization.