Genome duplication is thought to be central to the evolution of morphological complexity, and some polyploids enjoy a variety of capabilities that transgress those of their diploid progenitors. Comparison of genomic sequences from several tetraploid (A t D t ) Gossypium species and genotypes with putative diploid A-and D-genome progenitor species revealed that unidirectional DNA exchanges between homeologous chromosomes were the predominant mechanism responsible for allelic differences between the Gossypium tetraploids and their diploid progenitors. Homeologous gene conversion events (HeGCEs) gradually subsided, declining to rates similar to random mutation during radiation of the polyploid into multiple clades and species. Despite occurring in a common nucleus, preservation of HeGCE is asymmetric in the two tetraploid subgenomes. A t -to-D t conversion is far more abundant than the reciprocal, is enriched in heterochromatin, is highly correlated with GC content and transposon distribution, and may silence abundant A-genome-derived retrotransposons. D t -to-A t conversion is abundant in euchromatin and genes, frequently reversing losses of gene function. The long-standing observation that the nonspinnable-fibered D-genome contributes to the superior yield and quality of tetraploid cotton fibers may be explained by accelerated D t to A t conversion during cotton domestication and improvement, increasing dosage of alleles from the spinnable-fibered A-genome. HeGCE may provide an alternative to (rare) reciprocal DNA exchanges between chromosomes in heterochromatin, where genes have approximately five times greater abundance of D t -to-A t conversion than does adjacent intergenic DNA. Spanning exon-to-gene-sized regions, HeGCE is a natural noninvasive means of gene transfer with the precision of transformation, potentially important in genetic improvement of many crop plants.G ENOME duplication is a potentially rich source of genes with new (Stephens 1951;Ohno 1970) or modified functions (Lynch and Conery 2000), and is thought to be central to the evolution of morphological complexity (Freeling and Thomas 2006). Genome doubling may confer advantages to a polyploid (Comai 2005), via mechanisms such as increased gene dosage, "intergenomic heterosis" conferred by multiple alleles in a polyploid nucleus, or the evolution of novel gene functions (neofunctionalization) (Stephens 1951;Ohno 1970). Over time, duplicated genes may evolve subdivisions of ancestral functions (subfunctionalization) (Lynch and Force 2000) that render them interdependent. Subfunctionalization may sometimes lead to neofunctionalization (He and Zhang 2005).Polyploids have been suggested to enjoy a variety of capabilities that transgress those of their diploid progenitors. For example, the notion that polyploids may adapt better (Paterson et al. 2010) provides "natural replicates" for a variety of investigations. Study of the genes from three rounds of ancient whole genome duplications in Arabidopsis reveals a short phase of function relaxa...