Abstract
The genetic architecture underlying species differentiation is essential for understanding the mechanisms of speciation and post-zygotic reproductive barriers which exist between species. We undertook line-cross analysis of multiple hybrid (F1, F2 and backcrosses) and pure-species populations of two diploid eucalypt species from different subseries, Eucalyptus globulus and Eucalyptus nitens, to unravel the genetic architecture of their differentiation. The populations were replicated on two sites and monitored for growth and survival over a 14-year period. The hybrids exhibited severe outbreeding depression which increased with age. Of the composite additive, dominance and epistatic effects estimated, the additive × additive epistatic component was the most important in determining population divergence in both growth and survival. Significant dominance × dominance epistasis was also detected for survival at several ages. While favourable dominance and, in the case of survival, dominance × dominance epistasis could produce novel gene combinations which enhance hybrid fitness, at the population level, these effects were clearly overridden by adverse additive × additive epistasis which appears to be a major driver of overall outbreeding depression in the hybrid populations. The lack of model fit at older ages suggested that even high-order epistatic interactions may potentially have a significant contribution to outbreeding depression in survival. The estimated composite genetic parameters were generally stable across sites. Our results argue that the development of favourable epistasis is a key mechanism underlying the genetic divergence of eucalypt species, and epistasis is an important mechanism underlying the evolution of post-zygotic reproductive barriers.
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Acknowledgements
We thank Gunns Ltd (Ridgley) and Norske Skog (Tyenna) for provision of the trial sites. Funding for crossing and trial establishment was provided by the CRC for Temperate Hardwood Forestry. Trial measurements were supported by the CRC for Sustainable Production Forestry as well as an Australian Research Council Linkage grant (LP0884001) partnered by the Southern Tree Breeding Association and SeedEnergy Pty. Ltd. The financial support given by Fundação para a Ciência e Tecnologia (Lisboa, Portugal) through the Ciência 2007 initiative to João Costa e Silva and by the Australian Research Council Discovery grant (DP0986491) to Brad Potts is gratefully acknowledged and provided the opportunity to complete this long-term study.
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Costa e Silva, J., Potts, B.M. & Tilyard, P. Epistasis causes outbreeding depression in eucalypt hybrids. Tree Genetics & Genomes 8, 249–265 (2012). https://doi.org/10.1007/s11295-011-0437-8
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DOI: https://doi.org/10.1007/s11295-011-0437-8