Eucalypts are the world's most widely planted hardwood trees. Their outstanding diversity, adaptability and growth have made them a global renewable resource of fibre and energy. We sequenced and assembled .94% of the 640-megabase genome of Eucalyptus grandis. Of 36,376 predicted protein-coding genes, 34% occur in tandem duplications, the largest proportion thus far in plant genomes. Eucalyptus also shows the highest diversity of genes for specialized metabolites such as terpenes that act as chemical defence and provide unique pharmaceutical oils. Genome sequencing of the E. grandis sister species E. globulus and a set of inbred E. grandis tree genomes reveals dynamic genome evolution and hotspots of inbreeding depression. The E. grandis genome is the first reference for the eudicot order Myrtales and is placed here sister to the eurosids. This resource expands our understanding of the unique biology of large woody perennials and provides a powerful tool to accelerate comparative biology, breeding and biotechnology.A major opportunity for a sustainable energy and biomaterials economy in many parts of the world lies in a better understanding of the molecular basis of superior growth and adaptation in woody plants. Part of this opportunity involves species of Eucalyptus L'Hér, a genus of woody perennials native to Australia 1 . The remarkable adaptability of eucalypts coupled with their fast growth and superior wood properties has driven their rapid adoption for plantation forestry in more than 100 countries across six continents (.20 million ha) 2 , making eucalypts the most widely planted hardwood forest trees in the world. The subtropical E. grandis and the temperate E. globulus stand out as targets of breeding programmes worldwide. Planted eucalypts provide key renewable resources for the production of pulp, paper, biomaterials and bioenergy, while mitigating human pressures on native forests 3 . Eucalypts also have a large diversity and high concentration of essential oils (mixtures of mono-and sesquiterpenes), many of which have ecological functions as well as medicinal and industrial uses. Predominantly outcrossers 1 with hermaphroditic animal-pollinated flowers, eucalypts are highly heterozygous and display pre-and postzygotic barriers to selfing to reduce inbreeding depression for fitness and survival 4 .To mitigate the challenge of assembling a highly heterozygous genome, we sequenced the genome of 'BRASUZ1', a 17-year-old E. grandis genotype derived from one generation of selfing. The availability of annotated forest tree genomes from two separately evolving rosid lineages, Eucalyptus (order Myrtales) and Populus (order Malpighiales 5 ), in combination with genomes from domesticated woody plants (for example, Vitis, Prunus, Citrus), provides a comparative foundation for addressing

We used Arabidopsis thaliana mutants to examine how a photosynthetic regulatory process, the qE-type or DeltapH-dependent nonphotochemical quenching, hereafter named feedback de-excitation, influences plant fitness in different light environments. We show that the feedback de-excitation is important for plant fitness in the field and in fluctuating light in a controlled environment but that it does not affect plant performance under constant light conditions. Our findings demonstrate that the feedback de-excitation confers a strong fitness advantage under field conditions and suggest that this advantage is due to the increase in plant tolerance to variation in light intensity rather than tolerance to high-intensity light itself.

733I.733II.734III.735IV.735V.735VI.737VII.739VIII.740IX.742X.744745References745 Summary Plant secondary metabolites (PSMs) are ubiquitous in plants and play many ecological roles. Each compound can vary in presence and/or quantity, and the composition of the mixture of chemicals can vary, such that chemodiversity can be partitioned within and among individuals. Plant ontogeny and environmental and genetic variation are recognized as sources of chemical variation, but recent advances in understanding the molecular basis of variation may allow the future deployment of isogenic mutants to test the specific adaptive function of variation in PSMs. An important consequence of high intraspecific variation is the capacity to evolve rapidly. It is becoming increasingly clear that trait variance linked to both macro‐ and micro‐environmental variation can also evolve and may respond more strongly to selection than mean trait values. This research, which is in its infancy in plants, highlights what could be a missing piece of the picture of PSM evolution. PSM polymorphisms are probably maintained by multiple selective forces acting across many spatial and temporal scales, but convincing examples that recognize the diversity of plant population structures are rare. We describe how diversity can be inherently beneficial for plants and suggest fruitful avenues for future research to untangle the causes and consequences of intraspecific variation.

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BackgroundTerpenoids are abundant in the foliage of Eucalyptus, providing the characteristic smell as well as being valuable economically and influencing ecological interactions. Quantitative and qualitative inter- and intra- specific variation of terpenes is common in eucalypts.ResultsThe genome sequences of Eucalyptus grandis and E. globulus were mined for terpene synthase genes (TPS) and compared to other plant species. We investigated the relative expression of TPS in seven plant tissues and functionally characterized five TPS genes from E. grandis. Compared to other sequenced plant genomes, Eucalyptus grandis has the largest number of putative functional TPS genes of any sequenced plant. We discovered 113 and 106 putative functional TPS genes in E. grandis and E. globulus, respectively. All but one TPS from E. grandis were expressed in at least one of seven plant tissues examined. Genomic clusters of up to 20 genes were identified. Many TPS are expressed in tissues other than leaves which invites a re-evaluation of the function of terpenes in Eucalyptus.ConclusionsOur data indicate that terpenes in Eucalyptus may play a wider role in biotic and abiotic interactions than previously thought. Tissue specific expression is common and the possibility of stress induction needs further investigation. Phylogenetic comparison of the two investigated Eucalyptus species gives insight about recent evolution of different clades within the TPS gene family. While the majority of TPS genes occur in orthologous pairs some clades show evidence of recent gene duplication, as well as loss of function.Electronic supplementary materialThe online version of this article (doi:10.1186/s12864-015-1598-x) contains supplementary material, which is available to authorized users.

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The eucalypts, which include Eucalyptus, Angophora and Corymbia, are native to Australia and Malesia and include over 800 named species in a mixture of diverse and depauperate lineages. We assessed the fit of the eucalypt taxonomic classification to a phylogeny of 711 species scored for DNA sequences of plastid matK and psbA–trnH, as well as nuclear internal transcribed spacer and external transcribed spacer. Two broadly similar topologies emerge from both maximum likelihood and Bayesian analyses, showing Angophora nested within Corymbia, or Angophora sister to Corymbia. The position of certain species-poor groups on long branches fluctuated relative to the three major Eucalyptus subgenera, and positions of several closely related species within those subgenera were unstable and lacked statistical support. Most sections and series of Eucalyptus were not recovered as monophyletic. We calibrated these phylogenies against time, using penalised likelihood and constraints obtained from fossil ages. On the basis of these trees, most major eucalypt subgenera arose in the Late Eocene and Early Oligocene. All Eucalyptus clades with taxa occurring in south-eastern Australia have crown ages <20million years. Several eucalypt clades display a strong present-day geographic disjunction, although these clades did not have strong phylogenetic statistical support. In particular, the estimated age of the separation between the eudesmids (Eucalyptus subgenus Eudesmia) and monocalypts (Eucalyptus subgenus Eucalyptus) was consistent with extensive inland water bodies in the Eocene. Bayesian analysis of macroevolutionary mixture rates of net species diversification accelerated in five sections of Eucalyptus subgenus Symphyomyrtus, all beginning 2–3million years ago and associated with semi-arid habitats dominated by mallee and mallet growth forms, and with open woodlands and forests in eastern Australia. This is the first time that a calibrated molecular study has shown support for the rapid diversification of eucalypts in the recent past, most likely driven by changing climate and diverse soil geochemical conditions.