Diurnal and circadian regulation of wood formation in Eucalyptus trees

Abstract

Wood is one of the most important products of world trade, due to its countless uses as a source of timber, fibre, and renewable energy. In addition to its economic importance, the formation of wood represents a global carbon sink which reduces the excess atmospheric CO2 that contributes to global warming. The formation of wood or xylogenesis is a complex example of cell differentiation, controlled by multiple interacting environmental factors and the coordinated expression of hundreds of genes. Genomic studies have proved a valuable tool in identifying the genes associated with xylogenesis. The expression of these genes has been shown to under strict spatial regulation in a developmental-stage specific fashion. Despite recent advances in the understanding of this process, there remains much to learn about the cellular, molecular and developmental processes involved. While the spatial regulation of wood formation has been well described, less attention has been devoted to the temporal regulation of this process. Most organisms are known to match their activities to the daily oscillation of night and day in what is known as a diurnal rhythm. A subset of these diurnal rhythms are termed circadian rhythms, and persist in the absence of environmental time cues, with a period of approximately 24 hours. Circadian rhythms are endogenous in nature, being generated by a small number of central oscillator genes, and illustrate an organism's ability to measure time. Circadian rhythms are found across a wide taxonomic spectrum, and are believed to confer an adaptive benefit, possibly due to the ability to anticipate regular changes in the external environment. As wood formation is a major sink for the products of light driven photosynthesis, it represents a likely target for circadian control in plants. A large proportion of photosynthesis genes themselves are known to be under circadian control, as are several cell wall formation genes. Most studies of temporal rhythms in plants, however, have used the herbaceous model species Arabidopsis, which does not have a woody stem. It is likely, therefore, that the circadian control of many wood formation genes remains to be discovered. We used a spotted cDNA mIcroarray carryIng 2608 elements to quantitatively measure daily changes in transcript abundance in the wood-forming tissues of a fast growing, Eucalyptus hybrid. Eucalyptus is a large genus of tree species, many of which are of great economic importance, and are widely grown in plantations for solid timber and pulp production. We found that almost ten percent of the genes on the microarray showed significant daily changes in expression (-loglOP>3.74). These genes included Eucalyptusorthologues of the Arabidopsis central clock genes CCA1 (CIRCADIAN CLOCK ASSOCIATED 1) and GIGANTEA (GI) which cycled with a period and phase matching that seen in Arabidopsis. The remaining genes were involved in pathways including carbohydrate metabolism, hormone signalling, transcription regulation and wood formation. The types of genes that were seen to be diurnally influenced, suggests a role for circadian control of various important plant metabolic pathways, including aspects of carbon allocation to wood formation.