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Cold-Induced Photoinhibition, Pigment Chemistry, Growth and Nutrition of Eucalyptus Nitens and E

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Cold-Induced Photoinhibition, Pigment Chemistry, Growth and Nutrition of Eucalyptus Nitens and E Cold-induced photoinhibition, pigment chemistry, growth and nutrition of Eucalyptus nitens and E. globulus seedlings during establishment Dugald C. Close Submitted in fulfilment of the requirements for the degree of Doctor of Philosophy Schools of Agricultural and Plant Science .and CRC for Sustainable Production Forestry University of Tasmania Declarations This thesis contains no material which has been accepted for a degree or diploma by the University of Tasmania or any other institution. To the best of my knowledge and belief this thesis contains no material previously published or written by another person except where due acknowledgement is made in the text of the thesis. Dugald C. Close This thesis may be made available for loan and limited copying in accordance with the Copyright Act 1968. Dugald C. Close Abstract Australia is aiming to treble plantation wood production by 2020. Eucalyptus globulus Labill. and E. nitens (Deane and Maidem) Maiden are the predominant plantation species in southern Australia. This thesis describes physiological strategies employed by these species in response to cold-induced photoinhibition during seedling establishment. A series of experiments was conducted on seedlings pre- hardened in the nursery. Their physiological and growth responses after planting in the field was investigated. A field trial was established at 350 m asl in early spring 1997. Severe cold-induced photoinhibition caused photodamage which restricted growth of non-hardened E. globulus. Artificial shading alleviated cold-induced photoinhibition and photodamage in both E. globulus and E. nitens, and increased growth in E. globulus. Before planting, nutrient-starved E. nitens were photoinhibited and had high anthocyanin levels. Increased photoinhibition was not measured after planting because of sustained xanthophyll activity and/or light attenuation by high anthocyanin levels. In other treatments changes in anthocyanin levels were related to the severity of cold- induced photoinhibition. Relative to E. nitens, growth of E. globulus was more affected by cold-induced photoinhibition and photodamage. This was possibly due to inherently low levels of carotenoids and lack of acclimation to cold temperatures. The effects of shading on E. globulus and the absence of any effect of cold-hardening on E. nitens stresses the importance of incident light and pigment levels in cold-induced photoinhibition. II In a second field trial, an early winter planting of E. nitens was established at 700 m asl in June 1998. Shading may have increased biomass production because of alleviation of cold-induced photoinhibition. Growth in non-shaded than shaded seedlings was greater overall due to higher biomass production in spring and summer. Seedlings grew taller when shaded due to apical dominance. Fertilised seedlings produced more biomass in the field than non-fertilised seedlings. Low growth rates of E. nitens during winter at 700 m asl were associated with high NPQ and sustained xanthophyll activity; photooxidation of chlorophylls, xanthophylls and 13-carotene (which decreased light absorption), and increases in lutein and neoxanthin (which indicated an antioxidant role). In general fertilised seedlings had higher pigment levels which maintained higher levels of light utilisation and dissipation. A controlled environment experiment which induced cold-induced photoinhibition, confirmed that galloylglucoses and flavonoids can act as antioxidants during seedling establishment. Sideroxylonals were also implicated in this role. Anthocyanin kinetics during seedling establishment indicated absorption of irradiance between 400 and 590 nm during periods of greatest cold-induced photoinhibition. Chemical fractionation of leaf N and P indicated that E. globulus is more efficient at acquiring N and P than E. nitens. After planting, re-translocation of stored foliar protein N and inorganic P to roots occurred in both species. Greater amounts of re- translocation in fertilised seedlings may contribute to their superior growth in the field. Ill Planting seedlings involves risk. Planting nutrient-starved seedlings may decrease the risk of severe cold-induced photoinhibiiton and photodamage. However, this is at the expense of optimal growth performance. Planting altitude and season will determine whether fertilised or nutrient-starved seedlings should be planted. IV Acknowledgements During the course of this project I was supported by an Australian Postgraduate Award (APA) and a Co-operative Research Centre for Sustainable Production Forestry (CRC-SPF) scholarship. Substantial in-kind support was provided by North Forest Products (NFP) and North Eucalypt Technologies (NET) in the form of experimental sites, plant material and technical assistance. Many people have helped make this project a success: • Dr. Chris Beadle from whom I have learnt so much — thankyou; • Dr. Noel Davies for generous and expert assistance with HPLC and MS analysis; • Drs. Phil Brown, Mark Hovenden and Greg Holz for advice and support whenever requested; • Dr. Steve Wilson for advice during the first year of my project; • Ms. Ann Wilkinson and Mr. Petr Otahal for laboratory analysis assistance far beyond the call of duty; • Mr. Dale Worledge for weather station construction and operation, and electric fence engineering; • Dr. David Ratkowsky for expert statistical advice; • Ms. Maria Cherry and Mr. Sven Ladiges for assistance in field measurements; • Mr. Kelsey Joyce for assistance with frost tolerance testing, Mr. Chris Barnes for weed control advice, Ms. Renate Van Riet and Mr. Tim Hingston for seedling managment at the Ridgley nursery, Mr. Peter O'Neill and Mr. Terry Williams for seedling planting and tree shelter erection and Mr. Terry Williams for his excellent assistance in the construction of the electrified fence, all of North Eucalypt Technologies; • Messrs. Ian Ravenwood, Gavin Raison and Andrew Moore of North Forest Products for technical support at the North Forest Product's Somerset nursery; • Mr. Ian Cummings and Ms. Tracy Jackson for technical support at the plant science glasshouse complex; • Messrs. Phil Andrews and David Wilson for technical support at the agricultural science horticultural research station; • Ms. Karen Barry for the use of tetra-galloylglucose standard data; • Drs Bart Eschler and William Foley for sideroxylonal standard; • Mr. Trevor Bailey of the Australian Antarctic Division for the loan of their Pro 200 homogeniser; • Mr. Philip Coles for computers systems support; • Ms. Tara Simmul for manufacturing the porous cloth bags; • Mr. and Mrs. Greame Jones for providing accommodation in Burnie; • Mr. Allan Mills and Professor Robert Menary for expertly fuelling my interest in plant physiology. Primarily I have to thank my family: Owie (Kietha) Scan, Rob, Gail, Joc (Oliver), Rundy (David) and Bazza (Harriet) Close for being my family, particularly Rob whom assisted with time management and as a general sounding board and Owie for the many stimulating lunch time conversations. Kate Sice, who put up with the most, thankyou for being there. VI Publications Refereed journal articles Chapter 3: Close, D.C., Beadle, C.L., Brown, P.H. and Holz, G.K. (2000). Cold-induced photoinhibition affects establishment of Eucalyptus nitens (Deane and Maiden) Maiden and E. globulus Labill. Trees: Structure and Function 15: 32-41. Chapter 7: Close, D.C., Davies, N.W. and Beadle, C.L. Physiological antioxidants and compounds deterring herbivory and in light attenuation? Australian Journal of Plant Physiology 24: (In press). Chapter 9: Close, D.C., Beadle, C.L., Holz, G.K. and Ravenwood, I.C. (1999). A photobleaching event at the North Forest Products' Somerset nursery reduces growth of Eucalyptus globulus seedlings. Tasforests 11: 59-68. Manuscripts for refereed journals in internal CSIRO Review Chapter 3: Close DC, Beadle CL, Holz GK and Brown PH (2001). Effect of shadecloth tree shelters on cold-induced photoinhibition, foliar anthocyanin and growth of Eucalyptus globulus Labill. and E. nitens (Deane and Maiden) Maiden seedlings during establishment. Forest Ecology and Management: (internal CSIRO review). VII Chapter 5: Close, D.C., Beadle, C.L. and Hovenden, M.J. (2001). Cold-induced photoinhibition and foliar pigment dynamics of Eucalyptus nitens seedlings during establishment. Australian Journal of Plant Physiology: (internal CSLRO review). Chapter 5: Close, D.C., Beadle, C.L., Hovenden, M.J. and Brown P.H. (2001). Cold-induced photoinhibition, nutrition and growth analysis of Eucalyptus nitens (Deane and Maiden) Maiden seedlings during establishment. Tree Physiology: (internal CSIRO review). Chapter 6: Close, D.C., Beadle, C.L. and Hovenden, M.J. (2001). Sustained Xanthophyll Cycle Engagement is an overwintering strategy of Eucalyptus nitens (Deane and Maiden) Maiden seedlings. Oecologia: (internal CSIRO review). Chapter 8: Close, D.C., Beadle, C.L., Davies, N.W. and Hovenden, M.J. (2001). Effects of rapid induction and subsequent recovery from cold-induced photoinhibition on xanthophylls, anthocyanins, galloylglucoses, flavonols and sideroxylonals of Eucalyptus nitens seedling foliage. Plant, Cell and Environment: (internal CSIRO review). VIII Chapters 4, 5 and 8: Close, D.C. and Beadle, C.L. (2001). Partitioning of nitrogen and phosphorus chemical fractions in Eucalptus seedlings during spring and winter field trials and growth chamber trials. Communications in Soil Science and Plant Analysis: (internal CSIRO review). Conference publications Chapter 3: Close,
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