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Plant Respiration and Climate Change Effects

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Plant Respiration and Climate Change Effects Risø-R-1332(EN) Plant Respiration and Climate Change Effects Dan Bruhn Ph.D. thesis Plant Research Department Risø National Laboratory Botanical Institute University of Copenhagen Risø National Laboratory, Roskilde April 2002 Abstract The ongoing climate changes can affect many plant physiological processes. In turn, these effects on plants may result in a feedback between the climate change and the vegetation. Plant respiration is one of the key processes in terms of an understanding of plant growth and functioning in a future climate. Plant respiration can release up to about half of the assimilated carbon by photosynthesis on a daily basis. Thus potential effects of e.g. changed air temperature and atmos- pheric [CO2] are important to investigate for predictions of respiratory release of CO2 to the atmos- phere. This is further emphasised by the fact that global terrestrial respiratory CO2 release is about ten times as high as the antropogenic contribution of CO2 to the atmosphere on an annual basis. Plants therefore constitute an important component of the global carbon cycle. Short- and long-term (direct and indirect, respectively) effects of temperature and [CO2] on plant respiration were investigated in a number of plant species. The experiments tested effects of ei- ther temperature and/or [CO2] from the level of individual respiratory enzymes, isolated mitochondria, whole-tissue (above- and below-ground tissue), and up to the whole canopy level in forests. The short-term effects of elevated atmospheric [CO2] on plant respiration appeared to be less than suggested so far in the literature. This was true both at the tissue-level and for intact mitochon- dria. Respiratory enzymes can, however, be affected already at low [CO2]. Bicarbonate rather than CO2 was found to be the carbon species that affected one of the respiratory enzymes. These effects did not manifest itself at the tissue level, though, due to low degrees of control on the whole respiratory process exerted by the particular enzymes. Plant respiration on the other hand was affected by long-term growth at elevated atmospheric [CO2]. The findings of the reduced plant respiration at the leaf level were consistent with the literature and potential causes are discussed. Short-term effects of temperature on plant respiration were demonstrated to be dependent on the actual measurement temperature. Further, it is shown that mitochondrial leaf respiration in dark- ness and light differ substantially in the temperature sensitivity with the former being the far most sen- sitive. This has implications for modelling CO2 exchange between vegetation and atmosphere as dem- onstrated here, since this has so far been neglected. The underlying mechanisms per se call for further investigation as the much lower temperature sensitivity of leaf respiration in the light to a great degree may influence the daily carbon budget of plants. Long-term effects of temperature resulted in respiratory acclimation in a number of species. Respiratory acclimation appeared not to occur to any one single type of growth temperature. The im- plications of this finding in combination with the timing of acclimation are discussed for modelling respiratory CO2 release. In addition are some new methods for calculation of the degree of acclimation proposed. ISBN 87-550-3032-7 ISBN 87-550-3033-5 (Internet) ISSN 0106-2840 Print: Pitney Bowes Management Services Danmark A/S, 2002 Contents Abstract 2 Preface 4 1 Introduction 5 Climate Change in a Plant Ecophysiological Perspective (2000) In: Climate Change Research - Danish Contributions. Jørgensen A.M.K., Fenger J. & Halsnæs (eds.), DMI, Ministry of Transport, Gads Forlag. 2 Introduction to Plant Respiration 29 3 Phosphorus uptake by arbuscular mycorrhizal hyphae does not increase when the host plant grows under atmospheric CO2 enrichment 35 New Phytologist (2002) 154 In press. 4 Does the direct effect of atmospheric CO2 concentration on leaf respiration vary with temperature? Responses in two species of Plantago that differ in relative growth rate 45 Physiologia Plantarum 114: 57-64, 2002 5 Direct effects of [CO2] on enzyme activity and O2 uptake of isolated mitochondria and intact tissues: Comparing the response of Solanum tuberosum L. tubers and Glycine max L. cotyledons 53 Plant Physiology, submitted 6 Determining the temperature that leaf and root respiration acclimates to in several contrasting plant species differing in photosynthesis type and growth characteristics 69 Global Change Biology, submitted 7 Partial Respiratory Acclimation to Temperature by Plants - Discussion of Definitions, Calculations, and Applications 89 Prepared for Functional Ecology 8 The interaction of temperature and irradiance on leaf respiration is important to take into account when modeling CO2 flux between atmosphere and vegetation 101 Prepared for Tree Physiology 9 Supplementary Discussion, Conclusions, and Future Research 119 Appendix 133 Acknowledegements 140 Risø-R-1332(EN) 3 Preface This Ph.D.-thesis is submitted as part of the requirements for a Ph.D. degree at University of Copenhagen, Botanical Institute, Denmark. The project period was April 23rd 1999 to April 22nd 2002. The project has been carried out at the Plant Environment Interactions Programme, Plant Research Department, Risø National Laboratory, Roskilde, Denmark and Department of Biology, The University of York, York, UK. The Danish Research Academy, Risø National Laboratory, European Science Foun- dation, and Nordic Academy of Advanced Study have financially supported this project. The outline of the thesis is: Chapter 1 is an introduction to climate change and plant ecophysiology in gen- eral. Chapter 2 gives a brief introduction to plant respiration, its control, and methodological aspects. Chapter 3 to 5 deals mainly with effects of [CO2] on plant respiration. Chapter 3 treats the indirect (or long-term) effects and Chapter 4 & 5 treat the direct effects of [CO2]. Chapter 6 to 8 is about effects of tem- perature on plant respiration (Chapter 4 does also treat temperature effects to some extent). Chapter 6 is a study about the respiratory acclimation to tempera- ture. Chapter 7 treats respiratory acclimation to temperature in a theoretical manner. Chapter 8 is about direct (short-term) effects of temperature in combi- nation with light and the only chapter where effects are scaled up to the canopy level. Finally, Chapter 9 gives a personal supplementary discussion to the dis- cussion parts of Chapter 3 to 8. Chapter 9 also contains the conclusions from my own experiments separated out into each of the four main topics of this the- sis, i.e. short- and long-term effects of [CO2] and short- and long-term effects of temperature on plant respiration. In addition is presented a personal view on future research. In the appendix are included two Danish articles about climate change research within plant ecophysiology ongoing at Risø. These two articles are written for a broader forum than plant biologist. The thesis finishes with my acknowledgements to a number of people, all of who have helped me during this Ph.D. project at various stages. Research is both investigation and publication. Thus, the changing lay- out of the different chapters throughout the thesis reflects the stage of publica- tion of the individual chapters. Chapter 1 and 3 to 8 are all meant to be pub- lished separately. 4 Risø-R-1332(EN) 1 Introduction Risø-R-1332(EN) 5 6 Risø-R-1332(EN) Risø-R-1332(EN) 7 8 Risø-R-1332(EN) Risø-R-1332(EN) 9 10 Risø-R-1332(EN) Risø-R-1332(EN) 11 12 Risø-R-1332(EN) Risø-R-1332(EN) 13 14 Risø-R-1332(EN) Risø-R-1332(EN) 15 16 Risø-R-1332(EN) Risø-R-1332(EN) 17 18 Risø-R-1332(EN) Risø-R-1332(EN) 19 20 Risø-R-1332(EN) Risø-R-1332(EN) 21 22 Risø-R-1332(EN) Risø-R-1332(EN) 23 24 Risø-R-1332(EN) Risø-R-1332(EN) 25 26 Risø-R-1332(EN) Risø-R-1332(EN) 27 28 Risø-R-1332(EN) 2 Introduction to Plant Respiration This is an introduction to the plant physiological process, respiration – the piv- otal point in this thesis. Plant respiration was briefly introduced in Chapter 1 as one of several plant physiological processes it is necessary to understand to an- swer questions about climate change at the plant ecophysiological level. This is in order to at higher levels of integration address ecological challenges of the ongoing global climate change. Potential short-term and long-term effects by climate changes were also discussed in Chapter 1. The following chapter is lim- ited to dark (mitochondrial) respiration in plants. Photorespiration was shortly introduced in Chapter 1. Overview of Plant Respiration In respiration the relatively reduced organic compounds, carbohydrates, organic acids, and proteins are oxidized and some of the released energy is used for growth and maintenance. In plants the main substrate for respiration can be said to be the disaccharide, sucrose, which is the predominant sugar translocated via the phloem in most plants. The free energy released is transiently stored in ATP, which is the energy currency used for both cellular maintenance and growth. Sucrose may be fully oxidized to give CO2 and the net reaction is then: C12H22O11 + 12 O2 → 12 CO2 + 11 H2O Respiration takes place as a multistep process. Thereby the free energy is released in several steps preventing cellular damage. The three main stages are glycolysis, the TCA (tricarboxylic acid) cycle, and the oxidative phosphory- lation including the mitochondrial electron transport chain. The intermediates can enter the respiratory process at various points as substrates; equally can in- termediates leave the respiratory chain to function as precursors for biosynthesis of various compounds. Respiration is hereby coupled to many other metabolic pathways. Glycolysis takes place in the cytosol, but some glycolytic enzymes are also located in the plastids. Thus, substrates can also enter the respiratory proc- ess directly from both chloroplasts and amyloplasts.
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