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Diel Leaf Growth Cycles in Clusia Spp. Are Related to Changes Between C3 Photosynthesis and Crassulacean Acid Metabolism During Development and During Water Stress

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Diel Leaf Growth Cycles in Clusia Spp. Are Related to Changes Between C3 Photosynthesis and Crassulacean Acid Metabolism During Development and During Water Stress Plant, Cell and Environment (2008) 31, 484–491 doi: 10.1111/j.1365-3040.2008.01777.x Diel leaf growth cycles in Clusia spp. are related to changes between C3 photosynthesis and crassulacean acid metabolism during development and during water stress ACHIM WALTER1, MAJA M. CHRIST1, UWE RASCHER1, ULRICH SCHURR1 & BARRY OSMOND1,2 1Institute of Chemistry and Dynamics of the Geosphere ICG-3: Phytosphere, Research Center Jülich GmbH, 52425 Jülich, Germany and 2School of Biochemistry and Molecular Biology, Australian National University, Canberra, ACT 0200 Australia ABSTRACT elucidation of the signature metabolism of CAM, with malic acid synthesis and degradation in mature leaves and stems This study reports evidence that the timing of leaf growth of these succulent plants in relation to environment and responds to developmental and environmental constraints development (Kluge & Ting 1978; Osmond 1978; Black & in Clusia spp. We monitored diel patterns of leaf growth Osmond 2003; Lüttge 2004; Holtum & Winter 2005). Less -crassulacean acid metabolism (CAM) in the facultative C3 attention has been given to the regulation of carbohydrate species Clusia minor and in the supposedly obligate CAM metabolism and its relation to growth (Sutton 1975a,b; species Clusia alata using imaging methods and followed Holtum, Smith & Neuhaus 2005). The fundamental ques- diel patterns of CO exchange and acidification. Developing 2 tion of how CAM plants preserve carbohydrate reserves in leaves of well-watered C. minor showed a C -like diel 3 the light for acid synthesis in the dark, while at the same pattern of gas exchange and growth, with maximum relative time providing carbon for growth, remains enigmatic growth rate (RGR) in the early night period. Growth (Borland & Dodd 2002). In particular, the evidence for slowed when water was withheld, accompanied by noctur- discrete pools of carbohydrates engaged in dark CO2 nal CO2 exchange and the diel acid change characteristic of fixation and growth (Deléens & Garnier-Dardart 1977; CAM. Maximum leaf RGR shifted from early night to early Deléens, Garnier-Dardart & Querioz 1979; Borland et al. in the day when water was withheld. In well-watered C. 1994) challenges all present models of metabolic compart- alata, similar changes in the diel pattern of leaf growth mentation in photosynthetic tissues. occurred with the development of CAM during leaf ontog- The first detailed diel leaf carbon allocation budgets eny. We hypothesize that the shift in leaf growth cycle that of a CAM plant (Sedum telephium L.) were published by photosynthesis to CAM is accompanies the switch from C3 Borland (1996), followed by similar comparisons of several mainly caused by the primary demand of CAM for sub- species that switch between C3 metabolism and CAM fixation and acid synthesis, thus strates for nocturnal CO2 (Borland & Dodd 2002). The ‘conflict of interest’ for carbo- reducing the availability of carbohydrates for leaf growth at hydrate metabolism in CAM plants (Borland & Dodd night. Although the shift to leaf growth early in the light is 2002) presumably involves three major ‘decision points’ presumably associated with the availability of carbohy- (regulatory compromises): conservation of carbohydrates drates, source–sink relationships and sustained diurnal acid for CAM; for growth of young leaves; and for export and levels in young leaves of Clusia spp. need further evaluation growth of the plant as a whole.The diel relationships among in relation to growth processes. these processes are likely to be complex and further con- founded by shifting patterns during leaf development and Key-words: gas exchange; image analysis. plant growth. There have been few comprehensive studies of these relationships. For example, Wang & Nobel (1996) INTRODUCTION examined the enzymology of CO2 assimilation and regu- lated carbohydrate metabolism in Opuntia ficus-indica (L.) It is now clear that crassulacean acid metabolism (CAM) Mill., and went on to examine the relationship between is a complex ecological, physiological and biochemical local CAM and sink activity in growing cladodes and process that has been widely selected in diverse plant taxa carbon sources in mature cladodes (Wang, Zhang & Nobel to conserve water in arid habitats. From the earliest studies, 1998). Recently,it has become clear that diel patterns of leaf plant physiologists have been preoccupied with the growth can be examined in detail by imaging methods Correspondence: A. Walter. Fax +49 2461 61 2492; e-mail: a.walter@ (Schmundt et al. 1998; Walter & Schurr 2005). They can fz-juelich.de be related to diel patterns in leaf carbohydrate status © 2008 The Authors 484 Journal compilation © 2008 Blackwell Publishing Ltd Diel leaf growth cycle in Clusia spp. 485 (Walter et al. 2005) and to the expression of genes involved MATERIALS AND METHODS in cell division (Matsubara et al. 2006). Most annual C 3 Plant material plants investigated up to now show the strongest growth activity at night–day transitions or at night when it seems C. minor L. and C. alata Pl. and Tr. plants were raised from likely that growth is mainly driven by the mobilization of seedlings obtained of the Clusia collection of the Botanical chloroplast starch reserves (Walter & Schurr 2005). In Garden of the Darmstadt University of Technology, rapidly growing leaves of Populus deltoides Bartr. ex Marsh, Darmstadt, Germany. C. alata is an obligate CAM plant, transient dips in leaf relative growth rate (RGR) were while C. minor is a facultative CAM plant, i.e. it can switch associated with low leaf glucose levels in the after- between C3 and CAM photosynthesis, depending on the noon, possibly indicating that demand for sugar export external conditions (Haag-Kerwer, Franco & Lüttge 1992; dominated over local requirements for leaf growth (Walter Haag-Kerwer et al. 1996; Herzog et al. 1999). In the ex- et al. 2005). periments presented here, we controlled the mode of Clearly, the complex diel patterns of acid metabolism and photosynthesis of C. minor by withholding soil water. carbohydrate demand in CAM plants present a challenge to Plants were grown in the Phytec greenhouse of the Insti- the analysis of diel leaf growth patterns. Initial studies of tute of Phytosphere Research (ICG-3; Research Center ‘obligate’ CAM plants, i.e. those that do not show transi- Jülich, Jülich, Germany) that was designed to cultivate tions from C3 to CAM during drought stress or develop- plants under controlled environmental conditions similar ment, showed distinctive diel growth patterns that were to field conditions. For example, a high transparency for dominated by growth during the day in leaves of Kalanchoe photosynthetically active radiation (PAR) and ultraviolet and cladodes of Opuntia spp. (Gouws et al. 2005). This was (UV) radiation (up to 97% in visible light and up to 35% in marked contrast to the nocturnal maximum and diurnal UV-B transmittance) was achieved by installing a specially decline in rapidly growing leaves of Mesembryanthemum formulated microstructured glass (Centrosolar Glas, Fürth, crystallinum L. in C3 mode, and further studies of the effect Germany), which also led to a homogeneous illumination of of C3–CAM transitions immediately suggested themselves. the cultivation area. Plants were supplied with slow-release However, leaves of M. crystallinum become twisted or opti- fertilizer and watered automatically with tap water every cally complex (because of epidermal bladders) during the second day. During growth and during greenhouse transition to CAM and are not ideal for the imaging analy- experiments, mean midday irradiance (PAR) averaged sis of diel patterns of leaf growth. 800 mmol m-2 s-1, with maximal values of 1200 mmol m-2 s-1; The CAM literature is now replete with studies of devel- temperature was set to 21/19 °C day/night but reached opmentally and environmentally inducible CAM (Dodd 30 °C on sunny days; relative humidity was 60%. When et al. 2002), and these are nowhere more evident than in sunlight was lower than 130 mmol m-2 s-1 artificial illumina- the tropical tree genus Clusia (Lüttge 2007). For example, tion was provided by SON-T Agro 400 W (Philips, Köln, Schmitt, Lee & Lüttge (1988) showed that mature leaves of Germany) lamps. Clusia minor (incorrectly identified as Clusia rosea; Lüttge For the analyses of growth and gas exchange in con- 2006) responded rapidly and reversibly to drought stress, trolled climate conditions, plants were transferred to a converting from a C3 gas exchange pattern to CAM. Our growth chamber 2–3 d before the onset of analyses. goal was to relate diel changes in metabolism to diel Chamber relative humidity was kept at 60%, and PAR at changes in leaf growth. However, we soon recognized that the top leaves ranged between 600 and 800 mmol m-2 s-1 the bulk of literature on CAM deals with gas exchange and provided by a mixture of SON-T Agro 400 W and HPI-T metabolic profiles of fully expanded, mature leaves and plus 400 W (Philips) lamps (temperature and day length are their function as source leaves for the growth of the whole specified for each experiment in the Results section). plant. Thus, studies of the expression of CAM at different Drought stress was imposed by not watering the plants after stages of development in the field focused on the youngest transfer to the growth chamber. Leaf growth of these plants fully expanded leaves of C. minor L. (Borland et al. 1992). stopped completely within 1–2 weeks after the last irriga- Other Clusia species, such as Clusia alata and Clusia hilari- tion, depending on the relation between plant size and soil ana Schltdl. are reputedly ‘obligate’ CAM plants (Lüttge volume available in the pot. 1999), but again, fully expanded leaves of the same size on individual C. hilariana plants of different sizes have been Gas exchange measurements studied (Berg et al. 2004). As far as we know, there have been no previous investigations of the patterns of gas Net CO2 exchange and transpiration was recorded using exchange or acidification during the expansion growth of a CMS 400 minicuvette system of Walz (Effeltrich, young leaves in these C3-CAM plants.
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