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Leaf Photosynthesis and Simulated Carbon Budget of Gentiana Journal of Plant Ecology Leaf photosynthesis and simulated VOLUME 2, NUMBER 4, PAGES 207–216 carbon budget of Gentiana DECEMBER 2009 doi: 10.1093/jpe/rtp025 straminea from a decade-long Advanced Access published on 20 November 2009 warming experiment available online at www.jpe.oxfordjournals.org Haihua Shen1,*, Julia A. Klein2, Xinquan Zhao3 and Yanhong Tang1 1 Environmental Biology Division, National Institute for Environmental Studies, Onogawa 16-2, Tsukuba, Ibaraki, 305-8506, Japan 2 Department of Forest, Rangeland & Watershed Stewardship, Colorado State University, Fort Collins, CO 80523, USA and 3 Northwest Plateau Institute of Biology, Chinese Academy of Sciences, Xining, Qinghai, China *Correspondence address. Environmental Biology Division, National Institute for Environmental Studies, Onogawa 16-2, Tsukuba, Ibaraki, 305-8506, Japan. Tel: +81-29-850-2481; Fax: 81-29-850-2483; E-mail: [email protected] Abstract Aims 2) Despite the small difference in the temperature environ- Alpine ecosystems may experience larger temperature increases due ment, there was strong tendency in the temperature acclima- to global warming as compared with lowland ecosystems. Informa- tion of photosynthesis. The estimated temperature optimum of tion on physiological adjustment of alpine plants to temperature light-saturated photosynthetic CO2 uptake (Amax) shifted changes can provide insights into our understanding how these ;1°C higher from the plants under the ambient regime to plants are responding to current and future warming. We tested those under the OTCs warming regime, and the Amax was sig- the hypothesis that alpine plants would exhibit acclimation in pho- nificantly lower in the warming-acclimated leaves than the tosynthesis and respiration under long-term elevated temperature, leaves outside the OTCs. and the acclimation may relatively increase leaf carbon gain under 3) Temperature acclimation of respiration was large and sig- warming conditions. nificant: the dark respiration rates of leaves developed in the warming regime were significantly lower than leaves from the Methods ambient environments. Open-top chambers (OTCs) were set up for a period of 11 years to 4) The simulated net leaf carbon gain was significantly lower artificially increase the temperature in an alpine meadow ecosystem. in the in situ leaves under the OTCs warming regime than un- We measured leaf photosynthesis and dark respiration under differ- der the ambient open regime. However, in comparison with ent light, temperature and ambient CO2 concentrations for Gentiana the assumed non-acclimation leaves, the in situ warming- straminea, a species widely distributed on the Tibetan Plateau. Max- acclimated leaves exhibited significantly higher net leaf carbon imum rates of the photosynthetic electron transport ( Jmax), RuBP car- gain. boxylation (Vcmax) and temperature sensitivity of respiration Q10 5) The results suggest that there was a strong and significant were obtained from the measurements. We further estimated the leaf temperature acclimation in physiology of G. straminea in re- carbon budget of G. straminea using the physiological parameters sponse to long-term warming, and the physiological acclima- and environmental variables obtained in the study. tion can reduce the decrease of leaf carbon gain, i.e. increase relatively leaf carbon gain under the warming condition in the Important findings alpine species. 1) The OTCs consistently elevated the daily mean air temper- ature by ;1.6°C and soil temperature by ;0.5°C during the Keywords: alpine plant d acclimation d experimental growing season. warming d open-top chamber d photosynthesis d temperature Ó The Author 2009. Published by Oxford University Press on behalf of the Institute of Botany, Chinese Academy of Sciences and the Botanical Society of China. All rights reserved. For permissions, please email: [email protected] 208 Journal of Plant Ecology understood, though a few studies suggest that the acclimation INTRODUCTION response to increasing temperature is not so different among Global mean temperatures are increasing and are predicted to alpine and lowland herbaceous species (Arnone and Ko¨ rner increase further into the future (IPCC 2007). Temperature el- 1997; Collier 1996; Larigauderie and Ko¨ rner 1995). Our pri- evation will affect plant carbon budgets through its effects on mary aim was thus to assess physiological acclimation of alpine photosynthesis and respiration (e.g. Luo 2007). The change of plants and its consequence to leaf carbon gain under warming terrestrial vegetation carbon budgets can, in turn, directly conditions. We hypothesized that alpine plants would exhibit feedback to affect the CO2 concentration in the atmosphere photosynthetic and respiration acclimation to temperature el- (IPCC 2007; Luo 2007). Understanding the changes of plant evation, and the acclimation should increase the leaf carbon photosynthesis and respiration in responding to a warming en- gain. To test the hypothesis, we used one of the most common vironment is thus essential to further understand and predict species on the Qinghai-Tibetan Plateau, Gentiana straminea, the future carbon dynamics in the atmosphere (Atkin and growing within the open-top chambers (OTCs) for a period Tjoelker 2003; Dewar et al. 1999; Gunderson et al. 2000). of 11 years. Photosynthetic and respiration rates generally increase with a temporal increase of environmental temperature below the PLANT MATERIALS AND METHODS optimal temperature when other resources are not limiting Study site and plant materials (e.g. Berry and Bjo¨ rkman 1980; Hikosaka et al. 2006). The physiological processes can also result in acclimation to The field site was an alpine Kobresia humilis meadow (latitude long-term changes in the temperature environment (e.g. 37°37#N, longitude 101°12#E, altitude 3250 m), which is lo- cated at the northeastern edge of the Qinghai-Tibetan Plateau. Campbell et al. 2007; Larigauderie and Ko¨ rner 1995; Pearcy The annual mean air temperature of this site is 1.6°C and the 1977; Xiong et al. 2000; Zhou et al. 2007). The temperature sen- À annual precipitation is 562 mm (Klein et al. 2004). The OTCs, sitivity and acclimation degree of photosynthesis differ from each being 1.5 m diameter, 40 cm height and 1.0-mm thick that of respiration (Bruhn et al. 2007; Morison and Morecroft fiberglass (Solar Components Corporation, Manchester, 2006; Way and Sage 2008). Any changes in the two processes NH), were set up in 1997. Similar OTCs have been an impor- may eventually change the plant carbon budget, which alters tant method for simulating warming in the field, especially in the overall ecosystem carbon budget (Atkin et al. 2006; Dewar tundra or alpine ecosystems (Allen et al. 1992; Arft et al. 1999; et al. 1999; Loveys et al. 2002). The degree of acclimation in Ceulemans and Mousseau 1994; Walker et al. 2006). The OTCs photosynthesis and respiration is, therefore, an important de- elevated growing season averaged daily air temperature by terminant to predict the future response of ecosystems to ele- 0.6–2.0°C, and growing season averaged maximum daily air vated temperatures. Moreover, the plant-level response temperature by 1.9–7.3°C. For additional details on the study provides a mechanistic understanding that can be masked at site, the experimental design, the micro-climate effects of the the ecosystem level. treatments and the community and ecosystem responses, see Since the magnitude of photosynthetic and respiratory ac- Klein et al. (2004, 2005, 2007, 2008). climation varies with species and other environmental varia- Gentiana straminea Maximum. (Gentianaceae) is one of the bles (Berry and Bjo¨rkman 1980), these processes are still common forbs in the meadow and is a genus commonly found poorly understood, especially under field conditions and/or across the Tibetan Plateau and the Himalayan region. This spe- for long-term acclimation. Laboratory experiments, in which cies is also a medicinal plant that provides an important eco- the growth temperature often differs by 5–10°C, can provide system service in the region (Klein et al. 2008). Gentiana useful information for predicting the effects of global warming straminea grows its linear-shaped leaves diagonally from the on plant carbon budgets (see reviews from Hikosaka et al. 2006; soil surface to the top of the canopy. Its mature leaves are Medlyn et al. 2002). The information, however, can also be of 20–30 cm long. The inclination angle of fully expanded leaves limited use partly because the projected magnitude of global is 4969°, the ratio of length to width is approximately 4–5 (Cui warming in the short term is smaller than the experimental et al. 2004). conditions simulated in these laboratory studies. Therefore, understanding acclimation under field conditions with more Assessment of the micro-environmental parameter realistic temperature elevation and, if possible, under long- of inside and outside of OTCs term warming condition is needed to predict plant response We installed six quantum sensors (GaAsP Photodiode G1118, to potential climate change in the future. Hamamatsu City, Japan) on the horizontal surface at the The impact of global warming on terrestrial ecosystems has average height of the G. straminea leaves, three temperature been reported to be greater in arctic tundra and high mountain sensors (copper–constantan thermocouples) on the leaf regions than at low latitudes or altitudes (e.g. Beniston 2006; surface, three temperature sensors 20 cm above the soil Maxwell 1992; Mitchell et al. 1990). Nevertheless, plant surface and three temperature sensors (copper–constantan responses—especially ecophysiological responses to tempera- thermocouples) at a depth of 5 cm in the soil on the inside ture elevation in these cold ecosystems—have been scarcely and outside of the OTCs, respectively. All quantum sensors Shen et al. | Leaf photosynthesis response to experimental warming 209 were calibrated under sunlight and artificial shade against point in the absence of day respiration. Kc and Ko are Michaelis a standard quantum sensor (Li-Cor Model 190S, Li-Cor, constants for carboxylation and oxygenation, respectively. Lincoln, NE; Tang and Washitani 1995).
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