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Morphological Mechanism of Growth Response in Treeline Species Minjiang Fir to Elevated CO2 and Temperature

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Morphological Mechanism of Growth Response in Treeline Species Minjiang Fir to Elevated CO2 and Temperature Silva Fennica 45(2) research articles SILVA FENNICA www.metla.fi/silvafennica · ISSN 0037-5330 The Finnish Society of Forest Science · The Finnish Forest Research Institute Morphological Mechanism of Growth Response in Treeline Species Minjiang Fir to Elevated CO2 and Temperature Ying Hou, Jintao Qu, Zukui Luo, Chao Zhang and Kaiyun Wang Hou, Y., Qu, J., Luo, Z., Zhang, C. & Wang, K. 2011. Morphological mechanism of growth response in treeline species Minjiang fir to elevated CO2 and temperature. Silva Fennica 45(2): 181–195. To test whether and how morphological traits are linked with growth responses of plants to temperature and CO2 is important for understanding the mechanism underlying how plant growth will respond to global warming. In this study, using closed-top chambers to mimic future elevated CO2 and temperature, the growth response, morphological traits of Minjiang fir (Abies faxoniana Rehd.et Wils.) and the relationship of the two were investigated after two years of exposure to the single and combined elevation of CO2 and temperature. The results showed that biomass of Minjiang fir was 21%, 31%, and 35% greater than the control in elevated CO2, elevated temperature and the combination of elevated CO2 and temperature treatments, respectively. Elevated CO2 and temperature significantly affected the morphology of Minjiang fir, and a few morphological traits were highly correlated with growth responses. Larger branch angles at the upper layer, crown volume, and relative crown length contributed to positive growth responses to elevated CO2, while decreased specific leaf area (SLA) con- stricted any further growth response. Leaf morphological traits were more closely correlated with the response ratio than crown did in the elevated temperature, while in the combination of elevated CO2 and temperature, crown was more correlated with the response ratio than the leaf morphological traits. Thus, our results indicate that morphological traits may contribute differently to growth responses under different experimental conditions. Keywords Abies faxoniana, climate change, crown architecture, leaf morphology, response ratio Addresses Hou and Qu, Department of Life Sciences, Shangqiu Normal University, Shangqiu, China; Luo, School of Environment and Life Sciences, Kaili University, Kaili, China; Zhang and Wang, Shanghai Key Laboratory of Urbanization and Ecological Restoration, East China Normal University, Shanghai, China, and University of Eastern Finland, School of Forest Sciences, Joensuu, Finland E-mail (Wang) [email protected] Received 16 December 2010 Accepted 25 March 2011 Available at http://www.metla.fi/silvafennica/full/sf45/sf452181.pdf 181 Silva Fennica 45(2), 2011 research articles 1 Introduction enrichment through light interception, canopy atmosphere exchange processes, and nutrient and Plant growth is a complex phenomenon that inte- water absorption (Yokozawa et al. 1996, Gielen et grates many physiological and morphological al. 2002). Therefore, investigating the effects of processes. Growth responses of plants to elevated elevated CO2 and temperature on plant morphol- CO2 and temperature are closely related with ogy will lead us to gain a better understanding of their physiological and morphological adapta- how plant growth will respond to future changes tions. During the last several decades, many in climate. Such a study can also provide basic papers have been published describing the growth information for forest management and carbon response of plants to elevated CO2 and tempera- cycle models because some process-based growth ture (Gunderson and Wullschleger 1994, Idso and models include not only physiological indices, but Idso 1994, Pritchard et al. 1999, Ainsworth and also morphological indices, such as tree crown Long 2005 ). However, there have been highly size, leaf area, and branching (Landsberg and variable results which depend on the species, Waring 1997, Cropper 2000). growing conditions, and the duration of CO2 and Recently, some experiments were carried out temperature increase. There is abundant informa- to explore the relationship between morphology tion on the physiological mechanisms of growth and growth at elevated CO2 and temperature. For responses (Bazzaz and McConnaughay 1992, example, a study by Gielen et al. (2002) showed Idso and Kimball 1992, Wang et al. 2008), but that success of white poplar competing with black it remains unknown how and to what extent the poplar and euramerican poplar at elevated CO2 growth response is correlated with morphology. was due to longer, more horizontally orientated Although examining physiology is valuable in branches. Research on the relationship between the study of plant responses to elevated CO2 and crown structure and biomass revealed that crown temperature, the physiological parameters are a architecture affected tree growth through the relatively insensitive indicator of what is actually control of leaf area and its display for effec- happening to plant growth in terms of structure tive light capture and photosynthesis (Wang and and function (Stulen and Herton 1993, Sattler Jarvis 1990). The spatial distribution of leaf area and Rutishuser 1997). For example, differences and biomass within a crown might affect carbon in root depth and branching may influence pat- assimilation through its effect upon light intercep- terns of water and nutrient uptake independent of tion and net CO2 exchange rates for the whole total biomass (Tremmel and Bazzaz 1993). White tree (Reekie and Bazzaz 1989). However, the (1984) pointed out that in the vast literature of magnitude and direction of morphological effects plant responses to the environment, morphology on growth response to elevated CO2 and tempera- has been ignored. ture, or which component of morphological traits Morphology is the study of the size, shape, and are best related with growth response, remains structure of organisms or one of its parts (Sat- unanswered. tler 1996, Sattler and Rutishauser 1997). Various Our primary objective in this study was to environmental conditions make different pheno- investigate the morphological mechanism under- typic expression of the same genotype. Therefore, lying the growth response of Minjiang fir (Abies plant morphology has been recognized to be the faxoniana Rehd.et Wils.) to elevated CO2 and consequence of interactions between genotype, temperature. We first tested the effects of elevated environment and developmental history. Elevated CO2, elevated temperature and the combination CO2 alters plant structure through its effects on of the two on the growth and morphology of Min- both primary and secondary meristem of shoots jiang fir. Since structure determines the function, and roots. Similarly, elevated temperature may we predicted that the growth response would be contribute to developmental and morphological affected by its morphology. Thus, the relation- changes by influencing meristematic cells within ship between growth response and morphological the bud (Pritchard et al. 1999). These changes in traits was tested and discussed. We will address morphology, in turn, can influence the growth the following questions: 1) whether elevated CO2 responses of plants to CO2 and temperature and temperature affects the growth and morphol- 182 Hou et al. Morphological Mechanism of Growth Response in Treeline Species Minjiang Fir to Elevated CO2… ogy of Minjiang fir; 2) which and how morpho- out the growing season. In order to provide a logical traits are correlated with growth response comparison on the CON treatment and to detect to elevated CO2 and temperature. whether the chamber itself affects the morphology of plants, an “O” treatment was conducted with six trees and without a chamber. A chamber was used for each treatment, so four chambers was 2 Material and Methods used in total. There were six containers in each chamber. Containers rotated among chambers 2.1 Experimental Site and Facility once a week (simultaneously with a change in CO2 concentration and temperature) during the The experiment was conducted in a closed-top growing season, which ensured that all contain- chamber system situated in Maoxian Ecologi- ers spent approximately equal time within each cal Research Station (31°41´07˝N, 103°53´58˝E, chamber. This rotating averaged out any chamber 1800-m altitude) which belongs to the Chengdu differences (Gavazzi et al. 2000). Institute of Biology, Chinese Academy of Science. In this system, the hourly means of the 15 Mean annual precipitation was 919.5 mm and the second readings taken from April to October mean air temperature was 8.6 °C. The average in 2005 and 2007 indicated that the target tem- CO2 concentration over one growth season was perature and CO2 concentrations were achieved –1 around 360 µmol · mol . (Fig. 1). The mean ambient CO2 concentration 2 –1 Each chamber (3 m in height; 9.35 m in ground was 363.8 (± 34.2) µmol · mol . The CO2 con- area; 24.5 m3 in volume) was composed of a prism centration in EC and ECT was 347.1 (± 22.1) as its main body that was made up of a hendecahe- µmol · mol–1 and 352.8 (± 27.6) µmol · mol–1, dron and a corresponding partial sphere as its top. respectively, higher than in the CON. The CO2 Chambers were connected with a heating-cooling concentrations were within 650~800 µmol · mol–1 system, a CO2 flow system and a wind exchange for 91.32% (EC chamber) and 91.67% (ECT system, which were controlled by a computer. The chamber) of the exposure time. The tempera- computer-controlled heating and cooling
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