Environmental and physiological controls on diurnal and seasonal patterns of biogenic volatile organic compound emissions from five dominant woody species under field conditions☆
Graphical abstract
Introduction
Biogenic volatile organic compounds (BVOCs), with high reactivity in the atmosphere, have a larger impact on atmospheric chemistry, compared to anthropogenic volatile organic compounds (AVOCs) (Guenther et al., 2006; Peñuelas and Staudt, 2010). The global emissions of BVOC from terrestrial ecosystems have been estimated to range from 700 to 1000 Tg C yr−1 (Sindelarova et al., 2014). The largest emitter of BVOCs is thought to be forest ecosystems (Guenther et al., 1995). BVOCs are composed of a wide variety of chemical species, with isoprenoids (e.g., isoprene, monoterpenes, sesquiterpenes) and oxygenated VOCs dominated in general (Arneth et al., 2008). Previous studies have shown that the composition and emission magnitudes of BVOC can exhibit distinct variations among different species and ecosystems (Winters et al., 2009; Préndez et al., 2013; Aydin et al., 2014). The variations were mainly explained by different environmental conditions and also plant-specific characteristics.
It has been widely reported that conifer tree species such as Pinus pinea, Pinus sylvestris and Picea abies can emit significant amounts of monoterpenes (Keenan et al., 2009; Aydin et al., 2014), while the broad-leaved trees such as Populus deltoids, Populus alba, Eucalyptus sp., Quercus cerris and Acacia cyanophylla emit large amounts of isoprene (Padhy and Varshney, 2005; Loreto et al., 2007; Aydin et al., 2014; Baraldi et al., 2019). However, recent studies have shown that several broad-leaved tree species such as Quercus ilex, Quercus coccifera and Fagus sylvatica can also emit large quantity of monoterpenes (Keenan et al., 2009; Llusia et al., 2012, 2013; Sicard et al., 2018). It seems that the tree species growing in different forest ecosystems might have different emission patterns, potentially influenced by the local growing conditions. It is therefore important to assess the emission differences in BVOCs from species growing in different regions, potentially enriching the emission inventory we have for large scale modelling.
Temperature and light intensity are found to be the major environmental factors that can affect the emissions of isoprenoids. Temperature is considered to affect enzyme activity (i.e., for synthase) and influence both the volatility and diffusivity of mono- and sesquiterpenes stored in plant structures such as resin duct and glandular cells (Monson et al., 1995; Staudt and Bertin, 1998; Niinemets et al., 2002a). The light dependence of BVOCs is linked to the compound synthesis through photosynthesis (Niinemets et al., 1999). It has been previously thought that monoterpene emissions from conifers are light-independent, as monoterpenes mainly originate from compounds stored in storage organs such as resin ducts (Holzke et al., 2006; Lim et al., 2008). However, more and more studies have suggested that monoterpene emissions are a combination of emissions from stored and immediately synthesized compounds (Staudt et al., 1997; Guenther et al., 2012).
Many studies have previously investigated the relationship between BVOC emissions and leaf physiological parameters (Kesselmeier et al., 1997; Niinemets et al., 1999; Owen and Peñuelas, 2013; Ashworth et al., 2016; Pazouki and Niinemets, 2016; Li et al., 2017). For example, a recent study by Yuan et al. (2016) found that isoprene emissions were positively correlated to stomatal conductance. However, insignificant and/or negative relationships have also been reported for some of monoterpene compounds and stomatal conductance (Staudt et al., 1997; Niinemets et al., 2002a; Chen et al., 2019). Transpiration rate, as a substitute for stomatal resistance, has also been linked to the exchange rate of BVOCs (Jardine et al., 2008). Although a strong relationship between BVOC emission and transpiration rate has been often reported in the literature (Jardine et al., 2008; Winters et al., 2009), the relationship could vary among species and plant water conditions. Therefore, it is essential to explore the physiological control mechanism for BVOC emissions, which could be of importance for process-based modelling.
Several BVOC emission modelling approaches have been applied at regional and global scales for estimating temporal and spatial variations of BVOC emissions (Guenther et al., 2012; Grote et al., 2014; Tang et al., 2016). The modelling work is based on lab/field-based emission measurements, vegetation land-cover and the derived emission responses to environmental variables. The models often use the averaged emission capacity (standardized emission rates at 30 °C and 1000 μmol m−2 s−1) from published values. The levels of representativeness of these measured species and conditions in literature could lead to different levels of uncertainties in modelling results, as for the same region, local acclimation/adaptation might occur and cause different emission patterns. Therefore, considering within-species variations in emissions across different ecosystems can potentially improve the accuracy of model estimations.
In this study, we measured BVOC emissions from five dominant woody tree species (i.e., two coniferous and three broad-leaved species) in a temperate forest in the northern China. The objectives of this study are: 1) to evaluate if the temporal emission patterns differed within individuals and compared the measured results with those previously reported; 2) to get a better understanding of the driving factors (e.g., environmental variables and physiological parameters) for the observed BVOC variations.
Section snippets
Study area and tree species
The measurement site is located in the Jiufeng National Forest Park in the northwest Beijing (116°28′E, 39°54′N), and it is in the temperate humid monsoon climate zone, with an average annual temperature of 12.5 °C (Fig. 1). Five endemic tree species were investigated, including two evergreen conifers (P. tabuliformis and P. orientalis) and three broad-leaved deciduous trees (Q. variabilis, P. tomentosa and R. pseudoacacia). The tree density of each species has been assessed, which are 1377
Diurnal variations in isoprene and monoterpene emission rates
Large diurnal variations of isoprene and monoterpene emissions were observed (Fig. 2). In general, isoprene emissions peaked at 13:00 and 15:00 p.m. and there were higher observed emissions from the three broad-leaved tree species than the two conifers. Among the five tree species, the highest isoprene emitter was found to be P. tomentosa, with daily average and maximum of ERs of 13.26 ± 1.3 μg gdw−1 h−1 and 18.5 ± 3.2 μg gdw−1 h−1, respectively. The daily average isoprene emissions for
Discussion
In this study, the diurnal and annual emission patterns of isoprene and monoterpene emissions were investigated for the five dominated tree species growing in an artificial temperate forest in northern China. The observed emission diversity among different plant species could potentially suggest species-specific mechanisms of synthesis, storage and production. The linkage of the observed emissions with physiological parameters can further reveal the underlying processes regulating the emission
Conclusions
This study reported the field measured isoprene and monoterpene emissions from five dominant woody species in northern China and to the best of our knowledge, this is the first whole-year BVOC dataset from this region. The linkage of emissions with environmental factors and physiological parameters (including Pn, Tr, gs and Ci) has been explored at both diurnal and seasonal scales. The positive relationship between monoterpene emission with temperature, PAR and foliar photosynthetic
CRediT authorship contribution statement
Jungang Chen: Investigation, Data curation, Writing - original draft, Writing - review & editing. Jing Tang: Writing - review & editing. Xinxiao Yu: Supervision, Writing - review & editing, Funding acquisition.
Declaration of competing interest
The authors declare no competing interests.
Acknowledgments
This research project was supported by the National Natural Science Foundation of China (No. 31800377 and No. 8200904321), National Key R&D Program of China (2016YFC0500802), the Beijing Laboratory Project (2015BLUREE07), the Beijing Municipal Education Commission (CEFF- PXM2018_ 014207_ 000024) and the Special Foundation for Beijing Common Construction Project. Dr. Tang received funding from Swedish FORMAS mobility (grant no. 2016-01580). We also appreciate the constructive comments and
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