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Soil Moisture and Soluble Salt Content Dominate Changes in Foliar Δ13c and Δ15n of Desert Communities in the Qaidam Basin, Qinghai-Tibetan Plateau

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Soil Moisture and Soluble Salt Content Dominate Changes in Foliar Δ13c and Δ15n of Desert Communities in the Qaidam Basin, Qinghai-Tibetan Plateau fpls-12-675817 July 7, 2021 Time: 13:11 # 1 ORIGINAL RESEARCH published: 08 July 2021 doi: 10.3389/fpls.2021.675817 Soil Moisture and Soluble Salt Content Dominate Changes in Foliar d13C and d15N of Desert Communities in the Qaidam Basin, Qinghai-Tibetan Plateau Weiling Niu1, Hui Chen1,2* and Jianshuang Wu3,4* 1 Hebei Key Laboratory of Environmental Change and Ecological Construction, College of Resources and Environmental Sciences, Hebei Normal University, Shijiazhuang, China, 2 Hebei Technology Innovation Center for Remote Sensing Identification of Environmental Change, College of Resources and Environmental Sciences, Hebei Normal University, Shijiazhuang, China, 3 Institute of Environment and Sustainable Development in Agriculture, Chinese Academy of Agricultural Sciences, Beijing, China, 4 Theoretical Ecology, Institute of Biology, Freie Universität Berlin, Berlin, Germany Edited by: Changing precipitation and temperature are principal drivers for nutrient cycling Feng Cheng, 13 University of Nevada, Reno, dynamics in drylands. Foliar isotopic carbon (C) and nitrogen (N) composition (d C United States and d15N) are often used to describe the plant’s water use efficiency and nitrogen Reviewed by: use strategy in plant ecology research. However, the drivers and mechanisms under Xiangzhong Li, 13 15 Yunnan University, China differential foliar d C and d N among plant species and communities are largely Yuxin He, unknown for arid high-elevation regions. This study collected 462 leaf samples of ten Zhejiang University, China top-dominant plant species (two or three replicates per species) across 16 sites in 2005 *Correspondence: and 2010 to measure the community-weighted means (CWMs) of foliar d13C and d15N, Hui Chen [email protected] northeastern Qaidam Basin, Qinghai-Tibetan Plateau. Our results showed that the CWM Jianshuang Wu of foliar d15N was higher in 2005 than in 2010 and was lower in the warm-dry season [email protected] (July and August) than the cool-wet one (June and September) in 2010. Similarly, the 13 Specialty section: CWM of foliar d C was higher in 2005 than in 2010, but no difference between warm- This article was submitted to 13 dry and cool-wet seasons in 2010. C4 plants have higher d C and generally grow faster Functional Plant Ecology, 13 a section of the journal than C3 species under warm-wet weathers. This might be why the CWM of foliar d C Frontiers in Plant Science was high, while the CWM of foliar d15N was low in the wet sampling year (2010). The Received: 05 March 2021 general linear mixed models revealed that soil moisture was the most critical driver for Accepted: 17 June 2021 the CWM of foliar 15N, which explained 42.1% of the variance alone. However, the total Published: 08 July 2021 d soluble salt content was the crucial factor for the CWM of foliar 13C, being responsible Citation: d Niu W, Chen H and Wu J (2021) for 29.7% of the variance. Growing season temperature (GST) was the second most vital Soil Moisture and Soluble Salt factor and explained 28.0% and 21.9% of the variance in the CWMs of foliar d15N and Content Dominate Changes in Foliar 13 15 13 δ13C and δ15N of Desert d C. Meanwhile, remarkable differences in the CWMs of foliar d N and d C were also Communities in the Qaidam Basin, found at the species level. Specifically, Kalidium gracile and Salsola abrotanoides have Qinghai-Tibetan Plateau. higher foliar 15N, while Ephedra sinica and Tamarix chinensis have lower foliar 15N Front. Plant Sci. 12:675817. d d 13 doi: 10.3389/fpls.2021.675817 than other species. The foliar d C of Calligonum Kozlov and H. ammodendron was the Frontiers in Plant Science| www.frontiersin.org 1 July 2021| Volume 12| Article 675817 fpls-12-675817 July 7, 2021 Time: 13:11 # 2 Niu et al. d13C and d15N of Desert Plants highest among the ten species. Except for the foliar d13C of E. sinica was higher than Ceratoide latens between the two sampling years or between the cool-wet and warm- dry seasons, no significant difference in foliar d13C was found for other species. Overall, the CWMs of foliar d15N and d13C dynamics were affected by soil properties, wet-dry climate change, and species identity in high-elevation deserts on the Qinghai Tibetan Plateau. Keywords: carbon isotopic composition, environmental gradients, nitrogen isotopic composition, nitrogen utilization strategy, water use efficiency INTRODUCTION in tropical forests than temperate ones, indicating that plant NUE may increase with increasing temperature. In addition to Isotopic carbon (C) and nitrogen (N) composition (d13C temperature and precipitation, soil properties are also essential and d15N) can provide fundamental insights into ecosystem in controlling ecosystem N cycling (Booth et al., 2005; Chapin biogeochemical cycles (Handley et al., 1999). For example, foliar et al., 2011). For example, soil moisture can promote N 13 d C can infer intrinsic water use efficiency (WUE) of C3 plants mineralization and nitrification by affecting microbial activity (Hultine and Marshall, 2000; Warren et al., 2001; Qiang et al., (Butterbach-Bahl and Gundersen, 2011; Chapin et al., 2011). 2003) while d15N can reveal nitrogen use efficiency (NUE) and Soil pH can affect microbial nitrification and denitrification, fractionation during the N-uptake, transport, transform, and NH3 volatilization (Booth et al., 2005; Butterbach-Bahl and decomposition (Robinson, 2001). Therefore, both d13C and d15N Gundersen, 2011; Chapin et al., 2011). Soil texture and can be used to explore how plants respond to environmental mycorrhizal fungi can also significantly influence vegetation changes (Adams and Grierson, 2001; Canadell et al., 2002; d15N via plants’ N-uptake preference and fractionation during Dawson et al., 2002). the N-transfer between plant and mycorrhiza (Beyschlag et al., The d13C of terrestrial plant fossils is also increasingly used 2009; Klaus et al., 2013). Therefore, soil properties are also to reconstruct paleoclimate (Siegwolf, 2007; Werner et al., 2012) expected to affect vegetation d15N variation, especially at high- because the d13C of plant issues has recorded a series of climate elevation drylands. change information associated with plant growth (Wang et al., Plant d13C and d15N in response to changes in temperature 2005; Chen et al., 2007; Wang et al., 2010; Ma et al., 2012). and precipitation are species-specifically different (Robinson, Therefore, the d13C can also serve as a valuable indicator 2001; Golluscio and Oesterheld, 2007; Lazaro-Nogal et al., 2013). of plant physiology (Saurer et al., 1995; Loader et al., 2007; In addition to habitat conditions, species physiological traits can Dodd et al., 2008; Diefendorf et al., 2010). Precipitation and also regulate the variability of foliar d15N and d13C(Elmore temperature are the essential factors for plant growth, fitness, et al., 2017). For example, Gatica et al.(2017) found that the and performance in drylands (Huxman et al., 2004; Newman short-term interaction between environmental change and plant et al., 2006). Plant d13C declines with increasing mean annual functional traits may override temperature to affect plant d13C precipitation (MAP), likely due to the intrinsic WUE among and d15N in drylands. Current findings on the trait-regulating species (Golluscio and Oesterheld, 2007; Moreno-Gutierrez et al., effects on foliar d15N and d13C are mainly from temperate 2012). For example, Song et al.(2008) reported that foliar d13C (Garten et al., 2000; Peri et al., 2012) and tropical regions of dominant plants could describe alpine species differentiation (Powers and Schlesinger, 2002) but less from alpine biomes. Hgh- in response to water availability across the Tibetan Plateau. elevation deserts are sensitive to climate warming and wetting However, it is still under debates about how foliar d13C varies with (Yang et al., 2009; Lin et al., 2011; Lu et al., 2013). Warming- mean annual temperature (MAT). Foliar d13C has been reported induced C and N losses from alpine soils can even offset C and to be positively (Li et al., 2005; Wang and Schjoerring, 2012; N sequestration by vegetation, triggering positive feedback to Wang, 2018), negatively correlated with MAT (Song et al., 2008; climate warming (Tan et al., 2010; Lu et al., 2013). Therefore, a Zhou et al., 2011), and even no clear relation between them (Li better understanding of the mechanisms governing C- and N- et al., 2009). However, Yang et al.(2015) found that soil properties related processes is crucial in high-elevation ecosystems (Yang could explain more variance in d13C than climatic factors at et al., 2009; Averill et al., 2014). high-elevation grasslands. They even found a unimodal pattern This study explored how climate change (warm-wet vs cool- between foliar d13C and soil organic carbon (SOC) for alpine dry) affects foliar d13C and d15N of alpine desert plants in steppes on the Tibetan Plateau. Besides, high N availability in soils the northeastern Qaidam Basin, Qinghai-Tibetan Plateau. We could lead to high foliar d13C, primarily due to structural changes hypothesized that changes in precipitation and temperature in plant tissue under droughts (Bol et al., 2004). affect plant d13C and d15N differently among species genotypes. Similarly, foliar and soil d15N decreases with MAP in Specifically, we aim to (1) examine the differences in foliar d13C drylands (Handley et al., 1999; Aranibar et al., 2004), implying and d15N under the dry-wet change; (2) explore how foliar d13C ecosystem N cycling might be more open (Robinson, 2001). and d15N respond to changes in climate and soil factors; and Martinelli et al.(1999) found that foliar d15N was higher (3) to evaluate the relative contribution of environmental factors Frontiers in Plant Science| www.frontiersin.org 2 July 2021| Volume 12| Article 675817 fpls-12-675817 July 7, 2021 Time: 13:11 # 3 Niu et al. d13C and d15N of Desert Plants to changes in foliar d13C and d15N at both plant species and First, we chose an open flat area at each site where soil and community levels.
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