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Stable Carbon and Nitrogen Isotope Signatures of Root-Holoparasitic Cynomorium Songaricum and Its Hosts at the Tibetan Plateau A

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Stable Carbon and Nitrogen Isotope Signatures of Root-Holoparasitic Cynomorium Songaricum and Its Hosts at the Tibetan Plateau A This article was downloaded by: [Yueqin Yang] On: 01 May 2012, At: 03:11 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Isotopes in Environmental and Health Studies Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/gieh20 Stable carbon and nitrogen isotope signatures of root-holoparasitic Cynomorium songaricum and its hosts at the Tibetan plateau and the surrounding Gobi desert in China Yueqin Yang a b c , Xianfeng Yi c , Min Peng a & Yubi Zhou a a Key Laboratory of Adaptation and Evolution of Plateau Biota, Northwest Institute of Plateau Biology, Chinese Academy of Sciences, Xining, 810008, People's Republic of China b Graduate University of Chinese Academy of Sciences, Beijing, People's Republic of China c College of Agriculture, Henan University of Science and Technology, Luoyang, 471000, People's Republic of China Available online: 27 Apr 2012 To cite this article: Yueqin Yang, Xianfeng Yi, Min Peng & Yubi Zhou (2012): Stable carbon and nitrogen isotope signatures of root-holoparasitic Cynomorium songaricum and its hosts at the Tibetan plateau and the surrounding Gobi desert in China, Isotopes in Environmental and Health Studies, DOI:10.1080/10256016.2012.680593 To link to this article: http://dx.doi.org/10.1080/10256016.2012.680593 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Downloaded by [Yueqin Yang] at 03:11 01 May 2012 Isotopes in Environmental and Health Studies 2012, 1–11, iFirst Stable carbon and nitrogen isotope signatures of root-holoparasitic Cynomorium songaricum and its hosts at the Tibetan plateau and the surrounding Gobi desert in China YueqinYanga,b,c, Xianfeng Yic, Min Penga* and Yubi Zhoua aKey Laboratory of Adaptation and Evolution of Plateau Biota, Northwest Institute of Plateau Biology, Chinese Academy of Sciences, Xining 810008, People’s Republic of China; bGraduate University of Chinese Academy of Sciences, Beijing, People’s Republic of China; cCollege of Agriculture, Henan University of Science and Technology, Luoyang 471000, People’s Republic of China (Received 10 November 2011; final version received 26 February 2012) We first measured the δ13C and δ15N values of root holoparasite Cynomorium songaricum and its hosts from 19 sites across four provinces in northwest China, in an attempt to investigate their nutritional relationship at the Tibetan plateau and the surrounding Gobi desert. Our study showed that the δ13CofC. songaricum closely mirrored the values of its hosts, Nitraria tangutorum and N. sibirica across all sampling sites. C. songaricum was significantly depleted in 13C compared to host plants at the Tibetan plateau, showing an average parasite/host δ13C difference of −0.6 ‰. In contrast, 15NofC. songaricum was significantly enriched by +1.3 ‰ compared to the hosts, implying that these holoparasites had other nitrogen resources. Although no difference in the δ13C and δ15N values between holoparasites and hosts was detected, the δ13C and δ15N values of holoparasites were significantly correlated with those of their hosts at the Gobi desert. The δ13C versus δ15N values were significantly but negatively correlated for the hosts; however, holoparasite/host variation in δ13C was not correlated with the variation in δ15N. The δ13C versus δ15N values were negatively correlated in C. songaricum, and this relationship tended to be magnified along the increasing elevations independent of the host plants. C. songaricum at the Tibetan plateau exhibited different δ13C and δ15N signatures compared with those at the Gobi desert. Furthermore, both δ13C and δ15N values of C. songaricum and its host plants in salt marshes at the Tibetan plateau were different from those in sand sites at the Tibetan plateau and the Gobi desert. Our results indicate that the isotopic Downloaded by [Yueqin Yang] at 03:11 01 May 2012 difference depends on the different altitudes and habitats and is host-specific. Keywords: arid area; carbon-13; Gobi desert; holoparasites; isotope ecology; nitrogen-15; nutritional relationship; plants; Tibetan plateau 1. Introduction Parasitic plants can be classified as hemiparasites and holoparasites according to the extent to which they are unable to produce their own reduced carbon for growth and reproduction [1]. Unlike hemiparasites, holoparasites lack chlorophyll and are obligatory dependent on hosts for the organic and inorganic nutrition supplies through infecting both phloem and xylem of the hosts by the haustorium [2–4]. Although much literature has discussed the nutritional relationship of *Corresponding author. Email: [email protected] ISSN 1025-6016 print/ISSN 1477-2639 online © 2012 Taylor & Francis http://dx.doi.org/10.1080/10256016.2012.680593 http://www.tandfonline.com 2 Y. Yang et al. A B Host Nitraria spp. Holoparasite C. songaricum Figure 1. General features of the host plants Nitraria spp. and holoparasite C. songaricum at the Tibetan plateau and the surrounding Gobi desert. (a) Flowering C. songaricum below host plant Nitraria spp. (b) Immature C. songaricum parasitising deep roots of Nitraria spp. holoparasites and their hosts in differing environments [2,5], there still exists little knowledge on the ecological interactions of the root holoparasites and hosts in arid areas [6,7]. The root holoparasite Cynomorium songaricum (Cynomoriaceae, Saxifragales), known in Chinese herbal medicine as ‘suoyang’, is widely distributed at the Tibetan plateau and the surrounding Gobi desert. C. songaricum produces swollen tuberous haustorial root connections to its host plants [8], mainly Nitraria sibirica and N. tanguticum (Nitrariaceae, Sapindales) (Figure 1). Occasionally, it is also found to parasitise the roots of other host plants such as N. roborowskii, Zygophyllum xanthoxylum and Peganum harmala [9]. Interest in these plants has increased in recent years, and they are being extensively collected from wild populations for use in herbal medicines, causing a dramatic decline of this herbal resource. The chemical components and therapy functions of C. songaricum have been studied in the past three decades [10–17]. However, the nutritional relationship of C. songaricum with its hosts has been little studied. With the development of mass spectrometry, analyses of stable isotope ratios have been recog- nised as a powerful tool to investigate the carbon, water, and nutrient relations of parasitic plants and their hosts [18–26]. Stable carbon isotopes have been studied extensively to address various research questions and have provided valuable information about water use efficiency and carbon gain of parasitic plants [6,20,21,27]. However, identifying sources of nitrogen used by parasites needs further investigation [5]. Because holoparasites cannot produce reduced carbon through photosynthesis and other reduction processes, it has been suggested that identical or similar δ13C values should be observed in the hosts and parasites [19,20]. However, a number of environmental Downloaded by [Yueqin Yang] at 03:11 01 May 2012 factors, such as elevation, water availability, temperature and soil nutrition status, will have a strong influence on the stable isotope compositions of the hosts, especially for stable carbon isotopes [28–30]. Moreover, C. songaricum is often found to parasitise the deep roots of at least four water conservative host species distributed at the Tibetan plateau and the surrounding Gobi desert [9]. In this study, we analysed samples of C. songaricum and its hosts from 19 sites at the Tibetan plateau and the surrounding Gobi desert located at different elevations ranging from 1050 m to 3600 m in northwest China. We wished to investigate the nutritional relationship between C. songaricum and their hosts as well as the regional variations of this relationship among pop- ulations in different habitats (sand and salt marshes). We aimed to answer the question whether the patterns of stable isotopes in the holoparasite C. songaricum are identical along different environment gradients. 2. Materials and methods 2.1. Study sites To examine the trophic strategies of C. songaricum from a wide geographic area of its natural range, samples were collected from 19 sites at the Tibetan plateau and the surrounding Gobi desert Isotopes in Environmental and Health Studies 3 Figure 2. Map of sample collection across the Tibetan plateau and the surrounding Gobi desert. The characteristics of sites and IDs are listed in Table 1. Samples were collected from four provinces or autonomous regions: Qinghai province, Gansu province, Inner Mongolian autonomous region, and Ningxia Hui autonomous region. in four
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