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Xylem Anatomy and Hydraulic Traits of Two Co-Occurring Riparian Desert Plants

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Xylem Anatomy and Hydraulic Traits of Two Co-Occurring Riparian Desert Plants Ayup IAWAet al. –Journal Hydraulic 36 (1),traits 2015: of desert 69–83 plants 69 XYLEM ANATOMY AND HYDRAULIC TRAITS OF TWO CO-OCCURRING RIPARIAN DESERT PLANTS Mubarek Ayup1,2,*, Ya-Ning Chen1,2, Maina John Nyongesah3, Yuan-Ming Zhang1, Vishnu Dayal Rajput1,2 and Cheng-Gang Zhu1,2 1Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, China 2State Key Laboratory of Desert and Oasis Ecology, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, China 3School of Biological Sciences, Jaramogi Oginga Odinga University of Science and Technology, 210-40601 Bondo, Kenya *Corresponding author; e-mail: [email protected] ABSTRACT Populus euphratica Oliv. and Tamarix ramosissima Ledeb. are the dominant riparian plants in desert ecosystems in China, where they play a significant role in maintaining ecological balance. To obtain a better insight into the ecological adaptations of xylem structure and hydraulic traits in desert phreatophytes to extremely drought-stressed environments, we investigated various quantitative features of the vessels and intervessel pits, as well as the xylem hydraulic ef- ficiency (KS(MAX)) and native embolism rate (PLC, %), in the woody shoots and lateral roots (all c. 2–4.5 mm in diameter) of P. euphratica and T. ramosissima from natural populations in the Heihe River Basin, northwestern China. The relationships between xylem anatomy and hydraulic traits are also discussed. There were significant anatomical differences between lateral root and woody shoot xylem within individual species. For lateral roots , arithmetic, hydraulic and maximum vessel diameter (D, DH, DMAX), average vessel area (VA), interves- sel wall thickness (TVW), intervessel pit membrane and pit aperture areas (APM, APA), and intervessel pit membrane and pit aperture diameters (DPM, DPA), were larger than in woody shoots (P < 0.05).The mean KS(MAX) values in lateral roots were 6–11 times greater than in woody shoots for P. euphratica and T. ramosis- sima, respectively (P < 0.01). Woody shoots of T. ramosissima had higher na- tive PLC values (68%) than P. euphratica (39%).The different vessel group- ing patterns in the two species seemed to be related to their different native embolism level. It is possible that the lateral roots of these two riparian desert plants could be more resistant to embolism than the woody shoots, and that cavitation resistance in the root xylem of T. ramosissima is higher than that of P. euphratica. Keywords: Populus euphratica Oliv., Tamarix ramosissima Ledeb., xylem anatomy, hydraulic conductivity, native embolism, intervessel pits. © International Association of Wood Anatomists, 2015 DOI 10.1163/22941932-00000086 Published by Koninklijke Brill NV, Leiden Downloaded from Brill.com09/25/2021 08:47:28AM via free access 70 IAWA Journal 36 (1), 2015 INTRODUCTION Xylem hydraulic conductivity and cavitation are important factors that influence plant productivity and survival (Lens et al. 2011; Scholz et al. 2013a). Numerous studies have been conducted to understand wood anatomical variations and hydraulic func- tions at the inter-specific, intra-specific and intra-plant levels (Carlquist 1980; Mayr & Cochard 2003; Sano 2005; Lens et al. 2011; Scholz et al. 2013a). Most of these studies have demonstrated that within a species, root xylem has wider conduits (Zimmermann 1983; McElrone et al. 2004; Psaras & Sofroniou 2004; Domec et al. 2009) and higher conducting efficiencies (Kavanaugh et al. 1999; Martínez-Vilalta 2002; McElrone et al. 2004; Pratt et al. 2007) than stem xylem. However, in some species, root conduit diameter has been shown to be comparable to (Zimmermann 1983) or even smaller than stem xylem conduit diameter (Machado et al. 2007). While the root system accounts for 50% or more of the total resistance to water flow along the soil-to-leaf continuum (Nobel & Cui 1992; Domecet al. 2006a), the pit membrane ultra-structure has a significant influence on the total hydraulic resistance in the plant. Since the hydraulic conductivity and cavitation resistance of xylem cannot be fully understood without considering the influence of intervessel pits (Prattet al. 2007; Lens et al. 2011), various studies have been undertaken to probe how variations in pit morphology between root and stem xylem influence xylem hydraulic functions (Alder et al. 1996; Sperry & Ikeda 1997; Domec et al. 2006b; Schulte 2012). Studies have revealed that roots tend to have larger interconduit pits (Alder et al. 1996; Sperry & Ikeda 1997; Domec et al. 2006b; Hacke & Jansen 2009), a more porous pit membrane (Alder et al. 1996) and are more susceptible to embolisms than stem xylem (Sperry & Saliendra 1994; Sperry & Ikeda 1997; Hacke et al. 2000; McElrone et al. 2004). However, most of these studies were conducted on conifers (Sperry & Ikeda 1997; Domec et al. 2006b; Hacke & Jansen 2009), and only a few focused on the structural and quantitative differences in intervessel pits between the roots and aboveground organs of angiosperms (Alder et al. 1996). Deciduous P. euphratica trees and T. ramosissima shrubs are the dominant indigenous plant species that are widely distributed in forelands of the Heihe River Basin in an arid region of China. They play a protective role in blocking winds and stabilizing sands, and their growth and survival exclusively depend on their capacity to extend their root systems towards the groundwater (Gries et al. 2003; Zhu et al. 2009). However, due to extensive use of the water and land resources in the upper and central parts of the Heihe River, from the 1960s onwards, water resources discharged to the lower reaches have significantly decreased and thereby led to degradation of the riparian vegetation (Guo et al. 2009). In order to restore the ecosystem of the Heihe River, the Chinese government initiated and implemented the ecological water conveyance project (EWCP) (Chen et al. 2006). This project led to favorable conditions for the groundwater and plant communities. However, change was recorded at a small scale (100–400 m away from the water channel) (Guo et al. 2009). In this study, we assessed the hydraulic traits of xylem in the two aforementioned species, by measuring the hydraulic xylem efficiency and native embolism rate, and Downloaded from Brill.com09/25/2021 08:47:28AM via free access Ayup et al. – Hydraulic traits of desert plants 71 by analyzing the anatomy of xylem vessels and intervessel pits in both lateral roots and woody shoots (both 2–4.5 mm in diameter, including bark). Our objectives were to understand the variation in quantitative xylem vessel and intervessel pit features and hydraulic traits at the intra-plant level; and to discuss the possible relationships between xylem anatomy and hydraulic traits in these two species. This study should increase our understanding of the acclimatization strategy of riparian desert plant communities to severe drought stress. MATERIALS AND METHODS Study sites and plant materials Research was carried out in the lower reaches of the Heihe River Basin (42° 06' 012" N, 101° 03' 564" E) in Inner Mongolia, northwestern China. The Heihe River drains the second-largest inland arid catchment area in China, which is characterized by a fragile ecosystem with scarce water resources (Chen et al. 2006). Typical riparian desert forest grows along the river, where dominant species include Populus euphra- tica trees and Tamarix ramosissima shrubs (Xi et al. 2009). The region’s climate is extremely arid, with an average annual pan evaporation of 3755 mm. The mean an- nual precipitation is 42 mm where the minimum and maximum recorded precipitation is 7 and 103 mm, respectively. Average annual temperature ranges between 7.0 and 9.0 °C, with an absolute range from -36.4 to + 41.8 °C (Xi et al. 2009; Guo et al. 2009). We conducted xylem hydraulic measurements on lateral roots and woody shoots of 21–30 year old P. euphratica and older T. ramosissima during August, 2012. The tree age of P. euphratica was roughly estimated on the basis of trunk diameter (Lu 1978; Wang et al. 1996). Twelve P. euphratica trees (mean height of 9.3 ± 3.8 m, mean DBH of 21.5 ± 8.7 cm) and ten shrubs of T. ramosissima of comparable size (mean height of 2.8 ± 0.9 m, mean stem diameter of 10.0 ± 4.6 cm) were selected as target plants. Lateral root and woody shoot samples were collected from different individual trees. Three to four woody shoot samples (branches or twigs, 2–4.5 mm in diameter) were collected from the sun exposed upper canopy of each P. euphratica and T. ramosis- sima tree/shrub for 3 days in the morning between 05 : 30 am and 06 : 30 am. With the groundwater table being 1.5–3.5 m deep, lateral roots were excavated from the soil at a depth of 10 to 40 cm. From each target plant, hydraulic conductivity was measured in 5–8 lateral roots. The average gravimetric water content of the soil where these lateral roots were collected was 7.49 ± 3.18 (%) (mean ± SD). Detailed information about the sample numbers (N), hydraulic traits measurement sample length and diam- eter (including bark) is given in Table 1. Xylem hydraulic trait measurements The maximum hydraulic specific conductivity (or xylem hydraulic efficiency) (KS(MAX), kg/m/s/MPa) and native embolism rate (percentage loss of hydraulic con- ductivity, PLC, %) of the xylem were measured on lateral root and woody shoot (branch/ Downloaded from Brill.com09/25/2021 08:47:28AM via free access 72 IAWA Journal 36 (1), 2015 Table 1. Mean native embolism rate and hydraulic efficiency of different organs ofPopulus euphratica and Tamarix ramosissima. P-values are for the ANOVA between roots and woody shoots within the same species; differ- ent letters indicate significant differences between roots and woody shoots within the same plant (P < 0.05).
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