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Water Uptake and Hydraulic Redistribution Across Large Woody Plant, Cell and Environment (2010) 33, 2132–2148 doi: 10.1111/j.1365-3040.2010.02212.x Water uptake and hydraulic redistribution across large woody root systems to 20 m depthpce_2212 2132..2148 TIMOTHY M. BLEBY1, ANDREW J. MCELRONE2 & ROBERT B. JACKSON3 1School of Plant Biology, The University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia, 2USDA-ARS, Crops Pathology Genetics Research Unit, University of California, Davis, CA 95616, USA and 3Department of Biology & Nicholas School of the Environment, Duke University, Durham, NC 27708, USA ABSTRACT also occur during the day if the gradient in water potential is greater to dry soil than to the atmosphere. HR is a global Deep water uptake and hydraulic redistribution (HR) are phenomenon that occurs in a wide range of species across important processes in many forests, savannas and shru- many different biomes, from deserts to tropical rainforests blands. We investigated HR in a semi-arid woodland above (Caldwell, Dawson & Richards 1998; Jackson, Sperry & a unique cave system in central Texas to understand how Dawson 2000). deep root systems facilitate HR. Sap flow was measured in There is mounting evidence that HR has significant Quercus 9 trunks, 47 shallow roots and 12 deep roots of , potential to alter soil hydrology and provide ecological ben- Bumelia Prosopis and trees over 12 months. HR was exten- efits to at least some plants. The amount of water involved sive and continuous, involving every tree and 83% of roots, in HR is often small, but large trees and shrubs can move with the total daily volume of HR overa1month period relatively large amounts of water representing a consider- estimated to be approximately 22% of daily transpiration. able percentage of total daily water use (Ryel et al. 2002; During drought, deep roots at 20 m depth redistributed Brooks et al. 2006). Studies have shown that the water from water to shallow roots (hydraulic lift), while after rain, HR may delay the onset of soil drying during drought shallow roots at 0–0.5 m depth redistributed water among (Brooks et al. 2002; Hawkins et al. 2009), limit cavitation in other shallow roots (lateral HR). The main driver of HR fine roots (Domec et al. 2004), and moisten soil at the appeared to be patchy, dry soil near the surface, although surface or at depth to facilitate processes such as root or water may also have been redistributed to mid-level depths mycorrhizal growth (Huang 1999; Querejeta, Egerton- via deeper lateral roots. Deep roots contributed up to five Warburton & Allen 2003), decomposition (Aanderud & times more water to transpiration and HR than shallow Richards 2009) and nutrient acquisition (McCulley et al. roots during drought but dramatically reduced their contri- 2004). At large scales, HR may play an important role in bution after rain. Our results suggest that deep-rooted ecosystem water, carbon and nutrient cycling (Jackson et al. plants are important drivers of water cycling in dry ecosys- 2000), and it may also affect the energy balance and climate tems and that HR can significantly influence landscape of densely vegetated ecosystems (Lee et al. 2005). HR is hydrology. likely increasing globally as the abundance of deep-rooted woody plants increases through woody plant encroach- Key-words: caves; deep roots; ecohydrology; gum bumelia; ment, afforestation and other processes (Van Auken 2000; Heat Ratio Method; live oak; mesquite; woody plant Engel et al. 2005; Jackson, Jobbagy & Nosetto 2009). Studies encroachment. are clearly needed to predict the possible ecohydrological consequences of HR, particularly in water-limited environ- ments where deep-rooted plants can potentially tap and INTRODUCTION redistribute isolated water resources (Schulze et al. 1998; Seyfried et al. 2005; Scott et al. 2006; Lubczynski 2009). Plant roots are well known to transport water within the soil Plants with large, expansive woody root systems tend to profile via the process of hydraulic redistribution (HR), be the most effective redistributors of soil water. Large, defined as the passive movement of water from wet soil to deep root systems are advantageous for HR because they dry soil through roots, driven by gradients in soil water can connect multiple soil compartments and allow water to potential (Richards & Caldwell 1987; Burgess et al. 1998). move in virtually any direction dictated by soil water poten- The occurrence of HR depends on a suite of biological and tial gradients, including upwards (e.g. Burgess et al. 1998; physical variables, including root system size, soil texture, Moreira et al. 2003), downwards (e.g. Smith et al. 1999; and the degree of root-to-soil contact (Jackson et al. 2007). Burgess et al. 2001c) and laterally (e.g. Smart et al. 2005; It typically occurs at night when stomata are closed and Burgess & Bleby 2006). Deep roots in particular can allow plants become disconnected to the atmosphere, but it can access to deep water sources many tens of metres under- Correspondence: T. M. Bleby. E-mail: [email protected] ground (Schenk & Jackson 2002), and they can play a 2132 © 2010 Blackwell Publishing Ltd Hydraulic redistribution across deep root systems 2133 critical role in facilitating both hydraulic lift (e.g. Penuelas contribution of deep roots to HR is determined indirectly & Filella 2003) and downward HR (e.g. Hultine et al. 2003a) by monitoring sap flow in the upper portions of apparent in arid and semi-arid environments. tap or sinker roots (e.g. Hultine et al. 2003b; Oliveira et al. There is strong evidence that woody roots are well 2005), or by identifying lifted water in surface soil using designed to facilitate water transport driven by HR. Water stable isotopes (e.g. Penuelas & Filella 2003). While these flow through woody root xylem is generally unrestricted in approaches provide important information about the both directions (Schulte 2006), and HR-induced sap flow in dynamics of HR and the approximate location of deep intact lateral roots can be equal to or greater than sap flow roots and water sources, they give little or no information induced by transpiration (Burgess & Bleby 2006). Recent about specific depths or locations of water uptake or work also suggests that deep roots are naturally well actual rates of sap flow in deep roots. To overcome this adapted for water transport over long distances against the limitation, we used a novel cave system to access to woody force of gravity, in part due to their unique structural char- roots directly at 20 m depth. We targeted roots that had acteristics that promote high hydraulic efficiency, including been previously identified to parent trees above ground large xylem diameters (McElrone et al. 2004). Less well using DNA sequence variation (Jackson et al. 1999, 2002; understood is how large woody root systems function as an McElrone et al. 2007), which allowed us to monitor water integrated whole in space and time to facilitate HR. transport in the stem and in deep and shallow roots of the Understanding HR across large woody root systems same plant. requires knowledge of key attributes, including the role of Our objectives were threefold: (1) to describe the occur- roots of varying size and location in the root system (e.g. rence and frequency of HR for dominant evergreen and distal roots versus proximal roots), the behaviour of roots deciduous tree species in this ecosystem and how it changes connected to ephemeral and perennial soil water sources in response to environmental conditions; (2) to determine and sinks (e.g. shallow roots versus deep roots), the occur- the number of woody roots involved in HR and assess the rence of different types of HR (e.g. vertical HR versus degree of involvement of roots of varying size and depth; lateral HR), and the occurrence of HR under different and (3) to determine the specific contribution of deep roots environmental conditions (e.g. during drought versus after to HR, including their responses to surface soil moisture rain). These attributes are best studied using techniques conditions and their coordination with shallow roots to that can provide continuous data on the direction and quan- supply water for transpiration and HR. tity of water movement at specific locations.When roots are accessible, sap flow techniques are arguably the most pow- erful tool for studying HR, and a number of studies show MATERIALS AND METHODS that the best results are achieved when synchronous mea- Field site and study species surements of sap flow are obtained from as many roots, spanning as much of the root system, as possible (e.g. This study was conducted in the Edwards Plateau region of Hultine et al. 2004; Nadezhdina et al. 2006; Scholz et al. 2008; central Texas, USA.The region has karst geology character- Scott, Cable & Hultine 2008). ized by shallow, calcareous soils (frequently <20 cm depth) In this paper, we focus on the role of woody roots in HR overlying fractured Cretaceous limestone. The vegetation and the seasonal dynamics of HR in a water-limited savan- of the region is mainly savannah and woodland. We inves- nah populated by deep-rooted trees. We monitored stem tigated three dominant tree species: the evergreen Quercus and root sap flow using the Heat Ratio Method (HRM; fusiformis (Texas live oak), the deciduous Bumelia lanugi- Burgess et al. 2001b) to determine whole-plant water use, nosa (gum bumelia) and Prosopis glandulosa (honey mes- seasonal patterns of HR, and the water transport character- quite). These species are found throughout the Edwards istics of deep and shallow woody roots of Texas live oak Plateau and are considered relatively slow growing and (Quercus fusiformis Small), gum bumelia (Bumelia lanugi- drought tolerant (Auken et al. 1980). nosa Michx.), and honey mesquite (Prosopis glandulosa The study site was located near Menard, TX (30°55′ N, Torr.) in a semi-arid, karst system in central Texas, USA.
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