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Interactions Between Elevated Atmospheric CO2 and Defoliation on North American Rangeland Plant Species at Low and High N Availability

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Interactions Between Elevated Atmospheric CO2 and Defoliation on North American Rangeland Plant Species at Low and High N Availability Grass and Forage Science The Journal of the British Grassland Society The Official Journal of the European Grassland Federation Interactions between elevated atmospheric CO2 and defoliation on North American rangeland plant species at low and high N availability D. R. LeCain*, J. A. Morgan*, G. L. Hutchinson*, J. D. Reeder* and F. A. Dijkstra† *U.S. Department of Agriculture, Agricultural Research Service, Rangeland Resources Research Unit, Crops Research Laboratory, Fort Collins, CO, USA, and †Faculty of Agriculture, Food and Natural Resources, The University of Sydney, Level 4, Biomedical Building, 1 Central Avenue, Eveleigh, NSW 2015, Australia Abstract Keywords: semi-arid rangeland, CO2, defoliation, nitro- 15 Although common disturbances of grazing lands like gen, C3 grass, C4 grass, forb, root, biomass, N recov- plant defoliation are expected to affect their sensitivity ery, forage quality. to increasing atmospheric CO2 concentration, almost no research has been conducted to evaluate how important such effects might be on the direct responses of Introduction rangelands to CO2. This growth chamber experiment About 40% of terrestrial ecosystems are classified as subjected intact plant–soil cylinders from a Wyoming, rangelands (Suttie et al., 2005). Rangeland ecosystems USA, prairie to a 3-way factorial of CO2 (370 vs. are not characteristically productive lands, but they )1 720 lLL ), defoliation (non-clipped vs. clipped) and support most of the world’s managed livestock in )2 soil nitrogen (control vs. 10 g m added N) under addition to large herds of native ungulates (Campbell simulated natural climatic conditions. Above- and et al., 1997). The productivity of rangelands may be below-ground biomass and N dynamics of the func- slowly increasing owing to the fertilization effects of tional groups C3 grasses, C4 grasses and forbs were rising levels of atmospheric CO2 (Polley, 1997; Morgan investigated. CO2 and defoliation had independent et al., 2001a, 2004b). However, the consequences of influences on biomass and N parameters of these rising atmospheric CO2 concentrations on plant–herbi- rangeland plants. Growth under CO2-enriched condi- vore dynamics like defoliation, that influence the tions enhanced above-ground biomass 50% in C3 ecology of rangelands, are relatively unexplored. grasses alone, while shoot N concentration declined Elevated CO2 is well known to increase above- 15 16% in both C3 and C4 grasses. Plant-soil N uptake ground plant productivity (Korner, 2000; Morgan et al., was unaffected by CO2 treatment. In contrast, defolia- 2004a; De Graaff et al., 2006). Owing to greater photo- tion had no effect on biomass, but increased tissue N synthetic enhancement, C3 plant growth is stimulated concentration 29% across all functional groups. With- more by elevated CO2 than C4 species (Korner, 2000; out additional N, forage quality, which is in direct Reich et al., 2001). However, there are several reports of relation to N concentration, will decline under increas- improved productivity in C4 rangeland species (Read ing atmospheric CO2. Increased dominance of C3 1996; LeCain and Morgan, 1998; Wand et al., 1999; grasses plus reduced forage quality may necessitate Owensby et al., 1999; LeCain et al., 2003). This has been changes in grazing management practices in mixed- attributed to improved water-use efficiency and soil- species rangelands. water status resulting from reduced stomatal conduc- tance and plant transpiration under elevated CO2 (Volk et al., 2000; Nelson et al., 2003). This stomatal response Correspondence to: D. R. LeCain, U.S. Department of Agri- occurs similarly in both C3 and C4 grasses (Wand et al., culture, Agricultural Research Service, Rangeland Re- 1999) and is a major factor boosting productivity in sources Research Unit, Crops Research Laboratory, 1701 CO2-enriched semi-arid rangelands (Morgan et al., Centre Avenue, Fort Collins, CO 80526, USA. 2011). In many temperate rangelands (such as much E-mail: [email protected] of the USA Great Plains), C3 grasses are the major forage during spring while C4 grasses are the primary mid-late- Received 29 April 2011; revised 19 October 2011 season forage; forbs are typically a less important food 350 Ó 2012 Blackwell Publishing Ltd. Grass and Forage Science, 67, 350–360 doi: 10.1111/j.1365-2494.2011.00847.x CO2, defoliation and N interactions on rangeland plant species 351 source (Milchunas et al., 1995). Rangeland forb pro- expected to be a major factor in rangelands as many duction may be increased by higher atmospheric CO2, rangeland soils are N limiting for plant growth (Hungate presumably because most rangeland forbs are C3 pho- et al., 1997). Many studies report reduced plant-tissue N tosynthesis types (Reich et al., 2001; Teyssonneyre concentration under elevated CO2 (Wilsey et al., 1997; et al., 2002; Polley et al., 2003). In mixed-species Morgan et al., 2004a; Milchunas et al., 2005a; Reich rangelands, CO2-induced shifts in the balance of C3 et al., 2006) because of improved nitrogen-use effi- and C4 grasses and forbs have the potential to signif- ciency or reduced soil-N availability (King et al., 2004; icantly alter community structure and ecosystem pro- Wand and Midgley, 2004). Although productivity may cesses, including animal grazing processes (Campbell increase under elevated CO2, reduced tissue N and and Stafford Smith, 2000; Morgan et al., 2007). protein concentration will have negative consequences In most rangelands, there is more plant matter below for domestic and indigenous fauna (Campbell and than above ground (Milchunas and Lauenroth, 1993, Stafford Smith, 2000; Milchunas et al., 2005a). Con- 2001). Although there are many reports on the influ- versely, defoliation can increase N concentration in ence of elevated CO2 on root productivity, the results regrowth tissue (Milchunas et al., 1995, 2005a; Mikola vary widely between ecosystems and sampling methods et al., 2005), but in the long term, reduced soil-N and a conceptual understanding of root responses has availability under elevated CO2 may lessen this re- not emerged (Hebeisen et al., 1997; Wilsey et al., 1997; sponse. Morgan et al., 2001b; Milchunas et al., 2005b; De Graaff Our objective was to investigate how defoliation and et al., 2006). In a field study on the shortgrass steppe of soil-N availability interact to affect the above- and Colorado, we reported little change in root biomass below-ground productivity (assessed as plant biomass) after 5 years of elevated CO2 (LeCain et al., 2006). and tissue N responses of a semi-arid grassland to However, studies investigating the interaction of ele- growth under present-day [CO2] and CO2-enriched vated CO2 and grazing on root growth are rare conditions. In this experiment, we utilized perennial (Augustine et al., 2010). plant-intact soil microcosms extracted from a northern As it is difficult to evaluate how grazing might USA mixed-grass prairie and subjected them to manip- interact with CO2 levels in sparsely stocked rangelands, ulations of defoliation, CO2 and N. Responses were defoliation has been used to simulate grazing. Grazing evaluated for the three major plant functional groups, or defoliation alone can increase or decrease plant C3 grasses, C4 grasses and forbs. Our hypotheses were as productivity depending on grazing intensity and the follows: grazing tolerance of species (Hendon and Briske, 2002). H1: Elevated CO2 will increase shoot biomass in forbs Forbs are generally thought to be less grazing tolerant and C3 grasses, with little response observed in C4 than grasses because of herbivory-exposed meristems grasses and roots. Defoliation will reduce shoot (Heady and Child, 1994). While CO2 enrichment might biomass in forbs, but elevated CO2 will improve be expected to enhance plant recovery from defoliation, recovery from defoliation. because of more available CO2 and soil water (Wand H2: Enhanced shoot biomass under elevated CO2 will and Midgley, 2004), this has not always been found reduce tissue N concentration and N uptake (mea- 15 (Polley et al., 2011). Elevated CO2 and defoliation may sured by N) in all functional groups, particularly have opposing as well as additive effects on plant without N addition. Defoliation and N addition will productivity, species composition and allocation of counter the effect of elevated CO2 on these N resources (Morgan et al., 2001a; Harmens et al., 2004; parameters. Wand and Midgley, 2004; Lau and Tiffin, 2009; H3: Elevated CO2 will increase volumetric soil-water Augustine et al., 2010). Consequently, it is presently content (VWC) that will contribute to increased unclear how defoliation interacts with rising atmo- biomass of all functional groups. Defoliation will spheric CO2 to affect rangeland productivity and ecol- have minimal effect on VWC. ogy, particularly in a mixed-species ecosystem. In many arid and semi-arid rangelands, nitrogen (N) Materials and methods dynamics are second only to precipitation in impor- tance for plant–ecosystem processes (Burke et al., Site characteristics 1997). Responses of plant productivity to elevated CO2 and defoliation, and the resulting changes in Plant–soil cylinders for this experiment were extracted ecosystem functioning, are co-dependent on the cycling from the USDA-ARS High Plains Grasslands Research of N through the plant–soil–atmosphere. In the long Station (HPGRS), near Cheyenne, WY, USA (41°11¢ N term, plants will not sustain increased productivity lat, 104°54¢ W long). The HPGRS is within the under elevated CO2 when other factors become growth northern mixed-grass prairie, with elevations ranging limiting (De Graaff et al., 2006). Nitrogen limitation is from 1910 to 1950 m, an average 127-d growing Ó 2012 Blackwell Publishing Ltd. Grass and Forage Science, 67, 350–360 352 D. R. LeCain et al. season, and annual precipitation of 384 mm. Site Growth chamber conditions vegetation is mostly comprised of grasses, averaging 55% C3 grasses and 23% C4 grasses plus a variety of Environmental conditions in the growth chambers forbs, sedges and sub-shrubs. In our cylinders, there simulated the growing season of SE Wyoming.
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