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Rhizosphere interactions under elevated CO 2 : Impact on soil organic carbon dynamics. Shuijin Hu North Carolina State University Raleigh, NC 27695 Email: shuijin_hu@ncsu.edu. An Overview of Recent and Ongoing Research Projects. Microbes & plant competition.
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Rhizosphere interactions under elevated CO2: Impact on soil organic carbon dynamics Shuijin Hu North Carolina State University Raleigh, NC 27695 Email: shuijin_hu@ncsu.edu
An Overview of Recent and Ongoing Research Projects Microbes & plant competition Carbon & nitrogen dynamics in agroecosystems Plant-Microbe Interactions Microbial diversity & ecosystem stability Microbial responses to climate change
Air temperature has increased ca. 0.6 oC Air temperature is predicted to increase another 2-5 oC in the next 100 years
The increasing atmospheric CO2 is correlated with the temperature rise
One central goal of global change research is to understand: whether and how terrestrial ecosystems can sequester more organic C.
Why ecosystem C sequestration for mitigation of climate change? Active C pools on the Earth surface: • Air CO2-C: 750× 1015 g • 2. Biomass-C: 550-650 × 1015 g • Soil organic C: 1500-2100 × 1015 g
Elevated CO2 stimulates photosynthesis and net primary production – Increases short-term C inputs Herrick & Thomas. 2001
Prerequisites for long-term ecosystem C sequestration under elevated CO2 • Plants can effectively acquire available nutrients; • Mechanisms exist to sustain N supply for plants; • Microbial decomposition is “contained”. Plants are primarily nutrient-limited but microbes are C-limited
1. Can plants acquire available nutrients more effectively under elevated CO2? The prevailing paradigm in 1990’s was: Microbes outcompete plants for acquiring nutrients in soil.
Elevated CO2 Plants C inputs Available N Organic N Microbes Elevated CO2 alters the plant-microbial competition in favor of plant N utilization. Hu et al. 2001. Nature
Nutrient Limitation of Ecosystem C Sequestration Luo et al. 2004. Bioscience
Are there mechanisms that sustain N supply for plants under elevated CO2? • Can CO2-stimulation of plant growth be sustained over time? • To a large degree, it will depend on whether plants can acquire sufficient nutrients from the organic pool.
Elevated CO2 Plants C inputs Microbes Available N Elevated CO2 increased plant N acquisition from soil organic N pool. Hu et al. 2005. Global Ch. Biol. Zak et al. 2011. Ecology Letters. Drake et al. 2011. Ecology Letters + Organic N
The Summary The Summary Plants are more effective in nutrient acquisition under elevated than ambient CO2. Plants are more effective in nutrient acquisition under elevated than ambient CO2. Plants are more effective in nutrient acquisition under elevated than ambient CO2. Next Question How does elevated CO2 increase plant nutrient acquisition from soil? How does elevated CO2 increase plant nutrient acquisition from soil? How does elevated CO2 increase plant nutrient acquisition from soil? How does elevated CO2 increase plant nutrient acquisition from soil?
Elevated CO2 (15NH4)2SO4 NO3– NO3– NO3– bacteria PO43- fungi Ca2+ Residues K+ Mg2+ NH4+ PO43- Hu et al. 2005, Global Change Biol.
Indeed, one major finding over the last two decades is: Elevated CO2 increases soil fungi, particularly mycorrhizal fungi. Then the question is: Why? Treseder, 2004. New Phytologist
Ectomycorrhzae Arbuscular Mycorrhizae • Mycorrhizae are symbiotic associations between plant roots and fungi; • Over 80% of terrestrial plants form mycorrhizae with fungi; • Plants allocate up to 20% of photosynthates to mycorrizal fungi under ambient CO2 and up to 35-40% under elevated CO2.
AM fungi protect organic C from microbial attack Scanning electron micrograph of a VA mycorrhizal fungus with particles of clay firmly attached (left) and VA mycorrhizal fungi binding microaggregates into a stable macroaggregate (Tisdall and Oades 1979).
Elevated CO2 Plant Growth Mycorrhizae Extraradical Fungal Hyphae Cell Wall Materials (Chitin) Glomalin Polysaccharides Soil Aggregation Carbon Sequestration The current paradigm of elevated CO2 impact on soil C Rillig et al., 1999, Nature; Treseder & Allen, 2000, New Phytol. Antoninka et al. 2009, GCB. Wilson et al. 2009. Ecol. Letters
Major issues related to the current paradigm • The current paradigm is largely based on correlative information, rather than direct evidence; • Emerging evidence suggests that AM fungi may increase decomposition of organic residues (Hodge et al. 2001, Nature; PNAS 2010; Tu et al. 2006, Global Change Biology).
Can CO2-stimulation of AM fungi increase decomposition of organic matter in soil? Five stepsto assess the impact of CO2–enhancement of AM fungi on organic C decomposition
Step 1 A microcosm experiment to assess AMF-mediated organic C decomposition under different CO2 and N levels CSTR chambers Microcosm unit
Isolation of root contribution from fungal effects on organic C decomposition 13C/15N labeled materials
Step 2 A microcosm experiment to examine the impact of AMF identity on AMF-mediated organic C decomposition under different CO2 levels AM fungal species or assemblages A. Acaulospora morrowiae B. Gigaspora margarita C. Glomus clarum D. Assemblage A: The combination of A, B and C F. Assemblage B: Eight species from field, including A, B & C
Step 3 A field experiment to determine AMF-mediated organic C decomposition under elevated CO2 Open-top chambers used to simulate atmospheric CO2 concentrations under future climate scenarios
Result 1: Elevated CO2 increased mycorrhizal infection of roots and AMF biomass in soil Fig. S3. Elevated CO2 stimulated the growth of AMF in roots of Avena fatua and wheat, and in soil
Result 2: Higher AMF under elevated CO2 increases decomposition A: Microcosm Exp. 1; B. Microcosm Exp. 2; C. Field Exp. Cheng et al. 2012. Science
Why does elevated CO2 concentration increase organic C decomposition? Our initial hypothesis was: Elevated CO2 stimulates organic C decomposition because 1. N becomes more limiting, 2. plants under elevated CO2 need to obtain more N, and 3. plants allocate more carbohydrates to prime decomposition through stimulating saprotrophs. Does elevated CO2 really reduce N availability?
Result 3: Elevated CO2 reduces soil NH4+ in N-limiting soils but increases soil NO3- in the N-rich field soil A: Microcosm Exp. 1; B. Microcosm Exp. 2; C. Field Exp. Cheng et al. 2012. Science
These results led us to ask: • Why do plants not use the increased NO3- under elevated CO2? • 2. Does elevated CO2 lead to plant preference of soil NH4+ over soil NO3- ?
Fig. 1. Three methods for assessing nitrate absorption (Absorb) and assimilation (Assim.) in wheat and Arabidopsis plants in hydroponic solutions where the shoots were exposed to atmospheres containing 380-ppm CO2 and 21% O2, 720-ppm CO2 and 21% O2, or 380-ppm CO2 and 2% O2. Bloom et al. 2010. Science
Step 4 • A meta-analysis of elevated CO2 impact on soil N and plant N acquisition in the literature • Does elevated CO2 lead to plant preference of soil NH4+ over soil NO3- ?
Step 4 • A meta-analysis of elevated CO2 impact on soil N and plant N acquisition in the literature. • 38 studies that quantified the concentrations of soil NH4+ and NO3– and/or the capacity of plant use of NH4+ and NO3– under eCO2; • These studies encompassed more than 58 species of crop, grass, and tree species.
Result 4: Elevated CO2 reduced plant NO3- uptake and increased soil NO3- (Net effect %). 40 20 -20 -40 Cheng et al. 2012. Science
Step 5 • A field experiment to assess the impact of nitrification inhibition on AMF-mediated organic C decomposition under elevated CO2 X
Result 5: Inhibition of nitrification offsets CO2-enhancement of AMF-mediated organic C decomposition Cheng et al. 2012. Science
The Summary Fig. 4. A conceptual framework of AMF-mediated decomposition driven by CO2-enhancement of plant N acquisition. CO2-enhancement of AMF primes residue decomposition and ammonium (NH4+) release and optimizes NH4+ acquisition, while reducing nitrification. Cheng et al. 2012. Science
Potential Implications • The contribution of arbuscular mycorhizal fungi to soil C sequestration under future CO2 scenarios may have been over-estimated; • Increasing plant N use efficiency and reducing decomposition through effective management of soil N transformations are keys to facilitate soil C sequestration.
Acknowledgements Lab members (in the last 6 years): Sean Blosvies*, Xin Chen, Jared Chauncey, Lei Cheng, Mary Claire Garrison, Natalie Gross*, Anna Johnson, Marissa Lee, Lingli Liu, Karen Parker, Tomin Sa*, Qinghua Shi*, Cong Tu*, Jinping Wang*, Liang Wang, Yi Wang, Dolly Watson, Scotty Wells*, Li Zhang*, Yi Zhang*, Lishi Zhou* Major Collaborators NCSU: David Shew, Chris Reberg-Horton, Julie Grossman, Frank Louws, Mike Benson, David Bird, Mike Burton, Nancy Creamer, Marc Cubeta, Ralph Dean, Greg Hoyt, Paul Mueller, Jean Ristaino, David Ritchie, Tom Rufty, Michelle Schroeder, Wei Shi, Lane Tredway, Dolly Watson USDA-ARS: Fitz Booker, Kent Burkey Funding Agencies: USDA-NRI: Soil Processes, Pest Management Alternative, Managed Ecosystems USDA-SARE USDA_NIFA_ORG NC Center for Turfgrass Environmental Research & Education USDA-ARS Plant Research Unit