Rachel E. Gallery, Nicole Trahan, Emily Dynes, Dawson Fairbanks
Transcription
Rachel E. Gallery, Nicole Trahan, Emily Dynes, Dawson Fairbanks
Short-term responses of soil microbe communities to fire: functional shifts in high elevation mixed conifer forests Rachel E. Gallery, Nicole Trahan, Emily Dynes, Dawson Fairbanks, David J.P. Moore Summary • Fires and insect outbreaks are increasing in frequency and severity across the western U.S. with enormous and variable impacts on regional carbon, nitrogen, and water cycling. • Wildfires heat surface soils >200°C, changing soil physical, chemical, and biological traits. • In the short term, fire causes a wholesale transformations of organic compound substrate available for microbial metabolism; disturbance severity can alter succession trajectories. • In mixed conifer forests, shifts from carbon to nitrogen dominated enzyme activities in burned forests with alkaline soils suggest loss of microbial taxa and function. • Together, microbial biomass, substrate quality, and enzyme activity influence coupled biogeochemical cycles in ecosystems; responses to disturbance • Issue: Regional carbon models without explicit microbial processes may work well in equilibrium conditions but cannot replicate observed biogeochemical changes after disturbance. Approach: Quantify the post-fire response of microbial function in forest ecosystems to enable testing of microbial processes models. 2012 High Park Fire, CO A Enzymes link microbial community structure to biogeochemical process: Soil CO2 efflux, microbial activity, and organic substrates change post-fire UNBURNED BURNED Fig 3a. Fire alters soil respiration (Rs; soil efflux) rates that correspond with changes in soil chemistry such as pH. pH of unburned soils, which include MPB killed forests, were more acidic due to organic matter inputs, which correlates with higher average soil efflux rates. Soils from burned forests were more alkaline but also had higher range in pH likely due to differences in burn severity. Soil efflux rates were lower in burned forests reflecting low substrate and microbial populations. Fig 3b. The weak correlation between microbial biomass carbon (C) and soil efflux rates could be explained by the relative importance of other constituents of respiration (e.g., autotrophic), which are also affected by fire. Microbial biomass C is lower in burned soils though shifts in microbial community function (Fig 3c) may be more important drivers of Rs. 2013 Thompson Ridge Fire, NM B Fig 3c. β-glucosidase (cellulose decomposition; black) and leucine amino-peptidase (protein degradation; red) are the most widely measured glycosidase and aminopeptidase activities because they hydrolyze the most abundant C and N substrates respectively. β-glucosidase activity is lower in burned soils and correlates with microbial biomass C patterns (Fig 3b). Leucine amino-peptidase activity is higher in burned soils corresponding with higher protein relative abundance (Fig 4). Year of infestation 2000 2006 2001 2007 2002 2008 2003 2009 2004 2010 New and integrated data needed to test heterotrophic respiration models FEF Extended Area 0.6 High Park Fire Fig 1. (A) In the high elevation conifer forests of the Colorado Rocky Mountains, soil samples from 56 plots (3m radius) were sampled across 5 locations along a chronosequence of fire and mountain pine beetle (MPB) killed forests after the 2012 High Park Fire (black perimeter). (B) Soil samples from 22 soil pits across a gradient of burn severities in a mixed-conifer zero order basin from the Thompson Ridge Fire in the Jemez River Basin CZO were collected 18 days following the June 2013 fire. Soil CO2 efflux is correlated with microbial biomass in unburned forests Relative abundance (%) 2005 lignin phenol unknown protein 0.5 0.4 lipid Fig 4. Changes in microbial substrates can be N-bearing quantified using pyrolysis gas chromatographic polysaccharide mass spectroscopy (py-GC-MS). The relative abundance of different organic compounds by disturbance type is shown. Our preliminary estimates of labile polysaccharide pools and relative protein abundance appears to track observed β-glucosidase and leucine aminopeptidase activities (Fig 3c). 0.3 0.2 0.1 0 Fig 2. Across a gradient of forest plots containing live trees and those killed by either mountain pine beetle or by mechanical girdling, microbial biomass was a strong predictor of respiration (soil efflux; A) however spatial variation in soil efflux rates across the disturbance gradient was not well explained by coincident measurements of soil temperature or ,moisture (B and Table). In current Earth System Models (ESMs), heterotrophic respiration is typically modeled as a function of available carbon substrate, temperature and moisture (Shao et al. 2013). However when an ecological disturbance causes changes to the availability of substrate and the success of the microbial community, these simple predictors are unlikely to be sufficient. live beetle-kill Autotrophic turnover Dissolved Organic Carbon Uptake Vmax, Km Vmax, Km Catalyses Enzyme decay Microbial Biomass Carbon Soil Organic Carbon Enzymes Enzyme production Microbial turnover Modified from Allison et al. (2010). Methods: Soil efflux was measured using a LICOR LI-6400 infrared gas analyzer. Chloroform fumigated K2SO4 soil extracts from the chronosequences were quantified for microbial biomass carbon using the non-purge able-organic-C protocol on a Shimadzu TOC 5000 total organic carbon analyzer. Soil microbial biomass C was estimated by subtracting concentrations of Dissolved Organic Carbon in paired unfumigated samples and soil free controls. Potential activity of hydrolytic enzymes was measured using established fluorometric techniques with labeled substrate. β-glucosidase [BG] is a cellulase involved in cellulose decomposition to glucose. Leucine aminopeptidase [LAP] is involved in protein degradation. Fig 5b. Links between organic matter (C, N, P), microbial biomass C and N production, and exoenzyme activities, which are in part influenced by environmental cues. burn, beetle-kill After fire, changes in both microbial community function and organic substrate quality are evident. Both influence coupled biogeochemical cycles in ecosystems. Fig 5a. Soil microbes excrete exoenzymes to decompose polymers and acquire nutrients. These activities respond to disturbance and may indicate changing microbial function or soil quality. Rh estimates could be improved by considering microbial enzyme activity responses to effects of: temperature, moisture, microbial biomass C, and population dynamics. Polyphenols & hydrocarbons Lignin Tannin Cutin Suberin Fatty acids Humic substrate Organic nitrogen Protein Chitin Peptidoglycan Exoenzyme processing Polysaccharides Cellulose Hemicellulose Pectin Starch Chitin Peptidoglycan Organic phosphorus Nucleic acids phospholipids Inositol phosphates Exopolysaccharides CO2 Microbial Biomass C Production Assimilation ENVIRONMENT ENZYMES CELL Redrawn from Sinsabaugh and Shah (2012). X-CZO activities and CZ services: These results highlight the importance of incorporating a standard suite of microbial activity and community assays to enable comparisons across the CZO network and with other observatories, providing regional-scale microbial biodiversity and activity data in a standardized framework that can be used to consider CZ services, thus enabling more effective management and valuation of ecosystem services and improving ecosystem projections under global change scenarios. Acknowledgements: This research was funded by the U.S. National Science Foundation grants EAR-0724958 and EAR1331408 provided in support of the Catalina-Jemez Critical Zone Observatory. Funding to DJPM was provided by the U.S. Department Of Energy Terrestrial Ecosystem Science Program DES-C2365077 and by NSF RAPID 1262012. References: Shao et al. (2013); doi:10.1088/1748-9326/8/3/034034 Sinsabaugh and Shah (2012) DOI: 10.1146/annurevecolsys-071112-124414; Allison et al. (2010) doi:10.1038/ngeo846