Peter S. Curtis

THE META‐ANALYSIS OF RESPONSE RATIOS IN EXPERIMENTAL ECOLOGY

Meta-analysis provides formal statistical techniques for summarizing the results of independent experiments and is increasingly being used in ecology. The response ratio (the ratio of mean outcome in the experimental group to that in the control group) and closely related measures of proportionate change are often used as measures of effect magnitude in ecology. Using these metrics for meta-analysis requires knowledge of their statistical properties, but these have not been previously derived. We give the approximate sampling distribution of the log response ratio, discuss why it is a particularly useful metric for many applications in ecology, and demonstrate how to use it in meta-analysis. The meta-analysis of response-ratio data is illustrated using experimental data on the effects of increased atmospheric CO2 on plant biomass responses.

A meta-analysis of elevated CO2 effects on woody plant mass, form, and physiology

Abstract Quantitative integration of the literature on the effect of elevated CO2 on woody plants is important to aid our understanding of forest health in coming decades and to better predict terrestrial feedbacks on the global carbon cycle. We used meta-analytic methods to summarize and interpret more than 500 reports of effects of elevated CO2 on woody plant biomass accumulation and partitioning, gas exchange, and leaf nitrogen and starch content. The CO2 effect size metric we used was the log-transformed ratio of elevated compared to ambient response means weighted by the inverse of the variance of the log ratio. Variation in effect size among studies was partitioned according to the presence of interacting stress factors, length of CO2 exposure, functional group status, pot size, and type of CO2 exposure facility. Both total biomass (WT) and net CO2 assimilation (A) increased significantly at about twice ambient CO2, regardless of growth conditions. Low soil nutrient availability reduced the CO2 stimulation of WT by half, from +31% under optimal conditions to +16%, while low light increased the response to +52%. We found no significant shifts in biomass allocation under high CO2. Interacting stress factors had no effect on the magnitude of responses of A to CO2, although plants grown in growth chambers had significantly lower responses (+19%) than those grown in greenhouses or in open-top chambers (+54%). We found no consistent evidence for photosynthetic acclimation to CO2 enrichment except in trees grown in pots <0.5 l (−36%) and no significant CO2 effect on stomatal conductance. Both leaf dark respiration and leaf nitrogen were significantly reduced under elevated CO2 (−18% and −16% respectively, data expressed on a leaf mass basis), while leaf starch content increased significantly except in low nutrient grown gymnosperms. Our results provide robust, statistically defensible estimates of elevated CO2 effect sizes against which new results may be compared or for use in forest and climate model parameterization.

Effects of forest management on soil C and N storage: meta analysis

Abstract The effects of forest management on soil carbon (C) and nitrogen (N) are important to understand not only because these are often master variables determining soil fertility but also because of the role of soils as a source or sink for C on a global scale. This paper reviews the literature on forest management effects on soil C and N and reports the results of a meta analysis of these data. The meta analysis showed that forest harvesting, on average, had little or no effect on soil C and N. Significant effects of harvest type and species were noted, with sawlog harvesting causing increases (+18%) in soil C and N and whole-tree harvesting causing decreases (−6%). The positive effect of sawlog harvesting appeared to be restricted to coniferous species. Fire resulted in no significant overall effects of fire on either C or N (when categories were combined); but there was a significant effect of time since fire, with an increase in both soil C and N after 10 years (compared to controls). Significant differences among fire treatments were found, with the counterintuitive result of lower soil C following prescribed fire and higher soil C following wildfire. The latter is attributed to the sequestration of charcoal and recalcitrant, hydrophobic organic matter and to the effects of naturally invading, post-fire, N-fixing vegetation. Both fertilization and N-fixing vegetation caused marked overall increases in soil C and N.

NITROGEN ADDITIONS AND LITTER DECOMPOSITION: A META‐ANALYSIS

We conducted a meta-analysis of previously published empirical studies that have examined the effects of nitrogen (N) enrichment on litter decomposition. Our objective was to provide a synthesis of existing data that comprehensively and quantitatively evaluates how environmental and experimental factors interact with N additions to influence litter mass loss. Nitrogen enrichment, when averaged across all studies, had no statistically significant effect on litter decay. However, we observed significant effects of fertilization rate, site-specific ambient N-deposition level, and litter quality. Litter decomposition was inhibited by N additions when fertilization rates were 2-20 times the anthropogenic N- deposition level, when ambient N deposition was 5-10 kg N·ha 21 ·yr 21 , or when litter quality was low (typically high-lignin litters). Decomposition was stimulated at field sites exposed to low ambient N deposition (,5 kg N·ha 21 ·yr 21 ) and for high-quality (low-lignin) litters. Fertilizer type, litterbag mesh size, and climate did not influence the litter decay response to N additions.

Elevated atmospheric CO2 and feedback between carbon and nitrogen cycles

We tested a conceptual model describing the influence of elevated atmospheric CO2 on plant production, soil microorganisms, and the cycling of C and N in the plant-soil system. Our model is based on the observation that in nutrient-poor soils, plants (C3) grown in an elevated CO2 atmosphere often increase production and allocation to belowground structures. We predicted that greater belowground C inputs at elevated CO2 should elicit an increase in soil microbial biomass and increased rates of organic matter turnover and nitrogen availability. We measured photosynthesis, biomass production, and C allocation of Populus grandidentata Michx. grown in nutrient-poor soil for one field season at ambient and twice-ambient (i.e., elevated) atmospheric CO2 concentrations. Plants were grown in a sandy subsurface soil i) at ambient CO2 with no open top chamber, ii) at ambient CO2 in an open top chamber, and iii) at twice-ambient CO2 in an open top chamber. Plants were fertilized with 4.5 g N m−2 over a 47 d period midway through the growing season. Following 152 d of growth, we quantified microbial biomass and the availabilities of C and N in rhizosphere and bulk soil. We tested for a significant CO2 effect on plant growth and soil C and N dynamics by comparing the means of the chambered ambient and chambered elevated CO2 treatments.Rates of photosynthesis in plants grown at elevated CO2 were significantly greater than those measured under ambient conditions. The number of roots, root length, and root length increment were also substantially greater at elevated CO2. Total and belowground biomass were significantly greater at elevated CO2. Under N-limited conditions, plants allocated 50–70% of their biomass to roots. Labile C in the rhizosphere of elevated-grown plants was significantly greater than that measured in the ambient treatments; there were no significant differences between labile C pools in the bulk soil of ambient and elevated-grown plants. Microbial biomass C was significantly greater in the rhizosphere and bulk soil of plants grown at elevated CO2 compared to that in the ambient treatment. Moreover, a short-term laboratory assay of N mineralization indicated that N availability was significantly greater in the bulk soil of the elevated-grown plants. Our results suggest that elevated atmospheric CO2 concentrations can have a positive feedback effect on soil C and N dynamics producing greater N availability. Experiments conducted for longer periods of time will be necessary to test the potential for negative feedback due to altered leaf litter chemistry. ei]{gnH}{fnLambers} ei]{gnA C}{fnBorstlap}

Biometric and eddy-covariance based estimates of annual carbon storage in five eastern North American deciduous forests

Quantifying net carbon (C) storage by forests is a necessary step in the validation of carbon sequestration estimates and in assessing the possible role of these ecosystems in offsetting fossil fuel emissions. In eastern North America, five sites were established in deciduous forests to provide measurements of net ecosystem CO2 exchange (NEE) using micro-meteorological methods, and measures of major carbon pools and fluxes, using a combination of forest mensurational, eco-physiological, and other biometric methods. The five study sites, part of the AmeriFlux network, ranged across 10 ◦ of latitude and 18 ◦ of longitude, but were all of similar age, canopy height, and stand basal area. Here we present a cross-site synthesis of annual carbon storage estimates, comparing meteorological and biometric approaches, and also comparing biometric estimates based on analyses of autotrophic carbon pools and heterotrophic carbon fluxes (net ecosystem production, NEP) versus those based on measurements of change in two major carbon pools (� C). Annual above-ground net primary production (ANPP) varied nearly two-fold among sites and was strongly correlated with average annual temperature and with annual soil nitrogen mineralization (Nmin). Estimates of NEP ranged from 0.7 Mg C per hectare per year in northern lower Michigan to 3.5 Mg C per hectare per year in central Indiana, and were also well correlated with Nmin. There was less variation among sites in estimates

Meta-analysis in ecology

Abstract Meta-analysis is the statistical synthesis of the results of separate studies. It was adapted from other disciplines for use in ecology and evolutionary biology beginning in the early 1990s, and, at the turn of the century, has begun to have a substantial impact on the way data are summarized in these fields. We identify 119 studies concerned with meta-analysis in ecology and evolution, the earliest published in 1991 and the most recent in 2000. We introduce the statistical methods used in modern meta-analysis with references to the well-developed literature in the field. These formal, statistically defensible methods have been established to determine average treatment effects across studies when a common research question is being investigated, to establish confidence limits around the average effect size, and to test for consistency or lack of agreement in effect size as well as explanations for differences in the magnitude of the effect among studies. Problems with popular but statistically flawed methods for the quantitative summary of research results have been pointed out, and their use is diminishing. We discuss a number of challenges and threats to the validity of meta-analysis in ecology and evolution. In particular, we examine how difficulties resulting from missing data, publication bias, data quality and data exclusion, non-independence among observations, and the combination of dissimilar data sets may affect the perceived utility of meta-analysis in these fields and the soundness of conclusions drawn from its application. We highlight particular applications of meta-analysis in ecology and evolution, discuss several controversies surrounding individual meta-analyses, and outline some of the practical issues involved in carrying out a meta-analysis. Finally, we suggest changes that would improve the quality of data synthesis in ecology and evolutionary biology, and predict future directions for this emerging enterprise.

Belowground carbon allocation in forests estimated from litterfall and IRGA-based soil respiration measurements

Allocation of C to belowground plant structures is one of the most important, yet least well quantified fluxes of C in terrestrial ecosystems. In a literature review of mature forests worldwide, Raich and Nadelhoffer (1989) suggested that total belowground carbon allocation (TBCA) could be estimated from the difference between annual rates of soil respiration and aboveground litterfall. Here we analyze new measurements of soil respiration and litterfall, including data from the Ameriflux network. Our results generally agree with Raich and Nadelhoffer’s previous work. A regression analysis of data from mature forests produced the following relationship: annual soil respiration = 287 + 2.80 × annual litterfall. This regression slope indicates that, on average, soil respiration is roughly three times ab oveground litterfall-C, which further implies that TBCA is roughly twice annual aboveground litterfall-C. These inferences are based on the uncertain assumption of soil C stocks being at steady state. Nevertheless, changes in soil C would have to be very large to modify the conclusion that TBCA is generally much larger than litterfall. Among only mature temperate hardwood forests, however, the correlation between litterfall and soil respiration was poor, and the correlation among years for a single site was also poor. Therefore, the regression cannot be relied upon to provide accurate estimates of soil respiration or TBCA for individual sites. Moreover, interannual variation in TBCA, short-term changes in C stocks, or different temporal scales controlling leaf litter production and soil respiration may cause important deviations from the global average. The regression slope for data from young forests is steeper, possibly indicating proportionally greater TBCA, but the steady-state assumption is more problematic for young forests. This method

INTERACTIVE EFFECTS OF ATMOSPHERIC CO2 AND SOIL‐N AVAILABILITY ON FINE ROOTS OF POPULUS TREMULOIDES

The objective of this experiment was to understand how atmospheric carbon dioxide (CO2) and soil-nitrogen (N) availability influence Populus tremuloides fine-root growth and morphology. Soil-N availability may limit the growth response of forests to elevated CO2 and interact with atmospheric CO2 to alter litter quality and ecosystem carbon (C) and N cycling. We established a CO2 × N factorial field experiment and grew six genotypes of P. tremuloides for 2.5 growing seasons in 20 large open-top chamber/root-box experimental units at the University of Michigan Biological Station in northern lower Michigan (USA). In this paper we describe an integrated examination of how atmospheric CO2 and soil-N availability influence fine-root morphology, growth, mortality, and biomass. We also studied the relationship between root biomass and total soil respiration. Over 80% of the absorbing root length of P. tremuloides was accounted for by roots <0.4 mm in diameter, and specific root length (100–250 m/g) was much great...

ATMOSPHERIC CO2 AND THE COMPOSITION AND FUNCTION OF SOIL MICROBIAL COMMUNITIES

Elevated atmospheric CO2 has the potential to increase the production and alter the chemistry of organic substrates entering soil from plant production, the magnitude of which is constrained by soil-N availability. Because microbial growth in soil is limited by substrate inputs from plant production, we reasoned that changes in the amount and chemistry of these organic substrates could affect the composition of soil microbial com- munities and the cycling of N in soil. We studied microbial community composition and soil-N transformations beneath Populus tremuloides Michx. growing under experimental atmospheric CO2 (35.7 and 70.7 Pa) and soil-N-availability (low N 5 61 ng N·g 21 ·d 21 and high N 5 319 ng N·g 21 ·d 21 ) treatments. Atmospheric CO2 concentration was modified in large, open-top chambers, and we altered soil-N availability in open-bottom root boxes by mixing different proportions of A and C horizon material. We used phospholipid fatty-acid analysis to gain insight into microbial community composition and coupled this analysis to measurements of soil-N transformations using 15 N-pool dilution techniques. The infor- mation presented here is part of an integrated experiment designed to elucidate the phys- iological mechanisms controlling the flow of C and N in the plant-soil system. Our ob- jectives were (1) to determine whether changes in plant growth and tissue chemistry alter microbial community composition and soil-N cycling in response to increasing atmospheric CO2 and soil-N availability and (2) to integrate the results of our experiment into a synthesis of elevated atmospheric CO2 and the cycling of C and N in terrestrial ecosystems. After 2.5 growing seasons, microbial biomass, gross N mineralization, microbial im- mobilization, and nitrification (gross and net) were equivalent at ambient and elevated CO2, suggesting that increases in fine-root production and declines in fine-root N concentration were insufficient to alter the influence of native soil organic matter on microbial physiology; this was the case in both low- and high-N soil. Similarly, elevated CO2 did not alter the proportion of bacterial, actinomycetal, or fungal phospholipid fatty acids in low-N or high-N soil, indicating that changes in substrate input from greater plant growth under elevated CO2 did not alter microbial community composition. Our results differ from a substantial number of studies reporting increases and decreases in soil-N cycling under elevated CO 2. From our analysis, it appears that soil-N cycling responds to elevated atmospheric CO 2 in experimental situations where plant roots have fully colonized the soil and root-associated C inputs are sufficient to modify the influence of native soil organic matter on microbial physiology. In young developing ecosystems where plant roots have not fully exploited the soil, microbial metabolism appears to be regulated by relatively large pools of soil organic matter, rather than by the additional input of organic substrates under elevated CO 2.