Daniel B. Metcalfe

Specific root length as an indicator of environmental change

Abstract Specific root length (SRL, m g−1) is probably the most frequently measured morphological parameter of fine roots. It is believed to characterize economic aspects of the root system and to be indicative of environmental changes. The main objectives of this paper were to review and summarize the published SRL data for different tree species throughout Europe and to assess SRL under varying environmental conditions. Meta-analysis was used to summarize the response of SRL to the following manipulated environmental conditions: fertilization, irrigation, elevated temperature, elevated CO2, Al-stress, reduced light, heavy metal stress and physical disturbance of soil. SRL was found to be strongly dependent on the fine root classes, i.e. on the ectomycorrhizal short roots (ECM), and on the roots <0.5 mm, <1 mm, <2 mm and 1 – 2 mm in diameter SRL was largest for ECM and decreased with increasing diameter. Changes in soil factors influenced most strongly the SRL of ECM and roots <0.5 mm. The variation in the SRL components, root diameter and root tissue density, and their impact on the SRL value were computed. Meta-analyses showed that SRL decreased significantly under fertilization and Al-stress; it responded negatively to reduced light, elevated temperature and CO2. We suggest that SRL can be used successfully as an indicator of nutrient availability to trees in experimental conditions.

Effect of 7 yr of experimental drought on vegetation dynamics and biomass storage of an eastern Amazonian rainforest

*At least one climate model predicts severe reductions of rainfall over Amazonia during this century. Long-term throughfall exclusion (TFE) experiments represent the best available means to investigate the resilience of the Amazon rainforest to such droughts. *Results are presented from a 7 yr TFE study at Caxiuanã National Forest, eastern Amazonia. We focus on the impacts of the drought on tree mortality, wood production and above-ground biomass. *Tree mortality in the TFE plot over the experimental period was 2.5% yr(-1), compared with 1.25% yr(-1) in a nearby control plot experiencing normal rainfall. Differences in stem mortality between plots were greatest in the largest (> 40 cm diameter at breast height (dbh)) size class (4.1% yr(-1) in the TFE and 1.4% yr(-1) in the control). Wood production in the TFE plot was c. 30% lower than in the control plot. Together, these changes resulted in a loss of 37.8 +/- 2.0 Mg carbon (C) ha(-1) in the TFE plot (2002-2008), compared with no change in the control. *These results are remarkably consistent with those from another TFE (at Tapajós National Forest), suggesting that eastern Amazonian forests may respond to prolonged drought in a predictable manner.

Death from drought in tropical forests is triggered by hydraulics not carbon starvation

Drought threatens tropical rainforests over seasonal to decadal timescales, but the drivers of tree mortality following drought remain poorly understood. It has been suggested that reduced availability of non-structural carbohydrates (NSC) critically increases mortality risk through insufficient carbon supply to metabolism (‘carbon starvation’). However, little is known about how NSC stores are affected by drought, especially over the long term, and whether they are more important than hydraulic processes in determining drought-induced mortality. Using data from the world’s longest-running experimental drought study in tropical rainforest (in the Brazilian Amazon), we test whether carbon starvation or deterioration of the water-conducting pathways from soil to leaf trigger tree mortality. Biomass loss from mortality in the experimentally droughted forest increased substantially after >10 years of reduced soil moisture availability. The mortality signal was dominated by the death of large trees, which were at a much greater risk of hydraulic deterioration than smaller trees. However, we find no evidence that the droughted trees suffered carbon starvation, as their NSC concentrations were similar to those of non-droughted trees, and growth rates did not decline in either living or dying trees. Our results indicate that hydraulics, rather than carbon starvation, triggers tree death from drought in tropical rainforest.

The fate of assimilated carbon during drought: impacts on respiration in Amazon rainforests.

Interannual variations in CO2 exchange across Amazonia, as deduced from atmospheric inversions, correlate with El Niño occurrence. They are thought to result from changes in net ecosystem exchange and fire incidence that are both related to drought intensity. Alterations to net ecosystem production (NEP) are caused by changes in gross primary production (GPP) and ecosystem respiration (Reco). Here, we analyse observations of the components of Reco (leaves, live and dead woody tissue, and soil) to provide first estimates of changes in Reco during short-term (seasonal to interannual) moisture limitation. Although photosynthesis declines if moisture availability is limiting, leaf dark respiration is generally maintained, potentially acclimating upwards in the longer term. If leaf area is lost, then short-term canopy-scale respiratory effluxes from wood and leaves are likely to decline. Using a moderate short-term drying scenario where soil moisture limitation leads to a loss of 0.5 m2 m−2 yr−1 in leaf area index, we estimate a reduction in respiratory CO2 efflux from leaves and live woody tissue of 1.0 (±0.4) t C ha−1 yr−1. Necromass decomposition declines during drought, but mortality increases; the median mortality increase following a strong El Niño is 1.1% (n=46 tropical rainforest plots) and yields an estimated net short-term increase in necromass CO2 efflux of 0.13–0.18 t C ha−1 yr−1. Soil respiration is strongly sensitive to moisture limitation over the short term, but not to associated temperature increases. This effect is underestimated in many models but can lead to estimated reductions in CO2 efflux of 2.0 (±0.5) t C ha−1 yr−1. Thus, the majority of short-term respiratory responses to drought point to a decline in Reco, an outcome that contradicts recent regional-scale modelling of NEP. NEP varies with both GPP and Reco but robust moisture response functions are clearly needed to improve quantification of the role of Reco in influencing regional-scale CO2 emissions from Amazonia.

Are ectomycorrhizal fungi alleviating or aggravating nitrogen limitation of tree growth in boreal forests

Symbioses between plant roots and mycorrhizal fungi are thought to enhance plant uptake of nutrients through a favourable exchange for photosynthates. Ectomycorrhizal fungi are considered to play this vital role for trees in nitrogen (N)-limited boreal forests. We followed symbiotic carbon (C)-N exchange in a large-scale boreal pine forest experiment by tracing (13) CO(2) absorbed through tree photosynthesis and (15) N injected into a soil layer in which ectomycorrhizal fungi dominate the microbial community. We detected little (15) N in tree canopies, but high levels in soil microbes and in mycorrhizal root tips, illustrating effective soil N immobilization, especially in late summer, when tree belowground C allocation was high. Additions of N fertilizer to the soil before labelling shifted the incorporation of (15) N from soil microbes and root tips to tree foliage. These results were tested in a model for C-N exchange between trees and mycorrhizal fungi, suggesting that ectomycorrhizal fungi transfer small fractions of absorbed N to trees under N-limited conditions, but larger fractions if more N is available. We suggest that greater allocation of C from trees to ectomycorrhizal fungi increases N retention in soil mycelium, driving boreal forests towards more severe N limitation at low N supply.

Shifts in plant respiration and carbon use efficiency at a large‐scale drought experiment in the eastern Amazon

*The effects of drought on the Amazon rainforest are potentially large but remain poorly understood. Here, carbon (C) cycling after 5 yr of a large-scale through-fall exclusion (TFE) experiment excluding about 50% of incident rainfall from an eastern Amazon rainforest was compared with a nearby control plot. *Principal C stocks and fluxes were intensively measured in 2005. Additional minor components were either quantified in later site measurements or derived from the available literature. *Total ecosystem respiration (R(eco)) and total plant C expenditure (PCE, the sum of net primary productivity (NPP) and autotrophic respiration (R(auto))), were elevated on the TFE plot relative to the control. The increase in PCE and R(eco) was mainly caused by a rise in R(auto) from foliage and roots. Heterotrophic respiration did not differ substantially between plots. NPP was 2.4 +/- 1.4 t C ha(-1) yr(-1) lower on the TFE than the control. Ecosystem carbon use efficiency, the proportion of PCE invested in NPP, was lower in the TFE plot (0.24 +/- 0.04) than in the control (0.32 +/- 0.04). *Drought caused by the TFE treatment appeared to drive fundamental shifts in ecosystem C cycling with potentially important consequences for long-term forest C storage.

Factors controlling spatio‐temporal variation in carbon dioxide efflux from surface litter, roots, and soil organic matter at four rain forest sites in the eastern Amazon

change, though experimental precipitation exclusion over a one-hectare area was associated with a ten-fold reduction in litter respiration relative to unmodified sites. The estimated mean contribution of soil organic matter respiration fell from 49% during the wet season to 32% in the dry season, while root respiration contribution increased from 42% in the wet season to 61% during the dry season. Spatial variation in respiration from soil, litter, roots, and soil organic matter was not explained by volumetric soil moisture or temperature. Instead, spatial heterogeneity in litter and root mass accounted for 44% of observed spatial variation in soil respiration (p < 0.001). In particular, variation in litter respiration per unit mass and root mass accounted for much of the observed variation in respiration from litter and roots, respectively, and hence total soil respiration. This information about patterns of, and underlying controls on, respiration from different soil components should assist attempts to accurately model soil carbon dioxide fluxes over space and time.

Short‐term dynamics of abiotic and biotic soil 13CO2 effluxes after in situ 13CO2 pulse labelling of a boreal pine forest

Physical diffusion of isotopic tracers into and out of soil pores causes considerable uncertainty for the timing and magnitude of plant belowground allocation in pulse-labelling experiments. Here, we partitioned soil CO(2) isotopic fluxes into abiotic tracer flux (physical return), heterotrophic flux, and autotrophic flux contributions following (13)CO(2) labelling of a Swedish Pinus sylvestris forest. Soil CO(2) efflux and its isotopic composition from a combination of deep and surface soil collars was monitored using a field-deployed mass spectrometer. Additionally, (13)CO(2) within the soil profile was monitored. Physical (abiotic) efflux of (13)CO(2) from soil pore spaces was found to be significant for up to 48 h after pulse labelling, and equalled the amount of biotic label flux over 6 d. Measured and modelled changes in (13)CO(2) concentration throughout the soil profile corroborated these results. Tracer return via soil CO(2) efflux correlated significantly with the proximity of collars to trees, while daily amplitudes of total flux (including heterotrophic and autotrophic sources) showed surprising time shifts compared with heterotrophic fluxes. The results show for the first time the significance of the confounding influence of physical isotopic CO(2)-tracer return from the soil matrix, calling for the inclusion of meaningful control treatments in future pulse-chase experiments.

The linkages between photosynthesis, productivity, growth and biomass in lowland Amazonian forests

Understanding the relationship between photosynthesis, net primary productivity and growth in forest ecosystems is key to understanding how these ecosystems will respond to global anthropogenic change, yet the linkages among these components are rarely explored in detail. We provide the first comprehensive description of the productivity, respiration and carbon allocation of contrasting lowland Amazonian forests spanning gradients in seasonal water deficit and soil fertility. Using the largest data set assembled to date, ten sites in three countries all studied with a standardized methodology, we find that (i) gross primary productivity (GPP) has a simple relationship with seasonal water deficit, but that (ii) site-to-site variations in GPP have little power in explaining site-to-site spatial variations in net primary productivity (NPP) or growth because of concomitant changes in carbon use efficiency (CUE), and conversely, the woody growth rate of a tropical forest is a very poor proxy for its productivity. Moreover, (iii) spatial patterns of biomass are much more driven by patterns of residence times (i.e. tree mortality rates) than by spatial variation in productivity or tree growth. Current theory and models of tropical forest carbon cycling under projected scenarios of global atmospheric change can benefit from advancing beyond a focus on GPP. By improving our understanding of poorly understood processes such as CUE, NPP allocation and biomass turnover times, we can provide more complete and mechanistic approaches to linking climate and tropical forest carbon cycling.

The productivity, metabolism and carbon cycle of two lowland tropical forest plots in south-western Amazonia, Peru

Background: The forests of western Amazonia are known to be more dynamic that the better-studied forests of eastern Amazonia, but there has been no comprehensive description of the carbon cycle of a western Amazonian forest. Aims: We present the carbon budget of two forest plots in Tambopata in south-eastern Peru, western Amazonia. In particular, we present, for the first time, the seasonal variation in the detailed carbon budget of a tropical forest. Methods: We measured the major components of net primary production (NPP) and total autotrophic respiration over 3–6 years. Results: The NPP for the two plots was 15.1 ± 0.8 and 14.2 ± 1.0 Mg C ha−1 year−1, the gross primary productivity (GPP) was 35.5 ± 3.6 and 34.5 ± 3.5 Mg C ha−1 year−1, and the carbon use efficiency (CUE) was 0.42 ± 0.05 and 0.41 ± 0.05. NPP and CUE showed a large degree of seasonality. Conclusions: The two plots were similar in carbon cycling characteristics despite the different soils, the most notable difference being high allocation of NPP to canopy and low allocation to fine roots in the Holocene floodplain plot. The timing of the minima in the wet–dry transition suggests they are driven by phenological rhythms rather than being driven directly by water stress. When compared with results from forests on infertile forests in humid lowland eastern Amazonia, the plots have slightly higher GPP, but similar patterns of CUE and carbon allocation.