Magnus Lund

Joint control of terrestrial gross primary productivity by plant phenology and physiology

Significance Terrestrial gross primary productivity (GPP), the total photosynthetic CO2 fixation at ecosystem level, fuels all life on land. However, its spatiotemporal variability is poorly understood, because GPP is determined by many processes related to plant phenology and physiological activities. In this study, we find that plant phenological and physiological properties can be integrated in a robust index—the product of the length of CO2 uptake period and the seasonal maximal photosynthesis—to explain the GPP variability over space and time in response to climate extremes and during recovery after disturbance. Terrestrial gross primary productivity (GPP) varies greatly over time and space. A better understanding of this variability is necessary for more accurate predictions of the future climate–carbon cycle feedback. Recent studies have suggested that variability in GPP is driven by a broad range of biotic and abiotic factors operating mainly through changes in vegetation phenology and physiological processes. However, it is still unclear how plant phenology and physiology can be integrated to explain the spatiotemporal variability of terrestrial GPP. Based on analyses of eddy–covariance and satellite-derived data, we decomposed annual terrestrial GPP into the length of the CO2 uptake period (CUP) and the seasonal maximal capacity of CO2 uptake (GPPmax). The product of CUP and GPPmax explained >90% of the temporal GPP variability in most areas of North America during 2000–2010 and the spatial GPP variation among globally distributed eddy flux tower sites. It also explained GPP response to the European heatwave in 2003 (r2 = 0.90) and GPP recovery after a fire disturbance in South Dakota (r2 = 0.88). Additional analysis of the eddy–covariance flux data shows that the interbiome variation in annual GPP is better explained by that in GPPmax than CUP. These findings indicate that terrestrial GPP is jointly controlled by ecosystem-level plant phenology and photosynthetic capacity, and greater understanding of GPPmax and CUP responses to environmental and biological variations will, thus, improve predictions of GPP over time and space.

The uncertain climate footprint of wetlands under human pressure

Significance Wetlands are unique ecosystems because they are in general sinks for carbon dioxide and sources of methane. Their climate footprint therefore depends on the relative sign and magnitude of the land–atmosphere exchange of these two major greenhouse gases. This work presents a synthesis of simultaneous measurements of carbon dioxide and methane fluxes to assess the radiative forcing of natural wetlands converted to agricultural or forested land. The net climate impact of wetlands is strongly dependent on whether they are natural or managed. Here we show that the conversion of natural wetlands produces a significant increase of the atmospheric radiative forcing. The findings suggest that management plans for these complex ecosystems should carefully account for the potential biogeochemical effects on climate. Significant climate risks are associated with a positive carbon–temperature feedback in northern latitude carbon-rich ecosystems, making an accurate analysis of human impacts on the net greenhouse gas balance of wetlands a priority. Here, we provide a coherent assessment of the climate footprint of a network of wetland sites based on simultaneous and quasi-continuous ecosystem observations of CO2 and CH4 fluxes. Experimental areas are located both in natural and in managed wetlands and cover a wide range of climatic regions, ecosystem types, and management practices. Based on direct observations we predict that sustained CH4 emissions in natural ecosystems are in the long term (i.e., several centuries) typically offset by CO2 uptake, although with large spatiotemporal variability. Using a space-for-time analogy across ecological and climatic gradients, we represent the chronosequence from natural to managed conditions to quantify the “cost” of CH4 emissions for the benefit of net carbon sequestration. With a sustained pulse–response radiative forcing model, we found a significant increase in atmospheric forcing due to land management, in particular for wetland converted to cropland. Our results quantify the role of human activities on the climate footprint of northern wetlands and call for development of active mitigation strategies for managed wetlands and new guidelines of the Intergovernmental Panel on Climate Change (IPCC) accounting for both sustained CH4 emissions and cumulative CO2 exchange.

Annual CO2 balance of a temperate bog

Peatlands are generally small sinks for atmospheric CO2. However, the sustainability of this sink functioning is threatened in a changing climate. We measured the CO2 exchange in a temperate bog between August 2005 and July 2006 using the eddy covariance technique. During this period, the CO2 balance was –78.6 ± 20.0 g CO2 m-2 yr-1, which is a lower uptake than others have reported for comparable ecosystems, but in accordance with average Holocene uptake rates. Average winter emissions were small (11 μg CO2 m-2 s-1), but the accumulated non-growing season fluxes (121 g CO2m-2) represent a larger flux than the annual balance. Taking into consideration that wintertime temperatures in the northern hemisphere are expected to increase in the future, wintertime fluxes may become increasingly important. The highest instantaneous monthly average uptake occurred in June (–38 μg CO2 m-2 s-1), while the drier month of July showed a distinctly smaller sink (–22 μg CO2 m-2s-1). It is suggested that soil drying caused gross primary production to decline, while nighttime respiration showed a steadily increase. Our findings indicate that the effect of future warmer temperatures on the growing season CO2 exchange in peatlands is dependent on soil wetness.

Environmental controls on the CO2 exchange in north Europea mires

Net CO2 exchange measured under well-mixed atmospheric conditions in four different mires in Sweden and Finland were used to analyse which factors were controlling photosynthesis and respiration. The parameters of a light response function showed strong seasonal variations with similar behaviour for all mires. The half-monthly nighttime respiration rates in the central part of the growing season were about two times higher in the southernmost, warmest site, Fåje, as compared to the northernmost, coldest site, Kaamanen. However, Kaamanen had high photosynthesis rates, and this in combination with the long daylight periods in the middle of the summer caused Kaamanen to have the largest net ecosystem exchange (NEE) during the summer period. Fåje that showed the highest productivity had also the highest respiration and therefore, the lowest NEE during summer. Correlation between half-monthly components and different environmental variables showed the highest correlation between the components themselves. Thereafter came temperature except for Fåje where water table depth (WTD) explained most of the variance both for detrended and temperature-normalized components. All sites showed dependencies between WTD and the respective components during drying up periods. Temperature sensitivity was higher for productivity than for respiration indicating that CO2 uptake would increase during global warming.

Response of an arctic predator guild to collapsing lemming cycles

Alpine and arctic lemming populations appear to be highly sensitive to climate change, and when faced with warmer and shorter winters, their well-known high-amplitude population cycles may collapse. Being keystone species in tundra ecosystems, changed lemming dynamics may convey significant knock-on effects on trophically linked species. Here, we analyse long-term (1988–2010), community-wide monitoring data from two sites in high-arctic Greenland and document how a collapse in collared lemming cyclicity affects the population dynamics of the predator guild. Dramatic changes were observed in two highly specialized lemming predators: snowy owl and stoat. Following the lemming cycle collapse, snowy owl fledgling production declined by 98 per cent, and there was indication of a severe population decline of stoats at one site. The less specialized long-tailed skua and the generalist arctic fox were more loosely coupled to the lemming dynamics. Still, the lemming collapse had noticeable effects on their reproductive performance. Predator responses differed somewhat between sites in all species and could arise from site-specific differences in lemming dynamics, intra-guild interactions or subsidies from other resources. Nevertheless, population extinctions and community restructuring of this arctic endemic predator guild are likely if the lemming dynamics are maintained at the current non-cyclic, low-density state.

Thermal optimality of net ecosystem exchange of carbon dioxide and underlying mechanisms.

• It is well established that individual organisms can acclimate and adapt to temperature to optimize their functioning. However, thermal optimization of ecosystems, as an assemblage of organisms, has not been examined at broad spatial and temporal scales. • Here, we compiled data from 169 globally distributed sites of eddy covariance and quantified the temperature response functions of net ecosystem exchange (NEE), an ecosystem-level property, to determine whether NEE shows thermal optimality and to explore the underlying mechanisms. • We found that the temperature response of NEE followed a peak curve, with the optimum temperature (corresponding to the maximum magnitude of NEE) being positively correlated with annual mean temperature over years and across sites. Shifts of the optimum temperature of NEE were mostly a result of temperature acclimation of gross primary productivity (upward shift of optimum temperature) rather than changes in the temperature sensitivity of ecosystem respiration. • Ecosystem-level thermal optimality is a newly revealed ecosystem property, presumably reflecting associated evolutionary adaptation of organisms within ecosystems, and has the potential to significantly regulate ecosystem-climate change feedbacks. The thermal optimality of NEE has implications for understanding fundamental properties of ecosystems in changing environments and benchmarking global models.

Effects of drought conditions on the carbon dioxide dynamics in a temperate peatland

Drought is arguably the most important regulator of inter-annual variation in net ecosystem CO2 exchange (NEE) in peatlands. This study investigates effects of drought periods on NEE and its components, gross primary production (GPP) and ecosystem respiration (Reco), on the basis of eddy covariance measurements of land–atmosphere exchange of CO2 in 2006–2009 in a south Swedish nutrient-poor peatland. Two drought periods had dissimilar effects on the CO2 exchange. In 2006, there was a short but severe drought period in the middle of the growing season resulting in increased Reco rates, but no detectable effect on GPP rates. In contrast, in 2008 the drought period began early in the growing season and lasted for a longer period of time, resulting in reduced GPP rates, suggesting that GPP is most sensitive to drought during leaf out and canopy development compared with the full canopy stage. Both in 2006 and in 2008 the peatland acted as an annual source of atmospheric CO2, while in 2007 and 2009, when there were no drought periods, the peatland constituted a CO2 sink. It was concluded that the timing, severity and duration of drought periods regulate the effects on peatland GPP, Reco and NEE.

Trends in CO2 exchange in a high Arctic tundra heath, 2000–2010

We have measured the land-atmosphere CO2 exchange using the eddy covariance technique in a high Arctic tundra heath in northeast Greenland (Zackenberg). On the basis of 11 years of measurements (2000-2010), it was found that snow cover dynamics was important for the CO2 exchange. The start of CO2 uptake period correlated significantly with timing of snowmelt. Furthermore, for years with deep and long-lasting snowpacks, the following springs showed increased CO2 emission rates. In the first part of the study period, there was an increase of approximately 8 g C m(-2) yr(-1) in both accumulated gross primary production (GPP) and CO2 sink strength during summer. However, in the last few years, there were no significant changes in GPP, whereas ecosystem respiration (R-eco) increased (8.5 g C m(-2) yr(-1)) and ecosystem CO2 sink strength weakened (-4.1 g C m(-2) yr(-1)). It was found that temperature and temperature-related variables (maximum thaw depth and growing degree days) controlled the interannual variation in CO2 exchange. However, while R-eco showed a steady increase with temperature (5.8 g C m(-2) degrees C-1), the initial increase in GPP with temperature leveled off at the high end of observed temperature range. This suggests that future increases in temperature will weaken the ecosystem CO2 sink strength or even turn it into a CO2 source, depending on possible changes in vegetation structure and functioning as a response to a changing climate. If this trend is applicable also to other Arctic ecosystems, it will have implications for our current understanding of Arctic ecosystems dynamics. (Less)

Seasonal variation of photosynthetic model parameters and leaf area index from global Fluxnet eddy covariance data

Global vegetation models require the photosynthetic parameters, maximum carboxylation capacity (V(cm)), and quantum yield (alpha) to parameterize their plant functional types (PFTs). The purpose of this work is to determine how much the scaling of the parameters from leaf to ecosystem level through a seasonally varying leaf area index (LAI) explains the parameter variation within and between PFTs. Using Fluxnet data, we simulate a seasonally variable LAI(F) for a large range of sites, comparable to the LAI(M) derived from MODIS. There are discrepancies when LAI(F) reach zero levels and LAI(M) still provides a small positive value. We find that temperature is the most common constraint for LAI(F) in 55% of the simulations, while global radiation and vapor pressure deficit are the key constraints for 18% and 27% of the simulations, respectively, while large differences in this forcing still exist when looking at specific PFTs. Despite these differences, the annual photosynthesis simulations are comparable when using LAI(F) or LAIM (r(2) = 0.89). We investigated further the seasonal variation of ecosystem-scale parameters derived with LAI(F). V(cm) has the largest seasonal variation. This holds for all vegetation types and climates. The parameter alpha is less variable. By including ecosystem-scale parameter seasonality we can explain a considerable part of the ecosystem-scale parameter variation between PFTs. The remaining unexplained leaf-scale PFT variation still needs further work, including elucidating the precise role of leaf and soil level nitrogen.

Latent heat exchange in the boreal and arctic biomes

In this study latent heat flux (λE) measurements made at 65 boreal and arctic eddy-covariance (EC) sites were analyses by using the Penman-Monteith equation. Sites were stratified into nine different ecosystem types: harvested and burnt forest areas, pine forests, spruce or fir forests, Douglas-fir forests, broadleaf deciduous forests, larch forests, wetlands, tundra and natural grasslands. The Penman-Monteith equation was calibrated with variable surface resistances against half-hourly eddy-covariance data and clear differences between ecosystem types were observed. Based on the modeled behavior of surface and aerodynamic resistances, surface resistance tightly control λE in most mature forests, while it had less importance in ecosystems having shorter vegetation like young or recently harvested forests, grasslands, wetlands and tundra. The parameters of the Penman-Monteith equation were clearly different for winter and summer conditions, indicating that phenological effects on surface resistance are important. We also compared the simulated λE of different ecosystem types under meteorological conditions at one site. Values of λE varied between 15% and 38% of the net radiation in the simulations with mean ecosystem parameters. In general, the simulations suggest that λE is higher from forested ecosystems than from grasslands, wetlands or tundra-type ecosystems. Forests showed usually a tighter stomatal control of λE as indicated by a pronounced sensitivity of surface resistance to atmospheric vapor pressure deficit. Nevertheless, the surface resistance of forests was lower than for open vegetation types including wetlands. Tundra and wetlands had higher surface resistances, which were less sensitive to vapor pressure deficits. The results indicate that the variation in surface resistance within and between different vegetation types might play a significant role in energy exchange between terrestrial ecosystems and atmosphere. These results suggest the need to take into account vegetation type and phenology in energy exchange modeling.