The micrometeorological flux measurement technique known as relaxed eddy accumulation (REA) holds promise as a powerful new tool for ecologists. The more popular eddy covariance (eddy correlation) tech- nique requires the use of sensors that can respond at fast rates (10 Hz), and these are unavailable for many eco- logically relevant compounds. In contrast, the use of REA allows flux measurement with sensors that have much slower response time, such as gas chromatography and mass spectrometry. In this review, relevant micro- meteorological details underlying REA are presented, and critical analytical and system design details are discussed, with the goal of introducing the technique and its potential applications to ecologists. The validity of REA for measuring fluxes of isoprene, a photochemi- cally reactive hydrocarbon emitted by several plant species, was tested with measurements over an oak- hickory forest in the Walker Branch Watershed in eastern Tennessee. Concurrent eddy covariance mea- surements of isoprene flux were made using a newly available chemiluminesence instrument. Excellent agreement was obtained between the two techniques (r 2 a 0.974, n a 62), providing the first direct com- parison between REA and eddy covariance for mea- suring the flux rate of a reactive compound. The influ- ence of a bias in vertical wind velocity on the accuracy of REA was examined. This bias has been thought to be a source of significant error in the past. Measurements of normalized bias (w=rw) alone would lead us to think that a large potential error exists at this site. However, with our isoprene data and through simulations of REA with fast-response H2O and CO2 data, we conclude that accurate REA flux measurements can be made even in the presence of a bias in w.
Summary We conducted ecosystem carbon and water vapour exchange studies in an old‐growth Pinus ponderosa forest in the Pacific North‐west region of the United States. The canopy is heterogeneous, with tall multiaged trees and an open, clumped canopy with low leaf area. Carbon assimilation can occur throughout relatively mild winters, although night frosts can temporarily halt the process and physiological factors limit its efficiency. In contrast, carbon assimilation is often limited in the ‘growing season’ by stomatal closure associated with high evaporative demand ( D ) and soil water deficits. All of these factors present a challenge to effectively modelling ecosystem processes. Our objective was to generate an understanding of the controls on ecosystem processes across seasonal and annual cycles from a combination of fine‐scale process modelling, ecophysiological measurements, and carbon and water vapour fluxes measured by the eddy covariance method. Flux measurements showed that 50% and 70% of the annual carbon uptake occurred outside the ‘growing season’ (defined as bud break to senescence, ∼ days 125–275) in 1996 and 1997. On a daily basis in summer, net ecosystem productivity (NEP) was low when D and soil water deficits were large. Whole ecosystem water vapour fluxes (LE) increased from spring to summer (1.0–1.9 mm d−1) as conducting leaf area increased by 30% and as evaporative demand increased, while evaporation from the soil surface became a smaller portion of total LE as soil water deficits increased. The models underestimated soil evaporation, particularly following rain. In the SPA model, varying the temperature optimum for photosynthesis seasonally resulted in overestimation of carbon uptake in winter and spring, showing that in coniferous forests, assumptions about temperature optima are clearly important. Daily estimates of soil surface CO 2 flux from measurements and site meteorological data demonstrated that modelling of soil CO 2 flux based on an Arrhenius‐type equation in CANPOND overestimated CO 2 respired from the soil during drought and when temperatures were low.
Forestry – including afforestation (the planting of trees on land where they have not recently existed), reforestation, avoided deforestation, and forest management – can lead to increased sequestration of atmospheric carbon dioxide and has therefore been proposed as a strategy to mitigate climate change. However, forestry also influences land‐surface properties, including albedo (the fraction of incident sunlight reflected back to space), surface roughness, and evapotranspiration, all of which affect the amount and forms of energy transfer to the atmosphere. In some circumstances, these biophysical feedbacks can result in local climate warming, thereby counteracting the effects of carbon sequestration on global mean temperature and reducing or eliminating the net value of climate‐change mitigation projects. Here, we review published and emerging research that suggests ways in which forestry projects can counteract the consequences associated with biophysical interactions, and highlight knowledge gaps in managing forests for climate protection. We also outline several ways in which biophysical effects can be incorporated into frameworks that use the maintenance of forests as a climate protection strategy.
Stable isotopes of atmospheric carbon dioxide (CO 2 ) contain a wealth of information regarding biosphere‐atmosphere interactions. The carbon isotope ratio of CO 2 (δ 13 C) reflects the terrestrial carbon cycle including processes of photosynthesis, respiration, and decomposition. The oxygen isotope ratio (δ 18 O) reflects terrestrial carbon and water coupling due to CO 2 ‐H 2 O oxygen exchange. Isotopic CO 2 measurements, in combination with ecosystem‐isotopic exchange models, allow for the quantification of patterns and mechanisms regulating terrestrial carbon and water cycles, as well as for hypothesis development, data interpretation, and forecasting. Isotopic measurements and models have evolved significantly over the past two decades, resulting in organizations that promote model‐measurement networks, e.g., the U.S. National Science Foundation's Biosphere‐Atmosphere Stable Isotope Network, the European Stable Isotopes in Biosphere‐Atmosphere Exchange Network, and the U.S. National Environmental Observatory Network.
The plant in a field of energy, or in any considered area in which gradients exist, reacts passively to its surroundings . . . it should be clear that the sun, or even outer space, is not more removed from the plant than the next millimeter of air; it is the specific part of heat exchange being considered that determines what should be included in the plant’s environment.Klaus Raschke (1960)
The study of water exchange between soil, plants, and the atmosphere in response to seasonal or periodic droughts is critical to modeling the hydrologic cycle and biogeochemical processes in water‐controlled ecosystems. An essential step in such studies is to characterize changes in evaporation and transpiration under water stress. The objectives of this study are to investigate how soil moisture controls the evapotranspiration in a Californian oak savanna that experiences seasonal droughts, using multiyear field observations at the daily and stand scale, and to model these controls stochastically. The influence of soil moisture on evapotranspiration at the stand scale is studied using correlations between tower‐based evapotranspiration measurements and representative soil moisture obtained by aggregating point measurements. The observed pattern of this effect is found in agreement with an existing model that features a linear reduction of the evapotranspiration when soil moisture falls below a critical value. The model parameters are inferred using a Bayesian framework, and they are found to vary from year to year because of climate variability. The comparison between various aggregations of soil moisture at the stand scale from point measurements demonstrates that the spatial variability of the soil moisture and the water uptake capacity limited by the root biomass need be taken into account to produce a model that is most resistant to interannual variability. Finally, the parameterized model is used to predict the actual evapotranspiration with uncertainty estimates determined using the joint distribution of the parameters derived from the Bayesian framework. The satisfactory agreement between the predicted and measured evapotranspiration suggests that the calibrated model can be incorporated into water balance studies in the future.
We used gas exchange techniques to estimate maximum rate of carboxylation (V(cmax)), a measure of photosynthetic capacity, in the understory and upper crown of a closed deciduous forest over two seasons. There was extensive variability in photosynthetic capacity as a result of vertical canopy position, species type, leaf age and drought. Photosynthetic capacity was greater in oaks than in maples and greater in the overstory than in the understory. Parameter V(cmax) was maximal early in the season but declined slowly throughout most of the summer, and then more rapidly during senescence. There was also an apparent decline during drought in some trees. Variability in V(cmax) as a result of species or vertical canopy gradients was described well by changes in leaf nitrogen per unit area (N(a)). However, temporal changes in V(cmax) were often poorly correlated with leaf nitrogen, especially in spring and summer and during drought. This poor correlation may be the result of a seasonally dependent fractional allocation of leaf nitrogen to Rubisco; however, we could not discount Rubisco inactivation, patchy stomatal closure or changes in mesophyll resistance. Consequently, when a single annual regression equation of V(cmax) versus N(a) was used for this site, there were substantial errors in the temporal patterns in V(cmax) that will inevitably result in modeling errors.
During the growing season, nighttime ecosystem respiration emits 30–100% of the daytime net photosynthetic uptake of carbon, and therefore measurements of rates and understanding of its control by the environment are important for understanding net ecosystem exchange. Ecosystem respiration can be measured at night by eddy covariance methods, but the data may not be reliable because of low turbulence or other methodological problems. We used relationships between woody tissue, foliage, and soil respiration rates and temperature, with temperature records collected on site to estimate ecosystem respiration rates at six coniferous BOREAS sites at half‐hour or 1‐hour intervals, and then compared these estimates to nocturnal measurements of CO 2 exchange by eddy covariance. Soil surface respiration was the largest source of CO 2 at all sites (48–71%), and foliar respiration made a large contribution to ecosystem respiration at all sites (25–43%). Woody tissue respiration contributed only 5–15% to ecosystem respiration. We estimated error for the scaled chamber predictions of ecosystem respiration by using the uncertainty associated with each respiration parameter and respiring biomass value. There was substantial uncertainty in estimates of foliar and soil respiration because of the spatial variability of specific respiration rates. In addition, more attention needs to be paid to estimating foliar respiration during the early part of the growing season, when new foliage is growing, and to determining seasonal trends of soil surface respiration. Nocturnal eddy covariance measurements were poorly correlated to scaled chamber estimates of ecosystem respiration ( r 2 =0.06–0.27) and were consistently lower than scaled chamber predictions (by 27% on average for the six sites). The bias in eddy covariance estimates of ecosystem respiration will alter estimates of gross assimilation in the light and of net ecosystem exchange rates over extended periods.
Read moreAbstract Agricultural drainage of organic soils has resulted in vast soil subsidence and contributed to increased atmospheric carbon dioxide (CO 2 ) concentrations. The Sacramento‐San Joaquin Delta in California was drained over a century ago for agriculture and human settlement and has since experienced subsidence rates that are among the highest in the world. It is recognized that drained agriculture in the Delta is unsustainable in the long‐term, and to help reverse subsidence and capture carbon (C) there is an interest in restoring drained agricultural land‐use types to flooded conditions. However, flooding may increase methane (CH 4 ) emissions. We conducted a full year of simultaneous eddy covariance measurements at two conventional drained agricultural peatlands (a pasture and a corn field) and three flooded land‐use types (a rice paddy and two restored wetlands) to assess the impact of drained to flooded land‐use change on CO 2 and CH 4 fluxes in the Delta. We found that the drained sites were net C and greenhouse gas (GHG) sources, releasing up to 341 g C m −2 yr −1 as CO 2 and 11.4 g C m −2 yr −1 as CH 4 . Conversely, the restored wetlands were net sinks of atmospheric CO 2 , sequestering up to 397 g C m −2 yr −1 . However, they were large sources of CH 4 , with emissions ranging from 39 to 53 g C m −2 yr −1 . In terms of the full GHG budget, the restored wetlands could be either GHG sources or sinks. Although the rice paddy was a small atmospheric CO 2 sink, when considering harvest and CH 4 emissions, it acted as both a C and GHG source. Annual photosynthesis was similar between sites, but flooding at the restored sites inhibited ecosystem respiration, making them net CO 2 sinks. This study suggests that converting drained agricultural peat soils to flooded land‐use types can help reduce or reverse soil subsidence and reduce GHG emissions.
Read moreForest biomes are major reserves for terrestrial carbon, and major components of global primary productivity. The carbon balance of forests is determined by a number of component processes of carbon acquisition and carbon loss, and a small shift in the magnitude of these processes would have a large impact on the global carbon cycle. In this paper, we discuss the climatic influences on the carbon dynamics of boreal, temperate and tropical forests by presenting a new synthesis of micrometeorological, ecophysiological and forestry data, concentrating on three case‐study sites. Historical changes in the carbon balance of each biome are also reviewed, and the evidence for a carbon sink in each forest biome and its likely behaviour under future global change are discussed. We conclude that there have been significant advances in determining the carbon balance of forests, but there are still critical uncertainties remaining, particularly in the behaviour of soil carbon stocks.
Read moreUnderstanding terrestrial carbon metabolism is critical because terrestrial ecosystems play a major role in the global carbon cycle. Furthermore, humans have severely disrupted the carbon cycle in ways that will alter the climate system and directly affect terrestrial metabolism. Changes in terrestrial metabolism may well be as important an indicator of global change as the changing temperature signal. Improving our understanding of the carbon cycle at various spatial and temporal scales will require the integration of multiple, complementary and independent methods that are used by different research communities. Tools such as air sampling networks, inverse numerical methods, and satellite data (top-down approaches) allow us to study the strength and location of the global- and continental-scale carbon sources and sinks. Bottom-up studies provide estimates of carbon fluxes at finer spatial scales and examine the mechanisms that control fluxes at the ecosystem, landscape, and regional scales. Bottom-up approaches include comparative and process studies (for example, ecosystem manipulative experiments) that provide the necessary mechanistic information to develop and validate terrestrial biospheric models. An iteration and reiteration of top-down and bottom-up approaches will be necessary to help constrain measurements at various scales. We propose a major international effort to coordinate and lead research programs of global scope of the carbon cycle.
Read moreThe California Mediterranean savanna has harsh summer conditions with minimal soil moisture, high temperature, high incoming solar radiation and little or no precipitation. Deciduous blue oaks, Quercus douglasii Hook. and Arn., are winter-deciduous obligate phreatophytes, transpiring mostly groundwater throughout the summer drought. The objective of this work is to fully characterize the seasonal trends of photosynthesis in blue oaks as well as the mechanistic relationships between leaf structure and function. We estimate radiative load of the leaves via the FLiES model and perform in situ measurements of leaf water potential, leaf nitrogen content, an index of chlorophyll content (SPAD readings), photosynthetic and electron transport capacity, and instantaneous rates of CO2 assimilation and electron transport. We measured multiple trees over 3 years providing data from a range of conditions. Our study included one individual that demonstrated strong drought stress as indicated by changes in SPAD readings, leaf nitrogen and all measures of leaf functioning. In the year following severe environmental stress, one individual altered foliation patterns on the crown but did not die. In all other individuals, we found that net carbon assimilation and photosynthetic capacity decreased during the summer drought. SPAD values, electron transport rate (ETR) and quantum yield of photosystem II (PSII) did not show a strong decrease during the summer drought. In most individuals, PSII activity and SPAD readings did not indicate leaf structural or functional damage throughout the season. While net carbon assimilation was tightly coupled to stomatal conductance, the coupling was not as tight with ETR possibly due to contributions from photorespiration or other protective processes. Our work demonstrates that the blue oaks avoid structural damage by maintaining the capacity to convert and dissipate incoming solar radiation during the hot summer drought and are effective at fixing carbon by maximizing rates during the mild spring conditions.
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