Seasonality of temperate forest photosynthesis and daytime respiration.

Terrestrial ecosystems currently offset one-quarter of anthropogenic carbon dioxide (CO2) emissions because of a slight imbalance between global terrestrial photosynthesis and respiration. Understanding what controls these two biological fluxes is therefore crucial to predicting climate change. Yet there is no way of directly measuring the photosynthesis or daytime respiration of a whole ecosystem of interacting organisms; instead, these fluxes are generally inferred from measurements of net ecosystem-atmosphere CO2 exchange (NEE), in a way that is based on assumed ecosystem-scale responses to the environment. The consequent view of temperate deciduous forests (an important CO2 sink) is that, first, ecosystem respiration is greater during the day than at night; and second, ecosystem photosynthetic light-use efficiency peaks after leaf expansion in spring and then declines, presumably because of leaf ageing or water stress. This view has underlain the development of terrestrial biosphere models used in climate prediction and of remote sensing indices of global biosphere productivity. Here, we use new isotopic instrumentation to determine ecosystem photosynthesis and daytime respiration in a temperate deciduous forest over a three-year period. We find that ecosystem respiration is lower during the day than at night-the first robust evidence of the inhibition of leaf respiration by light at the ecosystem scale. Because they do not capture this effect, standard approaches overestimate ecosystem photosynthesis and daytime respiration in the first half of the growing season at our site, and inaccurately portray ecosystem photosynthetic light-use efficiency. These findings revise our understanding of forest-atmosphere carbon exchange, and provide a basis for investigating how leaf-level physiological dynamics manifest at the canopy scale in other ecosystems.

Tracking forest phenology and seasonal physiology using digital repeat photography: a critical assessment.

Digital repeat photography is becoming widely used for near-surface remote sensing of vegetation. Canopy greenness, which has been used extensively for phenological applications, can be readily quantified from camera images. Important questions remain, however, as to whether the observed changes in canopy greenness are directly related to changes in leaf-level traits, changes in canopy structure, or some combination thereof. We investigated relationships between canopy greenness and various metrics of canopy structure and function, using five years (2008­2012) of automated digital imagery, ground observations of phenological transitions, leaf area index (LAI) measurements, and eddy covariance estimates of gross ecosystem photosynthesis from the Harvard Forest, a temperate deciduous forest in the northeastern United States. Additionally, we sampled canopy sunlit leaves on a weekly basis throughout the growing season of 2011. We measured physiological and morphological traits including leaf size, mass (wet/dry), nitrogen content, chlorophyll fluorescence, and spectral reflectance and characterized individual leaf color with flatbed scanner imagery. Our results show that observed spring and autumn phenological transition dates are well captured by information extracted from digital repeat photography. However, spring development of both LAI and the measured physiological and morphological traits are shown to lag behind spring increases in canopy greenness, which rises very quickly to its maximum value before leaves are even half their final size. Based on the hypothesis that changes in canopy greenness represent the aggregate effect of changes in both leaf-level properties (specifically, leaf color) and changes in canopy structure (specifically, LAI), we developed a two end-member mixing model. With just a single free parameter, the model was able to reproduce the observed seasonal trajectory of canopy greenness. This analysis shows that canopy greenness is relatively insensitive to changes in LAI at high LAI levels, which we further demonstrate by assessing the impact of an ice storm on both LAI and canopy greenness. Our study provides new insights into the mechanisms driving seasonal changes in canopy greenness retrieved from digital camera imagery. The nonlinear relationship between canopy greenness and canopy LAI has important implications both for phenological research applications and for assessing responses of vegetation to disturbances.

Canopy nitrogen, carbon assimilation, and albedo in temperate and boreal forests: Functional relations and potential climate feedbacks.

The availability of nitrogen represents a key constraint on carbon cycling in terrestrial ecosystems, and it is largely in this capacity that the role of N in the Earth's climate system has been considered. Despite this, few studies have included continuous variation in plant N status as a driver of broad-scale carbon cycle analyses. This is partly because of uncertainties in how leaf-level physiological relationships scale to whole ecosystems and because methods for regional to continental detection of plant N concentrations have yet to be developed. Here, we show that ecosystem CO(2) uptake capacity in temperate and boreal forests scales directly with whole-canopy N concentrations, mirroring a leaf-level trend that has been observed for woody plants worldwide. We further show that both CO(2) uptake capacity and canopy N concentration are strongly and positively correlated with shortwave surface albedo. These results suggest that N plays an additional, and overlooked, role in the climate system via its influence on vegetation reflectivity and shortwave surface energy exchange. We also demonstrate that much of the spatial variation in canopy N can be detected by using broad-band satellite sensors, offering a means through which these findings can be applied toward improved application of coupled carbon cycle-climate models.

Identification of hydroxymethanesulfonate in fog water.

Previous studies have suggested that hydroxymethanesulfonate ion (HMSA) can be an important species in fog and cloud water. Formation of HMSA explains observed excesses of sulfur in the S(IV) state (+4 oxidation state) and formaldehyde (CH(2)O) in fogs and clouds. HMSA was determined in fog water by a novel ion-pairing chromatographic technique. Concentrations in samples collected in Bakersfield, California, within 5 kilometers of major sources of sulfur dioxide (SO(2)), were as high as 300 micromoles per liter. Total CH(2)O and S(IV) concentrations, which were measured independently, ranged from 10 to 200 and 5 to more than 300 micromoles per liter, respectively. Concentrations of CH(2)O, S(IV), and HMSA at Buttonwillow, California, which is 15 kilometers from the nearest source of SO(2), were less than those at Bakersfield but not absent. These data confirm that HMSA forms in atmospheric water droplets and can reach appreciable concentrations. HMSA represents an important source of acidity for water droplets and may also play a role in long-distance transport and transformation of SO(2).

Chemical composition of Acid fog.

Fog water collected at three sites in Los Angeles and Bakersfield, California, was found to have higher acidity and higher concentrations of sulfate, nitrate, and ammonium than previously observed in atmospheric water droplets. The pH of the fog water was in the range of 2.2 to 4.0. The dominant processes controlling the fog water chemistry appear to be the condensation and evaporation of water vapor on preexisting aerosol and the scavenging of gas-phase nitric acid.

Net Exchange of CO2 in a Mid-Latitude Forest.

The eddy correlation method was used to measure the net ecosystem exchange of carbon dioxide continuously from April 1990 to December 1991 in a deciduous forest in central Massachusetts. The annual net uptake was 3.7 +/- 0.7 metric tons of carbon per hectare per year. Ecosystem respiration, calculated from the relation between nighttime exchange and soil temperature, was 7.4 metric tons of carbon per hectare per year, implying gross ecosystem production of 11.1 metric tons of carbon per hectare per year. The observed rate of accumulation of carbon reflects recovery from agricultural development in the 1800s. Carbon uptake rates were notably larger than those assumed for temperate forests in global carbon studies. Carbon storage in temperate forests can play an important role in determining future concentrations of atmospheric carbon dioxide.

Factors controlling long- and short-term sequestration of atmospheric CO2 in a mid-latitude forest.

Net uptake of carbon dioxide (CO2) measured by eddy covariance in a 60- to 80-year-old forest averaged 2.0 +/- 0.4 megagrams of carbon per hectare per year during 1993 to 2000, with interannual variations exceeding 50%. Biometry indicated storage of 1.6 +/- 0.4 megagrams of carbon per hectare per year over 8 years, 60% in live biomass and the balance in coarse woody debris and soils, confirming eddy-covariance results. Weather and seasonal climate (e.g., variations in growing-season length or cloudiness) regulated seasonal and interannual fluctuations of carbon uptake. Legacies of prior disturbance and management, especially stand age and composition, controlled carbon uptake on the decadal time scale, implying that eastern forests could be managed for sequestration of carbon.

Atmospheric deposition, CO2, and change in the land carbon sink.

Concentrations of atmospheric carbon dioxide (CO2) have continued to increase whereas atmospheric deposition of sulphur and nitrogen has declined in Europe and the USA during recent decades. Using time series of flux observations from 23 forests distributed throughout Europe and the USA, and generalised mixed models, we found that forest-level net ecosystem production and gross primary production have increased by 1% annually from 1995 to 2011. Statistical models indicated that increasing atmospheric CO2 was the most important factor driving the increasing strength of carbon sinks in these forests. We also found that the reduction of sulphur deposition in Europe and the USA lead to higher recovery in ecosystem respiration than in gross primary production, thus limiting the increase of carbon sequestration. By contrast, trends in climate and nitrogen deposition did not significantly contribute to changing carbon fluxes during the studied period. Our findings support the hypothesis of a general CO2-fertilization effect on vegetation growth and suggest that, so far unknown, sulphur deposition plays a significant role in the carbon balance of forests in industrialized regions. Our results show the need to include the effects of changing atmospheric composition, beyond CO2, to assess future dynamics of carbon-climate feedbacks not currently considered in earth system/climate modelling.

An ecosystem-scale perspective of the net land methanol flux: synthesis of micrometeorological flux measurements.

Methanol is the second most abundant volatile organic compound in the troposphere and plays a significant role in atmospheric chemistry. While there is consensus about the dominant role of living plants as the major source and the reaction with OH as the major sink of methanol, global methanol budgets diverge considerably in terms of source/sink estimates reflecting uncertainties in the approaches used to model, and the empirical data used to separately constrain these terms. Here we compiled micrometeorological methanol flux data from eight different study sites and reviewed the corresponding literature in order to provide a first cross-site synthesis of the terrestrial ecosystem-scale methanol exchange and present an independent data-driven view of the land-atmosphere methanol exchange. Our study shows that the controls of plant growth on the production, and thus the methanol emission magnitude, and stomatal conductance on the hourly methanol emission variability, established at the leaf level, hold across sites at the ecosystem-level. Unequivocal evidence for bi-directional methanol exchange at the ecosystem scale is presented. Deposition, which at some sites even exceeds methanol emissions, represents an emerging feature of ecosystem-scale measurements and is likely related to environmental factors favouring the formation of surface wetness. Methanol may adsorb to or dissolve in this surface water and eventually be chemically or biologically removed from it. Management activities in agriculture and forestry are shown to increase local methanol emission by orders of magnitude; they are however neglected at present in global budgets. While contemporary net land methanol budgets are overall consistent with the grand mean of the micrometeorological methanol flux measurements, we caution that the present approach of simulating methanol emission and deposition separately is prone to opposing systematic errors and does not allow taking full advantage of the rich information content of micrometeorological flux measurements.

Environmental controls on the photosynthesis and respiration of a boreal lichen woodland: a growing season of whole-ecosystem exchange measurements by eddy correlation.

Measurements of net ecosystem CO2 exchange by eddy correlation, incident photosynthetically active photon flux density (PPFD), soil temperature, air temperature, and air humidity were made in a black spruce (Picea mariana) boreal woodland near Schefferville, Quebec, Canada, from June through August 1990. Nighttime respiration was between 0.5 and 1.5 kg C ha-1 h-1, increasing with temperature. Net uptake of carbon during the day peaked at 3 kg C ha-1 h-1, and the daily net uptake over the experiment was 12 kg C ha-1 day-1. Photosynthesis dropped substantially at leaf-to-air vapor pressure deficit (VPD) greater than 7 mb, presumably as a result of stomatal closure. The response of ecosystem photosynthesis to incident PPFD was markedly non-linear, with an abrupt saturation at 600 µmol m-2 s-1. This sharp saturation reflected the geometry of the spruce canopy (isolated conical crowns), the frequently overcast conditions, and an increase in VPD coincident with high radiation. The ecosystem light-use efficiency increased markedly during overcast periods as a result of a more even distribution of light across the forest surface. A mechanistic model of forest photosynthesis, parameterized with observations of leaf density and nitrogen content from a nearby stand, provided accurate predictions of forest photosynthesis. The observations and model results indicated that ecosystem carbon balance at the site is highly sensitive to temperature, and relatively insensitive to cloudiness.