The net ecosystem exchange of carbon dioxide was measured by eddy covariance methods for 3 years in two old-growth forest sites near Santarém, Brazil. Carbon was lost in the wet season and gained in the dry season, which was opposite to the seasonal cycles of both tree growth and model predictions. The 3-year average carbon loss was 1.3 (confidence interval: 0.0 to 2.0) megagrams of carbon per hectare per year. Biometric observations confirmed the net loss but imply that it is a transient effect of recent disturbance superimposed on long-term balance. Given that episodic disturbances are characteristic of old-growth forests, it is likely that carbon sequestration is lower than has been inferred from recent eddy covariance studies at undisturbed sites.
Coupled climate-carbon cycle models suggest that Amazon forests are vulnerable to both long- and short-term droughts, but satellite observations showed a large-scale photosynthetic green-up in intact evergreen forests of the Amazon in response to a short, intense drought in 2005. These findings suggest that Amazon forests, although threatened by human-caused deforestation and fire and possibly by more severe long-term droughts, may be more resilient to climate changes than ecosystem models assume.
a b s t r a c tWe investigated the seasonal patterns of Amazonian forest photosynthetic activity, and the effects thereon of variations in climate and land-use, by integrating data from a network of ground-based eddy flux towers in Brazil established as part of the 'Large-Scale Biosphere Atmosphere Experiment in Amazonia' project. We found that degree of water limitation, as indicated by the seasonality of the ratio of sensible to latent heat flux (Bowen ratio) predicts seasonal patterns of photosynthesis. In equatorial Amazonian forests (5• N-5 • S), water limitation is absent, and photosynthetic fluxes (or gross ecosystem productivity, GEP) exhibit high or increasing levels of photosynthetic activity as the dry season progresses, likely a consequence of allocation to growth of new leaves. In contrast, forests along the southern flank of the Amazon, pastures converted from forest, and mixed forest-grass savanna, exhibit dry-season declines in GEP, consistent with increasing degrees of water limitation. Although previous work showed tropical ecosystem evapotranspiration (ET) is driven by incoming radiation, GEP observations reported here surprisingly show no or negative relationships with photosynthetically active radiation (PAR). Instead, GEP fluxes largely followed the phenology of canopy photosynthetic capacity (Pc), with only deviations from this primary pattern driven by variations in PAR. Estimates of leaf flush at three * Corresponding author. Tel.: +1 520 6261500; fax: +1 520 621 9190. 182-183 (2013) 128-144 129 non-water limited equatorial forest sites peak in the dry season, in correlation with high dry season light levels. The higher photosynthetic capacity that follows persists into the wet season, driving high GEP that is out of phase with sunlight, explaining the negative observed relationship with sunlight. Overall, these patterns suggest that at sites where water is not limiting, light interacts with adaptive mechanisms to determine photosynthetic capacity indirectly through leaf flush and litterfall seasonality. These mechanisms are poorly represented in ecosystem models, and represent an important challenge to efforts to predict tropical forest responses to climatic variations.
 The Amazon Basin is crucial to global circulatory and carbon patterns due to the large areal extent and large flux magnitude. Biogeophysical models have had difficulty reproducing the annual cycle of net ecosystem exchange (NEE) of carbon in some regions of the Amazon, generally simulating uptake during the wet season and efflux during seasonal drought. In reality, the opposite occurs. Observational and modeling studies have identified several mechanisms that explain the observed annual cycle, including: (1) deep soil columns that can store large water amount, (2) the ability of deep roots to access moisture at depth when near-surface soil dries during annual drought, (3) movement of water in the soil via hydraulic redistribution, allowing for more efficient uptake of water during the wet season, and moistening of near-surface soil during the annual drought, and (4) photosynthetic response to elevated light levels as cloudiness decreases during the dry season. We incorporate these mechanisms into the third version of the Simple Biosphere model (SiB3) both singly and collectively, and confront the results with observations. For the forest to maintain function through seasonal drought, there must be sufficient water storage in the soil to sustain transpiration through the dry season in addition to the ability of the roots to access the stored water. We find that individually, none of these mechanisms by themselves produces a simulation of the annual cycle of NEE that matches the observed. When these mechanisms are combined into the model, NEE follows the general trend of the observations, showing efflux during the wet season and uptake during seasonal drought.
Abstract. We used eddy covariance to measure the net exchange of CO2 between the atmosphere and an old-growth tropical forest in Pari, Brazil from 1 July 2000 to 1 July 2001. The mean air temperature and daily temperature range varied little year-round; the rainy season lasted from late December to around July. Daytime CO2 uptake under high irradiance averaged 16-19 mol.m-2.s-1. Light was the main controller of CO2 exchange, accounting for 48% of the half-hour-to-half-hour variance. The rate of canopy photosynthesis at a given irradiance was lower in the afternoon than the morning. This photosynthetic inhibition was probably caused by high evaporative demand, high temperature, an intrinsic circadian rhythm, or a combination of the three. Wood increment increased from January to May, suggesting greater rates of carbon sequestration during the wet season. However, the daily net CO2 exchange measured by eddy covariance revealed the opposite trend, with greater carbon accumulation during the dry season. A reduction in respiration during the dry season was an important cause of this seasonal pattern. The surface litter was desiccated in the dry season, and the seasonal pattern of respiration appears to be a direct result of reduced forest floor decomposition during drought. In contrast, canopy photosynthesis was not directly reduced by the dry season, probably because deep rooting allows the forest to avoid drought stress
We used the eddy covariance technique from July 2000 to July 2001 to measure the fluxes of sensible heat, water vapor, and CO2 between an old‐growth tropical forest in eastern Amazonia and the atmosphere. Precipitation varied seasonally, with a wet season from mid‐December 2000 to July 2001 characterized by successive rainy days, wet soil, and, relative to the dry season, cooler temperatures, greater cloudiness, and reduced incoming solar and net radiation. Average evapotranspiration decreased from 3.96 ± 0.65 mm/d during the dry season to 3.18 ± 0.76 mm/d during the wet season, in parallel with decreasing radiation and decreasing water vapor deficit. The average Bowen ratio was 0.17 ± 0.10, indicating that most of the incoming radiation was used for evaporation. The Bowen ratio was relatively low during the early wet season (December–March), as a result of increased evaporative fraction and reduced sensible heat flux. The seasonal decline in Bowen ratio and increase in evaporative fraction coincided with an increase in ecosystem CO2 assimilation capacity, which we attribute to the growth of new leaves. The evaporative fraction did not decline as the dry season progressed, implying that the forest did not become drought stressed. The roots extracted water throughout the top 250 cm of soil, and water redistribution, possibly by hydraulic lift, partially recharged the shallow soil during dry season nights. The lack of drought stress during the dry season was likely a consequence of deep rooting, and possibly vertical water movement, which allowed the trees to maintain access to soil water year round.
 We investigated the seasonal patterns of water vapor and sensible heat flux along a tropical biome gradient from forest to savanna. We analyzed data from a network of flux towers in Brazil that were operated within the Large-Scale Biosphere-Atmosphere Experiment in Amazonia (LBA). These tower sites included tropical humid and semideciduous forest, transitional forest, floodplain (with physiognomies of cerrado), and cerrado sensu stricto. The mean annual sensible heat flux at all sites ranged from 20 to 38 Wm À2, and was generally reduced in the wet season and increased in the late dry season, coincident with seasonal variations of net radiation and soil moisture. The sites were easily divisible into two functional groups based on the seasonality of evaporation: tropical forest and savanna. At sites with an annual precipitation above 1900 mm and a dry season length less than 4 months (Manaus, Santarem and Rondonia), evaporation rates increased in the dry season, coincident with increased radiation. Evaporation rates were as high as 4.0 mm d À1 in these evergreen or semidecidous forests. In contrast, ecosystems with precipitation less than 1700 mm and a longer dry season (Mato Grosso, Tocantins and São Paulo) showed clear evidence of reduced evaporation in the dry season. Evaporation rates were as low as 2.5 mm d À1 in the transitional forests and 1 mm d À1 in the cerrado. The controls on evapotranspiration seasonality changed along the biome gradient, with evaporative demand (especially net radiation) playing a more important role in the wetter forests, and soil moisture playing a more important role in the drier savannah sites.
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