Precipitation Pattern Determines the Inter-annual Variation of Herbaceous Layer and Carbon Fluxes in a Phreatophyte-Dominated Desert Ecosystem

Arid and semi-arid ecosystems dominated by shrubby species are an important component in the global carbon cycle but are largely under-represented in studies of the effect of climate change on carbon flux. This study synthesizes data from long-term eddy covariance measurements and experiments to assess how changes in ecosystem composition, driven by precipitation patterns, affect inter-annual variability of carbon flux and their components in a halophyte desert community dominated by deep-rooted shrubs (phreatophytes, which depend on groundwater as their primary water source). Our results demonstrated that the carbon balance of this community responded strongly to precipitation variations. Both pre-growing season precipitation and growing season precipitation frequency significantly affected inter-annual variations in ecosystem carbon flux. Heavy pre-growing season precipitation (November–April, mostly as snow) increased annual net ecosystem carbon exchange, by facilitating the growth and carbon assimilation of shallow-rooted annual plants, which used spring and summer precipitation to increase community productivity. Sufficient pre-growing season precipitation led to more germination and growth of shallow-rooted annual plants. When followed by high-frequency growing season precipitation, community productivity of this desert ecosystem was lifted to the level of grassland or forest ecosystems. The long-term observations and experimental results confirmed that precipitation patterns and the herbaceous component were dominant drivers of the carbon dynamics in this phreatophyte-dominated desert ecosystem. This study illustrates the importance of inter-annual variations in climate and ecosystem composition for the carbon flux in arid and semi-arid ecosystems. It also highlights the important effect of changing frequency and seasonal pattern of precipitation on the regional and global carbon cycle in the coming decades.     

Assessing canopy performance using carbonyl sulfide measurements

Carbonyl sulfide (COS) is a tracer of ecosystem photosynthesis that can advance carbon cycle research from leaf to global scales; however, a range of newly reported caveats related to sink/source strength of various ecosystem components hinder its application. Using comprehensive eddy‐covariance and chamber measurements, we systematically measure ecosystem contributions from leaf, stem, soil, and litter and were able to close the ecosystem COS budget. The relative contributions of nonphotosynthetic components to the overall canopy‐scale flux are relatively small (~4% during peak activity season) and can be independently estimated based on their responses to temperature and humidity. Converting COS to photosynthetic CO₂ fluxes based on the leaf relative uptake of COS/CO₂, faces challenges due to observed daily and seasonal changes. Yet, this ratio converges around a constant value (~1.6), and the variations, dominated by light intensity, were found unimportant on a flux‐weighted daily time‐scale, indicating a mean ratio of daytime gross‐to‐net primary productivity of ~2 in our ecosystem. The seasonal changes in the leaf relative uptake ratio may indicate a reduction in mesophyll conductance in winter, and COS‐derived canopy conductance permitted canopy temperature estimate consistent with radiative skin temperature. These results support the feasibility of using COS as a powerful and much‐needed means of assessing ecosystem function and its response to change.     

Seasonal variations of CO2 and water vapour exchange rates over a temperate deciduous forest

Long-term and direct measurements of CO₂ and water vapour exchange are needed over forested ecosystems to determine their net annual fluxes of carbon dioxide and water. Such measurements are also needed to parameterize and test biogeochemical, ecological and hydrological assessment models. Responding to this need, eddy covariance measurements of CO₂ and water vapour were made ever a deciduous forest growing near Oak Ridge, TN, between April 1993 and April 1994. Periodic measurements were made of leaf area index, stomatal resistance, soil moisture and pre-dawn leaf water potential to characterize the gas exchange capacity of the canopy. Four factors had a disproportionate influence on the seasonal variation of CO₂ flux densities. These factors were photon flux densities (during the growing season), temperature (during the dormant season), leaf area index and the occurrence of drought The drought period occurred during the peak of the growing season and caused a significant decline in daily and hourly CO₂ flux densities, relative to observations over the stand when soil moisture was plentiful. The annual net uptake of carbon was calculated by integrating flux measurements and filling missing and spurious data with the relations obtained between measured CO₂ fluxes and environmental forcing variables. The net flux of carbon for the period between April 1993 and April 1994 was -525 g C m^?2 y^?1. This value represents a net flux of carbon from the atmosphere and into the forest. The net annual carbon exchange of this southern temperate broadleaved forest exceeded values measured over a northern temperate forest (which experiences a shorter growing season and has less leaf area) by 200 g C m^?2 y^?1 (cf. Wofsy et al 1993). The seasonal variation of canopy evaporation (latent heat flux) was controlled mostly by changes in leaf area and net radiation. A strong depression in evaporation rates was not observed during the drought Over a broadleaved forest large vapour pressure deficits promote evaporation and trees in a mixed stand are able to tap a variety of deep and shallow water sources.     

基于EALCO模型对中亚热带人工针叶林CO2通量季节变异的模拟

 EALCO模型是一个基于生理生态学过程,模拟生态系统下垫面与大气之间水、热和碳通量交换的综合模型。将该模型应用在亚热带常绿针叶林, 对其生态系统过程进行了模拟,以深入探讨季节性干旱对生态系统过程的影响。对EALCO模型进行了参数化与初始化并对模型的光合作用时段和 落叶机制进行了改进,以更好地模拟亚热带人工针叶林生态系统。千烟洲通量观测站自2002年底开始应用涡度相关技术对中亚热带人工针叶林 生态系统进行通量观测,该站点2003年经历了一次较严重的季节性干旱(由高温与少雨综合作用造成),降水量仅为多年平均值的65%,而2004年 的年降水量与多年平均值较为接近,利用该站点2003和2004年特殊的气候条件,使用其通量观测数据对模型的模拟效果进行检验。从模拟结果 的总体趋势来看,模型能较好地从半小时、日及年尺度上反映两年内土壤-植被-大气之间的碳交换状况。总初级生产力(Gross primary production, GPP)在一年中呈现单峰型变化,遇高温及干旱胁迫GPP值下降。由于受到干旱胁迫的影响,2003年GPP值比2004年偏低12.9%。模拟 结果显示,2003年GPP值比2004年偏低11.2%。观测数据与模拟结果均显示,水分胁迫期间净碳交换量(Net ecosystem production, NEP)模拟值 与实测值的日变化均呈现一种“偏态",即一天中生态系统碳交换量最大值出现在上午某一时刻,之后逐渐降低。 模拟结果显示,水分匮缺对 光合能力的影响比对生态系统呼吸作用的影响更为强烈,因而导致了净生态系统生产力的降低。进一步分析表明,水分匮缺期间,晴天正午之 前,深层土壤( >20 cm) 水分的匮缺抑制了光合作用能力,正午之后,高温与深层土壤水分匮缺共同削弱光合作用能力,影响各占一半。     

Modelling Seasonal and Inter-annual Variations in Carbon and Water Fluxes in an Arid-Zone <Emphasis Type="Italic">Acacia</Emphasis> Savanna Woodland, 1981–2012

Changes in climatic characteristics such as seasonal and inter-annual variability may affect ecosystem structure and function, hence alter carbon and water budgets of ecosystems. Studies of modelling combined with field experiments can provide essential information to investigate interactions between carbon and water cycles and climate. Here we present a first attempt to investigate the long-term climate controls on seasonal patterns and inter-annual variations in water and carbon exchanges in an arid-zone savanna-woodland ecosystem using a detailed mechanistic soil–plant–atmosphere model (SPA), driven by leaf area index (LAI) simulated by an ecohydrological model (WAVES) and observed climate data during 1981–2012. The SPA was tested against almost 3 years of eddy covariance flux measurements in terms of gross primary productivity (GPP) and evapotranspiration (ET). The model was able to explain 80 and 71% of the variability of observed daily GPP and ET, respectively. Long-term simulations showed that carbon accumulation rates and ET ranged from 20.6 g C m^?2 mon^?1 in the late dry season to 45.8 g C m^?2 mon^?1 in the late wet season, respectively, primarily driven by seasonal variations in LAI and soil moisture. Large climate variations resulted in large seasonal variation in ecosystem water-use efficiency (eWUE). Simulated annual GPP varied between 146.4 and 604.7 g C m^?2 y^?1. Variations in annual ET coincided with that of GPP, ranging from 110.2 to 625.8 mm y^?1. Annual variations in GPP and ET were driven by the annual variations in precipitation and vapour pressure deficit (VPD) but not temperature. The linear coupling of simulated annual GPP and ET resulted in eWUE having relatively small year-to-year variation.     

Linking drought legacy effects across scales: From leaves to tree rings to ecosystems

Severe drought can cause lagged effects on tree physiology that negatively impact forest functioning for years. These “drought legacy effects” have been widely documented in tree‐ring records and could have important implications for our understanding of broader scale forest carbon cycling. However, legacy effects in tree‐ring increments may be decoupled from ecosystem fluxes due to (a) postdrought alterations in carbon allocation patterns; (b) temporal asynchrony between radial growth and carbon uptake; and (c) dendrochronological sampling biases. In order to link legacy effects from tree rings to whole forests, we leveraged a rich dataset from a Midwestern US forest that was severely impacted by a drought in 2012. At this site, we compiled tree‐ring records, leaf‐level gas exchange, eddy flux measurements, dendrometer band data, and satellite remote sensing estimates of greenness and leaf area before, during, and after the 2012 drought. After accounting for the relative abundance of tree species in the stand, we estimate that legacy effects led to ~10% reductions in tree‐ring width increments in the year following the severe drought. Despite this stand‐scale reduction in radial growth, we found that leaf‐level photosynthesis, gross primary productivity (GPP), and vegetation greenness were not suppressed in the year following the 2012 drought. Neither temporal asynchrony between radial growth and carbon uptake nor sampling biases could explain our observations of legacy effects in tree rings but not in GPP. Instead, elevated leaf‐level photosynthesis co‐occurred with reduced leaf area in early 2013, indicating that resources may have been allocated away from radial growth in conjunction with postdrought upregulation of photosynthesis and repair of canopy damage. Collectively, our results indicate that tree‐ring legacy effects were not observed in other canopy processes, and that postdrought canopy allocation could be an important mechanism that decouples tree‐ring signals from GPP.     

A test of the optimality approach to modelling canopy properties and CO2 uptake by natural vegetation

Photosynthesis provides plants with their main building material, carbohydrates, and with the energy necessary to thrive and prosper in their environment. We expect, therefore, that natural vegetation would evolve optimally to maximize its net carbon profit (NCP), the difference between carbon acquired by photosynthesis and carbon spent on maintenance of the organs involved in its uptake. We modelled N(CP) for an optimal vegetation for a site in the wet-dry tropics of north Australia based on this hypothesis and on an ecophysiological gas exchange and photosynthesis model, and compared the modelled CO2 fluxes and canopy properties with observations from the site. The comparison gives insights into theoretical and real controls on gas exchange and canopy structure, and supports the optimality approach for the modelling of gas exchange of natural vegetation. The main advantage of the optimality approach we adopt is that no assumptions about the particular vegetation of a site are required, making it a very powerful tool for predicting vegetation response to long-term climate or land use change.     

Seasonal carbon dynamics and water fluxes in an Amazon rainforest

Satellite‐based observations indicate that seasonal patterns in canopy greenness and productivity in the Amazon are negatively correlated with precipitation, with increased greenness occurring during the dry months. Flux tower measurements indicate that the canopy greening that occurs during the dry season is associated with increases in net ecosystem productivity (NEP) and evapotranspiration (ET). Land surface and terrestrial biosphere model simulations for the region have predicted the opposite of these observed patterns, with significant declines in greenness, NEP, and ET during the dry season. In this study, we address this issue mainly by developing an empirically constrained, light‐controlled phenology submodel within the Ecosystem Demography model version 2 (ED2). The constrained ED2 model with a suite of field observations shows markedly improved predictions of seasonal ecosystem dynamics, more accurately capturing the observed patterns of seasonality in water, carbon, and litter fluxes seen at the Tapajos National Forest, Brazil (2.86°S, 54.96°W). Long‐term simulations indicate that this light‐controlled phenology increases the resilience of Amazon forest NEP to interannual variability in climate forcing.     

Estimating canopy photosynthetic parameters in maize field based on multi-spectral remote sensing

Aims Determination of canopy photosynthetic parameters is key to accurate simulation of ecosystem function by using remote sensing methods. Currently, remote estimation of vegetation canopy structure characteristics has been widely adopted. However, directly estimating photosynthetic variables (photosynthetic capacity and efficiency) at canopy scale based on field spectrometry combined with CO₂ flux measurements is rare.
Methods In this study, we remotely estimated solar radiation use efficiency (εN, net ecosystem CO₂ exchange/absorbed photosynthetically active radiation (NEE_CO2/APAR); εG, gross primary productivity/absorbed photosynthetically active radiation (GPP/APAR); α, apparent quantum efficiency) and photosynthetic capacity (P_max) based on in situ measurements of spectral reflectance and ecosystem CO₂ fluxes, along with observational data on micrometeorological factors during the entire growing season for a maize canopy in Northeast China.
Important findings Results showed that the seasonal variations in P_max and α exhibited a single peak; whereas the values of εN and εG were higher at the start of vegetative stage and then rapidly decreased with the development of maize until displaying a single peak at the intermediate and late stages of the growing season, coinciding with the occurrence of peak values in P_max. A comparison was made on the predictive performance based on normalized difference vegetation index (NDVI), ratio vegetation index (RVI), wide dynamic range vegetation index (WDRVI), 2-band enhanced vegetation index (EVI2), and chlorophyll index (CI) in estimating four canopy photosynthetic parameters with any combination of two separate wavelengths at the range of 400–1 300 nm, which showed that EVI2 was most closely and linearly related to photosynthetic capacity and efficiency. This study demonstrates that multi-spectral remote sensing information is sensitive to the variations in canopy photosynthetic parameters in maize field and can be used to quantitatively monitor seasonal dynamics of canopy photosynthesis, and to accurately assess crop productivity and ecosystem CO₂ exchange capacity.     

Physiology-phenology interactions in a productive semi-arid pine forest

This study explored possible advantages conferred by the phase shift between leaf phenology and photosynthesis seasonality in a semi-arid Pinus halepensis forest system, not seen in temperate sites. Leaf-scale measurements of gas exchange, nitrogen and phenology were used on daily, seasonal and annual time-scales. Peak photosynthesis was in late winter, when high soil moisture, mild temperatures and low leaf vapour pressure deficit (D(L)) allowed high rates associated with high water- and nitrogen-use efficiencies. Self-sustained new needle growth through the dry and hot summer maximized photosynthesis in the following wet season, without straining carbon storage. Low rates of water loss were associated with increasing sensitivity of stomatal conductance (g(s)) to soil moisture below a relative extractable water (REW) of 0.4, and decreased g(s )sensitivity to D(L) below REW of approx. 0.2. This response was captured by the modified Ball-Berry (Leuning) model. While most physiological parameters and responses measured were typical of temperate pines, the photosynthesis-phenological phasing contributed to high productivity under warm-dry conditions. This contrasts with reported effects of short-term periodical droughts and could lead to different predictions of the effect of warming and drying climate on pine forest productivity.     

Do dynamic global vegetation models capture the seasonality of carbon fluxes in the Amazon basin? A data‐model intercomparison

To predict forest response to long‐term climate change with high confidence requires that dynamic global vegetation models (DGVMs) be successfully tested against ecosystem response to short‐term variations in environmental drivers, including regular seasonal patterns. Here, we used an integrated dataset from four forests in the Brasil flux network, spanning a range of dry‐season intensities and lengths, to determine how well four state‐of‐the‐art models (IBIS, ED2, JULES, and CLM3.5) simulated the seasonality of carbon exchanges in Amazonian tropical forests. We found that most DGVMs poorly represented the annual cycle of gross primary productivity (GPP), of photosynthetic capacity (Pc), and of other fluxes and pools. Models simulated consistent dry‐season declines in GPP in the equatorial Amazon (Manaus K34, Santarem K67, and Caxiuanã CAX); a contrast to observed GPP increases. Model simulated dry‐season GPP reductions were driven by an external environmental factor, ‘soil water stress’ and consequently by a constant or decreasing photosynthetic infrastructure (Pc), while observed dry‐season GPP resulted from a combination of internal biological (leaf‐flush and abscission and increased Pc) and environmental (incoming radiation) causes. Moreover, we found models generally overestimated observed seasonal net ecosystem exchange (NEE) and respiration (R_e) at equatorial locations. In contrast, a southern Amazon forest (Jarú RJA) exhibited dry‐season declines in GPP and R_e consistent with most DGVMs simulations. While water limitation was represented in models and the primary driver of seasonal photosynthesis in southern Amazonia, changes in internal biophysical processes, light‐harvesting adaptations (e.g., variations in leaf area index (LAI) and increasing leaf‐level assimilation rate related to leaf demography), and allocation lags between leaf and wood, dominated equatorial Amazon carbon flux dynamics and were deficient or absent from current model formulations. Correctly simulating flux seasonality at tropical forests requires a greater understanding and the incorporation of internal biophysical mechanisms in future model developments.     

Seasonal Variation in Respiratory and Photosynthetic Parameters in Three Mosses from the Maritime Antarctic

Carbon fixation under controlled conditions was measured inthree mosses from the maritime Antarctic using an infra-redgas analysis system. Gas exchange parameters were determinedduring each season in 1993 and 1994 using the Arrhenius equationand a hyperbolic tangent function applied to respiration andphotosynthesis, respectively. Environmental data was collectedin 1994 for comparison. All seasonal variations were greaterinBrachythecium than in the species from less hydric habitats.Respiration rates were highest in summer and lowest in winterat all temperatures inBrachythecium, but there was little changeinChorisodontium orAndreaea . There was some seasonal variationin the initial slope (K_p) of the photosynthesis-irradiance curvein all species, although the environmental data suggested thatthis was of little ecological importance. In all species seasonalchanges in the maximum rates of photosynthesis (GP_max, NP_max)were observed, generally with a pattern of summer maxima, althoughthere were some interannual differences. These changes are consideredto be the most important in affecting the overall annual productivityof the mosses. There were no seasonal variations in the optimumtemperatures for either gross or net photosynthesis, or forthe irradiance at the onset of light saturation (I_k). The resultshave important implications for the use of models to estimatethe productivity of the Antarctic flora based upon present orpredicted climate data. Antarctica; bryophytes; mosses; carbon exchange; irradiance; photosynthesis; productivity models; respiration; seasonal variation; temperature     

Cool-dry season depression in gas exchange of canopy leaves and water flux of tropical trees at the northern limit of Asian tropics

Trees on the northern boundary of Asian tropics experience hot-humid and cool-dry seasons, but little is known about their seasonal dynamics in canopy physiology. We used a canopy crane to reach the canopy of nine tropical tree species and measured canopy leaf gas exchange, water status, and trunk sap flux during the hot-humid and cool-dry seasons in Xishuangbanna, China. We found that most tree species exhibited significant reductions in maximum photosynthetic rate (A_max), stomatal conductance (g_smax), predawn and midday leaf water potentials, and maximum sap flux density in the cool-dry season. Compared to the hot-humid season, A_max declined by 19–60%, and maximum water flux declined by ?14% (an increase) to 42%. The cool-dry season decline in A_max of four species can be partly explained by an increased stomatal limitation (decreased g_smax and intercellular CO₂ concentrations). Therefore, a predicted increase in drought in this region may decrease the carbon sequestration and productivity of these forests. We did not find a tradeoff between performance (A_max in the hot-humid season) and persistence through the cool-dry season; species with higher A_max in the hot-humid season did not show higher percent seasonal declines in the cool-dry season. A_max was significantly and positively associated with the trunk sap flux for both seasons, but the association was weaker in the cool-dry season. Thus, our results suggest that some tradeoffs and trait associations are environment dependent. Our results are important for understanding carbon and water fluxes of seasonal tropical forests and their responses to environmental changes.

     

Carbon assimilation in crops at high temperatures

Global temperatures are rising, and higher rates of temperature increase are projected over land areas that encompass the globe's major agricultural regions. In addition to increased growing season temperatures, heat waves are predicted to become more common and severe. High temperatures can inhibit photosynthetic carbon gain of crop plants and thus threaten productivity, the effects of which may interact with other aspects of climate change. Here, we review the current literature assessing temperature effects on photosynthesis in key crops with special attention to field studies using crop canopy heating technology and in combination with other climate variables. We also discuss the biochemical reactions related to carbon fixation that may limit crop photosynthesis under warming temperatures and the current strategies for adaptation. Important progress has been made on several adaptation strategies demonstrating proof‐of‐concept for translating improved photosynthesis into higher yields. These are now poised to test in important food crops.     

Spectral vegetation indices as the indicator of canopy photosynthetic productivity in a deciduous broadleaf forest

Aims Understanding of the ecophysiological dynamics of forest canopy photosynthesis and its spatial and temporal scaling is crucial for revealing ecological response to climate change. Combined observations and analyses of plant ecophysiology and optical remote sensing would enable us to achieve these studies. In order to examine the utility of spectral vegetation indices (VIs) for assessing ecosystem-level photosynthesis, we investigated the relationships between canopy-scale photosynthetic productivity and canopy spectral reflectance over seasons for 5 years in a cool, temperate deciduous broadleaf forest at 'Takayama' super site in central Japan.Methods Daily photosynthetic capacity was assessed by in situ canopy leaf area index (LAI), (LAI × V cmax [single-leaf photosynthetic capacity]), and the daily maximum rate of gross primary production (GPP max) was estimated by an ecosystem carbon cycle model. We examined five VIs: normalized difference vegetation index (NDVI), enhanced vegetation index (EVI), green–red vegetation index (GRVI), chlorophyll index (CI) and canopy chlorophyll index (CCI), which were obtained by the in situ measurements of canopy spectral reflectance.Important findings Our in situ observation of leaf and canopy characteristics, which were analyzed by an ecosystem carbon cycling model, revealed that their phenological changes are responsible for seasonal and interannual variations in canopy photosynthesis. Significant correlations were found between the five VIs and canopy photosynthetic capacity over the seasons and years; four of the VIs showed hysteresis-type relationships and only CCI showed rather linear relationship. Among the VIs examined, we applied EVI–GPP max relationship to EVI data obtained by Moderate Resolution Imaging Spectroradiometer to estimate the temporal and spatial variation in GPP max over central Japan. Our findings would improve the accuracy of satellite-based estimate of forest photosynthetic productivity in fine spatial and temporal resolutions, which are necessary for detecting any response of terrestrial ecosystem to meteorological fluctuations.     

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

Photosynthesis and isoprene emission from trees along an urban–rural gradient in Texas

Isoprene emission is an important mechanism for improving the thermotolerance of plant photosystems as temperatures increase. In this study, we measured photosynthesis and isoprene emission in trees along an urban–rural gradient that serves as a proxy for climate change, to understand daily and seasonal responses to changes in temperature and other environmental variables. Leaf‐level gas exchange and basal isoprene emission of post oak (Quercus stellata) and sweet gum (Liquidambar styraciflua) were recorded at regular intervals over an entire growing season at urban, suburban, and rural sites in eastern Texas. In addition, the temperature and atmospheric carbon dioxide concentration experienced by leaves were experimentally manipulated in spring, early summer, and late summer. We found that trees experienced lower stomatal conductance and photosynthesis and higher isoprene emission, at the urban and suburban sites compared to the rural site. Path analysis indicated a daily positive effect of isoprene emission on photosynthesis, but unexpectedly, higher isoprene emission from urban trees was not associated with improved photosynthesis as temperatures increased during the growing season. Furthermore, urban trees experienced relatively higher isoprene emission at high CO₂ concentrations, while isoprene emission was suppressed at the other sites. These results suggest that isoprene emission may be less beneficial in urban, and potentially future, environmental conditions, particularly if higher temperatures override the suppressive effects of high CO₂ on isoprene emission. These are important considerations for modeling future biosphere–atmosphere interactions and for understanding tree physiological responses to climate change.     

Measurements of gross and net ecosystem productivity and water vapour exchange of a Pinus ponderosa ecosystem,and an evaluation of two generalized models

Net ecosystem productivity (NEP), net primary productivity (NPP), and water vapour exchange of a mature Pinus ponderosa forest (44°30′ N, 121°37′ W) growing in a region subject to summer drought were investigated along with canopy assimilation and respiratory fluxes. This paper describes seasonal and annual variation in these factors, and the evaluation of two generalized models of carbon and water balance (PnET‐II and 3‐PG) with a combination of traditional measurements of NPP, respiration and water stress, and eddy covariance measurements of above‐and below‐canopy CO₂ and water vapour exchange. The objective was to evaluate the models using two years of traditional and eddy covariance measurements, and to use the models to help interpret the relative importance of processes controlling carbon and water vapour exchange in a water‐limited pine ecosystem throughout the year. PnET‐II is a monthly time‐step model that is driven by nitrogen availability through foliar N concentration, and 3‐PG is a monthly time‐step quantum‐efficiency model constrained by extreme temperatures, drought, and vapour pressure deficits. Both models require few parameters and have the potential to be applied at the watershed to regional scale. There was 2/3 less rainfall in 1997 than in 1996, providing a challenge to modelling the water balance, and consequently the carbon balance, when driving the models with the two years of climate data, sequentially. Soil fertility was not a key factor in modelling processes at this site because other environmental factors limited photosynthesis and restricted projected leaf area index to ～1.6. Seasonally, GEP and LE were overestimated in early summer and underestimated through the rest of the year. The model predictions of annual GEP, NEP and water vapour exchange were within 1–39% of flux measurements, with greater disparity in 1997 because soil water never fully recharged. The results suggest that generalized models can provide insights to constraints on productivity on an annual basis, using a minimum of site data.     

Measuring and modelling seasonal variation of carbon dioxide and water vapour exchange of a Pinus ponderosa forest subject to soil water deficit

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 CO₂ flux from measurements and site meteorological data demonstrated that modelling of soil CO₂ flux based on an Arrhenius‐type equation in CANPOND overestimated CO₂ respired from the soil during drought and when temperatures were low.