Drivers of terrestrial plant production across broad geographical gradients

Terrestrial net primary production (NPP) varies across global climate gradients, but the mechanisms through which climate drives this variation remain subject to debate. Specifically, it is debatable whether NPP is primarily influenced by ‘direct’ effects of climate on the kinetics of plant metabolism or ‘indirect’ effects of climate on plant size, stand biomass, stand age structure and growing season length. We clarify several issues in this debate by presenting multiple lines of evidence that support a primarily indirect influence of climate on global variation in NPP across broad geographical gradients. First, we highlight > 60 years of research that suggests leaf area, growing season length, plant biomass and/or plant age are better predictors of NPP than climate or latitude. Second, we refute recent claims that using biomass and age as predictors of NPP represents circular reasoning. Third, we illustrate why effects of climate on the kinetics of plant production must be evaluated using instantaneous (not annualized) rates of productivity. Fourth, we review recent analyses showing that the effects of biomass and age on NPP are much stronger than the effects of climate. Fifth, we present new analyses of a high‐quality NPP dataset that demonstrate further that biomass, age and growing season length are better predictors of global variation in NPP than climate variables. Our results are consistent with the hypothesis that variation in NPP across global climate gradients primarily reflects the influence of climate on growing season length and stand biomass, as well as stand age, rather than the effects of temperature and precipitation on the kinetics of metabolism. However, this hypothesis should be evaluated further using larger, high‐quality observational and experimental datasets spanning multiple geographical scales.     

Deriving a new phenological indicator of interannual net carbon exchange in contrasting boreal deciduous and evergreen forests

Phenology is an important variable affecting the annual net ecosystem production (NEP) of terrestrial ecosystems. A new phenological indicator was proposed based on the ratio of respiration season length and growing season length (respiration–growth length ratio, RGR). Validation of this new phenological indicator was conducted using continuous flux measurements at contrasting boreal deciduous and evergreen forests in Canada. Analyses based on yearly anomalies of both annual NEP and phenological indicators indicated that the RGR can explain more proportion of interannual NEP variability compared to existing phenological metrics, including the carbon uptake period and the autumn lag. A multivariate regression model was used to predict the respiration–growth length ratio anomaly using anomalies of spring air temperature, autumn radiation and soil water content (SWC), which serves as a prerequisite for this indicator being scaled up for regional applications where flux data were unavailable. By normalization growing season length, interannual NEP showed comparable sensitivity to RGR variations of different plant functional types, which is a great advantage over other phenological indicators. The high potential of RGR in explaining interannual NEP variability may highlight the importance of respiration process in controlling annual NEP, which has probably been overlooked or underestimated in existing phenological studies. The comparable sensitivity of RGR to annual NEP observed at different plant functional types would favor its application in tracking interannual variability of NEP regionally and complementary to existing indices to promote our understanding of carbon sequestration with future climate change.     

The influence of meteorology and phenology on net ecosystem exchange in an eastern Siberian boreal larch forest

Aims Boreal forests play an important role in the global carbon cycle. Compared with the boreal forests in North America and Europe, relatively few research studies have been conducted in Siberian boreal forests. Knowledge related to the role of Siberian forests in the global carbon balance is thus essential for a full understanding of global carbon cycle.Methods This study investigated the net ecosystem exchange (NEE) during growing season (May–September) in an eastern Siberian boreal larch forest for a 3-year period in 2004–2006 with contrasting meteorological conditions.Important findings The study found that the forest served as a carbon sink during all of the 3 studied years; in addition, the meteorological conditions essentially influenced the specific annual value of the strength of the carbon sinks in each year. Although 2005 was the warmest year and much wetter than 2004, 2005 also featured the greatest amount of ecosystem respiration, which resulted in a minimum value of NEE. The study also found that the phenological changes observed during the three study years had a relatively small effect on annual NEE. Leaf expansion was 26 days earlier in 2005 than in the other 2 years, which resulted in a longer growing season in 2005. However, the NEE in 2005 was counterbalanced by the large rate of ecosystem respiration that was caused by the higher temperatures in the year. This study showed that meteorological variables had larger influences on the interannual variations in NEE for a Siberian boreal larch forest, as compared with phenological changes. The overall results of this study will improve our understanding of the carbon balance of Siberian boreal larch forests and thus can help to forecast the response of these forests to future climate change.     

Aboveground net production and dry matter allocation ofPinus pumila forests in the Kiso mountain range,central Japan

Aboveground net production rates of the subalpine stone pine (Pinus pumila) forests in central Japan were estimated by the summation method; net production was defined as the sum of annual biomass increment and annual loss due to death. In the two pine stands of different scrub heights, P1 (200 cm) and P2 (140 cm), aboveground biomass reached 177 and 126 ton ha⁻¹, respectively. Leaf biomass was about 14 ton ha⁻¹ in each stand. The estimates of aboveground net production during the 2 year period (1987–1989) averaged 4.1 and 3.7 ton ha⁻¹ y⁻¹ in P1 and P2, respectively, which were at the lowest among the pine forests in the world. Two indices of efficiency of energy fixation, that is, the ratio of net production to the total radiation during a growing season and the ratio of net production to total radiation per unit of leaf weight, were evaluated. Both efficiency indices for the twoP. pumila stands fell in the range obtained for other Japanese evergreen conifer forests. This suggested that the low annual net production of the stone pine stands were mainly due to a limitation in the length of the growing season. The pine forests were also characterized by a small allocation (about 17%) of aboveground net production into biomass increment, in comparison with other evergreen conifer forest types. Annual net carbon gain in theP. pumila forests was suggested to be largely invested in leaf production at the expense of the growth of woody parts.     

Nitrogen controls the net primary production of an alpine Kobresia meadow in the northern Qinghai‐Tibet Plateau

Net primary production (NPP) is a fundamental property of natural ecosystems. Understanding the temporal variations of NPP could provide new insights into the responses of communities to environmental factors. However, few studies based on long‐term field biomass measurements have directly addressed this subject in the unique environment of the Qinghai‐Tibet plateau (QTP). We examined the interannual variations of NPP during 2008–2015 by monitoring both aboveground net primary productivity (ANPP) and belowground net primary productivity (BNPP), and identified their relationships with environmental factors with the general linear model (GLM) and structural equation model (SEM). In addition, the interannual variation of root turnover and its controls were also investigated. The results show that the ANPP and BNPP increased by rates of 15.01 and 143.09 g/m² per year during 2008–2015, respectively. BNPP was mainly affected by growing season air temperature (GST) and growing season precipitation (GSP) rather than mean annual air temperature (MAT) or mean annual precipitation (MAP), while ANPP was only controlled by GST. In addition, available nitrogen (AN) was significantly positively associated with BNPP and ANPP. Root turnover rate averaged 30%/year, increased with soil depth, and was largely controlled by GST. Our results suggest that alpine Kobresia meadow was an N‐limited ecosystem, and the NPP on the QTP might increase further in the future in the context of global warming and nitrogen deposition.     

The effect of a dry spring on seasonal carbon allocation and vegetation dynamics in a poplar bioenergy plantation

In this study the seasonal variation in carbon, water and energy fluxes as well as in net primary productivity (NPP) of different tree components is presented for a 2‐year‐old poplar (Populus spp.) plantation. A thorough ecophysiological study was performed at ecosystem scale, at tree and at leaf level, in this high‐density bioenergy plantation. Seasonal variation in NPP and fluxes was analysed in relation to meteorological parameters at the field site. The growing season length in terms of carbon uptake was controlled by leaf area development until the maximum leaf area index (LAI_max) was reached. Afterwards, a shift to belowground carbon allocation was observed. A dry period in spring caused a reduced leaf area production as well as a decrease in net ecosystem exchange and gross primary production (GPP) due to stomatal closure. Water use efficiency and fine root growth increased in response to limiting soil water availability in the root zone. When soil water availability was not limiting, GPP was controlled by a decrease in solar radiation and air temperature. The results of this study indicate that the productivity of recently established bioenergy plantations with fast‐growing trees is very sensitive to drought. The interaction between soil water availability and factors controlling ecosystem GPP is crucial in assessing the CO₂ mitigation potential under future climate conditions.     

Climate change and the effect on the biomass growth in steppe communities in Kherlon River Basin, Northern China

The bacterial community composition and diversity in rock varnish of Turpan Basin were investigated by restriction fragment length polymorphism (RFLP) and clone library of the 16S rRNA gene. 114 positive clones were screened, which could be grouped into 28 phylotypes and then further divided into 23 different operational taxonomic units (OTUs). These were affiliated into 5 phyla (Acidobacteria, Proteobacteria, Chloroflexi, Firmicutes and Cyanobacteria). Clones from actinobacteria were the dominant, accounting for 67.5% of total clones in the library, followed by Proteobacteria (15.8%), Chloroflexi (13.2%), Firmicutes (2.6%) and Cyanobacteria (0.9%). Rubrobacter (accounts for 35%) in the phylum Actinobacteria was the dominant genus and contained many species which might be resistant to gamma radiation. A 70% of the library clone sequences showed less 97% similarity to 16S rRNA gene sequences of standard strains obtained by pure culture. Shannon–Wiener index value of this study is 2.52 and is lower than deep-sea sediments, soils, lakes and other environments. Results of this study showed that bacterial diversity in rock varnishes of Turpan Basin was low, but maybe exist a large number of new unknown taxons, especially species that could well adapted to drought and resist radiation.     

Alpine grassland plants grow earlier and faster but biomass remains unchanged over 35 years of climate change

Satellite data indicate significant advancement in alpine spring phenology over decades of climate warming, but corresponding field evidence is scarce. It is also unknown whether this advancement results from an earlier shift of phenological events, or enhancement of plant growth under unchanged phenological pattern. By analyzing a 35‐year dataset of seasonal biomass dynamics of a Tibetan alpine grassland, we show that climate change promoted both earlier phenology and faster growth, without changing annual biomass production. Biomass production increased in spring due to a warming‐induced earlier onset of plant growth, but decreased in autumn due mainly to increased water stress. Plants grew faster but the fast‐growing period shortened during the mid‐growing season. These findings provide the first in situ evidence of long‐term changes in growth patterns in alpine grassland plant communities, and suggest that earlier phenology and faster growth will jointly contribute to plant growth in a warming climate.     

Temporal changes in the relationship between tree-ring growth and net primary production in northern Japan: a novel approach to the estimation of seasonal photosynthate allocation to the stem

Tree-ring (TR) observations provide important data on long-term forest dynamics and their underlying ecophysiological mechanisms. To elucidate the seasonal link between photosynthetic carbon acquisition and TR growth, we analyzed the correlation between observed TR data (carbon sink) and model-estimated net primary production (NPP; carbon source). Temporal trends of the TR–NPP correlation over the last century were also analyzed to identify influences of past climate changes. We used TR data from Picea glehnii at seven sites on Hokkaido Island, Japan, which were obtained from the International Tree-Ring Data Bank. At each site, NPP was estimated using the Vegetation Integrative Simulator for Trace gases model, which was driven by long-term (1900–2010) meteorological data. Site-mean tree-ring width index (TRWI) chronologies were analyzed to reveal any relationship with the current or previous year’s annual or monthly NPP. We found moderate to strong correlations between TRWIs and model-estimated monthly NPP from April to June, especially in June of the current year, but no clear spatial trend was observed. During the twentieth century, the TRWI–NPP correlation increased for February, March, April, and July NPP of the current year and for October NPP of the previous year. Ecophysiologically, the period from April to June corresponds to the season when tree cambial cells are formed in the study area. Our findings suggest that photosynthate produced during this cambial growth season is allocated to stem growth and that this source allocation season has become longer due to past environmental changes.     

Impact of precipitation dynamics on net ecosystem productivity

Net ecosystem productivity (NEP) was measured on shortgrass steppe (SGS) vegetation at the USDA Central Plains Experimental Range in northeastern Colorado from 2001 to 2003. Large year‐to‐year differences were observed in annual NEP, with >95% of the net carbon uptake occurring during May and June. Low precipitation during the 2002 April to June time period greatly reduced annual net carbon uptake. Large precipitation events (>10 mm day^?1) promoted carbon uptake, while small precipitation events (<10 mm day^?1) enhanced heterotrophic respiration and resulted in a net loss of carbon from the system. Large precipitation event enhanced carbon uptake was attributed to increased soil water content (SWC), which promotes plant photosynthesis. The large precipitation events which occurred from July to October have lower increases in daytime net CO₂ uptake (NEP_d) due to the presence of low live plant biomass compared to earlier in the growing season. Live aboveground plant biomass (AGB), solar radiation, and SWC were the major variables that controlled NEP_d, while AGB, SWC, and relative humidity control nighttime respiration losses (NEP_n). Aboveground plant biomass is the most important variable for controlling both NEP_d and NEP_n dynamics. These results suggest that the major factor controlling growing season NEP_n is the amount of carbon fixed via photosynthesis during the day. Heterotrophic soil respiration is greatly enhanced for one to 2 days following rainfall events with daily rainfall events >5 mm having a similar increase in respiration (>3.00 g m Cm^?2 day^?1). In addition, the size of the heterotrophic respiration pulse is independent of both the amount of time since the last rainfall event and the time of occurrence during the growing season.     

Nonvascular contribution to ecosystem NPP in a subarctic heath during early and late growing season

Bryophytes and lichens abound in many arctic ecosystems and can contribute substantially to the ecosystem net primary production (NPP). Because of their growth seasonality and their potential for growth out of the growing season peak, bryophyte and lichen contribution to NPP may be particularly significant when vascular plants are less active and ecosystems act as a source of carbon (C). To clarify these dynamics, nonvascular and vascular aboveground NPP was compared for a subarctic heath during two contrasting periods of the growing season, viz. early-mid summer and late summer-early autumn. Nonvascular NPP was determined by assessing shoot biomass increment of three moss species (Hylocomium splendens, Pleurozium schreberi and Dicranum elongatum) and by scaling to ecosystem level using average standing crop. For D. elongatum, these estimates were compared with production estimates obtained from measurements of shoot length increase. Vascular NPP was determined by harvesting shrub and herb apical growth and considering production due to stem secondary growth of shrubs. Hylocomium splendens and Pleurozium schreberi showed highest biomass growth in late summer, whereas for D. elongatum this occurred in early summer. Maximum relative growth rates were ca. 0.003–0.007 g g⁻¹ d⁻¹. For D. elongatum, production estimates from length growth differed from estimations from biomass growth, likely because of an uncoupling between length growth and biomass shoot growth. Nonvascular NPP was 0.37 and 0.46 g dry weight m⁻² d⁻¹, in early and late summer, respectively, whereas in the same periods vascular NPP was 3.6 and 1.1 g dry weight m⁻² d⁻¹. The contribution of nonvascular NPP to total aboveground NPP was therefore minor in early summer but substantial in late summer, when 25% of the C accumulated by the vegetation was incorporated into nonvascular plant tissue. The expected global change-induced reduction of nonvascular plant biomass in subarctic heath is likely therefore to enhance C release during the late part of the growing season.     

Interannual variability in net CO2 exchange of a native tallgrass prairie

Year‐round eddy covariance flux measurements were made in a native tallgrass prairie in north‐central Oklahoma, USA during 1997–2000 to quantify carbon exchange and its interannual variability. This prairie is dominated by warm season C₄ grasses. The soil is a relatively shallow silty clay loam underlined with a heavy clay layer and a limestone bedrock. During the study period, the prairie was burned in the spring of each year, and was not grazed. In 1997 there was adequate soil moisture through the growing season, but 1998 had two extended periods of substantially low soil moisture (with concurrent high air temperatures and vapor pressure deficits), one early and one later in the growing season. There was also moisture stress in 1999, but it was less severe and occurred later in the season. The annual net ecosystem CO₂ exchange, NEE (before including carbon loss during the burn) was 274, 46 and 124 g C m ^{? 2} yr ^{? 1} in 1997, 1998, and 1999, respectively (flux toward the surface is positive), and the associated variation seemed to mirror the severity of moisture stress. We also examined integrated values of NEE during different periods (e.g. day/night; growing season/senescence). Annually integrated carbon dioxide uptake during the daytime showed the greatest variability from year to year, and was primarily linked to the severity of moisture stress. Carbon loss during nighttime was a significant part of the annual daytime NEE, and was fairly stable from year to year. When carbon loss during the burn (estimated from pre‐ and post‐burn biomass samples) was incorporated in the annual NEE, the prairie was found to be approximately carbon neutral (i.e. net carbon uptake/release was near zero) in years with no moisture stress (1997) or with some stress late in the season (1999). During a year with severe moisture stress early in the season (1998), the prairie was a net source of carbon. It appears that moisture stress (severity as well as timing of occurrence) was a dominating factor regulating the annual carbon exchange of the prairie.     

Leaf Phenology and Seasonal Carbon Gain in the Invasive Plant, Bunias orientalis L.

Abstract: In two potentially competing herbaceous plants, the invasive Bunias orientalis L. (Brassicaceae) and the native Picris hieracioides L. (Asteraceae), seasonal changes in leaf CO₂ gas exchange and plant growth were studied over an entire growing season from February 1998 to December 1998 in two experimental fields. The study was motivated by the hypothesis that pre-adaptive phenological displacement of alien species relative to the native flora may be an important reason for the observed expansion of B. orientalis in central Europe. We quantified the importance of phenological differences for annual carbon gain in both species by estimating total leaf carbon gain from the results of leaf CO₂ exchange and changes in plant leaf area. Bunias orientalis achieved almost half of its annual carbon gain in the time between early September and December, when competition for light by other species, like P. hieracioides , is low. Our quantitative approach corroborates the notion that the phenological shift of a relatively poor competitor, such as B. orientalis , could be of great importance for the success as an invasive species.     

基于森林资源清查资料的落叶松林生物量和净生长量估算模式

 丰富的森林资源清查资料是了解各类森林材积准确信息的重要途径,如果能将这些资源用于估算森林生物量和生产力的动态变化,不仅对于科学地指导森林的经营管理,而且对于全球变化的研究,特别是区域尺度的生产力模型验证,都具有重要意义。根据我国落叶松(Larix)林生物量和材积的实际调查资料,探讨了基于森林资源清查资料(森林材积V和林龄A)估算森林生物量和生产力的方法,指出无论是人工林还是天然林,落叶松林的生物量与其蓄积量、生产力与其年均净生物生产量(B/A)和年均净蓄积生产量(V/A)均呈双曲线关系,但落叶松林的生产力与其生物量(B)关系不明显,并分别建立了人工和天然落叶松林的相关模型;所建模型克服了将森林生物量与其蓄积量之比作为常数的不足,并考虑了林龄对于森林生产力的影响。     

锡林河流域一个放牧羊草群落中碳素平衡的初步估计

 野外调查与历史资料相结合,对内蒙古锡林河流域一个放牧羊草（Leymus chinensis）草原群落的碳素贮量、主要流量和周转速度等进行了估计,在此基础上对放牧情况下该群落的碳素收支进行了概算。结果表明：1）该群落中地上部净初级生产固碳量的两年平均值为78.2 gC·m－2·a－1, 根系碳素输入量的平均值为322.5 gC·m－2·a－1, 碳素输入总量为400.7 gC·m－2·a－1; 2）土壤净呼吸量为343.7 gC·m－2·a－1,家畜采食量为49.7 gC·m－2·a－1,动物（昆虫）采食量为14.7 gC·m－2·a－1,地上立枯阶段的淋溶与光化学分解损失为3.2 gC·m－2·a－1,碳素输出总量为411.3 gC·m－2·a－1; 3）该群落中碳素输出略大于输入,净释放速率为10.6 gC·m－2·a－1,0～30 cm土壤中的碳素周转速率为6.2%,周转时间为16年。     

秦岭山地植被净初级生产力及对气候变化的响应

基于1999~2009年的NDVI数据和气象数据,利用CASA模型对秦岭山地植被净初级生产力（Net primary productivity,NPP）进行模拟估算,并分析了秦岭NPP的时空变化特征及其对气候变化的响应。结果表明：1999~2009年11年间秦岭山地的平均年NPP为542.24 gC·m^-2·a^-1;研究期内秦岭NPP呈显著增长趋势（P<0.01）,2008年最高（718.77 gC·m^-2·a^-1）,2001年最低（471.78 gC·m^-2·a^-1）;四季对全年NPP的贡献率大小依次为夏季（49.90%）>春季（26.16%）>秋季（18.87%）>冬季（5.07%）;月NPP与温度和降水都显著相关,但与温度的相关性更高,月水平上温度对NPP的影响比降水大;生长季期间NPP与温度和降水的相关性在空间分布上都以正相关为主。     

Gross primary production controls the subsequent winter CO2 exchange in a boreal peatland

In high‐latitude regions, carbon dioxide (CO₂) emissions during the winter represent an important component of the annual ecosystem carbon budget; however, the mechanisms that control the winter CO₂ emissions are currently not well understood. It has been suggested that substrate availability from soil labile carbon pools is a main driver of winter CO₂ emissions. In ecosystems that are dominated by annual herbaceous plants, much of the biomass produced during the summer is likely to contribute to the soil labile carbon pool through litter fall and root senescence in the autumn. Thus, the summer carbon uptake in the ecosystem may have a significant influence on the subsequent winter CO₂ emissions. To test this hypothesis, we conducted a plot‐scale shading experiment in a boreal peatland to reduce the gross primary production (GPP) during the growing season. At the growing season peak, vascular plant biomass in the shaded plots was half that in the control plots. During the subsequent winter, the mean CO₂ emission rates were 21% lower in the shaded plots than in the control plots. In addition, long‐term (2001–2012) eddy covariance data from the same site showed a strong correlation between the GPP (particularly the late summer and autumn GPP) and the subsequent winter net ecosystem CO₂ exchange (NEE). In contrast, abiotic factors during the winter could not explain the interannual variation in the cumulative winter NEE. Our study demonstrates the presence of a cross‐seasonal link between the growing season biotic processes and winter CO₂ emissions, which has important implications for predicting winter CO₂ emission dynamics in response to future climate change.     

NET PRIMARY PRODUCTION AND PHENOLOGY ON A SOUTHERN APPALACHIAN WATERSHED

Net primary production (NPP) is an important function of plant communities which has not often been examined seasonally in a forested ecosystem. The major objective of the study was to measure above-ground NPP seasonally and relate it to phenological activity on a hardwood forest watershed at Coweeta Hydrologic Laboratory, North Carolina. NPP was estimated as the increase in biomass, estimated from regression equations on diameter. Diameter increases were measured by vernier tree bands. Phenological observations were made on bud break, leaf emergence, flowering, mature fruit, leaf senescence, and leaf fall. The species studied intensively were Acer rubrum, Quercus prinus, Carya glabra, Cornus florida, and Liriodendron tulipifera. Liriodendron was found to be the most productive species per individual, but Quercus prinus was the most productive per unit ground area. The total watershed estimate of aboveground NPP was 8,754 kg ha^-1 yr^-1 and included 47.9% leaves, 33.2% wood, 7.8% bark, 4.8% reproductive tissues, 4.2% loss to consumers, and 2.1% twigs. Increases in leaf biomass were most rapid in the spring, but woody tissue production peaked in June and continued through August. Since leaf production peaked in the spring, the plants' photosynthetic machinery was activated early in the growing season to support woody tissue production, which followed the period of rapid leaf growth, and reproductive activity. Flowering occurred during the leaf expansion period except for Acer rubrum, which flowered before leaf emergence. Fruit maturation occurred during late summer to early fall, when there were no additional biomass increases. Acer rubrum was an exception as its fruit matured during the period of leaf expansion.     

Net primary productivity mapped for Canada at 1-km resolution

Aim To map net primary productivity (NPP) over the Canadian landmass at 1‐km resolution. Location Canada. Methods A simulation model, the Boreal Ecosystem Productivity Simulator (BEPS), has been developed. The model uses a sunlit and shaded leaf separation strategy and a daily integration scheme in order to implement an instantaneous leaf‐level photosynthesis model over large areas. Two key driving variables, leaf area index (every 10 days) and land cover type (annual), are derived from satellite measurements of the Advanced Very High Resolution Radiometer (AVHRR). Other spatially explicit input data are also prepared, including daily meteorological data (radiation, precipitation, temperature, and humidity), available soil water holding capacity (AWC) and forest biomass. The model outputs are compared with ground plot data to ensure that no significant systematic biases are created. Results The simulation results show that Canada’s annual net primary production was 1.22 Gt C year^?1 in 1994, 78% attributed to forests, mainly the boreal forest, without considering the contribution of the understorey. The NPP averaged over the entire landmass was ~140 g C m^?2 year^?1 in 1994. Geographically, NPP varied greatly among ecozones and provinces/territories. The seasonality of NPP is characterized by strong summer photosynthesis capacities and a short growing season in northern ecosystems. Conclusions This study is the first attempt to simulate Canada‐wide NPP with a process‐based model at 1‐km resolution and using a daily step. The statistics of NPP are therefore expected to be more accurate than previous analyses at coarser spatial or temporal resolutions. The use of remote sensing data makes such simulations possible. BEPS is capable of integrating the effects of climate, vegetation, and soil on plant growth at a regional scale. BEPS and its parameterization scheme and products can be a basis for future studies of the carbon cycle in mid‐high latitude ecosystems.     

Photosynthesis and carbon balance of a Sahelian fallow savanna

Eddy-covariance measurements of CO₂ exchange above a Sahelian savanna consisting of small shrubs over a near-continuous herb layer were made during the HAPEX-Sahel experiment in Niger, West Africa. The measurements were made near-continuously during an 8-week period, covering the main part of the rainy season and three weeks at the beginning of the dry season. The measurements were corrected for in-canopy storage of CO₂ and the night-time measurements used to derive respiration functions for the soil, roots and above-ground plant material. Photosynthetic CO₂ uptake was estimated and compared to simulations using a biochemical photosynthesis model in a simple, ‘big-leaf’, implementation. The model satisfactorily reproduced the measurements (coefficient of determination 0.80) using parameters defined from the literature and based on soil nutrient concentrations. When the quantum yield (α) and rubisco capacity (V_mr) were fitted to the data with allowance for physiological changes through the season, an excellent agreement between model and measurements was obtained (coefficient of determination 0.93, RMS error 1.46 μmol m^–2 s^–1). The fitted photosynthesis and respiration model was used to estimate the carbon balance of the savanna site during the growing season of 1992 and for the complete calendar year. Harvest estimates of net plant biomass accumulation during the growing season and annual wood accumulation agreed well with modelled net photosynthesis and annual net carbon accumulation, respectively. Peak instantaneous ecosystem CO₂ uptake was comparable to peak values observed in other biomes, but annual photosynthesis and carbon sequestration were considerably lower than observed elsewhere.