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.

Respiratory carbon losses and the carbon-use efficiency of a northern hardwood forest, 1999-2003.

Quantitative assessment of carbon (C) storage by forests requires an understanding of climatic controls over respiratory C loss. Ecosystem respiration can be estimated biometrically as the sum (R Sigma) of soil (Rs), leaf (Rl) and wood (Rw) respiration, and meteorologically by measuring above-canopy nocturnal CO2 fluxes (Fcn). Here we estimated R Sigma over 5 yr in a forest in Michigan, USA, and compared R Sigma and Fcn on turbulent nights. We also evaluated forest carbon-use efficiency (Ec = P(NP)/P(GP)) using biometric estimates of net primary production (P(NP)) and R Sigma and Fcn-derived estimates of gross primary production (P(GP)). Interannual variation in R Sigma was modest (142 g C m(-2) yr(-1)). Mean annual R Sigma was 1425 g C m(-2) yr(-1); 71% from Rs, 18% from Rl, and 11% from Rw. Hourly R Sigma was well correlated with Fcn, but 11 to 58% greater depending on the time of year. Greater R Sigma compared with Fcn resulted in higher estimated annual P(GP) and lower annual Ec (0.42 vs 0.54) using biometric and meteorological data, respectively. Our results provide one of the first multiyear estimates of R Sigma in a forested ecosystem, and document the responses of component respiratory C losses to major climatic drivers. They also provide the first assessment of Ec in a deciduous forest using independent estimates of P(GP).

Predicting the onset of net carbon uptake by deciduous forests with soil temperature and climate data: a synthesis of FLUXNET data.

We tested the hypothesis that the date of the onset of net carbon uptake by temperate deciduous forest canopies corresponds with the time when the mean daily soil temperature equals the mean annual air temperature. The hypothesis was tested using over 30 site-years of data from 12 field sites where CO(2) exchange is being measured continuously with the eddy covariance method. The sites spanned the geographic range of Europe, North America and Asia and spanned a climate space of 16 degrees C in mean annual temperature. The tested phenology rule was robust and worked well over a 75 day range of the initiation of carbon uptake, starting as early as day 88 near Ione, California to as late as day 147 near Takayama, Japan. Overall, we observed that 64% of variance in the timing when net carbon uptake started was explained by the date when soil temperature matched the mean annual air temperature. We also observed a strong correlation between mean annual air temperature and the day that a deciduous forest starts to be a carbon sink. Consequently we are able to provide a simple phenological rule that can be implemented in regional carbon balance models and be assessed with soil and temperature outputs produced by climate and weather models.

Environmental controls on sap flow in a northern hardwood forest.

Our objective was to gain a detailed understanding of how photosynthetically active radiation (PAR), vapor pressure deficit (D) and soil water interact to control transpiration in the dominant canopy species of a mixed hardwood forest in northern Lower Michigan. An improved understanding of how these environmental factors affect whole-tree water use in unmanaged ecosystems is necessary in assessing the consequences of climate change on the terrestrial water cycle. We used continuously heated sap flow sensors to measure transpiration in mature trees of four species during two successive drought events. The measurements were scaled to the stand level for comparison with eddy covariance estimates of ecosystem water flux (Fw). Photosynthetically active radiation and D together explained 82% of the daytime hourly variation in plot-level transpiration, and low soil water content generally resulted in increased stomatal sensitivity to increasing D. There were also species-specific responses to drought. Quercus rubra L. showed low water use during both dry and wet conditions, and during periods of high D. Among the study species, Acer rubrum L. showed the greatest degree of stomatal closure in response to low soil water availability. Moderate increases in stomatal sensitivity to D during dry periods were observed in Populus grandidentata Michx. and Betula papyrifera Marsh. Sap flow scaled to the plot level and Fw demonstrated similar temporal patterns of water loss suggesting that the mechanisms controlling sap flow of an individual tree also control ecosystem evapotranspiration. However, the absolute magnitude of scaled sap flow estimates was consistently lower than Fw. We conclude that species-specific responses to PAR, D and soil water content are key elements to understanding current and future water fluxes in this ecosystem.

Interactive effects of soil temperature, atmospheric carbon dioxide and soil N on root development, biomass and nutrient uptake of winter wheat during vegetative growth.

Nutrient requirements for plant growth are expected to rise in response to the predicted changes in CO(2) and temperature. In this context, little attention has been paid to the effects of soil temperature, which limits plant growth at early stages in temperate regions. A factorial growth-room experiment was conducted with winter wheat, varying soil temperature (10 degrees C and 15 degrees C), atmospheric CO(2) concentration (360 and 700 ppm), and N supply (low and high). The hypothesis was that soil temperature would modify root development, biomass allocation and nutrient uptake during vegetative growth and that its effects would interact with atmospheric CO(2) and N availability. Soil temperature effects were confirmed for most of the variables measured and 3-factor interactions were observed for root development, plant biomass components, N-use efficiency, and shoot P content. Importantly, the soil temperature effects were manifest in the absence of any change in air temperature. Changes in root development, nutrient uptake and nutrient-use efficiencies were interpreted as counterbalancing mechanisms for meeting nutrient requirements for plant growth in each situation. Most variables responded to an increase in resource availability in the order: N supply >soil temperature >CO(2).

Family- and population-level responses to atmospheric CO2 concentration: gas exchange and the allocation of C, N, and biomass in Plantago lanceolata (Plantaginaceae).

To ascertain the inheritance of responses to changing atmospheric CO(2) content, we partitioned response to elevated CO(2) in Plantago lanceolata between families and populations in 18 families in two populations. Plants were grown in 35 Pa and 71 Pa partial pressure of CO(2) (pCO(2)) in open-top chambers. We measured above- and belowground mass, carbon (C), nitrogen (N), hexose sugar, and gas exchange properties in both CO(2) treatments. Families within populations differed in mass, mass allocation, root : shoot ratios, aboveground percentage N, C : N ratio, and gas exchange properties. The CO(2) × family interaction is the main indicator of potential evolutionary responses to changing CO(2). Significant CO(2) × family interactions were observed for N content, C : N ratio, and photosynthetic rate (A: instantaneous light-saturated carbon assimilation capacity), intercellular CO(2) concentration, transpiration rate (E), and water use efficiency (WUE = A/E), but not for stomatal conductance. Families differed significantly in acclimation across time. The ratio of A in elevated vs. ambient growth CO(2), when measured at a common internal CO(2) partial pressure was 0.79, indicating down-regulation of A under CO(2) enrichment. Mass, C : N ratio, percentage, C (%C), and soluble sugar all increased significantly but overall %N did not change. Increases in %C and sugar were significant and were coincident with redistribution of N aboveground. The observed variation among populations and families in response to CO(2) is evidence of genetic variation in response and therefore of the potential for novel evolutionary trajectories with rising atmospheric CO(2).

Stomatal conductance of forest species after long-term exposure to elevated CO2 concentration: a synthesis.

• Data from 13 long-term (> 1 yr), field-based studies of the effects of elevated CO2 concentration ([CO2 ]) on European forest tree species were analysed using meta-analysis and modelling. Meta-analysis was used to determine mean responses across the data sets, and data were fitted to two commonly used models of stomatal conductance in order to explore response to environmental conditions and the relationship with assimilation. • Meta-analysis indicated a significant decrease (21%) in stomatal conductance in response to growth in elevated [CO2 ] across all studies. The response to [CO2 ] was significantly stronger in young trees than old trees, in deciduous compared to coniferous trees, and in water stressed compared to nutrient stressed trees. No evidence of acclimation of stomatal conductance to elevated [CO2 ] was found. • Fits of data to the first model showed that growth in elevated [CO2 ] did not alter the response of stomatal conductance to vapour pressure deficit, soil water content or atmospheric [CO2 ]. Fits of data to the second model indicated that conductance and assimilation responded in parallel to elevated [CO2 ] except when water was limiting. • Data were compared to a previous meta-analysis and it was found that the response of gs to elevated [CO2 ] was much more consistent in long-term (> 1 yr) studies, emphasising the need for long-term elevated [CO2 ] studies. By interpreting data in terms of models, the synthesis will aid future modelling studies of responses of forest trees to elevated [CO2 ].

Atmospheric CO(2) and mycorrhiza effects on biomass allocation and nutrient uptake of nodulated pea (Pisum sativum L.) plants.

The effect of ambient and elevated atmospheric CO(2) on biomass partitioning and nutrient uptake of mycorrhizal and non-mycorrhizal pea plants grown in pots in a controlled environment was studied. The hypothesis tested was that mycorrhizae would increase C assimilation by increasing photosynthetic rates and reduce below-ground biomass allocation by improving nutrient uptake. This effect was expected to be more pronounced at elevated CO(2) where plant C supply and nutrient demand would be increased. The results showed that mycorrhizae did not interact with atmospheric CO(2) concentration in the variables measured. Mycorrhizae did not affect photosynthetic rates, had no effect on root weight or root length density and almost no effect on nutrient uptake, but still significantly increased shoot weight and reduced root/shoot ratio at harvest. Elevated CO(2) increased photosynthetic rates with no evidence for down-regulation, increased shoot weight and nutrient uptake, had no effect on root weight, and actually reduced root/shoot ratio at harvest. Non-mycorrhizal plants growing at both CO(2) concentrations had lower shoot weight than mycorrhizal plants with similar nutritional status and photosynthetic rates. It is suggested that the positive effect of mycorrhizal inoculation was caused by an enhanced C supply and C use in mycorrhizal plants than in non-mycorrhizal plants. The results indicate that plant growth was not limited by mineral nutrients, but partially source and sink limited for carbon. Mycorrhizal inoculation and elevated CO(2) might have removed such limitations and their effects on above-ground biomass were independent, positive and additive.

Genotypic variation in physiological and growth responses of Populus tremuloides to elevated atmospheric CO2 concentration.

Physiological and biomass responses of six genotypes of Populus tremuloides Michx., grown in ambient t (357 micromol mol(-1)) or twice ambient (707 micromol mol(-1)) CO2 concentration ([CO2]) and in low-N or high-N soils, were studied in 1995 and 1996 in northern Lower Michigan, USA. There was a significant CO2 x genotype interaction in photosynthetic responses. Net CO2 assimilation (A) was significantly enhanced by elevated [CO2] for five genotypes in high-N soil and for four genotypes in low-N soil. Enhancement of A by elevated [CO2] ranged from 14 to 68%. Genotypes also differed in their biomass responses to elevated [CO2], but biomass responses were poorly correlated with A responses. There was a correlation between magnitude of A enhancement by elevated [CO2] and stomatal sensitivity to CO2. Genotypes with low stomatal sensitivity to CO2 had a significantly higher A at elevated [CO2] than at ambient [CO2], but elevated [CO2] did not affect the ratio of intercellular [CO2] to leaf surface [CO2]. Stomatal conductance and A of different genotypes responded differentially to recovery from drought stress. Photosynthetic quantum yield and light compensation point were unaffected by elevated [CO2]. We conclude that P. tremuloides genotypes will respond differentially to rising atmospheric [CO2], with the degree of response dependent on other abiotic factors, such as soil N and water availability. The observed genotypic variation in growth could result in altered genotypic representation within natural populations and could affect the composition and structure of plant communities in a higher [CO2] environment in the future.

Genotypic variation for condensed tannin production in trembling aspen (POPULUS TREMULOIDES, salicaceae) under elevated CO2 and in high- and low-fertility soil.

The carbon/nutrient balance hypothesis suggests that leaf carbon to nitrogen ratios influence the synthesis of secondary compounds such as condensed tannins. We studied the effects of rising atmospheric carbon dioxide on carbon to nitrogen ratios and tannin production. Six genotypes of Populus tremuloides were grown under elevated and ambient CO(2) partial pressure and high- and low-fertility soil in field open-top chambers in northern lower Michigan, USA. During the second year of exposure, leaves were harvested three times (June, August, and September) and analyzed for condensed tannin concentration. The carbon/nutrient balance hypothesis was supported overall, with significantly greater leaf tannin concentration at high CO(2) and low soil fertility compared to ambient CO(2) and high soil fertility. However, some genotypes increased tannin concentration at elevated compared to ambient CO(2), while others showed no CO(2) response. Performance of lepidopteran leaf miner (Phyllonorycter tremuloidiella) larvae feeding on these plants varied across genotypes, CO(2), and fertility treatments. These results suggest that with rising atmospheric CO(2), plant secondary compound production may vary within species. This could have consequences for plant-herbivore and plant-microbe interactions and for the evolutionary response of this species to global climate change.

Modelling the growth, survival and death of microorganisms in foods: the UK food micromodel approach.

Techniques for the development of mathematical models in the area of predictive microbiology have greatly improved recently, allowing better and more accurate descriptions of microbial responses to particular environmental conditions, thus enabling predictions of those responses to be made with greater confidence. Recognising the potential value of applying these techniques in the food industry, the Ministry of Agriculture, Fisheries and Food (MAFF) initiated a nationally coordinated five-year programme of research into the growth and survival of microorganisms in foods, with the aim of developing a computerised Predictive Microbiology Database in the UK. This initiative has resulted in the systematic generation of data, through protocols which ensure consistency of methodology, so that data in the database are truly comparable and compatible, and lead to reliable predictive models. The approaches taken by scientists involved in this programme are described and the various stages in the development of mathematical models summarized. It is hoped that this initiative and others being developed in the USA, Australia, Canada and other countries, will encourage a more integrated approach to food safety which will influence all stages of food production and, eventually, result in the development of an International Predictive Microbiology Database.

Growth and senescence in plant communities exposed to elevated CO2 concentrations on an estuarine marsh.

Three high marsh communities on the Chesapeake Bay were exposed to a doubling in ambient CO2 concentration for one growing season. Open-top chambers were used to raise CO2 concentrations ca. 340 ppm above ambient over monospecific communities of Scirpus olneyi (C3) and Spartina patens (C4), and a mixed community of S. olneyi, S. patens, and Distichlis spicata (C4). Plant growth and senescence were monitored by serial, nondestructive censuses. Elevated CO2 resulted in increased shoot densities and delayed sensecence in the C3 species. This resulted in an increase in primary productivity in S. olneyi growing in both the pure and mixed communities. There was no effect of CO2 on growth in the C4 species. These results demonstrate that elevated atmospheric CO2 can cause increased aboveground production in a mature, unmanaged ecosystem.

Nitrogen and carbon dynamics in C3 and C4 estuarine marsh plants grown under elevated CO2 in situ.

Carbon dioxide concentrations were elevated in three estuarine communities for an entire growing season. Open top chambers were used to raise CO2 concentrations ca. 336 ppm above ambient in monospecific communities of Scirpus olneyi (C3) and Spartina patens (C4), and a mixed community of S. olneyi, S. patens and Distichlis spicata (C4). Nitrogen and carbon concentration (% wt) of aboveground tissue was followed throughout growth and senescence. Green shoot %N was reduced and %C was unchanged under elevated CO2 in S. olneyi. This resulted in a 20%-40% increase in tissue C/N ratio. There was no effect of CO2 on either C4 species. Maximum aboveground N (g/m2) was unchanged in S. olneyi, indicating that increased productivity under elevated CO2 was dependent on reallocation of stored N. There was no change in the N recovery efficiency of S. olneyi in pure stand and a decrease in the mixed community. Litter C/N ratio was not affected by elevated CO2 suggesting that decomposition and N mineralization rates will also remain unchanged. Continued growth responses to elevated CO2 could, however, be limited by the ability of S. olneyi to increase the total aboveground N pool.