A meta-analysis of leaf gas exchange and nitrogen in trees grown under elevated carbon dioxide

The response of trees to rising atmospheric CO₂ concentration ([CO₂]) is of concern to forest ecologists and global carbon modellers and is the focus of an increasing body of research work. I review studies published up to May 1994, and several unpublished works, which reported at least one of the following: net CO₂ assimilation (A), stomatal conductance (g_s), leaf dark respiration (R_d) leaf nitrogen or specific leaf area (SLA) in woody plants grown at <400 μmol mol^?1 CO₂ or at 600–800 μmol mol^?1 CO₂. The resulting data from 41 species were categorized according to growth conditions (unstressed versus stressed), length of CO₂ exposure, pot size and exposure facility [growth chamber (GC), greenhouse (GH), or open-top chamber (OTC)] and interpreted using meta-analytic methods. Overall, A showed a large and significant increase at elevated [CO₂] but length of CO₂ exposure and the exposure facility were important modifiers of this response. Plants exposed for < 50 d had a significantly greater response, and those from GCs had a significantly lower response than plants from longer exposures or from OTC studies. Negative acclimation of A was significant and general among stressed plants, but in unstressed plants was influenced by length of CO₂ exposure, the exposure facility and/or pot size. Growth at elevated [CO₂] resulted in moderate reductions in gs in unstressed plants, but there was no significant effect of CO₂ on gs in stressed plants. Leaf dark respiration (mass or area basis) was reduced strongly by growth at high [CO₂] > while leaf N was reduced only when expressed on a mass basis. This review is the first meta-analysis of elevated CO₂ studies and provides statistical confirmation of several general responses of trees to elevated [CO₂]. It also highlights important areas of continued uncertainty in our understanding of these responses.     

Carbon isotope fractionation in tree ring early and late wood in relation to intra-growing season water balance

We determined the stable carbon isotope composition (δ^1.3C) of cellulose extracted from early and late wood in Douglas fir [Pseudotsuga menziexii (Mirb.) Franco] tree rings. Data were obtained for the period 1962 to 1981, at the start of which the trees were 20 years old. A water balance model was used to calculate daily stand transpiration and water deficit. The model incorporates site factors (soil water availability, slope and aspect) and environmental variables (solar radiation, air temperature and rainfall). There was far greater variability in late wood than in early wood δ^1.3C. In wet years, late wood δ^1.3C was significantly lighter (by as much as 2δ) than early wood δ^1.3C but in dry years this difference was reversed. Differences between spring and summer cumulative transpiration accounted for almost 60δ of the variability in differences between early and late wood δ^1.3C. We found excellent correspondence between summer cumulative transpiration and late wood δ^1.3C, with estimates of transpiration accounting for up to 93% of the variability in δ^1.3C. Correlations between early wood δ^1.3C and spring transpiration were generally poor (r2<0.4), but we were able to identify those exceptional years in which there had been a very dry spring. Our results indicate that, while tree ring δ^1.3C correlates reasonably well with basal area increment, it is a far better indicator of inter- and intra-annual variability in water availability than radial growth.     

Carbon isotope composition of C4 grasses is influenced by light and water supply

The carbon isotope composition of C⁴ grasses has the potential to be used as an indicator of changes in the isotopic composition and concentration of atmospheric CO2, especially for climate reconstruction. The usefulness of C₄ grasses for this purpose hinges on the assumption that their photosynthetic discrimination against ¹³C remains constant in a wide range of environmental conditions. We tested this assumption by examining the effects of light and water stress on the carbon isotope composition of C₄ grasses using different biochemical subtypes (NADP-ME, NAD-ME, PCK) in glasshouse experiments. We grew 14 different C₄ grass species in four treatments: sun-watered, sun-drought, shade-watered and shade-drought. Carbon isotope discrimination (Δ) rarely remained constant. In general, Δ values were lowest in sun-watered grasses, greater for sun-drought plants and even higher for plants of the shade-watered treatment. The highest Δ values were generally found in the most stressed grasses, the shade-drought plants. Grasses of the NADP-ME subtype were the least influenced by a change in environmental variables, followed by PCK and NAD-ME subtypes. Water availability affected the carbon isotope discrimination less than light limitation in PCK and NAD-ME subtypes, but similarly in NADP-ME subtypes. In another experiment, we studied the effect of increasing light levels (150 to 1500 μmol photons m^?2 s^?1) on the Δ values of 18 well-watered C₄ grass species. Carbon isotope discrimination remained constant until photon flux density (PFD) was less than 700 μmol photons m^?2 s^?1. Below this light level, Δ values increased with decreasing irradiance for all biochemical subtypes. The change in A was less pronounced in NADP-ME and PCK than in NAD-ME grasses. Grasses grown in the field and in the glasshouse showed a similar pattern. Thus, caution should be exercised when using C4 plants under varying environmental conditions to monitor the concentration or carbon isotopic composition of atmospheric CO2 in field/glasshouse studies or climate reconstruction.     

The response of isoprene emission rate and photosynthetic rate to photon flux and nitrogen supply in aspen and white oak trees

Isoprene is the primary biogenic hydrocarbon emitted from temperate deciduous forest ecosystems. The effects of varying photon flux density (PFD) and nitrogen growth regimes on rates of isoprene emission and net photosynthesis in potted aspen and white oak trees are reported. In both aspen and oak trees, whether rates were expressed on a leaf area or dry mass basis, (1) growth at higher PFD resulted in significantly higher rates of isoprene emission, than growth at lower PFD, (2) there is a significant positive relationship between isoprene emission rate and leaf nitrogen concentration in both sun and shade trees, and (3) there is a significant positive correlation between isoprene emission rate and photosynthetic rate in both sun and shade trees. The greater capacity for isoprene emission in sun leaves was due to both higher leaf mass per unit area and differences in the biochemical and/or physiological properties that influence isoprene emission. Positive correlations between isoprene emission rate and leaf nitrogen concentration support the existence of mechanisms that link leaf nitrogen status to isoprene synthase activity. Positive correlations between isoprene emission rate and photosynthesis rate support previous hypotheses that isoprene emission plays a role in protecting photosynthetic mechanisms during stress.     

Geochemistry of aquatic humic substances in the Lake Fryxell Basin, Antarctica

Dissolved organic carbon (DOC) in Lake Fryxell, 10 streams flowing into the lake, and the moat surrounding the lake was studied to determine the influence of sources and biogeochemical processes on its distribution and chemical nature. Lake Fryxell is an amictic, permanently ice-covered lake in the McMurdo Dry Valleys which contains benthic and planktonic microbial populations, but receives essentially no input of organic material from the ahumic soils of the watershed. Biological activity in the water column does not appear to influence the DOC depth profile, which is similar to the profiles for conservative inorganic constituents. DOC values for the streams varied with biomass in the stream channel, and ranged from 0.2 to 9.7 mg C/L. Fulvic acids in the streams were a lower percentage of the total DOC than in the lake. These samples contain recent carbon and appear to be simpler mixtures of compounds than the lake samples, indicating that they have undergone less humification. The fulvic acids from just above the sediments of the lake have a high sulfur content and are highly aliphatic. The main transformations occurring as these fractions diffuse upward in the water column are 1) loss of sulfur groups through the oxycline and 2) decrease in aliphatic carbon and increase in the heterogeneity of aliphatic moieties. The fraction of modem¹⁴C content of the lake fulvic acids range from a minimum of 0.68 (approximately 3000 years old) at 15m depth to 0.997 (recent material) just under the ice. The major processes controlling the DOC in the lake appear to be: 1) The transport of organic matter by the inflow streams resulting in the addition of recent organic material to the moat and upper waters of the lake; 2) The diffusion of organic matter composed of relict organic material and organic carbon resulting from the degradation of algae and bacteria from the bottom waters or sediments of the lake into overlying glacial melt water, 3) The addition of recent organic matter to the bottom waters of the lake from the moat.     

The origin and recycling of sedimented biogenic debris in a subalphine eutrophic lake (Lake Bled, Slovenia)

The areal distribution of organic C contents, ¹³C values, total N and P and biogenic Si contents in surficial sediments were used to study the distribution, origin and diagenetic transformations of sedimented biogenic debris in the eutrophic subalpine Lake Bled (Slovenia), which for most of the yearhas an anoxic hypolimnion. The influence of an allochthonous input, restricted to the western basin, was clearly traced by higher organic C and total N and P contents, higher ¹³C values, and higher sedimentation rate in comparison to the eastern basin. The low ¹³C values of sedimentary organic matter in the major part of the lake, lower than the ¹³C values of different types of organic matter, suggest that this sedimentary organic matter is most probably the product of a microbial community and not a residue of primary production.The temporal variation of benthic diffusive fluxes of NH₄, Si and PO₄, derived from modelling the pore water profiles, was related to sedimentation of phytoplanktonic blooms, while the PO₄ fluxes were also dependent on changing redox conditions at the sediment-water interface in the period of the winter-spring overtum. The removal of PO₄ in pore waters is probably due to the adsorption of phosphate and precipitation of apatite and vivianite. The budget of C, N and P at the sediment-water interface revealed a high recycling efficiency (>70%), also confirmed by the rather uniform (or only slightly decreasing) vertical profiles of organic C, total N and P in sediment cores and C/N and C/P ratios. The percentage of biogenic Si recycling is low (<10%), suggesting its removal in sediments.     

Effects of elevated CO2 and increased nitrogen deposition on photosynthesis and growth of understory plants in spruce model ecosystems

We studied the effects of atmospheric CO₂ enrichment (280, 420 and 560 l CO₂ l^–1) and increased N deposition (0,30 and 90 kg ha^–1 year^–1) on the spruce-forest understory species Oxalis acetosella, Homogyne alpina and Rubus hirtus. Clones of these species formed the ground cover in nine 0.7 m² model ecosystems with 5-year-old Picea abies trees (leaf area index of approx 2.2). Communities grew on natural forest soil in a simulated montane climate. Independently of N deposition, the rate of light-saturated net photosynthesis of leaves grown and measured at 420 l CO₂ l^–1 was higher in Oxalis and in Homogyne, but was not significantly different in Rubus compared to leaves grown and measured at the pre-industrial CO₂ concentration of 280 l l^–1. Remarkably, further CO₂ enrichment to 560 l l^–1 caused no additional increase of CO₂ uptake. With increasing CO₂ supply concentrations of non-structural carbohydrates in leaves increased and N concentrations decreased in all species, whereas N deposition had no significant effect on these traits. Above-ground biomass and leaf area production were not significantly affected by elevated CO₂ in the more vigorously growing species O. acetosella and R. hirtus, but the slow growing H. alpina produced almost twice as much biomass and 50% more leaf area per plant under 420 l CO₂ l^–1 compared to 280 l l^–1 (again no further stimulation at 560 l l^–1). In contrast, increased N addition stimulated growth in Oxalis and Rubus but had no effect on Homogyne. In Oxalis (only) biomass per plant was positively correlated with microhabitat quantum flux density at low CO₂, but not at high CO₂ indicating carbon saturation. On the other hand, the less shade-tolerant Homogyne profited from CO₂ enrichment at all understory light levels facilitating its spread into more shady micro-habitats under elevated CO₂. These species-specific responses to CO₂ and N deposition will affect community structure. The non-linear responses to elevated CO₂ of several of the traits studied here suggest that the largest responses to rising atmospheric CO₂ are under way now or have already occurred and possible future responses to further increases in CO₂ concentration are likely to be much smaller in these understory species.     

Exchange of NO and NO2 between wheat canopy monoliths and the atmosphere

The fluxes of NO and NO₂ between wheat canopy monoliths and the atmosphere were investigated with the dynamic chamber technique. For this purpose monoliths were dug out at different plant growth stages from a field site, transported to the institute, and placed in an environmental growth chamber. The wheat canopy monoliths were exposed over a period of four days to the average ratios of atmospheric NO₂ and NO measured at the field site, i.e. NO₂ concentration of about 18 mL L^-1 plus NO concentration lower than 0.5 nL L^-1. Under these conditions NO emission into the atmosphere and NO₂ deposition into canopy monoliths was observed. Both fluxes showed diurnal variation with maximum rates during the light and minimum rates during darkness. NO₂ fluxes correlated with soil temperature as well as with light intensity. NO fluxes correlated with soil temperature but not with light intensity. From the investigation performed the diurnal variation of the NO and NO₂ compensation points, the maximum rates of NO and NO₂ emission, and the total resistances of NO and NO₂ fluxes were calculated. Under the assumption that the measured data are representative for the whole vegetation period, annual fluxes of NO and NO₂ were estimated. Annual NO emission into the atmosphere amounted to 87 mg N m^-2 y^-1 (0.87 kg ha^-1 y^-1), annual NO₂ deposition into canopy monoliths amounted to 1273 mg N m^-2 y^-1 (12.73 kg ha^-1 y^-1). Apparently, the uptake of atmospheric nitrogen by the wheat field from NO₂ deposition is about 15 times higher than the loss of nitrogen from NO emission. It can therefore be assumed that even in rural areas wheat fields are a considerable sink for atmospheric nitrogen. The annual sink strength estimated in the present study is ca. 12 kg N ha^-1 y^-1. The possible origin of the NO emitted and the fate of atmospheric NO₂ taken up by the wheat canopy monoliths are discussed.Preliminary results of this paper were presented at the Joint Workshop COST 611/Working Party 3 and EUROTRAC in Delft, The Netherlands (Ludwig et al., 1991).     

Effect of maximal arm exercise on skin blood flux in the paralyzed lower limbs in persons with spinal cord injury

The purpose of the present study was to examine the effect of maximal arm exercise on the skin blood circulation of the paralyzed lower limbs in persons with spinal cord injury (PSCI). Eight male PSCI with complete lesions located between T3 and L1 performed graded maximal arm-cranking exercise (MACE) to exhaustion. The skin blood flux at the thigh (SBFT) and that at the calf (SBFC) were monitored using laser-Doppler flowmeter at rest and for 15 s immediately after the MACE. The subject's mean peak oxygen uptake and peak heart rate was 1.41 ± 0.22 1·min⁻¹ and 171.6 ± 19.2 beats·min⁻¹, respectively. No PSCI showed any increase in either SBFT or SBFC after the MACE, when compared with the values at rest. These results suggest that the blood circulation of the skin in the paralyzed lower limbs in PSCI is unaffected by the MACE.     

太湖全新世沉积物有机联同位素的分布及其古气侯意义

通过太湖两个钻孔岩芯有机碳同位素（１３Ｃ／１２Ｃ）的分析,发现δ１３Ｃ的垂直分布与太湖地区１６０００ａＢ．Ｐ．以来古气候波动相关。根据δ１３Ｃ垂直分布曲线可推测,大约在１１０００—６０００ａＢ．Ｐ,太湖地区处于温暖湿润期,气温高于现在平均温度。另一方面,根据太湖两岩芯沉积物的δ１３Ｃ值的对比发现,大约在１１０００—６０００ａＢ．Ｐ,西太湖（Ｗ１Ｂ）岩芯沉积物的δ１３Ｃ平均值明显高于东太湖（Ｅ２Ｂ）。据此可认为,在这一期间西太湖很可能有过海水侵入。