ALBERT

Notes: A model is presented which solves simultaneously for leaf-scale stomatal conductance, CO2 assimilation and the energy balance as a function of leaf position within canopies of well-watered vegetation. Fluxes and conductances were calculated separately for sunlit and shaded leaves. A linear dependence of photosynthetic capacity on leaf nitrogen content was assumed, while leaf nitrogen content and light intensity were assumed to decrease exponentially within canopies. Separate extinction coefficients were used for diffuse and direct beam radiation. An efficient Gaussian integration technique was used to compute fluxes and mean conductances for the canopy. The multilayer model synthesizes current knowledge of radiation penetration, leaf physiology and the physics of evaporation and provides insights into the response of whole canopies to multiple, interacting factors. The model was also used to explore sources of variation in the slopes of two simple parametric models (nitrogen- and light-use efficiency), and to set bounds on the magnitudes of the parameters.For canopies low in total N, daily assimilation rates are ∼10% lower when leaf N is distributed uniformly than when the same total N is distributed according to the exponentially decreasing profile of absorbed radiation. However, gains are negligible for plants with high N concentrations. Canopy conductance, Gc should be calculated as Gc=Aσ(fslgsl+fshgsh), where Δ is leaf area index, fsi and fsh are the fractions of sunlit and shaded leaves at each level, and gsi and gsh are the corresponding stomatal conductances. Simple addition of conductances without this weighting causes errors in transpiration calculated using the ‘big-leaf’ version of the Penman-Monteith equation. Partitioning of available energy between sensible and latent heat is very responsive to the parameter describing the sensitivity of stomata to the atmospheric humidity deficit. This parameter also affects canopy conductance, but has a relatively small impact on canopy assimilation.Simple parametric models are useful for extrapolating understanding from small to large scales, but the complexity of real ecosystems is thus subsumed in unexplained variations in parameter values. Simulations with the multilayer model show that both nitrogen- and radiation-use efficiencies depend on plant nutritional status and the diffuse component of incident radiation, causing a 2- to 3-fold variation in these efficiencies.

Notes: Higher rates of nitrate assimilation are required to support faster growth in enhanced carbon dioxide. To investigate how this is achieved, tobacco plants were grown on high nitrate and high light in ambient and enhanced (700 μmol mol–1) carbon dioxide. Surprisingly, enhanced carbon dioxide did not increase leaf nitrate reductase (NR) activity in the middle of the photoperiod. Possible reasons for this anomalous result were investigated. (a) Measurements of biomass, nitrate, amino acids and glutamine in plants fertilized once and twice daily with 12 mol m–3 nitrate showed that enhanced carbon dioxide did not lead to a nitrate limitation in these plants. (b) Enhanced carbon dioxide modified the diurnal regulation of NR activity in source leaves. The transcript for nia declined during the light period in a similar manner in ambient and enhanced carbon dioxide. The decline of the transcript correlated with a decrease of nitrate in the leaf, and was temporarily reversed after re-irrigating with nitrate in the second part of the photoperiod. The decline of the transcript was not correlated with changes of sugars or glutamine. NR activity and protein decline in the second part of the photoperiod, and NR is inactivated in the dark in ambient carbon dioxide. The decline of NR activity was smaller and dark inactivation was partially reversed in enhanced carbon dioxide, indicating that post-transcriptional or post-translational regulation of NR has been modified. The increased activation and stability of NR in enhanced carbon dioxide was correlated with higher sugars and lower glutamine in the leaves. (c) Enhanced carbon dioxide led to increased levels of the minor amino acids in leaves. (d) Enhanced carbon dioxide led to a large decrease of glycine and a small decrease of serine in leaves of mature plants. The glycine:serine ratio decreased in source leaves of older plants and seedlings. The consequences of a lower rate of photorespiration for the levels of glutamine and the regulation of nitrogen metabolism are discussed. (e) Enhanced carbon dioxide also modified the diurnal regulation of NR in roots. The nia transcript increased after nitrate fertilization in the early and the second part of the photoperiod. The response of the transcript was not accentuated in enhanced carbon dioxide. NR activity declined slightly during the photoperiod in ambient carbon dioxide, whereas it increased 2-fold in enhanced carbon dioxide. The increase of root NR activity in enhanced carbon dioxide was preceded by a transient increase of sugars, and was followed by a decline of sugars, a faster decrease of nitrate than in ambient carbon dioxide, and an increase of nitrite in the roots. (f) To interpret the physiological significance of these changes in nitrate metabolism, they were compared with the current growth rate of the plants. (g) In 4–5-week-old plants, the current rate of growth was similar in ambient and enhanced carbon dioxide (≈ 0·4 g–1 d–1). Enhanced carbon dioxide only led to small changes of NR activity, nitrate decreased, and overall amino acids were not significantly increased. (h) Young seedlings had a high growth rate (0·5 g–1 d–1) in ambient carbon dioxide, that was increased by another 20% in enhanced carbon dioxide. Enhanced carbon dioxide led to larger increases of NR activity and NR activation, a 2–3-fold increase of glutamine, a 50% increase of glutamate, and a 2–3-fold increase in minor amino acids. It also led to a higher nitrate level. It is argued that enhanced carbon dioxide leads to a very effective stimulation of nitrate uptake, nitrate assimilation and amino acid synthesis in seedlings. This will play an important role in allowing faster growth rates in enhanced carbon dioxide at this stage.

Notes: Root exudates were sampled from detopped root systems of castor bean (Ricinus communis). Different volume flux rates were imposed by changing the pneumatic pressure around the root system using a Passioura-type pressure chamber. The concentrations of cations, anions, amino acids, organic acids and abscisic acid decreased hyperbolically when flux rates increased from pure root exudation up to values typical for transpiring plants. Concentrations at low and high fluxes differed by up to 40 times (phosphate) and the ratio of substances changed by factors of up to 10. During the subsequent reduction of flux produced by lowering the pneumatic pressure in the root pressure chamber, the concentrations and ratios of substances deviated (at a given flux rate) from those found when flux was increased. The flux dependence of exudate composition cannot therefore be explained by a simple dilution mechanism. Xylem sap samples from intact, transpiring plants were collected using a Passioura-type root pressure chamber. The concentrations of the xylem sap changed diurnally. Substances could be separated into three groups: (1) calcium, magnesium and amino acid concentrations correlated well with the values expected from their concentration-flux relationships, whereas (2) the concentrations of sulphate and phosphate deviated from the expected relationships during the light phase, and (3) nitrate and potassium concentrations in intact plants varied in completely the opposite manner from those in isolated root systems. Abscisic acid concentrations in the root exudate were dependent on the extent of water use and showed strong diurnal variations in the xylem sap of intact plants even in droughtstressed plants. Calculations using root exudates overestimated export from the root system in intact plants, with the largest deviation found for proton flux (a factor of 10). We conclude that root exudate studies cannot be used as the sole basis for estimating fluxes of substances in the xylem of intact plants. Consequences for studying and modelling xylem transport in whole plants are discussed.

Notes: We studied the effects of variations of water flux through the plant, of diurnal variation of water flux, and of variation of vapour pressure deficit at the leaf on compensation pressure in the Passioura-type pressure chamber, the composition of the xylem sap and leaf conductance in Ricinus communis. The diurnal pattern of compensation pressure showed stress relaxation during the night hours, while stress increased during the day, when water limitation increased. Thus compensation pressure was a good measure of the momentary water status of the root throughout the day and during drought. The bulk soil water content at which predawn compensation pressure and abscisic acid concentration in the xylem sap increased and leaf conductance decreased, was high when the water usage of the plant was high. For all xylem sap constituents analysed, variations in concentrations during the day were larger than changes in mean concentrations with drought. Mean concentrations of phosphate and the pH of the xylem sap declined with drought, while nitrate concentration remained constant. When the measurement leaf was exposed to a different VPD from the rest of the plant, leaf conductance declined by 400mmol m−2 s−1 when compensation pressure increased by 1 MPa in all treatments. The compensation pressure needed to keep the shoot turgid, leaf conductance and the abscisic acid concentration in the xylem were linearly related. This was also the case when the highly dynamic development of stress was taken into account.

Notes: Based on review and original data, this synthesis investigates carbon pools and fluxes of Siberian and European forests (600 and 300 million ha, respectively). We examine the productivity of ecosystems, expressed as positive rate when the amount of carbon in the ecosystem increases, while (following micrometeorological convention) downward fluxes from the atmosphere to the vegetation (NEE = Net Ecosystem Exchange) are expressed as negative numbers. Productivity parameters are Net Primary Productivity (NPP=whole plant growth), Net Ecosystem Productivity (NEP = CO2 assimilation minus ecosystem respiration), and Net Biome Productivity (NBP = NEP minus carbon losses through disturbances bypassing respiration, e.g. by fire and logging). Based on chronosequence studies and national forestry statistics we estimate a low average NPP for boreal forests in Siberia: 123 gC m–2 y–1. This contrasts with a similar calculation for Europe which suggests a much higher average NPP of 460 gC m–2 y–1 for the forests there. Despite a smaller area, European forests have a higher total NPP than Siberia (1.2–1.6 vs. 0.6–0.9 × 1015 gC region–1 y–1). This arises as a consequence of differences in growing season length, climate and nutrition. For a chronosequence of Pinus sylvestris stands studied in central Siberia during summer, NEE was most negative in a 67-y old stand regenerating after fire (– 192 mmol m–2 d–1) which is close to NEE in a cultivated forest of Germany (– 210 mmol m–2 d–1). Considerable net ecosystem CO2-uptake was also measured in Siberia in 200- and 215-y old stands (NEE:174 and – 63 mmol m–2 d–1) while NEP of 7- and 13-y old logging areas were close to the ecosystem compensation point. Two Siberian bogs and a bog in European Russia were also significant carbon sinks (– 102 to – 104 mmol m–2 d–1). Integrated over a growing season (June to September) we measured a total growing season NEE of – 14 mol m–2 summer–1 (– 168 gC m–2 summer–1) in a 200-y Siberian pine stand and – 5 mol m–2 summer–1 (– 60 gC m–2 summer–1) in Siberian and European Russian bogs. By contrast, over the same period, a spruce forest in European Russia was a carbon source to the atmosphere of (NEE: + 7 mol m–2 summer–1 = + 84 gC m–2 summer–1). Two years after a windthrow in European Russia, with all trees being uplifted and few successional species, lost 16 mol C m–2 to the atmosphere over a 3-month in summer, compared to the cumulative NEE over a growing season in a German forest of – 15.5 mol m–2 summer–1 (– 186 gC m–2 summer–1; European flux network annual averaged – 205 gC m–2 y–1). Differences in CO2-exchange rates coincided with differences in the Bowen ratio, with logging areas partitioning most incoming radiation into sensible heat whereas bogs partitioned most into evaporation (latent heat). Effects of these different surface energy exchanges on local climate (convective storms and fires) and comparisons with the Canadian BOREAS experiment are discussed. Following a classification of disturbances and their effects on ecosystem carbon balances, fire and logging are discussed as the main processes causing carbon losses that bypass heterotrophic respiration in Siberia. Following two approaches, NBP was estimated to be only about 13–16 mmol m–2 y–1 for Siberia. It may reach 67 mmol m–2 y–1 in North America, and about 140–400 mmol m–2 y–1 in Scandinavia. We conclude that fire speeds up the carbon cycle, but that it results also in long-term carbon sequestration by charcoal formation. For at least 14 years after logging, regrowth forests remain net sources of CO2 to the atmosphere. This has important implications regarding the effects of Siberian forest management on atmospheric concentrations. For many years after logging has taken place, regrowth forests remain weaker sinks for atmospheric CO2 than are nearby old-growth forests.

Notes: Abstract  Until now, many extracellular matrix proteins, e.g. osteopontin and osteonectin, have been used to determine a cell’s osteogenic maturation. The disadvantage in evaluation of these proteins is their relative wide-ranging appearance throughout the osteogenic differentiation process. Thus, the aim of this study was to establish an immunohistochemical setup using E11, a marker that binds selectively to cells of the late osteogenic cell lineage. In addition, the histochemical expression of the bone matrix proteins osteonectin, osteopontin and fibronectin was compared to that of E11 using monoclonal antibodies. For light microscopical detection of osteogenic markers in cultured cells we developed a simple paraffin technique using a fibrin glue as embedding medium. This allows the handling of cultured cells such as a tissue sample and includes the use of stored biological specimens for further immunohistochemical experiments. We used newborn rat calvariae for whole tissue preparations and for isolation and cultivation of bone cells. In addition, we included the rat osteosarcoma cell line ROS 17/2.8 in this study. For the first time, we have localised E11 in osteocytes of rat calvaria preparations at the electron microscopical level. E11 was detected at plasma membranes of osteocytes and their processes, but not at those of osteoblasts. Accompanying experiments with cultured newborn rat calvaria cells and ROS 17/2.8 cells revealed E11 reactivity on a subset of cells. The results obtained confirm the suitability of the differentiation marker E11 as a sensitive instrument for the characterisation of bone cell culture systems.

Notes: Summary Experiments with Prunus armeniaca were carried out under conditions of constant temperature but varying air humidity. Experiments were also contucted with a constant water vapor difference between the evaporating sites in a leaf and the air, but with varying leaf temperature. These served as a basis for predicting the daily course of total diffusion resistance under the natural climatic conditions of a desert. For the simulation, the rsults of the experiments at constant conditions with only one variable factor are fitted with empirical equations which serve as “calibration curves” to predict the change in diffusion resistance caused by a change in humidity and temperature calculated from the meteorological data of a desert day. The simulation shows that for P. armeniaca humidity and temperature are the dominating factors in controlling the daily course of diffusion resistance. For meteorologically very different days the simulation allows the increase in diffusion resistance in the morning to be predicted with an accuracy of 90%–105% as compared to directly observed measurements. In the afternoon, especially after extreme climatic conditions during the morning, the deviation between predicted and observed values of diffusion resistance may be greater, but not more than -20% to -30%. This possibly indicates the existence of an additional factor of significance which was not included in the simulation. The two peaked curves of net photosynthesis and transpiration characteristic of plants living under arid conditions can be explained in this species by the humidity-and temperature-controlled stomatal response. This stomatal regulation leads to a decreasing total daily transpirational water loss on a dry day as compared to a moist one. The significance of this controlling mechanism for the primary production and the water relations of P. armeniaca is discussed.

Notes: Summary During the dry season in the Negev desert (Israel) Artemisia herbaalba in its natural habitat has a very low water content. It shows values of negative hydrostatic pressure in the xylem down to -163 bars and an extreme of osmotic potential in the leaves of -92 bars. The diurnal water stress does not decrease strongly in the night. Under these conditions Artemisia is still photosynthetically active for a few hours of the day during the whole dry period.

Notes: Summary A digital registration system used with temperature- and humidity-controlled cuvettes for net photosynthesis and transpiration measurements in the field is described. The associated errors of the measured parameters and calculated data are estimated. The digitalization is based on an analogue registration which is of primary importance in the control of experimental conditions in the cuvettes. The digital system is connected to the analogue registration in series. The error associated with digitalization is 0.1% across 70% of the scale. This error increases to 0.2% between 3 and 30% on the scale due to a minor lack of linearity. The reproducibility of the digitalization is ±0.024%. The error associated with data transfer in the digitalization and the errors of the analogue registration are estimated for temperature and humidity measurements (error of air and leaf temperature is ±0.1° C; error of the dew point temperature is ±1.1° C dew point). The effect of these errors on the calculation of relative humidity and the water vapour difference between the leaf and the air is determined using the progressive error law. At 30° C and 50% relative humidity, the error in relative humidity is ±7.4%, the error for the water vapour difference is ±6.6%. The dependence of these errors on temperature and humidity is shown. The instrument error of the net photosynthesis measurement is calculated to be ±4.2%. Transpiration measurements have an average inaccuracy of ±8.3%. The total diffusion resistance which is calculated from values of transpiration and the water vapour difference has an average error of ±10.9%. The sizeable influence of errors in humidity and temperature measurements on the calculated diffusion resistance is demonstrated. The additional influence of biological errors associated with field measurements is discussed.