ALBERT

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: 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: Abstract Above-and belowground biomass distribution, isotopic composition of soil and xylem water, and carbon isotope ratios were studied along an aridity gradient in Patagonia (44–45°S). Sites, ranging from those with Nothofagus forest with high annual rainfall (770 mm) to Nothofagus scrub (520 mm), Festuca (290 mm) and Stipa (160 mm) grasslands and into desert vegetation (125 mm), were chosen to test whether rooting depth compensates for low rainfall. Along this gradient, both mean above-and belowground biomass and leaf area index decreased, but average carbon isotope ratios of sun leaves remained constant (at-27‰), indicating no major differences in the ratio of assimilation to stomatal conductance at the time of leaf growth. The depth of the soil horizon that contained 90% of the root biomass was similar for forests and grasslands (about 0.80–0.50 m), but was shallower in the desert (0.30 m). In all habitats, roots reached water-saturated soils or ground water at 2–3 m depth. The depth profile of oxygen and hydrogen isotope ratios of soil water corresponded inversely to volumetric soil water contents and showed distinct patterns throughout the soil profile due to evaporation, water uptake and rainfall events of the past year. The isotope ratios of soil water indicated that high soil moisture at 2–3 m soil depth had originated from rainy periods earlier in the season or even from past rainy seasons. Hydrogen and oxygen isotope ratios of xylem water revealed that all plants used water from recent rain events in the topsoil and not from water-saturated soils at greater depth. However, this study cannot explain the vegetation zonation along the transect on the basis of water supply to the existing plant cover. Although water was accessible to roots in deeper soil layers in all habitats, as demonstrated by high soil moisture, earlier rain events were not fully utilized by the current plant cover during summer drought. The role of seedling establishment in determining species composition and vegetation type, and the indirect effect of seedling establishment on the use of water by fully developed plant cover, are discussed in relation to climate change and vegetation modelling.

Notes: Abstract The depth at which plants are able to grow roots has important implications for the whole ecosystem hydrological balance, as well as for carbon and nutrient cycling. Here we summarize what we know about the maximum rooting depth of species belonging to the major terrestrial biomes. We found 290 observations of maximum rooting depth in the literature which covered 253 woody and herbaceous species. Maximum rooting depth ranged from 0.3 m for some tundra species to 68 m for Boscia albitrunca in the central Kalahari; 194 species had roots at least 2 m deep, 50 species had roots at a depth of 5 m or more, and 22 species had roots as deep as 10 m or more. The average for the globe was 4.6±0.5 m. Maximum rooting depth by biome was 2.0±0.3 m for boreal forest. 2.1±0.2 m for cropland, 9.5±2.4 m for desert, 5.2±0.8 m for sclerophyllous shrubland and forest, 3.9±0.4 m for temperate coniferous forest, 2.9±0.2 m for temperate deciduous forest, 2.6±0.2 m for temperate grassland, 3.7±0.5 m for tropical deciduous forest, 7.3±2.8 m for tropical evergreen forest, 15.0±5.4 m for tropical grassland/savanna, and 0.5±0.1 m for tundra. Grouping all the species across biomes (except croplands) by three basic functional groups: trees, shrubs, and herbaceous plants, the maximum rooting depth was 7.0±1.2 m for trees, 5.1±0.8 m for shrubs, and 2.6±0.1 m for herbaceous plants. These data show that deep root habits are quite common in woody and herbaceous species across most of the terrestrial biomes, far deeper than the traditional view has held up to now. This finding has important implications for a better understanding of ecosystem function and its application in developing ecosystem models.

Notes: Abstract Understanding and predicting ecosystem functioning (e.g., carbon and water fluxes) and the role of soils in carbon storage requires an accurate assessment of plant rooting distributions. Here, in a comprehensive literature synthesis, we analyze rooting patterns for terrestrial biomes and compare distributions for various plant functional groups. We compiled a database of 250 root studies, subdividing suitable results into 11 biomes, and fitted the depth coefficient β to the data for each biome (Gale and Grigal 1987). β is a simple numerical index of rooting distribution based on the asymptotic equation Y=1-βd, where d = depth and Y = the proportion of roots from the surface to depth d. High values of β correspond to a greater proportion of roots with depth. Tundra, boreal forest, and temperate grasslands showed the shallowest rooting profiles (β=0.913, 0.943, and 0.943, respectively), with 80–90% of roots in the top 30 cm of soil; deserts and temperate coniferous forests showed the deepest profiles (β=0.975 and 0.976, respectively) and had only 50% of their roots in the upper 30 cm. Standing root biomass varied by over an order of magnitude across biomes, from approximately 0.2 to 5 kg m-2. Tropical evergreen forests had the highest root biomass (5 kg m-2), but other forest biomes and sclerophyllous shrublands were of similar magnitude. Root biomass for croplands, deserts, tundra and grasslands was below 1.5 kg m-2. Root/shoot (R/S) ratios were highest for tundra, grasslands, and cold deserts (ranging from 4 to 7); forest ecosystems and croplands had the lowest R/S ratios (approximately 0.1 to 0.5). Comparing data across biomes for plant functional groups, grasses had 44% of their roots in the top 10 cm of soil. (β=0.952), while shrubs had only 21% in the same depth increment (β=0.978). The rooting distribution of all temperate and tropical trees was β=0.970 with 26% of roots in the top 10 cm and 60% in the top 30 cm. Overall, the globally averaged root distribution for all ecosystems was β=0.966 (r 2=0.89) with approximately 30%, 50%, and 75% of roots in the top 10 cm, 20 cm, and 40 cm, respectively. We discuss the merits and possible shortcomings of our analysis in the context of root biomass and root functioning.

Notes: Abstract The study presents a data set of above-ground biomass (AGB), structure, spacing and fire regime, for 24 stands of pristine Siberian Scots pine (Pinus sylvestris) forests with lichens (n = 20) or Vaccinium/mosses (n = 4) as ground cover, along four chronosequences. The stands of the “lichen” site type (LT) were stratified into three chronosequences according to stand density and fire history. Allometric equations were established from 90 sample trees for stem, coarse branch, fine branch, twig and needle biomass. The LT stands exhibited a low but sustained biomass accumulation until a stand age of 383 years. AGB reached only 6–10 kgdw m−2 after 200 years depending on stand density and fire history compared to 20 kgdw m−2 in the “Vaccinium” type (VT) stands. Leaf area index (LAI) in the LT stands remained at 0.5–1.5 and crown cover was 30–60%, whereas LAI reached 2.5 and crown cover was 〉100% in the VT stands. Although nearest-neighbour analyses suggested the existence of density-dependent mortality, fire impact turned out to have a much stronger effect on density dynamics. Fire scar dating and calculation of mean and initial fire return intervals revealed that within the LT stands differences in structure and biomass were related to the severity of fire regimes, which in turn was related to the degree of landscape fragmentation by wetlands. Self-thinning analysis was used to define the local carrying capacity for biomass. A series of undisturbed LT stands was used to characterise the upper self-thinning boundary. Stands that had experienced a moderate fire regime were positioned well below the self-thinning boundary in a distinct fire-thinning band of reduced major axis regression slope −0.26. We discuss how this downward shift resulted from alternating phases of density reduction by fire and subsequent regrowth. We conclude that biomass in Siberian Scots pine forests is strongly influenced by fire and that climate change will affect ecosystem functions predominantly via changes in fire regimes.

Notes: Abstract The grass flora of Namibia (374 species in 110 genera) shows surprisingly little variation in δ13C values along a rainfall gradient (50–600 mm) and in different habitat conditions. However, there are significant differences in the δ13C values between the metabolic types of the C4 photosynthetic pathway. NADP-ME-type C4 species exhibit the highest δ13C values (−11.7 ‰) and occur mainly in regions with high rainfall. NAD-ME-type C4 species have significantly lower δ13C values (−13.4 ‰) and dominate in the most arid part of the precipitation regime. PCK-type C4 species play an intermediate role (−12.5 ‰) and reach a maximum abundance in areas of intermediate precipitation. This pattern is also evident in genera containing species of different metabolic types. Within the same genus NAD species reach more negative δ13C values than PCK species and δ13C values decreased with rainfall. Also in Aristida, with NADP-ME-type photosynthesis, δ13C values decreased from −11 ‰ in the inland region (600 mm precipitation) to −15 ‰ near the coast (150 mm precipitation), which is a change in discrimination which is otherwise associated by a change in metabolism. The exceptional C3 species Eragrostis walteri and Panicum heterostachyum are coastal species experiencing 50 mm precipitation only. Many of the rare species and monotypic genera grow in moist habitats rather than in the desert, and they are not different in their carbon isotope ratios from the more common flora. The role of species diversity with respect to habitat occupation and carbon metabolism is discussed.

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: Abstract The most widely distributed coniferous forests in the world are the larch forests. In the Russian Federation they occupy 27.6 × 106 ha. In Siberia, the larch species Larix russica generally grows west of the Yenissei River, and Larix gmelinii grows to the east. The morphological and physiological features of L. gmelinii make it possible for this species to grow in the far north of eastern Siberia, where climate conditions are more severe: The range of air temperature fluctuations in this region is more than 100°C, from 38°C down to 64°C below zero. One of the major adaptions to unfavorable soil conditions is provided by a specific feature of root formation in L. gmelinii, in which the apex central root dies off at the permafrost border and a root system develops in upper soil layers. The major larch vulnerability factors are natural and anthropogenic fires and damage caused by insects, which become more frequent with hot and dry weather. The consequences of projected global warming could be both positive and negative for larch forests. Permafrost melting may result in improved soil nutrition in the areas the larch forests occupy, yet the frequency of forest fires and damage by pathogens are likely to increase. Global warming is expected to cause forest die back and increased areas of steppe in the southern regions of eastern Siberia.