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.
We investigated the daily exchange of CO 2 between undisturbed Larix gmelinii (Rupr.) Rupr. forest and the atmosphere at a remote Siberian site during July and August of 1993. Our goal was to measure and partition total CO 2 exchanges into aboveground and belowground components by measuring forest and understory eddy and storage¯uxes and then to determine the relationships between the environmental factors and these observations of ecosystem metabolism. Maximum net CO 2 uptake of the forest ecosystem was extremely low compared to the forests elsewhere, reaching a peak of only $5 mmol m À2 s À1 late in the morning. Net ecosystem CO 2 uptake increased with increasing photosynthetically active photon ux density (PPFD) and decreased as the atmospheric water vapor saturation de®cit (D) increased. Daytime ecosystem CO 2 uptake increased immediately after rain and declined sharply after about six days of drought. Ecosystem respiration at night averaged $2.4 mmol m À2 s À1 with about 40% of this coming from the forest¯oor (roots and heterotrophs). The relationship between the understory eddy¯ux and soil temperature at 5 cm followed an Arrhenius model, increasing exponentially with temperature (Q 10 $2.3) so that on hot summer afternoons the ecosystem became a source of CO 2 . Tree canopy CO 2 exchange was calculated as the difference between above and below canopy eddy¯ux. Canopy uptake saturated at $6 mmol CO 2 m À2 s À1 for a PPFD above 500 mmol m À2 s À1 and decreased with increasing D. The optimal stomatal control model of Ma Èkela È et al. (1996) was used as a`big leaf' canopy model with parameter values determined by the non-linear least squares. The model accurately simulated the response of the forest to light, saturation de®cit and drought. The precision of the model was such that the daily pattern of residuals between modeled and measured forest exchange reproduced the component storagē ux. The model and independent leaf-level measurements suggest that the marginal water cost of plant C gain in Larix gmelinii is more similar to values from deciduous or desert species than other boreal forests. During the middle of the summer, the L. gmelinii forest ecosystem is generally a net sink for CO 2 , storing $0.75 g C m À2 d À1. Published by Elsevier Science B.V.
Mineral nutrition of Norway spruce (Picea abies (L.) Karst.) and beech (Fagus sylvatica L.) was investigated along a transect extending from northern Sweden to central Italy. Nitrogen (N) concentrations of needles and leaves in stands growing on acid soils did not differ significantly between central Italy and southern Sweden (1.0 +/- 0.1 mmol N g(-1) for needles and 1.9 +/- 0.14 mmol N g(-1) for leaves). In both species, foliar N concentrations were highest in Germany (1.2 mmol N g(-1) for needles and 2.0 mmol N g(-1) for leaves) and decreased by 50% toward northern Sweden (0.5 mmol N g(-1)). Both species showed constant S/N and P/N ratios along the transect. Calcium, K and Mg concentrations generally reflected local soil conditions; however, Mg concentrations reached deficiency values in Germany. Leaf area per unit dry weight varied significantly along the transect with lowest values for Norway spruce recorded in northern Sweden and Italy (3.4 m(2) kg(-1)) and a maximum in central Europe (4.7 m(2) kg(-1)). A similar pattern was observed for beech. Despite the low variation in foliar N concentrations on the large geographic scale, local and regional variations in N concentrations equalled or exceeded the variation along the entire continental transect. Furthermore, nutrient contents (i.e., nutrient concentration x dry weight per needle or leaf) showed a greater variation than nutrient concentrations along the transect. Nitrogen contents of Norway spruce needles reached minimum values in northern Sweden (2.4 micro mol N needle(-1)) and maximum values in Denmark (5.0 micro mol N needle(-1)). The N content of beech leaves was highest in Denmark (242 micro mol N leaf(-1)). At the German site, foliar N content rather than N concentration reflected the seasonal dynamics of foliar growth and N storage of the two species. During foliage expansion, there was an initial rapid increase in N content and a decrease in N concentration. This pattern lasted for about 2 weeks after bud break and was followed by 6 weeks during which dry weight and N content of the foliage increased, resulting in a further decrease in N concentration. During summer, dry weight and N content of mature needles of Norway spruce increased further to reach a maximum in autumn, whereas N concentration remained constant. In spring, reallocation of N from 1- and 2-year-old needles was 1.5 and 1.0 micro mol N needle(-1), respectively. This remobilized N was a major source of N for the development of new needles, which had an N content of 1.5 micro mol N needle(-1) after bud break. The seasonal remobilization of N from old foliage decreased with increasing needle age. Needle N content and dry weight decreased progressively with age (1 micro mol N needle(-1) between age classes 2 and 5), whereas N concentrations remained constant. For Norway spruce, annual stemwood production was correlated with needle N content but not with foliar N concentration or with the total amount of N in the canopy. Interspecific and geographical differences in plant nutrition are d...
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