Photosynthetic temperature adaptation of Pinus cembra within the timberline ecotone of the Central Austrian Alps

? Temperature is suggested to determine the upper limit of tree life. Therefore, future climate warming may be of importance for tree distribution within the European Alps, where low temperatures limit carbon metabolism.

? We focused on the effects of air and soil temperature on net photosynthesis (P _n) of Pinus cembra an evergreen climax species of the timberline ecotone of the Central Austrian Alps. Light response and temperature response curves were estimated along an altitudinal gradient ranging from the forest limit up to the krummholz limit in both summer and fall.

? In general, P _n was significantly lower in fall as compared to summer. Nevertheless, independent from season mean P _n values tended to increase with elevation and were positively correlated with root zone temperatures. The specific leaf area by contrast declined with increasing elevation. Furthermore, the temperature optimum of net photosynthesis declined with increasing elevation and was positively correlated with the mean maximum air temperature of the 10 days prior the date of measurement.

? Thus, our findings appear to reflect a long-term adaptation of the photosynthetic apparatus of Pinus cembra to the general temperature conditions with respect to elevation combined with a short term acclimation to the prevailing temperature regime.

     

Spatial and seasonal variations in mobile carbohydrates in Pinus cembra in the timberline ecotone of the Central Austrian Alps

To test whether the altitudinal limit of tree growth is determined by carbons shortage or by a limitation in growth we investigated non structural carbohydrates and their components starch and total soluble sugars in Pinus cembra trees along an elevational gradient in the timberline ecotone of the Central Austrian Alps. NSC contents in needles, branches, stems, and coarse roots were measured throughout an entire growing season. At the tissue level NSC contents were not significantly more abundant in treeline trees as compared to trees at lower elevations. Along our 425 m elevational transect from the closed forest to the treeline we failed to find a stable elevational trend in the total NSC pool of entire trees and observed within season increases in the tree's NSC pool that can be attributed to an altitudinal increase in leaf mass as needles contained the largest NSC fraction of the whole tree NSC pool. Furthermore, whole tree NSC contents were positively correlated with net photosynthetic capacity. Although our observed NSC characteristics do not support the hypothesis that tree life at their upper elevational limit is determined by an insufficient carbon balance we found no consistent confirmation for the sink limitation hypothesis.     

Seasonal variation of soil respiration rates in a secondary forest and agroforestry systems

Agroforestry systems are widely practiced in tropical forests to recover degraded and deforested areas and also to balance the global carbon budget. However, our understanding of difference in soil respiration rates between agroforestry and natural forest systems is very limited. This study compared the seasonal variations in soil respiration rates in relation to fine root biomass, microbial biomass, and soil organic carbon between a secondary forest and two agroforestry systems dominated by Gmelina arborea and Dipterocarps in the Philippines during the dry and the wet seasons. The secondary forest had significantly higher (p < 0.05) soil respiration rate, fine root biomass and soil organic matter than the agroforestry systems in the dry season. However, in the wet season, soil respiration and soil organic matter in the G. arborea dominated agroforestry system were as high as in the secondary forest. Whereas soil respiration was generally higher in the wet than in the dry season, there were no differences in fine root biomass, microbial biomass and soil organic matter between the two seasons. Soil respiration rate correlated positively and significantly with fine root biomass, microbial biomass, and soil organic C in all three sites. The results of this study indicate, to some degree, that different land use management practices have different effects on fine root biomass, microbial biomass and soil organic C which may affect soil respiration as well. Therefore, when introducing agroforestry system, a proper choice of species and management techniques which are similar to natural forest is recommended.     

Carbon dioxide gas exchange of cembran pine (Pinus cembra) at the alpine timberline during winter

Winter CO(2) gas exchange of the last three flushes of cembran pine (Pinus cembra L.) was studied under ambient conditions at the alpine timberline, an ecotone with strong seasonal changes in climate. During the coldest months of the year, December to March, gas exchange was almost completely suppressed and even the highest irradiances and temperatures did not cause a significant increase in net photosynthesis compared to spring and fall. In general, daily CO(2) balance was negative between December and March except during extended warm periods in late winter. However, because twig respiration was also reduced to a minimum during the December-March period, daily carbon losses were minimal. Total measured carbon loss during the winter months was small, equalling the photosynthetic production of one to two warm days in spring or summer when average air temperature was above 6 degrees C.     

The carbon budget of an adult <Emphasis Type="Italic">Pinus cembra</Emphasis> tree at the alpine timberline in the Central Austrian Alps

We investigated carbon (C) uptake and respiratory losses of an adult Pinus cembra tree at the alpine timberline throughout an entire year by means of an automated, multiplexing gas exchange system. These chamber measurements were then combined with biomass data for scaling up the C budget to the tree level. Integrated over an entire year, the cumulative C gain of the tree under study was 23.5 kg of C in 2002. The daily C balance was negative for 5 months and the estimated total wintertime respiratory losses were 9% of the amount of C fixed during the growing season. The total annual C loss of the tree consumed 55% of the annual net C gain and the remaining surplus was stored in new tissues (36%) and used for fine root growth (9%). Thus, the overall C budget of P. cembra at the upper timberline is balanced fairly well, although the C sink strength in fine roots is strongly limited owing to low root zone temperatures when compared to conifers at lower elevation sites.     

Winter frost resistance of Pinus cembra measured in situ at the alpine timberline as affected by temperature conditions

Winter frost resistance (WFR), midwinter frost hardening and frost dehardening potential of Pinus cembra L. were determined in situ by means of a novel low-temperature freezing system at the alpine timberline ecotone (1950 m a.s.l., Mt Patscherkofel, Innsbruck, Austria). In situ liquid nitrogen (LN?)-quenching experiments should check whether maximum WFR of P. cembra belonging to the frost hardiest conifer group, being classified in US Department of Agriculture climatic zone 1, suffices to survive dipping into LN? (-196 °C). Viability was assessed in a field re-growth test. Maximum in situ WFR (LT??) of leaves was <-?75 °C and that of buds was less (-70.3 °C), matching the lowest water contents. In midwinter, in situ freezing exotherms of leaves, buds and the xylem were often not detectable. Ice formed in the xylem at a mean of -2.8 °C and in leaves at -3.3 °C. In situ WFR of P. cembra was higher than that obtained on detached twigs, as reported earlier. In situ LN?-quenching experiments were lethal in all cases even when twigs of P. cembra were exposed to an in situ frost hardening treatment (12 days at -20 °C followed by 3 days at -50 °C) to induce maximum WFR. Temperature treatments applied in the field significantly affected the actual WFR. In January a frost hardening treatment (21 days at -20 °C) led to a significant increase of WFR (buds: -62 °C to <-?70 °C; leaves: -59.6 °C to -65.2 °C), showing that P. cembra was not at its specific maximum WFR. In contrast, simulated warm spells in late winter led to premature frost dehardening (buds: -32.6 °C to -10.2 °C; leaves: -32.7 to -16.4 °C) followed by significantly earlier bud swelling and burst in late winter. Strikingly, both temperature treatments, either increased air temperature (+10.1 °C) or increased soil temperature (+6.5 °C), were similarly effective. This high readiness to frost harden and deharden in winter in the field must be considered to be of great significance for future winter survival of P. cembra. Determination of WFR in field re-growth tests appears to be a valuable tool for critically judging estimates of WFR obtained on detached twigs in an ecological context.     

Influence of climate on radial growth of Pinus cembra within the alpine timberline ecotone

Radial growth variability and response to interannual climate variation of Cembran pine (Pinus cembra L.) were studied in the timberline ecotone on Mt. Patscherkofel (2246 m a.s.l.). The study area, which is in the inner alpine dry region of the Central Austrian Alps, is characterized by a continental climate with minimum precipitation in winter (about 150 mm during December-February) and frequent occurrence of warm dry winds (F?hn) in early spring. The hypothesis that spatial and temporal variability of radial growth is caused by site-related differences in sensitivity to winter stress (i.e., desiccation) was examined by applying dendroclimatological techniques. Ordination methods applied to tree ring time series revealed that spatial variability in radial growth is influenced by the local site factors elevation and slope aspect. Growth-climate relationships were explored using Pearson product-moment correlation coefficients and multiple regression analysis. Radial growth at the timberline was positively correlated with temperature in July and was also strongly correlated with mild temperatures in the previous autumn and high precipitation in winter (January-March). At the tree line, temperatures in the previous autumn and precipitation in late winter (March) also controlled radial growth, whereas July temperature was not significantly correlated with ring width. Because previous autumn temperature and winter precipitation were the main growth-determining factors at the timberline and the tree line, and both of these climate variables are known to influence susceptibility of trees to winter stress, the results support the working hypothesis. Analysis of climatic conditions in extreme growth years confirmed the high sensitivity of tree ring growth to precipitation in late winter (March) at the tree line plots. Furthermore, extent of growth reduction and release varied spatially and temporarily, with south- and west-facing stands showing a higher sensitivity to climate variation in the most recent decade (1990s) than the north-facing stand. This aspect-related change in sensitivity to climate may be associated with effects of climate warming on cambial activity.     

Effects of atmospheric and climate change at the timberline of the Central European Alps

? This review considers potential effects of atmospheric change and climate warming within the timberline ecotone of the Central European Alps. After focusing on the impacts of ozone (O₃) and rising atmospheric CO₂ concentration, effects of climate warming on the carbon and water balance of timberline trees and forests will be outlined towards conclusions about changes in tree growth and treeline dynamics.

? Presently, ambient ground-level O₃ concentrations do not exert crucial stress on adult conifers at the timberline of the Central European Alps. In response to elevated atmospheric CO₂ Larix decidua showed growth increase, whereas no such response was found in Pinus uncinata. Overall climate warming appears as the factor responsible for the observed growth stimulation of timberline trees.

? Increased seedling re-establishment in the Central European Alps however, resulted from invasion into potential habitats rather than upward migration due to climate change, although seedlings will only reach tree size upon successful coupling with the atmosphere and thus loosing the beneficial microclimate of low stature vegetation.

? In conclusion, future climate extremes are more likely than the gradual temperature increase to control treeline dynamics in the Central European Alps.

     

Effects of the lack of forest management on spatiotemporal dynamics of a subalpine Pinus cembra forest

Knowledge about the stand structure and dynamics of subalpine forests is crucial to preserve their multifunctionality. In the present study, we reconstructed the spatiotemporal dynamics of a subalpine Pinus cembra forest in the eastern Italian Alps in response to natural disturbances and forest management. We adopted a concurrent point pattern, dendroecological and growth dominance (GD) analysis. We mapped and measured all trees of Pinus cembra and Larix decidua in a 1?ha plot. We analyzed intra- and interspecific spatial patterns and spatial autocorrelation of tree size and age. We explored establishment dynamics and shifts in competition by analyzing growth suppression/release patterns and GD trends. Results showed a clumped, uneven-aged, multilayered structure where pine was dominant. The synergic action of ecological and human-induced factors is discussed to explain the prevalence of pine over time. Spatial pattern and autocorrelation analyses suggest a different colonization strategy of the two species, in which pine established after small-scale perturbations and experienced a stronger inter- and intra-specific competition. The interruption of tree establishment and shift in GD toward large trees resulting from the lack of forest management are the most important findings of this research. This highlights the importance of an active management to avoid the homogenization of the forest structure that is generally associated with a reduction in biodiversity and protective ability of forests.     

Seasonal dynamics of soil carbon dioxide efflux and simulated rhizosphere respiration in a beech forest

Respiration of the rhizosphere in a beech (Fagus sylvatica L.) forest was calculated by subtracting microbial respiration associated with organic matter decomposition from daily mean soil CO2 efflux. We used a semi-mechanistic soil organic matter model to simulate microbial respiration, which was validated against "no roots" data from trenched subplots. Rhizosphere respiration exhibited pronounced seasonal variation from 0.2 g C m(-2) day(-1) in January to 2.3 g C m(-2) day(-1) in July. Rhizosphere respiration accounted for 30 to 60% of total soil CO2 efflux, with an annual mean of 52%. The high Q10 (3.9) for in situ rhizosphere respiration was ascribed to the confounding effects of temperature and changes in root biomass and root and shoot activities. When data were normalized to the same soil temperature based on a physiologically relevant Q10 value of 2.2, the lowest values of temperature-normalized rhizosphere respiration were observed from January to March, whereas the highest value was observed in early July when fine root growth is thought to be maximal.     

长白山白桦林地土壤呼吸的季节变化

2004年5月至9月,研究了长白山白桦林土壤呼吸以及根系呼吸对土壤呼吸的贡献随土壤温度和土壤湿度的季节变化,研究结果表明：土壤总呼吸、断根土壤呼吸和根系呼吸在生长季内有相似的季节变化趋势,夏季潮湿而且温度较高,呼吸速率也较高,春季和秋季温度较低,呼吸速率也较低。2004年5月至9月,土壤总呼吸、断根土壤呼吸和根系呼吸的平均值分别为4.44,2.30和2.14μmol&#183;m^-2s^-1,三者与土壤温度均呈指数相关,与土壤湿度呈线性相关,三者的Q10值分别为2.82,2.59和3.16,这与其他学者的结果相似。根系呼吸是土壤呼吸的一个重要组成部分,2004年5月至9月,根系呼吸对土壤总呼吸的贡献在29.3～58.7%之间。根据Q10模型估算的土壤总呼吸、断根土壤呼吸和根系呼吸的全年平均值分别为1.96、1.08和0.87μmol&#183;m^-2s^-1,即741.73、408.71和329.24gC&#183;m^-2&#183;a^-1,全年根系对土壤总呼吸的贡献为44.4%。土壤呼吸和土壤温度之间的关系模型是了解和预测长白山白桦林生态系统潜在的随森林管理和气候变化而变化的有用工具。     

No diurnal variation in rate or carbon isotope composition of soil respiration in a boreal forest

Characterization of soil respiration rates and delta(13)C values of soil-respired CO(2) are often based on measurements at a particular time of day. A study by Gower et al. (2001) in a boreal forest demonstrated diurnal patterns of soil CO(2) flux using transparent measurement chambers that included the understory vegetation. It is unclear whether these diurnal patterns were solely the result of photosynthetic CO(2) uptake during the day by the understory or whether there were underlying trends in soil respiration, perhaps driven by plant root allocation, as recently demonstrated in Mediterranean oak savannah. We undertook intensive sampling campaigns in a boreal Picea abies L. Karst. forest to investigate whether diurnal variations in soil respiration rate and stable carbon isotope ratio (delta(13)C) exist in this ecosystem when no understory vegetation is present in the measurement chamber. Soil respiration rates and delta(13)C were measured on plots in which trees were either girdled (to terminate the fraction of soil respiration directly dependent on recent photosynthate from the trees), or not girdled, every 4 h over two 48-hour cycles during the growth season of 2004. Shoot photosynthesis and environmental parameters were measured concurrently. No diurnal patterns in soil respiration rates and delta(13)C were observed in either treatment, despite substantial variations in climatic conditions and shoot photosynthetic rates in non-girdled trees. Consequently, assessment of daily soil respiration rates and delta(13)C in boreal forest systems by single, instantaneous daily measurements does not appear to be confounded by substantial diurnal variation.     

Heterotrophic respiration causes seasonal hysteresis in soil respiration in a warm-temperate forest

To assess the effect of changes in organic litter stock on seasonal changes in heterotrophic respiration (R _H), soil respiration (R _S), and total ecosystem respiration (R _E), we measured seasonal changes in leaf litter respiration (R _LL) by the chamber method and estimated the seasonal change in total R _H using the RothC model in a warm-temperate mixed deciduous?Cevergreen forest in Japan. Both R _E and R _S had seasonal hysteresis and were higher in spring than at the same temperature during autumn. Under warm and humid conditions, the rate of decomposition of newly supplied leaf litter in one?year was high (60% loss). Consequently, R _LL and R _H were higher in spring after leaf drop, when more fresh material was available, than in autumn. In this study, 42 and 88% of the difference in R _E and R _S between spring and autumn (soil temperature 16?C18°C) could be accounted for by the difference in R _H, respectively, and 71% of the difference in R _H could be accounted for by the difference in R _LL. This study showed that seasonal changes in heterotrophic respiration (R _LL and R _H) could be a major factor in the seasonal hysteresis of R _E and R _S.     

Sap flow characteristics and whole-tree water use of Pinus cembra across the treeline ecotone of the central Tyrolean Alps

Whole-tree water use of nine Pinus cembra trees was estimated in the treeline ecotone of the Central Austrian Alps. Sap flow density using Granier-type thermal dissipation probes and environmental parameters was monitored along an elevational gradient from the forest limit up to treeline and finally mediating the krummholz limit throughout two growing seasons. Normalized sap flow density (Q _s) was significantly correlated with solar radiation (R _s) and vapor pressure deficit (D) throughout the treeline ecotone. Multiple regression analysis indicated that at the forest limit and at treeline, D had a similar effect on Q _s than R _s. At the krummholz limit by contrast, D had a greater effect on Q _s than R _s due to partially stomatal closure and wind-induced clustering of the needles, which impaired their response to available irradiance. Whole-tree water use scaled to crown surface area estimated for an entire growing (172 days) declined from 449 mm at the forest limit to 274 mm at treeline and was 251 mm at the krummholz limit, which is within the values estimated for other European forest ecosystems. Nevertheless, the observation above the forest limit in the central Tyrolean Alps tree transpiration scaled to crown surface area is comparable to the water use of adjacent low-stature vegetation, which should also be taken into account when forecasting potential effects of global change on the water balance of the treeline ecotone.     

温度对杉木林土壤呼吸的影响(英文)

Soil samples collected from the surface soil (0(10 cm) in an 88-year-old Chinese fir (Cunninghamia lanceolata) forest in Nanping, Fujian, China were incubated for 90 days at the temperatures of 15°C, 25°C and 35°C in laboratory. The soil CO2 evolution rates were measured at the incubation time of 2, 5, 10, 15, 20, 25, 30, 35, 40, 50, 60, 70, 80 and 90 days. The results showed that CO2 evolution rates of soil samples varied significantly with incubation time and temperature during the incubation period. Mean CO2 evolution rate and cumulative amount of CO2 evolution from soil were highest at 35°C, followed by those at 25°C, and 15°C. Substantial differences in CO2 evolution rate were found in Q10 values calculated for the 2nd and 90th day of incubation. The Q10 value for the average CO2 evolution rate was 2.0 at the temperature range of 15-25°C, but it decreased to 1.2 at 25- 35°C. Soil CO2 evolution rates decreased with the incubation time. The cumulative mineralized C at the end of incubation period (on the 90th day) was less than 10% of the initial C amounts prior to incubation.     

Seasonal variation in ethylene concentration in the wood of Pinus sylvestris L

Ethylene concentrations were determined in gas samples extracted from sealed holes made in the sapwood and heartwood of stems of 70-100-year-old Scots pine (Pinus sylvestris L). Gas could be collected from the heartwood holes by lowering the pressure by means of a large syringe. However, attempts to extract gas from air spaces in the sapwood with the same technique failed, presumably because of lack of an interconnected system of gas-filled canals. High ethylene concentrations, usually in excess of 1 ppm, accumulated in the sapwood holes within one day after sealing. Ethylene concentrations in the sapwood rose to 3-7 ppm during the growing season, and decreased to 0.1-0.3 ppm during the winter. In response to extreme drought, sapwood ethylene concentration increased to 30 ppm, followed by a rapid decrease after the onset of rain. Ethylene concentrations in gas samples from the heartwood were consistently lower than 1 ppm. The lowest values, about 0.1 ppm, were found during the autumn and early winter, whereas values around 0.5 ppm were typical from February to August.     

Effects of the extreme drought in 2003 on soil respiration in a mixed forest

We present a field study on the drought effects on total soil respiration (SR_t) and its components, i.e., “autotrophic” (SR_a: by roots/mycorrhizosphere) and “heterotrophic” respiration (SR_h: by microorganisms and soil fauna in bulk soil), in a mature European beech/Norway spruce forest. SR_a and SR_h were distinguished underneath groups of beech and spruce trees using the root exclusion method. Seasonal courses of SR_a and SR_h were studied from 2002 to 2004, with the summer of 2003 being extraordinarily warm and dry in Central Europe. We (1) analyzed the soil temperature (T _s) and moisture sensitivity of SR_a and SR_h underneath both tree species, and (2) examined whether drought caused differential decline of SR_a between spruce and beech. Throughout the study period, SR_a of beech accounted for 45–55% of SR_t, independent of the soil water regime; in contrast, SR_a was significantly reduced during drought in spruce, and amounted then to only 25% of SR_t. In parallel, fine-root production was decreased during 2003 by a factor of six in spruce (from 750 to 130 mg l⁻¹ a⁻¹), but remained at levels similar to those in 2002 in beech (about 470 mg l⁻¹ a⁻¹). This species-specific root response to drought was related to a stronger decline of SR_a in spruce (by about 70%) compared to beech (by about 50%). The sensitivity of SR_a and SR_h to changing T _s and available soil water was stronger in SR_a than SR_h in spruce, but not so in beech. It is concluded that SR_a determines the effect of prolonged drought on the C efflux from soil to a larger extent in spruce than beech, having potential implications for respective forest types. This article belongs to the special issue "Growth and defence of Norway spruce and European beech in pure and mixed stands."     

Temporal variation in nutrient status of a floodplain forest soil

Seasonal variation in nutrient status of a floodplain soil was studied in a silver maple (Acer saccharinum L.) forest in central Illinois. Statistically significant temporal differences were measured in the concentration of NH₄-N, NO₃-N, and extractable P. Patterns of variation in NH₄-N and NO₃-N were related to organic C content and mineralization processes in the soil. Variation in extractable P was affected by flood-induced anaerobic conditions and soil pH. Temporal variation in soil nutrient status was one of the most dynamic features of nutrient cycling in the floodplain forest ecosystem. The potential for seasonal variation in the availability of nutrients should be considered when chemical data are used to characterize forest soil fertility and site productivity. Samples should be collected at several points in time if possible.     

使用Li-6400测定森林土壤呼吸应注意的几个问题

正确使用Li-6400进行森林土壤呼吸测定是保证测定精度的前提.本文以落叶松林为例,对Li-6400使用过程应该注意的问题进行了研究.结果表明:塑料圈至少应该测定前12小时进行设置,以减少土壤CO2涌出效应对呼吸的影响,而且塑料圈插入深度对测定结果影响很大:当塑料圈插入过浅时横向气体扩散及塑料圈不稳定而造成测定时CO2的再涌出效应使土壤呼吸测定值偏高;而当插入深度过深时,切断根系导致根系呼吸下降,最活跃的土壤表层呼吸受塑料圈阻隔导致的土壤微生物呼吸测定值下降,这些现象共同造成显著低估土壤呼吸测定值.此外,在晴天白天大部分时间内目标[CO2]值设定为外界相应时段的[CO2]平均值,可以保证测定误差小于5%,而在清晨和傍晚进行测定时,则应该及时调整目标值为外界[CO2]保证测量准确性.这一测定原则在实际测定中具有较高的实用性.