Nitrogen Fixation in the Canopy of Temperate Forest Trees: A Re-examination

¹⁵N₂ studies and acetylene reduction assays of leaves and shootsof Douglas fir and other forest trees do not confirm previousreports that extensive nitrogen fixation occurs on leaf surfacesand it is concluded that the importance of nitrogen fixationin the canopy of forest trees has been exaggerated. The presenceof nitrogen-fixing bacteria on the leaves of trees is confirmed,however, and they have been identified as Enterobacter agglomerans,Clostridium butyricum and Bacillus sp. Their distribution onleaves is fortuitous since dead oak leaves and artificial leavesbecome colonized to the same extent as living oak leaves. nitrogen fixation, acetylene reduction, Enterobacter agglomerans, Clostridium butyricum, Bacillus sp, Douglas fir, Pseudotsuga menziensii, larch, Larix x oak, Quercus petraea.     

落叶松、水曲柳的氮营养行为及其种间分异

对已郁闭的12年生带状混交林和盆栽苗木的研究表明,落叶松和水曲柳在吸收空间的利用和吸收N素的时间(季节)、数量、形态诸方面都有不同的行为模式,二者存在着显著的种间差异.水曲柳根系发达,与落叶松混交能够保持固有的扎根习性不变,吸收根主要集中在土壤上层(0～20cm);落叶松根系在土壤上层受到水曲柳排斥,吸收根大量向下层(20～60cm)发展.在整个生长季中,两树种都有明显的N吸收峰期,水曲柳峰期出现在6月中下旬,落叶松峰期则出现在7月中下旬,二者是明显错开的.在吸收N素的数量上,水曲柳吸收量大,N利用效率较低;落叶松吸收量相对较小,N利用效率较高(比水曲柳高60.8%).在对N素养分化学形态的偏向选择方面,水曲柳较喜NO₃^--N,落叶松则对NO₃^--N和NH⁺-N无明显的偏好性.     

Landscape history in the subalpine karst region of Moncodeno (Lombardy Prealps, Northern Italy)

In situ larch stumps have been found above the present forest line in the karst region of Moncodeno, Grigna Settentrionale, northern Italy, an area where very few trees currently grow. Samples from living trees at the treeline and in the forest nearby were collected to develop a reference chronology for cross-dating the stumps. The latter span the period between 1218 and 1900, and together with the living trees, which cover the last 150 years, they have led to the development of a 784-year larch tree-ring chronology (1218–2001). Age and location of the stumps reflect past human activity. Deforestation and grazing have intensified the erosion processes in the area. The soil has thinned out and fragmented allowing karst landforms, previously buried by the soil, to emerge. Presently, the rocky outcrops limit the establishment of young larches so that only few scattered trees are currently found.     

The relationship between the growth and mineral composition of the foliage of Japanese larch (Larix leptolepis,Murr.)

Summary Investigations on young Japanese larch trees growing on a heterogeneous site, have revealed significant linear correlations between the height of the trees and concentrations of nitrogen, phosphorus, potassium and ash in the needles, but only doubtfully so in the case of calcium. Analysis of the multiple regression of height on the concentrations of these nutrients suggests that only the contributions made by nitrogen and potassium are significant and a regression equation relating height to the concentration of these two nutrients in the needles has been calculated. The function 123.27% N + 188.69% K – 180.91 provides an estimate of tree height with a standard error of ± 40.9 cm and with a highly significant multiple correlation coefficient of .916. It is argued from these results that, under the conditions of the experiment and the range of concentrations encountered, the height growth of the trees is limited by deficiencies in both nitrogen and potassium.     

Postpollination drop production in hybrid larch is not related to the diurnal pattern of xylem water potential

The hourly production of postpollination drops in the ovules of three hybrid larch trees (Larix x. marschlinsii Coaz) was examined and compared with the measured diurnal pattern of xylem water potential of the same trees under non-stressed conditions during two reproductive seasons. There was no consistent relationship between xylem water potential and ovular drop production in hybrid larch. Individual trees that showed a diurnal drop production in one year did not follow a similar pattern the other year. One tree produced drops in over half of its ovules hourly, regardless of the measured xylem water potential. Variation in number of drops produced per ovule and drop mass was observed between the three trees both years, and between years for two of the trees. This variation was not in response to xylem water potential or environmental conditions. Here we report the first simultaneous measurements of branch water status and the production of ovular secretions for any conifer. We conclude that the postpollination drop production of hybrid larch is not regulated by a tree's overall water status, but is under the control of localized structures such as the cone or the ovule.     

Recent drought stress leads to growth reductions in Larix sibirica in the western Khentey,Mongolia

Trends in air temperature and precipitation in the forest‐steppe ecotone of the western Khentey, northern Mongolia were studied and related to stem increment and shoot water relations in Mongolia's most common tree species, Siberian larch (Larix sibirica). The area has been subject to a significant increase of summer temperature and a decrease of summer precipitation during the last 47 years. Tree‐ring width series from >400 larch trees show a strongly decreasing annual increment since the 1940s. The onset of this decrease is independent of the age of the trees and, therefore, can be attributed to the increasing aridity in the 20th century. Simultaneously to the declining annual increment, regeneration of Siberian larch decreased as well; today regeneration is virtually lacking in the larch forests on mountain slopes of the western Khentey. Measurements of shoot water potentials during the growing season exhibited daily minimum water potentials close to the point of zero turgor for extended periods. The drought stress indicated by these results is in line with the current low annual increment. Trees in the forest interior were more severely stressed and grow more slowly than trees at the forest line to steppe. This is attributable to the recent increase in aridity, as the stand density and probably also the trees themselves in the forest interior are adapted to moister conditions, whereas the trees at the forest edge have always been exposed to a more extreme microclimate. The progressing increase in aridity during the 21st century that is predicted for the western Khentey, suggests a future decline of larch forests. A widespread increase of aridity predicted for most parts of the Mongolian forest belt, suggests even a supra‐regional decline of larch.     

Individual sensitivity of Larix sibirica L. in open woodland of the Shira forest-steppe

Some indices of individual variability in annual growth of Siberian larch in open woodlands of Khakassia are compared. In the Shira forest-steppe the Siberian larch exhibits high sensitivity of annual growth and diverse responses to changes in vegetative conditions. Stabilizing selection maintains dominance of trees with moderately sensitive annual growth. Such trees are the most vital. It has been shown that the adaptive norm of population’s response and the individual variability of annual growth positively correlate with the total annual precipitation. The sensitivity index of annual growth is a better indicator of the genetic structure of the population than the coefficient of variability of annual growth indices.     

Water shortage affects the water and nitrogen balance in Central European beech forests

Whilst forest policy promotes cultivation and regeneration of beech dominated forest ecosystems, beech itself is a highly drought sensitive tree species likely to suffer from the climatic conditions prognosticated for the current century. Taking advantage of model ecosystems with cool-moist and warm-dry local climate, the latter assumed to be representative for future climatic conditions, the effects of climate and silvicultural treatment (different thinning regimes) on water status, nitrogen balance and growth parameters of adult beech trees and beech regeneration in the understorey were assessed. In addition, validation experiments with beech seedlings were carried out under controlled conditions, mainly in order to assess the effect of drought on the competitive abilities of beech. As measures of water availability xylem flow, shoot water potential, stomatal conductance as well as delta (13)C and delta (18)O in different tissues (leaves, phloem, wood) were analysed. For the assessment of nitrogen balance we determined the uptake of inorganic nitrogen by the roots as well as total N content and soluble N compounds in different tissues of adult and young trees. Retrospective and current analysis of delta (13)C, growth and meteorological parameters revealed that beech growing under warm-dry climatic conditions were impaired in growth and water balance during periods with low rain-fall. Thinning affected water, N balance and growth mostly of young beech, but in a different way under different local climatic conditions. Under cool, moist conditions, representative for the current climatic and edaphic conditions in beech forests of Central Europe, thinning improves nutrient and water status consistent to published literature and long-term experience of forest practitioners. However, beech regeneration was impaired as a result of thinning at higher temperatures and under reduced water availability, as expected in future climate.     

Heterosis in hybrid larch (Larix decidua x leptolepis)

Summary Individual 33-year-old forest trees of the deciduous conifer speciesLarix decidua, Larix leptolepis andLarix decidua x leptolepis were investigated with respect to the phenomenon of stem heterosis in hybrid larch; the first part of this study compares the gas exchange responses of leaves. CO₂ assimilation per leaf area was similar in the three larch species, but on a dry weight basis the nitrogen content of the needles and maximum CO₂ assimilation rate (A_max) were slightly higher in the hybrid. This increase was accompanied by a higher protein content than in the Japanese and a lower specific leaf weight than in the European larch. All three species were similar in terms of the photosynthetic nitrogen use and stomatal conductance atA _max. The similar slopes of the area-related steady-state responses of gas exchange against irradiance, evaporative demand and internal CO₂ concentration led to similar rates of CO₂ uptake under ambient conditions. The natural combinations and variability of the environmental factors also reduced the small dry weight-related difference inA _max between hybrid larch and the parent species, such that all trees achieved similar daily carbon gains. Thus, the ecological significance of small interspecific differences in the metabolism of leaves has very little effect under the natural habitat conditions of a temperate climate. The second part of the study will investigate the effect of growth characteristics on the heterosis of hybrid larch.     

The influence of micrografting in vitro on tissue culture behavior and vegetative propagation of old European larch trees

The influence of a new micrografting method in vitro was tested by using long shoot meristems of adult 140 year old European larch trees grafted onto larch seedling rootstocks in sterile peat pellets. Five months after grafting and transfer of grafts to ex vitro conditions, new shoots from sprouting scions were re-established into tissue culture. The propagation behavior of shoots derived from these explants was compared with shoots established via bud culture from the same donor trees. Shoot explants from micrografts multiplied more rapidly than shoots explants from the original adult donor trees. Rooting experiments with cuttings from six micrografted clones resulted in 49.9 ± 11.9% rooting. Cuttings from donor trees invariably showed no root formation. The results confirm a rejuvenating influence of micrografting in larch.     

Plasticity in gas‐exchange physiology of mature Scots pine and European larch drive short‐ and long‐term adjustments to changes in water availability

Adjustment mechanisms of trees to changes in soil‐water availability over long periods are poorly understood, but crucial to improve estimates of forest development in a changing climate. We compared mature trees of Scots pine (Pinus sylvestris) and European larch (Larix decidua) growing along water‐permeable channels (irrigated) and under natural conditions (control) at three sites in inner‐Alpine dry valleys. At two sites, the irrigation had been stopped in the 1980s. We combined measurements of basal area increment (BAI), tree height and gas‐exchange physiology (Δ¹³C) for the period 1970–2009. At one site, the Δ¹³C of irrigated pine trees was higher than that of the control in all years, while at the other sites, it differed in pine and larch only in years with dry climatic conditions. During the first decade after the sudden change in water availability, the BAI and Δ¹³C of originally irrigated pine and larch trees decreased instantly, but subsequently reached higher levels than those of the control by 2009 (15 years afterwards). We found a high plasticity in the gas‐exchange physiology of pine and larch and site‐specific responses to changes in water availability. Our study highlights the ability of trees to adjust to new conditions, thus showing high resilience.     

Influence of Gmelin larch trees on postfire restoration of forest communities in the permafrost zone of Central Siberia

This article shows the phytocoenotic role of individual postfire trees of the Gmelin larch under conditions of the permafrost zone of Central Siberia. On the basis of the vital state of the trees, the limits of their influence on the recovery of the species of the lower layers of phytocenosis are defined and the formation of the new stand generation is shown.     

Comparative Reaction of Larch (<Emphasis Type="Italic">Larix sibirica</Emphasis> Ledeb.) Radial Increment on Climate Change in the Forest Steppe and Highlands of Southern Siberia

The influence of climate changes on larch (Larix sibirica Ledeb.) radial increment under conditions of a limited (forest steppe) and sufficient (high-altitude Kuznetsk Alatau, floodplain stands) humidification is considered. The relationship between growth index of larch trees (N = 257) and ecological and climatic variables is analyzed. In the forest steppe, with the onset of warming, a decrease in the aridity of the climate, an increase in the duration of the growing season (1980s), and an increase in the larch growth index followed by its depression in the 1990s have been observed. Radial-increment depression is caused by an increase in vapor-pressure deficit and arid climate due to a rising air temperature. In the 2000s, radial-increment fluctuations with average values not exceeding those before the beginning of current climate warming period occurred. In the highlands, since the 1970s, there has been a general increase in the larch radial increment closely associated with the main limiting factor of growth—air temperature. At the same time, in arid years, the radial-increment depression of larch trees in highland and floodplain larch forests is also noted. When implementing “hard” climate scenarios (RCP 6.0 and RCP 8.5), it is likely that the larch growth index in a forest steppe will decrease further and its increase in areas of sufficient moisture will be observed.     

Contrasting responses of seedling and sapling densities to livestock density in the Mongolian forest-steppe

The past and present regeneration of Siberian larch (Larix sibirica) was studied in the forest-steppe of the Mongolian Altai, an area which has experienced an increase in annual mean temperature by 2.1 °C since 1940 and is subjected to grazing by mixed herds of livestock owned by pastoral nomads. Past regeneration was reconstructed from tree rings and present regeneration was analyzed by surveying seedlings, sapling-sized trees and the occurrence of viable larch seeds in the soil seed bank. Forest regeneration occurred throughout the twentieth and the early twenty-first centuries in the forest interiors, but ceased after the late 1970s at forest edges. The density of sapling-sized larch trees decreased with livestock density linearly in the interior and exponentially at the edge. Most sapling-sized trees had visible damage from livestock browsing, which also manifested in wood-anatomical anomalies. By contrast, the densities of 1- and 2-year old seedlings increased with livestock density at the forest edge, suggesting that seedlings in this habitat benefitted from the reduction in competition intensity due to livestock grazing. This relationship also suggests that larch seedlings, in contrast to sapling-sized trees, were avoided by the livestock, as otherwise removal by the herbivores should have counteracted the promotion due to reduced competition. Near-consistency of the correlations of total livestock and goat densities with sapling and seedling densities suggests that the control of larch regeneration is primarily a function of goat density, which have tripled in the Mongolian livestock during the past 20 years for economic reasons.     

Light,nitrogen, and phosphorus limitation of edaphic algae in a delaware salt marsh

The effects on edaphic algae associated with a pure stand of the cord grass, Spartina alterniflora Loisel of manipulating light intensity and additions of inorganic nitrogen and phosphorus as fertilizers to the marsh surface have been investigated for one year. The standing crop of edaphic algae as measured by chlorophyll a production was limited only by phosphorus supplies during fall and winter, by both phosphorus and nitrogen in spring, and only by nitrogen during the summer. Since the responses were in phase with the seasonal fluctuations in the concentration of nitrogen and phosphorus, it is concluded that the flood tide is the major source of nitrogen and phosphorus compounds for edaphic algal growth. Reduction in the quantity of light reaching the edaphic algae by Spartina cover is always a limiting factor for the standing crop. A gradient in the composition of the algal flora is directly related to light intensity, and indicates that this factor determines the relative contribution of diatoms and filamentous algae to the community. The interaction of light intensity and nutrients in regulating the production of edaphic algae and cord grass on the marsh under study over a yearly cycle has also been investigated.     

Wood transformation in dead-standing trees in the forest-tundra of Central Siberia

Changes in the composition of wood organic matter in dead-standing spruce and larch trees depending on the period after their death have been studied in the north of Central Siberia. The period after tree death has been estimated by means of cross-dating. The results show that changes in the composition of wood organic matter in 63% of cases are contingent on tree species. Wood decomposition in dead-standing trees is accompanied by an increase in the contents of alkali-soluble organic compounds. Lignin oxidation in larch begins approximately 80 years after tree death, whereas its transformation in spruce begins not earlier than after 100 years. In the forest-tundra of Central Siberia, the rate of wood organic matter transformation in dead-standing trees is one to two orders of magnitude lower than in fallen wood, which accounts for their role as a long-term store of carbon and mineral elements in these ecosystems.     

Tree-ring growth of Gmelin larch under contrasting local conditions in the north of Central Siberia

While the forest-tundra zone in Siberia, Russia has been dendroclimatologically well-studied in recent decades, much less emphasis has been given to a wide belt of northern taiga larch forests located to the south. In this study, climate and local site conditions are explored to trace their influence on radial growth of Gmelin larch (Larix gmelinii (Rupr.) Rupr.) trees developed on permafrost soils in the northern taiga. Three dendrochronological sites characterized by great differences in thermo-hydrological regime of soils were established along a short (ca. 100 m long) transect: on a river bank (RB), at riparian zone of a stream (RZ) and on a terrace (TER). Comparative analysis of the rate and year-to-year dynamics of tree radial growth among sites revealed considerable difference in both raw and standardized tree-ring width (TRW) chronologies obtained for the RZ site, characterized by shallow soil active layer depth and saturated soils. Results of dendroclimatic analysis indicated that tree-ring growth at all the sites is mostly defined by climatic conditions of a previous year and precipitation has stronger effect on TRW chronologies in comparison to the air temperatures. Remarkably, a great difference in the climatic response of TRW chronologies has been obtained for trees growing within a very short distance from each other. The positive relation of tree-ring growth with precipitation, and negative to temperature was observed in the dry site RB. In contrary, precipitation negatively and temperature positively influenced tree radial growth of larch at the water saturated RZ. Thus, a complicate response of northern Siberian larch forest productivity to the possible climate changes is expected due to great mosaic of site conditions and variability of environmental factors controlling tree-ring growth at different sites. Our study demonstrates the new possibilities for the future dendroclimatic research in the region, as various climatic parameters can be reconstructed from tree-ring chronologies obtained for different sites.     

Niche breadth and range area in North American trees

Identifying factors affecting species distribution is a longstanding goal in ecology and evolution that is accentuated by our need to anticipate climate change impacts. We sought to test whether any phylogenetic effect can be detected in either the environmental characteristics or range attributes of North American trees, and to explore the existence of a general interspecific pattern in the environmental factors influencing species range size. To do so we tested prevailing hypotheses relating climatic and edaphic characteristics to species range size in the North American arboflora (n = 598), using spatial null models to test for the relevance of observed patterns. We found that interspecific variation in the range area of North American trees is strongly related to the environmental regimes characteristic of the species range. Linear models and phylogenetic regressions involving six environmental characteristics explained 83% of the variance in species range area, and affirmed a positive relationship between niche breadth and range size. Tree species that can tolerate a larger variability in local climatic conditions, deal with harsher edaphic conditions, and weak levels of environmental energy tend to have larger range area; this can account for the greater geographic range of species at higher latitudes, the Rapoport effect. There is a significant phylogenetic signal for both range area and limits in North American trees, and for climatic limits, but not for energy or edaphic characteristics associated with species range. These findings highlight the possibility that species with small geographic ranges may be more sensitive to the effects of climate change.     

Tree-ring analysis of larch sawfly (Pristiphora erichsonii (Hartig)) defoliation events and hydrological growth suppression in a peatland

Insect defoliation events are a major forest disturbance in the boreal forest in Canada. Reconstructing previous events are crucial to understanding natural factors that lead to insect defoliation periods, improving our ability to predict future infestations and increasing the reliability of forest management plans and pest control programs. Researchers have often been limited in their ability to draw accurate conclusions regarding the history of larch sawfly (Pristiphora erichsonii (Hartig)) infestation events in North America. It is well known that floods can affect survival of larch sawfly populations, as well as suppress radial growth of eastern larch (Larix laricina (Du Roi) K. Koch) trees. Eastern larch often inhabits peatlands where high water-table levels can lead to a decrease in tree-ring widths. Water-table level increases result in similar radial-growth patterns to when trees are defoliated by larch sawfly, making accurate diagnoses of larch sawfly events a challenge. This fact becomes more accentuated when non-host species used for standard dendroecological analyses (often black spruce (Picea mariana (Mill.) Britton) present an opposite relationship to water-table variability, enhancing chances of drawing erroneous conclusions when using program OUTBREAK.This paper introduces a novel case study to advance the understanding of the complex landscapes where these larch sawfly events are taking place. It stresses the necessity of using upland sites less influenced by the hydrological regime to accurately reconstruct larch sawfly defoliation events. We found that where investigations are taking place solely in peatlands or lowlands, and where they are heavily influenced by hydrological conditions, the use of other lines of evidence such as pale rings and long-term hydrological records are crucial to understand the dynamics in the system. In our case study, program OUTBREAK identified five defoliation events in a peatland and upland site; however, hydrological data and pale ring evidence indicated that the last event identified in the peatland site by the host/non-host analysis was confounded by hydrological growth suppression.     

Comparison of foliar water use efficiency among 17 provenances of Larix gmelinii in the Mao’ershan area

 研究环境变化下的树木水分利用效率对探讨森林生态系统碳水耦联关系及其对气候变化的响应和适应对策具有重要意义。落叶松(Larix gmelinii)为我国北方森林的建群种之一。将水热条件不同的17个种源落叶松种植在帽儿山森林生态系统研究站的同质园内30年后, 测定其针叶水分利用效率(WUE)及其相关因子。结果表明: WUE、净光合速率(P_n)、蒸腾速率(T_r)、气孔导度(G_s)、比叶面积(SLA)和叶片氮含量(N_L)均存在显著的种源差异(p < 0.05)。WUE和G_s呈显著指数相关关系, 当G_s < 0.2 时WUE随G_s的增大而明显增大, 而当G_s > 0.2时WUE趋于稳定。WUE和SLA及N_L分别呈线性负相关和正相关关系, 且随种源原地的干燥度指数(AI)的增大其相关性明显增强。WUE和种源原地年平均气温、平均年降水量及AI分别呈线性负相关、负相关和正相关关系, 并且相关系数依次增大; T_r则仅和种源原地年平均气温呈线性正相关关系, 而P_n和种源原地AI呈线性正相关关系。不同种源落叶松由于对种源原地环境条件的适应而存在针叶结构和生理特征的显著差异, 并因此引起针叶水分利用效率的差异。