Physiological responses of birch (Betula pendula) to ozone: a comparison between open-soil-grown trees exposed for six growing seasons and potted seedlings exposed for one season

Physiological responses of 4-year-old potted saplings of an O3-tolerant clone of Betula pendula Roth to short-term ozone (O3) exposure (one growing season) were compared with those of 6-year-old open-soil-grown trees of the same clone fumigated with O3 for six growing seasons. In the 2001 growing season, both groups of plants were exposed to ambient (control) and 1.6x ambient (elevated) O3 concentration under similar microclimatic conditions in a free air O3 exposure facility. Growth, net photosynthesis, stomatal conductance, stomatal density, visible foliar injury, starch and nutrient concentrations, bud formation and differences in O3 responses between lower, middle and upper sections of the canopy were determined. The potted saplings were unaffected by elevated O3 concentration, whereas the open-soil-grown trees showed a 3-38% reduction in shoot growth, a 22% reduction in number of overwintering buds, a 26-65% decrease in autumnal net photosynthesis, 30% and 20-23% reductions in starch and nitrogen concentrations of senescing leaves, respectively, and disturbances in stomatal conductance. The greater O3 sensitivity of open-soil-grown trees compared with potted saplings was a result of senescence-related physiological factors. First, a lower net photosynthesis to stomatal conductance ratio in open-soil-grown trees at the end of the season promoted O3 uptake and decreased photosynthetic gain, leading to the onset of visible foliar injuries. Second, decreased carbohydrate reserves may have resulted in deleterious carry-over effects arising from the reduced formation of over-wintering buds. Finally, the leaf-level O3 load was higher for open-soil-grown trees than for potted saplings because of slower leaf senescence in the trees. Thus, O3 sensitivity in European white birch increases with increasing exposure time and tree size.     

不同混交比例的人工针阔带状混交林生物量变化动态

通过对20世纪80年代营造的针阔混交林生物量调查表明,红松阔叶混交林生物量比红松纯林高4.9%~21%,落叶松阔叶混交林生物量比落叶松纯林高7.6%~21.3%,红松阔叶混交林单株生物量比红松纯林高120%~148%,落叶松阔叶混交林单株生物量比落叶松纯林高108%~151%,在不同混交林型中,灌木层和草本层植物种类丰富、数量多,所以其生物量大于纯林。     

Carbon gain and bud physiology in Populus tremuloides and Betula papyrifera grown under long-term exposure to elevated concentrations of CO2 and O3

Paper birch (Betula papyrifera Marsh.) and three trembling aspen clones (Populus tremuloides Michx.) were studied to determine if alterations in carbon gain in response to an elevated concentration of CO(2) ([CO(2)]) or O(3) ([O(3)]) or a combination of both affected bud size and carbohydrate composition in autumn, and early leaf development in the following spring. The trees were measured for gas exchange, leaf size, date of leaf abscission, size and biochemical characteristics of the overwintering buds and early leaf development during the 8th-9th year of free-air CO(2) and O(3) exposure at the Aspen FACE site located near Rhinelander, WI. Net photosynthesis was enhanced 49-73% by elevated [CO(2)], and decreased 13-30% by elevated [O(3)]. Elevated [CO(2)] delayed, and elevated [O(3)] tended to accelerate, leaf abscission in autumn. Elevated [CO(2)] increased the ratio of monosaccharides to di- and oligosaccharides in aspen buds, which may indicate a lag in cold acclimation. The total carbon concentration in overwintering buds was unaffected by the treatments, although elevated [O(3)] decreased the amount of starch by 16% in birch buds, and reduced the size of aspen buds, which may be related to the delayed leaf development in aspen during the spring. Elevated [CO(2)] generally ameliorated the effects of elevated [O(3)]. Our results show that both elevated [CO(2)] and elevated [O(3)] have the potential to alter carbon metabolism of overwintering buds. These changes may cause carry-over effects during the next growing season.     

Simulation study of the vegetation structure and function in eastern Siberian larch forests using the individual-based vegetation model SEIB-DGVM

The global ecosystem model SEIB-DGVM was adapted for an eastern Siberian larch forest through incorporation of empirical rules of allometry, allocation, and phenology developed for a larch stand at the Spasskaya-pad tower site, Yakutsk, Russia. After calibration, the model reconstructed post-fire successional patterns of forest structure and carbon cycling. It also reconstructed seasonal changes in carbon, water, and energy cycling in a mature larch forest. Sensitivity analysis showed that simulated functional properties of forest (LAI, NPP, carbon pools, and water runoff) are mainly determined by climatic environment, and population dynamic parameters (i.e., parameters for establishment and mortality) plays only minor role on them. Sensitivity analysis also showed that plant productivity and biomass were mainly limited by available water at Spasskaya-pad, where mean annual precipitation is only 257 mm. In the model, higher air temperature increases plant productivity via extension of growing season, and decreases plant productivity via causing drought and higher respiration. We found that the net effect is reduction of productivity, suggesting a possibility that global warming induces decrement of plant productivity in eastern Siberian larch forests.     

Timing, light availability and vigour determine the response of Abies alba saplings to leader shoot browsing

Herbivore browsing on tree saplings is a common phenomenon that can cause damage particularly on preferred species. In this study, the combined effects of light availability and timing of browsing on the response of 9-year-old Abies alba saplings were tested experimentally. Leader shoot clipping was applied before budburst, shortly after budburst or in autumn on saplings grown in full light or under artificial shade. Timing of clipping, light availability and tree vigour (expressed as height and tree ring width before clipping) had an effect on the height after clipping. After clipping in autumn or before budburst, fast-growing fir saplings bent up twigs to form new leader shoots and overcompensated height loss; saplings characterised by intermediate growth rates formed new shoots out of regular visible lateral buds; and slow-growing saplings had no new shoot in the first year after clipping, such that the clipping-induced height difference even increased over time. Saplings clipped shortly after budburst elongated the remaining part of the shoot in the first year and developed shoots out of the most distal lateral buds in the second growing season, leading to complete height compensation. Multi-trunking was typical for all clipped trees. We conclude that the microscale conditions under which a tree is growing (i.e. which affect tree vigour) are highly important for determining whether the height reduction imposed by browsing is offset by overcompensation or increases over time relative to unclipped trees. This response can partly be influenced by forest management via enhancing tree vigour via the light regime.     

水曲柳落叶松混交林中细根空间分布

采用根钻取样方法对年生水曲柳落叶松混交林中细根空间分布状况进行了研究。结果表明,水曲柳落叶松地下生物量的空间分配差异显著。在林分水平上,水曲柳的根生物量密度高于落叶松(分别为4442.3和2234.9g/m3)。两树种在相邻区域中分配的细根生物量较高,表明种间根系竞争较弱。落叶松行间的水曲柳细根生物量密度和根长密度均高于水曲柳行间的落叶松细根,表明水曲柳地下部分具有较强能力。根系的空间分布有利于混交林中水曲柳的生长。图1表4参19。     

Effects of winter buds on winter nitrogen uptake and allocation in Pinus densiflora saplings

Studies of nitrogen (N) use by plants have confirmed some winter N uptake; however, the mode of regulation of plant N use in winter is unknown. The regulation of N use by plants during winter may differ from that in the growing season, as plant growth strongly affects N use. We investigated the effects of winter buds on winter N use by Japanese red pine (Pinus densiflora), as a previous study demonstrated that N absorbed during winter contributes significantly to leaf growth in the following spring. We conducted a bud pruning experiment during winter to examine the effects of winter buds on winter N uptake and allocation among plant organs using ¹⁵N labeling. Over a three-week labeling period, the ¹⁵N content in roots increased to 0.20 ± 0.12 mg N g DW^?1, which is equivalent to 1.8 ± 1.1 % of the total N content in the roots. However, this absorbed ¹⁵N rarely appeared in needles and buds. Bud pruning did not affect ¹⁵N uptake and allocation. On the other hand, significant total N retranslocation was found within the crowns of saplings without bud pruning, but N was not retranslocated in bud-pruned plants. The bud pruning experiment indicated that N was retranslocated from needles into winter buds. Since soil N availability changes dramatically and is unstable in many forest ecosystems, N contained in needles would be a more stable source of N than newly absorbed N.     

Integrated individual tree biomass simultaneous equations for two larch species in northeastern and northern China

Forest biomass estimation at large scale has become an important topic in the background of facing global climate change, and it is fundamental to develop individual tree biomass equations suitable for large-scale estimation. Based on the measured data of biomass components and stem volume from 100 sample trees of two larch species (Larix gmelinii and L. principis-rupprechtii) in northeastern and northern China, an integrated equation system including individual tree biomass equations, stem volume equation and height–diameter regression model were constructed using the dummy variable model and error-in-variable simultaneous equations. In the system, all the parameters of equations were estimated simultaneously, so that the aboveground biomass equation was compatible to stem volume equation and biomass conversion factor (B_CF) function; the belowground biomass equation was compatible to root-to-shoot ratio (R_SR) function; and stem wood, stem bark, branch and foliage biomass equations were additive to aboveground biomass equation. In addition, the system also ensured the compatibility between one- and two-variable models. The results showed that: (1) whether aboveground biomass equations or belowground biomass equations and stem volume equations, the estimates for larch in northeastern China were greater than those in northern China; (2) B_CF of a larch tree decreased with the growing diameter while R_SR increased with the growing diameter; (3) the proportion of stem wood biomass to aboveground biomass increased with the growing diameter while those of stem bark, branch, and foliage biomass decreased.     

Growth, allocation and tissue chemistry of Picea abies seedlings affected by nutrient supply during the second growing season

One-year-old Norway spruce (Picea abies (L.) Karst.) seedlings were grown hydroponically in a growth chamber to investigate the effects of low and high nutrient availability (LN; 0.25 mM N and HN; 2.50 mM N) on growth, biomass allocation and chemical composition of needles, stem and roots during the second growing season. Climatic conditions in the growth chamber simulated the mean growing season from May to early October in Flakaliden, northern Sweden. In the latter half of the growing season, biomass allocation changed in response to nutrient availability: increased root growth and decreased shoot growth led to higher root/shoot ratios in LN seedlings than in HN seedlings. At high nutrient availability, total biomass, especially stem biomass, increased, as did total nonstructural carbohydrate and nitrogen contents per seedling. Responses of stem chemistry to nutrient addition differed from those of adult trees of the same provenance. In HN seedlings, concentrations of alpha-cellulose, hemicellulose and lignin decreased in the secondary xylem. Our results illustrate the significance of retranslocation of stored nutrients to support new growth early in the season when root growth and nutrient uptake are still low. We conclude that nutrient availability alters allocation patterns, thereby influencing the success of 2-year-old Norway spruce seedlings at forest planting sites.     

植物激素对黄檗幼树生长性状的影响及生物量模型的构建

以黄檗5 a生幼树为材料,对叶面喷施不同组合的植物激素,利用正交设计,研究其对生长性状的影响,并建立生物量模型,结果表明:栽植当年和翌年秋季幼树的成活率与保存率分别为97%和95%,平茬与未平茬在定植2年后未发现生长性状有显著差异;植物激素不同组合处理对幼树树高、地径生长量影响较为明显,3种植物激素对树高和地径生长量影响大小依次为NAA6-BAGA3;有效促进树高生长的最佳植物激素配比为NAA200 mg·L-1+6-BA400 mg·L-1+GA3400 mg·L-1,有效促进地径生长的最佳植物激素配比为NAA200 mg·L-1+6-BA400 mg·L-1+GA3200 mg·L-1。地径是影响生物量的主因子,拟合程度最高的一元生物量方程为Y=0.061-0.083X+0.047X2。     

Height growth of young larch (Larix kaempferi) in relation to the frequency of deer browsing damage in Hokkaido, Japan

The effect of browsing frequency by Sika deer (Cervus nippon yezoensis) on the height growth of Japanese larch (Larix kaempferi) in Hokkaido, Japan, was analyzed. Tree heights at the time of the census and at the start of the current growing season were determined for 100 trees selected randomly at each site. The tallest shoot on each tree was identified and the number of browsing occurrences on the current-year shoot was determined. Young larches browsed by deer repeatedly develop new shoots during a growing season. Larch is a browsing-tolerant species, and the average height growth of the most heavily browsed trees was >15 cm. Although resprouting partly compensated for decreased height growth caused by browsing, the annual increase in height decreased with increased browsing frequency. Thus, browsing frequency during a growing season best reflects the intensity of browsing damage to larch trees.     

Ectomycorrhizae Reduce Damage to Russian Larch by Otiorhyncus Larvae

The effect of ectomycorrhizae on damage caused by Otiorhyncus larvae and on plant growth was evaluated in a 3-yr field experiment. Russian larch seedlings, inoculated with ectomycorrhizal fungi, were compared with uninoculated plants. Assessment of larval damage at the end of the first growing season showed a significant effect of treatment on damage intensity, but not on damage incidence: 11.2% of uninoculated seedlings were severely damaged, but only 3.5% of inoculated plants. There was a significant effect of treatment on plant mortality. The accumulated mortality at the end of the third growing season was 34.5% for uninoculated plants, but 17.6% for inoculated plants. The height increment of inoculated plants (4.5 cm) was significantly greater than that of uninoculated plants (2.7 cm) in the first growing season. No difference in height increment between treatments was observed during the second and third growing seasons.     

Nitrogen availability modifies the ozone responses of Scots pine seedlings exposed in an open-field system

Three-year-old Scots pine (Pinus sylvestris L.) seedlings were exposed to either ambient or elevated (1.5-1.6 x ambient) ozone concentration ([O3]) for three growing seasons in an open-field fumigation facility where they were irrigated during the growing season with a nutrient solution providing nitrogen (N) at 70 (LN treatment), 100 (control) or 150% (HN treatment) of the optimum supply rate. Treatment effects were most evident during the third year of exposure, when the ambient [O3] + HN treatment enhanced whole-plant biomass, root/shoot dry weight ratio, needle pigment concentrations and the number of chloroplast plastoglobuli in the mesophyll cells in current-year (C) needles, whereas it reduced starch accumulation in C needles and abscission of 2-year-old (C+2) needles. In the control fertilization, 3 years of exposure to elevated [O3] decreased stem-base diameter and increased K concentration and electron density of chloroplast stroma in C needles. Plants in the HN treatment exposed for 3 years to elevated [O3] had significantly lower heights, current-year main shoot length and root/shoot dry mass ratio than control plants, and increased abscission of C+2 needles. In contrast, O3-induced changes in the ultrastructure of mesophyll cells were most evident in seedlings grown for 3 years in the LN treatment. We conclude that, in Scots pine, a relatively O3-tolerant species, chronic O3 exposure leads to cumulative growth reduction, increased needle abscission and changes in carbon allocation that are strongly influenced by plant N availability.     

Effects of partial defoliation on carbon and nitrogen partitioning and photosynthetic carbon uptake by two-year-old cork oak (Quercus suber) saplings

At the end of the growing season in late July, 20-month-old cork oak (Quercus suber L.) saplings were partially defoliated (63% of leaf area) to evaluate their ability to recover leaf area after defoliation. At 18 and 127 days after defoliation, changes in starch and nitrogen pools were determined in leaves and perennial organs, and variations in photosynthetic carbon uptake were investigated. To determine the role of stored nitrogen in regrowth after defoliation, plant nitrogen was labeled in the previous winter by enriching the nutrient solution with 15N. Plants recovered the lost leaf area in 127 days. Although there was remobilization of starch and nitrogen from leaves and perennial organs, the availability of resources for growth in the following spring was not decreased by defoliation. On the contrary, starch concentration in coarse roots was higher in defoliated saplings than in control saplings, presumably as a result of the higher net CO2 exchange rate in newly developed leaves compared with pre-existing leaves.     

不同种源非洲桃花心木苗期抗寒能力和生长节律研究

2008年1月上旬至2009年7月下旬,对非洲引进的3个桃花心木种源的幼树进行每周1次的观测.结果显示:种源间抗寒能力、树高生长和地径生长以及速生期都存在明显差异.种源3-1324与4-1325在苗期表现出一定的抗寒性,种源1-1320苗期对低温敏感.总体上,非洲桃花心木幼树1年抽梢4～7次,1～2月为树高生长休眠期,...     

日本落叶松容器苗不同控释肥生长效应

以落叶松网袋容器播种苗为对象,施用不同配方控释肥,进行落叶松苗期生长量、生物量、根冠比、叶绿素含量的测定,采用模糊数学隶属值计算方法选出适合日本落叶松育苗的专用配方控释肥。结果表明:按播种后天数,日本落叶松苗期生长阶段为:出苗期(7 39 d),生长初期(40 54 d),速生期(55 114 d)及生长后期(115 135d);不同配方控释肥对落叶松苗高、地径、单株生物量、地上生物量、地下生物量及叶绿素含量的影响差异极显著,而不同配方控释肥间根冠比差异不显著。控释肥配方2综合效果优于其他配方控释肥及从美国进口控释肥爱贝施,可以作为日本落叶松育苗专用肥。     

Morphological acclimation to understory environments in Abies amabilis, a shade- and snow-tolerant conifer species of the Cascade Mountains, Washington, USA

Light-related plasticity in a variety of crown morphology and within-tree characteristics was examined in sun and shade saplings of Abies amabilis Dougl. ex J. Forbes growing in two late-successional forests with different snow regimes in the Cascade Mountains of Washington, USA. Compared with sun saplings, shade saplings typically had broad flat crowns as a result of acclimation at several scales (needle, shoot, branch, crown and whole sapling). Shoots of shade saplings had a smaller needle mass per unit of stem length than shoots of sun saplings, a feature that enhances light-interception efficiency by reducing among-needle shading. The low annual rate of needle production by shade saplings was associated with a longer needle lifespan and slower needle turnover. Reduced needle production within a shoot was reflected at the branch level, with lateral branches of shade saplings having a smaller needle mass than branches of the same length of sun saplings. Reduced allocation to needles permits greater investment in branches and stems, which is necessary to support the horizontally expanding branch system characteristic of shade saplings. Mean branch age of shade saplings was significantly higher than that of sun saplings. Shade saplings had lower needle mass per unit of trunk biomass or total biomass, reflecting greater investment in the trunk as a support organ. Increased investment in support organs in shade was more evident in the snowier habitat. The observed morphological acclimation makes A. amabilis highly shade and snow-tolerant and thus able to dominate in many late-successional forests in snowy coastal mountain regions.     

Whole-plant water flux in understory red maple exposed to altered precipitation regimes

Sap flow gauges were used to estimate whole-plant water flux for five stem-diameter classes of red maple (Acer rubrum L.) growing in the understory of an upland oak forest and exposed to one of three large-scale (0.64 ha) manipulations of soil water content. This Throughfall Displacement Experiment (TDE) used subcanopy troughs to intercept roughly 30% of the throughfall on a "dry" plot and a series of pipes to move this collected precipitation across an "ambient" plot and onto a "wet" plot. Saplings with a stem diameter larger than 10 cm lost water at rates 50-fold greater than saplings with a stem diameter of 1 to 2 cm (326 versus 6.4 mol H(2)O tree(-1) day(-1)). These size-class differences were driven largely by differences in leaf area and cross-sectional sapwood area, because rates of water flux expressed per unit leaf area (6.90 mol H(2)O m(-2) day(-1)) or sapwood area (288 mol H(2)O dm(-2) day(-1)) were similar among saplings of the five size classes. Daily and hourly rates of transpiration expressed per unit leaf area varied throughout much of the season, as did soil matrix potentials, and treatment differences due to the TDE were observed during two of the seven sampling periods. On July 6, midday rates of transpiration averaged 1.88 mol H(2)O m(-2) h(-1) for saplings in the "wet" plot, 1.22 mol H(2)O m(-2) h(-1) for saplings in the "ambient" plot, and 0.76 mol H(2)O m(-2) h(-1) for saplings in the "dry" plot. During the early afternoon of August 28, transpiration rates were sevenfold lower for saplings in the "dry" plot compared to saplings in the "wet" plot and 2.5-fold lower compared to saplings in the "ambient" plot. Treatment differences in crown conductance followed a pattern similar to that of transpiration, with values that averaged 60% lower for saplings in the "dry" plot compared to saplings in the "wet" plot and 35% lower compared to saplings in the "ambient" plot. Stomatal and boundary layer conductances were roughly equal in magnitude. Estimates of the decoupling coefficient (Omega) ranged between 0.64 and 0.72 for saplings in the three TDE treatment plots. We conclude that red maple saplings growing in the understory of an upland oak forest are responsive to their edaphic and climatic surroundings, and because of either their small stature or their shallow root distribution, or both, are likely to be impacted by precipitation changes similar to those predicted by global climate models.     

Effects of long-term,elevated ultraviolet-B radiation on phytochemicals in the bark of silver birch (Betula pendula)

Long-term outdoor experiments were conducted to investigate the effects of elevated ultraviolet-B (UV-B, 280-320 nm) radiation on secondary metabolites (phenolics and terpenoids) and the main soluble sugars (sucrose, raffinose and glucose) in the bark of silver birch (Betula pendula Roth) saplings. Saplings were exposed to a constant 50% increase in erythemal UV irradiance (UV-B(CIE); based on the CIE (International Commission on Illumination) erythemal action spectrum) and a small increase in UV-A radiation (320-400 nm) for three growing seasons in an irradiation field in central Finland. Two control groups were used: saplings exposed to ambient radiation and saplings exposed to slightly increased UV-A radiation. Concentrations of sucrose, raffinose and glucose in bark were higher in UV-treated saplings than in saplings grown in ambient radiation, indicating that stem carbohydrate metabolism was changed by long-term elevated UV radiation. Saplings in the elevated UV-A + UV-B radiation treatment and the UV-A radiation control treatment had significantly increased concentrations of certain UV-absorbing phenolics, such as salidroside, 3,4'-dihydroxypropiophenone-3-glucoside, (+)-catechin and (-)-epicatechin compared with saplings in ambient radiation. In contrast, the radiation treatments had no effect on the non-UV-B-absorbing terpenoids, papyriferic acid and deacetylpapyriferic acid. We conclude that plant parts, in addition to leaves, accumulate specific phenolic UV-filters in response to UV radiation exposure.     

Bud set and autumn coloration of Larix sibirica and Larix gmelinii provenances in a field trial in southern Finland

The main aim of this study was to analyse the differences in the bud set, autumn coloration and needle shedding of seedlings in 20 Russian larch provenances and five comparison entries in field conditions in southern Finland over two growing seasons. Furthermore, it was studied if the differences could be explained by the geographic and climatic conditions of the provenance origins. We found that the provenances from cold northern climates developed both their terminal buds first and formed autumn colour as well as shed their needles much earlier than southern provenances. The timing of the bud set occurred earlier in 2008 compared to 2009, which was probably caused by lower than average temperatures in July and August of 2008. Thus, in addition to photoperiod, temperature also seemed to affect the bud set. The southern Dahurian larch provenances were able to utilise the length of the growing season the most effectively. The climatic adaptation, growth and properties of the provenances should be, however, studied over a longer time period. Therefore, currently used Raivola origin is still the safest choice for larch forestry in Finland.