The importance of biotic filtering on boreal conifer recruitment at alpine treeline

Treeline, the ecotone where forest transitions to alpine or tundra ecosystems, is considered the thermal limit to tree growth and survival. Despite temperature increases across mountainous areas and high latitudes globally, there has been no ubiquitous change in treeline position. The process of range expansion must initially depend on increased recruitment at, or beyond current range limits and recruitment limitations have been hypothesized as a mechanism for the variable response of treeline position to climate warming. We conducted a unique series of observational and experimental studies to quantify early-life stage constraints, from seed production to seedling establishment, on black spruce Picea mariana and tamarack Larix laricina recruitment at a model alpine treeline in Newfoundland, Canada. We found recruitment at treeline to be simultaneously seed and establishment limited. The treeline population produced fewer seeds than the forest population and black spruce seeds produced at treeline were less viable. Tamarack was more seed limited than black spruce where seed viability was low regardless of altitudinal position. Post-dispersal seed predation greatly constrained recruitment across the altitudinal gradient; however, black spruce seeds experienced the lowest levels of invertebrate seed predation on the lichen mat at treeline. If seeds were not consumed, individuals at treeline were establishment limited where germination and seedling establishment was both less abundant and delayed on lichen substrate. Our study highlights the need for multiple factors to align temporally for significant recruitment at treeline to occur.     

Carbon, nitrogen and phosphorus in volcanic soils following afforestation with native birch (Betula pubescens) and introduced larch (Larix sibirica) in Iceland

Afforestation has become an important tool for soil protection and land reclamation in Iceland. Nevertheless, the harsh climate and degraded soils are growth-limiting for trees, and little is know about changes in soil nutrients in maturing forests planted on the volcanic soils. In the present chronosequence study, changes in C, N and total P in soil (0–10 and 10–20 cm depth) and C and N in foliar tissue were investigated in stands of native Downy birch (Betula pubescens Enrh.) and the in Iceland introduced Siberian larch (Larix sibirica Ledeb.). The forest stands were between 14 and 97 years old and were established on heath land that had been treeless for centuries. Soils were Andosols derived from basaltic material and rhyolitic volcanic ash. A significant effect of tree species was only found for the N content in foliar tissue. Foliar N concentrations were significantly higher and foliar C/N ratios significantly lower in larch needles than in birch leaves. There was no effect of stand age. Changes in soil C and the soil nutrient status with time after afforestation were little significant. Soil C concentrations in 0–10 cm depth in forest stands older than 30 years were significantly higher than in heath land and forest stands younger than 30 years. This was attributed to a slow accumulation of organic matter. Soil N concentrations and soil P_tot were not affected by stand age. Nutrient pools in the two soil layers were calculated for an average weight of soil material (400 Mg soil ha⁻¹ in 0–10 cm depth and 600 Mg soil ha⁻¹ in 10–20 cm depth, respectively). Soil nutrient pools did not change significantly with time. Soil C pools were in average 23.6 Mg ha⁻¹ in the upper soil layer and 16.9 Mg ha⁻¹ in the lower soil layer. The highest annual increase in soil C under forest compared to heath land was 0.23 Mg C ha⁻¹ year⁻¹ in 0–10 cm depth calculated for the 53-year-old larch stand. Soil N pools were in average 1.0 Mg N ha⁻¹ in both soil layers and did not decrease with time despite a low N deposition and the uptake and accumulation of N in biomass of the growing trees. Soil P_tot pools were in average 220 and 320 kg P ha⁻¹ in the upper and lower soil layer, respectively. It was assumed that mycorrhizal fungi present in the stands had an influence on the availability of N and P to the trees. Responsible Editor: Hans Lambers.     

Carbon allocation in shoots of alpine treeline conifers in a CO<Subscript>2</Subscript> enriched environment

With a new approach we assessed the relative contribution of stored and current carbon compounds to new shoot growth in alpine treeline conifers. Within a free air CO₂ enrichment experiment at the alpine treeline in Switzerland, ¹³C-depleted fossil CO₂ was used to trace new carbon in the two tree species Larix decidua L. and Pinus uncinata Ramond over two subsequent years. The deciduous L. decidua was found to supply new shoot growth (structural woody part) by 46% from storage. Surprisingly, the evergreen P. uncinata, assumed to use current-year photosynthates, also utilized a considerable fraction of storage (42%) for new wood growth. In contrast, the needles of P. uncinata were built up almost completely from current-year photosynthates. The isotopic composition of different wood carbon fractions revealed a similar relative allocation of current and stored assimilates to various carbon fractions. Elevated CO₂ influenced the composition of woody tissue in a species-specific way, e.g. the water soluble fraction decreased in pine in 2001 but increased in larch in 2002 compared to ambient CO₂. Heavy defoliation applied as an additional treatment factor in the second year of the experiment decreased the lipophilic fraction in current-year wood in both species compared to undefoliated trees. We conclude that storage may play an important role for new shoot growth in these treeline conifers and that altered carbon availability (elevated CO₂, defoliation) results in significant changes in the relative amount of mobile carbon fractions in woody tissue. In particular, stored carbon seems to be of greater importance in the evergreen P. uncinata than has been previously thought.     

Temporal variation of δ<Superscript>13</Superscript>C of larch leaves from a montane boreal forest in Mongolia

This paper reports the temporal variation (2002–2004) in foliar δ¹³C values, which are indicative of long-term integrated photosynthetic and water use characteristics, of Siberian larch (Larix sibirica Ledeb.) trees in a montane forest at Mongonmorit, NE Mongolia. At the stand, the δ¹³C value for understory shaded leaves was more negative by 2‰ on average than that for sunlit leaves sampled concurrently from open and sun-exposed environments in a forest gap. The δ¹³C value of both sunlit and shaded leaves showed pronounced intra- but relatively small inter-seasonal variations. The δ¹³C value was more positive for juvenile than mature leaves. We conjecture that juvenile leaves may derive carbon reserves in woody tissues (e.g., stems). Regardless of leaf habitats, the δ¹³C value was also affected by insect herbivores occurred in mid summer of 2003, being more negative in newly emerging leaves from the twigs after defoliation than in non-defoliated mature leaves. This pattern seems to contrast with that for the juvenile leaves in the early growing season. We surmise that the newly emerging leaves used stored organic carbon that was depleted due to fractionation during remobilization and translocation for leaf regrowth. There was also intra- and inter-seasonal variation in the foliar N concentrations and C:N ratios. A good positive (negative) correlation between the foliar δ¹³C values and N concentrations (C:N ratios) was also observed for both sunlit and shaded leaves, suggesting that the relationship between water and nitrogen use is a crucial factor affecting the plant carbon–water relationship in this mid latitude forest with a cold semiarid climate. Our isotopic data demonstrate that the larches in NE Mongolia exhibits relatively higher water use efficiency with a distinct within-season variability.     

Exogenous succinic acid mediates responses of Larix olgensis A. Henry to cadmium stress

Abstract

Trace metal contamination of soil is an increasing problem. Organic acid application can restore trace metal elements such as cadmium (Cd) in contaminated soil. Changbai larch (Larix olgensis A. Henry) is an economically important forestry species in northeast China; however, growth is inhibited by severe Cd contamination. We investigated the effects of different concentrations of exogenous succinic acid (SA) on Cd tolerance and physiological and morphological toxicity in L. olgensis seedlings. Seedlings were planted in pots containing Cd-contaminated or uncontaminated Haplic Cambisol. Seedlings in Cd-contaminated soil were treated daily with SA solution at 0, 0.04, 0.2, 1.0, and 2.0?mmol kg^?1 of soil for 10, 20 or 30?days. Cd treatment induced seedling damage and significantly increased the relative conductivity and malondialdehyde content of the leaves, inhibiting soluble protein and proline contents, superoxide dismutase and peroxidase activity, chlorophyl fluorescence and pigment content. Decreases in the length, surface area, volume of roots and leaves, and specific root length were also observed. Effects increased in control plants with time. SA treatment also reduced the Cd content of the fine roots and leaves and Mg, K, and Ca contents. Moreover, plant growth was significantly promoted and damage was reversed, especially at 5.0 and 10.0?mmol?L^?1 SA for 30?days. SA therefore alleviated Cd-induced injury, improving tolerance to Cd stress. SA application combined with afforestation could therefore help restore Cd-contaminated soil in northeast China. Further studies aimed at determining the detoxification mechanism of L. olgensis seedlings are now required.     

帽儿山地区兴安落叶松人工林树木年轮气候学研究

通过帽儿山兴安落叶松（Larix gmelinii）人工林树木年轮样本和气象资料,对该地区兴安落叶松进行了树木年轮气候学研究,结果表明：过去50年年均温度上升达到了显著水平（p<0.05）,平均温度每10年约上升0.4℃,年平均最高气温每10年约上升0.3℃,年平均最低气温每10年约上升0.5℃,但是年降水量随着年份变化不显著（p>0.05）。从月均温度来看,所有月份均出现明显上升趋势,其中冬季2月份温度上升最为明显,达到0.9~1℃/10年,而夏季（6~8月）上升的较小,达到0.2~0.7℃/10年;多数月份降雨量随年龄变化不显著（p>0.05）。在这一气候变暖过程中,早材及总年轮宽度生长随着夏季（6~7月）温度上升而下降,春季(5月)温度的升高而升高,晚材随着秋季（9月）温度上升而增加,导致在年水平上,年轮生长随着年均温的变化不显著（p>0.05）。降雨量在未来气候变化过程中,没有稳定的变化趋势,但是对年轮影响明显,在年水平上,早材与年轮的生长均受降水量的影响较大(p<0.05)。如果未来东北地区气候变暖趋势明显,而降水量变化不明显,春季和秋季温度升高导致的年轮生长增加会被夏季过高温度抑制年轮生长所抵消,因此,落叶松林径向生长受到的影响可能不大。     

气候暖化对解冻期不同纬度兴安落叶松林土壤氧化亚氮释放的影响

春季解冻期北方森林土壤释放出大量的N₂O,是大气温室气体的主要来源之一.本研究将分布于塔河(52°31′ N)、松岭(50°43′ N)、孙吴（49°13′ N）和带岭（47°05′ N）4个纬度上林分状况相似的兴安落叶松林样地（包括幼树、地被物和A、B层土壤）移置至其自然分布区南缘,模拟气候暖化,分析春季解冻期土壤N2O通量的时间动态及其影响因子.结果表明：各处理的土壤N2O释放高峰均出现在解冻中后期.带岭、孙吴、松岭和塔河样地土壤解冻期的土壤N₂O平均通量分别为（66.5±9.3）、（54.3±5.6）、（44.3±5.3）和（33.5±3.7） μg·m^-2·h^-1;土壤N₂O通量与5 cm土壤温度和0～10 cm土壤微生物生物量碳呈显著正相关,但与土壤含水率的相关性不显著.解冻期4个处理的土壤N₂O释放差异显著,其平均通量和累积释放量均随纬度升高而下降.这种差异主要是由于不同纬度土壤的微生物活性和土壤物理性质的差异引起的.     

太白红杉林斑块结构与群落特征

根据森林循环理论, 森林群落的动态是处于不同发育阶段的镶嵌系统。通过调查太白山亚高山针叶林带太白红杉 (Larix chinensis) 林的结构, 确定出4种斑块阶段 (林窗阶段、建立阶段、成熟阶段和衰退阶段), 研究了森林斑块动态变化和生物多样性变化规律, 并测定分析了不同斑块类型内光照和温度的日变化规律。结果表明:1) 群落内不同斑块类型的比例分别是:林窗阶段40.3%, 建立阶段34.0%, 成熟阶段17.2%, 退化阶段8.5%。2) 不同斑块类型内, 环境因子 (光照和温度等) 的日变化差异明显。其中, 在林窗阶段的光照强度和土壤表面温度要比其他3个阶段变化更大。3) 森林循环过程中, 不同高度和不同径级个体的密度存在着明显差异。4) 平均胸径、高度和平均基面积、个体平均体积和立地材积均随着森林的循环而增加。5) 森林循环过程中, 生物多样性的变化为波形的。运用自然干扰与斑块动态理论, 解释了太白红杉林循环过程中不同斑块之间生物多样性存在差异的可能原因。     

13CO2 pulse-labelling of photoassimilates reveals carbon allocation within and between tree rings

Post-photosynthetic fractionation processes during translocation, storage and remobilization of photoassimilate are closely related to intra-annual sigma13C of tree rings, and understanding how these processes affect tree-ring sigma13C is therefore indispensable for improving the quality of climate reconstruction. Our first objective was to study the relationship between translocation path and phloem grain. We pulse-labelled a branch of Larix gmelinii (Rupr.) Rupr. and later analysed the sigma13C distribution in the stem. A 13C spiral translocation path closely related to the spiral grain was observed. Our second objective was to study the use of remobilized storage material for earlywood formation in spring, which is a suspected cause of the autocorrelation (correlation of ring parameters to the climate in the previous year) observed in (isotope) dendroclimatology. We pulse-labelled whole trees to study how spring, summer and autumn photoassimilate is later used for both earlywood and latewood formation. Analysis of intra-annual sigma13C of the tree rings formed after the labelling revealed that earlywood contained photoassimilate from the previous summer and autumn as well as from the current spring. Latewood was mainly composed of photoassimilate from the current year's summer/autumn, although it also relied on stored material in some cases. These results emphasize the need for separating earlywood and latewood for climate reconstruction work with narrow boreal tree rings.