基于日均温度的华山松径向生长敏感温度研究

树干木质部形成周期与温度密切相关,存在影响形成层活动的敏感温度,但是尚未研究尝试从树木年轮中探索影响径向生长的敏感温度。华山主峰的华山松径向生长对气候变化较敏感,且华山西峰的气象站记录了1953年以来气象资料,这为探索树木径向生长与温度的关系提供了宝贵的材料。以华山主峰的华山松年轮资料和日值温度资料为基础,通过分析历年日平均温度的变化过程和年轮宽度、早材宽度、晚材宽度、最小密度以及最大密度的相关性,尝试探索华山松径向生长与日均温度变化的关系。结果表明春季一定温度的初日时间对华山松径向生长有较大的影响,其中3 ℃和8 ℃初日时间和持续时间对年轮宽度的促进作用最明显,而3 ℃对早材宽度影响较大,8℃初日时间对晚材的影响较大;高于11 ℃的温度会对华山松的径向生长造成限制,其中以11 ℃的作用最明显;而温度的终日时间对年轮特征影响较小。说明3 ℃是早材形成的敏感温度,8 ℃是晚材形成的敏感温度,温度高于11 ℃会对华山松的径向生长构成胁迫。这证明华山松年轮特征中包含了较多的气候信息,形成层活动和木质部的生长存在阈值温度,通过分析不同温度的生长期与年轮特征的关系可以找到影响华山松径向生长的敏感温度。这些敏感的温度是通过什么生理过程影响木质部的形成尚无法得知,但是这为解释树木生长对全球温度升高的响应以及利用华山松年轮重建历史气候提供了重要依据。     

Jack pine of all trades: Deciphering intraspecific variability of a key adaptive trait at the rear edge of a widespread fire-embracing North American conifer

Premise

Understanding mechanisms fostering long-term persistence of marginal populations should provide key insights about species resilience facing climate change. Cone serotiny is a key adaptive trait in Pinus banksiana (jack pine), which shows phenotypic variation according to the fire regime. Compared to range-core populations within the fire-prone boreal forest, low and variable serotiny in rear-edge populations suggest local adaptation to uncommon and unpredictable wildfire regime. We assessed environmental/physiological factors that might modulate intraspecific variation in cone serotiny.

Methods

We experimentally subjected closed cones to incrementing temperatures, then tested seed germination to determine whether and how various ecological factors (cone age, branch height, tree size, tree age) are related to cone dehiscence and seed viability in jack pines from rear-edge and range-core populations in eastern Canada.

Results

Cones from rear-edge populations dehisce at a lower opening temperature, which increases with cone age. Cones from range-core stands open at a more constant, yet higher temperature. Cones from rear-edge stands take between 13 and 27 years to reach the level of serotiny achieved at the range core. At the rear edge, seed viability is steady (51%), whereas it decreases from 70% to 30% in 20 years at the range core.

Conclusions

We inferred the mechanisms of a bet-hedging strategy in rear-edge populations, which ensures steady recruitment during fire-free intervals and successful postfire regeneration. This capacity to cope with infrequent and unpredictable fire regime should increase the resilience of jack pine populations as global changes alter fire dynamics of the boreal forest.     

Wood properties and ring width responses to long-term atmospheric CO2 enrichment in field-grown loblolly pine (Pinus taeda L.)

Loblolly pine (Pinus taeda L.) were grown in the field, under non-limiting nutrient conditions, in open-top chambers for 4 years at ambient CO₂ partial pressures (pCO₂) and with a CO₂-enriched atmosphere (+ 30 Pa pCO₂ compared to ambient concentration). A third replicate of trees were grown without chambers at ambient pCO₂. Wood anatomy, wood density and tree ring width were analysed using stem wood samples. No significant differences were observed in the cell wall to cell lumen ratio within the latewood of the third growth ring formed in 1994. No significant differences were observed in the density of resin canals or in the ratio of resin canal cross-sectional area to xylem area within the same growth ring. Ring widths were significantly wider in the CO₂-enrichment treatment for 3 of 4 years compared to the ambient chamber control treatment. Latewood in the 1995 growth ring was significantly wider than that in the ambient control and represented a larger percentage of the total growth-ring width. Carbon dioxide enrichment also significantly increased the total wood specific gravity (determined by displacement). However, when determined as total sample wood density by X-ray densitometry, the density of enriched samples was not significantly higher than that of the ambient chamber controls. Only the 1993 growth ring of enriched trees had a significantly higher maximum latewood density than that of trees grown on non-chambered plots or ambient chambered controls. No significant differences were observed in the minimum earlywood density of individual growth rings between chambered treatments. These results show that the most significant effect of CO₂ enrichment on wood production in loblolly pine is its influence on radial growth, measured as annual tree ring widths. This influence is most pronounced in the first year of growth and decreases with age.     

农田防护林木材材质物理特性研究

对早柳和小青杨的年轮宽度、年轮密度、木材纤维长度和干缩率及其径向变异进行了研究．结果表明,木材物理特性受年轮年龄的影响显著,而纤维长度不受年轮宽度影响．综合材质物理特性的径向变异规律,可将旱柳和小青杨整个生长期划分为４个时期：幼林期、速生期、成熟期和更新期,确定了各生长期的年龄值．     

What are the consequences of growth selection on wood density in the French maritime pine breeding programme?

Volume and stem straightness were the main selection criteria for the first two generations of the French maritime pine (Pinus pinaster Ait.) breeding programme. In this article, we investigate the consequences of this selection on wood quality. Wood density, as a predictor of wood quality, is studied both in the breeding populations and in commercial varieties. Phenotypic and genetic correlations between wood density and growth traits are investigated in successive breeding populations with three genetic field experiments of respectively 30, 29 and 12 years old. Correlation estimates were either slightly negative or non-significantly different from zero depending on the test considered. Consequently, a low impact of growth selection on wood quality should be expected in improved seed sources. However, we observed a significant wood density decrease in two improved varieties as compared to unimproved seed sources at age 15. In addition to this first effect on wood density, growth improvement is also expected to reduce the rotation age and thus increase the proportion of juvenile wood, which is known as having a lower density than mature wood. This change was studied and quantified using a growth model. Finally, a wood density decrease reaching up to 6% was predicted in the improved varieties compared to unimproved material, when both the observed decrease in wood density and the predicted increase in juvenile wood proportion were taken into account. Implications for the breeding programme were considered.     

长白山北坡不同年龄红松年表及其对气候的响应

运用树木年轮气候学方法,研究了长白山北坡红松(Pinus koraiensis)不同年龄年表特征及其与气候因子间的关系,以期揭示年龄因素对年表的潜在影响。结果表明,平均年龄为63a的红松低龄年表与平均年龄为184a的高龄年表对气候的响应明显不同:低龄红松径向生长与当年1、2月月平均温度负相关(P<0.05),同时也受到上年及当年多个月份的月平均最高温度或最低温度的影响,但与降水的相关性未达到显著水平;高龄红松径向生长则与月平均温度间的关系不明显,而与当年1、2、4、6、7、9月的月平均最高温度正相关,与当年4月、9月的月平均最低温度负相关,同时受到上年5月及当年5月月总降水量的影响。因此,年龄因素对红松年表的气候响应方面存在一定影响,且高龄年表对气候响应的敏感性更高,包含有更多的气候信息。     

Phenotypic selection on growth rhythm in whitebark pine under climatic conditions warmer than seed origins

Growth rhythm that is well synchronized with seasonal changes in local climatic conditions is understood to enhance fitness; however, rapid ongoing climate change threatens to disrupt this synchrony. To evaluate phenotypic selection on growth rhythm under expected warmer and drier future climate, seedlings from 49 populations of whitebark pine (Pinus albicaulis Engelm.) were grown and measured over more than 10 years in two common garden field experiments on sites that approximate the projected future climate of the seed origins. Selection on growth rhythm was assessed by relating individual plant fitness to timing and rate of shoot elongation. Differential survival clearly evidenced selection on growth rhythm. We detected directional and stabilizing selection that varied in magnitude between experimental sites and among years. The observed phenotypic selection supports the interpretation of clinal variation among populations within tree species as reflecting adaptive variation in response to past natural selection mediated by climate. To the extent that growth rhythm is heritable, results of the present study suggest evolution of whitebark pine toward a more distinct timing of shoot elongation and generally more rapid elongation in the immediate next generation under ongoing climate change in environments similar to the study sites.     

Climate driven trends in tree biomass increment show asynchronous dependence on tree-ring width and wood density variation

Tree growth is a key ecosystem function supporting climate change mitigation strategies. However climate change may induce feedbacks on radial growth and wood density, affecting the carbon sequestration capacity of forests. Using a mixed modeling technique long-term trends in radial growth, wood density and above-ground biomass, defined as the product of the annual basal area growth with the wood density, of common beech (Fagus sylvatica) and sessile oak (Quercus petraea) in the Belgian Ardennes, were determined and explained using climate drivers of change. This modeling strategy allowed us to determine if the same conclusions can be drawn when only BAI is considered, as is assumed in most carbon sequestration studies, when looking at long-term trends in carbon sequestration. The models indicate that above-ground biomass increment changes over time are more driven by changes in radial growth than by changes in wood density. Nevertheless, the assumption of constant wood density in most carbon sequestration studies is incorrect. Ignoring wood density results in an underestimation of long-term trends in above-ground biomass increment for beech, and an overestimation of above-ground biomass increment for oak. Interesting is that radial growth is mostly driven by climate variables of the current year, whereas wood density is more driven by the climate variables of the previous year. Beech radial growth and wood density is found to be negatively influenced by drought and positively by water availability. Oak radial growth and wood density is negatively affected by late frost and positively by water availability. The findings of this study suggest that radial growth in combination with wood density should be used in carbon sequestration studies as different climate driven long-term trends in radial growth and wood density are found.     

Local environment prevails over population variations in growth-climate relationships of Pinus pinaster provenances

In view of the projected decrease in precipitation and increase in temperature, a better understanding on growth-climate responses in different populations of tree species is needed to improve and enhance the conservation and management strategies for major forest tree species. In this study, we assessed differences in growth traits (i.e., stem diameter, tree height, and stem radial growth) and analysed climate-growth relationships in five provenances of maritime pine (Pinus pinaster Ait.) grown in four replicated common gardens in Sardinia (Italy). Stem radial growth increased under a positive water balance in late winter and early spring. Conversely, high temperature and low precipitation in summer had a negative impact on stem radial growth. At age 40, none of the considered provenances (Portugal, Corsica, Tuscany, Sardinia-Telti, Sardinia-Limbara) showed a substantial advantage in more than one common garden site for tree height and stem diameter. Nevertheless, differences were found among common garden sites in terms of dendrometric parameters, stem radial growth, and growth-climate responses (including sensitivity to summer drought), suggesting a greater site dependent over tree growth. Although in juvenile stages the Portuguese provenance (in particular) showed greater growth than the Sardinian ones, this study showed that, with tree age, the differences among the five provenances tends to narrow. Therefore, irrespective of the seed source, tree growth patterns and growth-climate responses were similar at age 40. This result can be important for implementing forest management strategies to balance adaptation and mitigation potential of maritime pine plantations in harsh environmental conditions.     

Modelling variability of wood density in beech as affected by ring age,radial growth and climate

Although it has been recognized as a key parameter of wood quality and a good source of information on growth, annual wood density has been little studied within diffuse-porous trees such as beech ( Fagus sylvatica Liebl.). In this paper we examine the variability encountered in beech ring density series and analyze the influences of ring age, ring width, climate and between-tree variability on density. Thirty ring sequences were sampled from 55-year- old dominant beech trees growing within the same stand; ring density and width were measured using radiography. Ring density proved to be less variable through time than ring width. The relationship between these two variables was less than observed in ring-porous trees and it showed great variation between trees. The sensitivity of ring width and density to climate was also different; width was strongly linked to soil water deficit whereas density was correlated to temperature and August rainfall. Unlike ring width, wood density showed sensitivity towards climatic characteristics of the late growing season. A large part of annual density variability remains unexplained, even using advanced modelled water balance variables. We hypothesize that a significant part of the tree ring is under internal control. We also demonstrated great inter-tree variability (the tree effect) in ring density, which has an influence on density but not on trees response to climate.     

伊春地区红松和红皮云杉径向生长对气候变化的响应

树木生长-气候关系对准确评估气候变化对森林生态系统影响、预测森林生产力与植被动态及揭示树木对气候变化的响适应策略至关重要。在全球变暖背景下,升温可能会对树木的生长产生影响,从而改变区域森林生态系统的生产力或碳储量。本研究利用生长-气候响应函数、滑动相关分析等树木年轮学方法,探讨伊春地区阔叶红松林内红松和红皮云杉径向生长的主要限制因子及两者径向生长对快速升温(1980年后)响应的异同。结果表明:1980年前红松径向生长有明显加速的趋势,红皮云杉上升趋势较弱;而1980年后红松径向生长趋势显著下降,红皮云杉则下降不明显。红皮云杉径向生长与上一年9月及当年6月平均气温显著负相关,而红松径向生长与上一年12月及当年1月、4月和6月最低气温显著正相关。1980年快速升温后,高温对两树种生长的抑制作用增强,尤其是红松。生长季末(9月)降水对红松和红皮云杉的限制作用由升温前的负相关转变为升温后的显著正相关。温度是限制红松和红皮云杉径向生长的主要气候因子,降水影响相对较弱;其中红松径向生长对气候变化的响应比红皮云杉更敏感。快速升温后,红松和红皮云杉生长-气候关系的变化可能与升温导致的暖干旱化有关。若气候变暖持续或加剧,二者径向生长的气候限制因子也将由温度转变为水分;红松和红皮云杉会出现生长衰退,尤其是红松。     

不同林龄、树种落叶松人工林径向生长与气候变化的关系

研究人工林径向生长与气候变化的关系对全球气候变暖背景下人工林合理经营有着重要的意义。该文对在辽东山区广泛栽培的黄花落叶松(Larix olgensis)和日本落叶松(Larix kaempferi)人工林, 运用树木年轮气候学方法建立了辽宁草河口和湾甸子林场落叶松人工林年表, 分析了落叶松径向生长对气候变化的响应以及气候条件、树种、立地条件和林分因子(林龄、密度、蓄积量等)的相对影响程度。结果发现在影响年轮-气候关系的因素中, 气象因子的潜在蒸发散(PET)的影响力最大; 林龄、密度和蓄积量同时也具有重要的影响作用。中龄落叶松人工林径向生长主要与气温呈正相关关系, 成熟落叶松人工林径向生长主要与气温呈负相关关系; 而其他因素, 如树种、立地条件等的影响作用不大。这表明在气候变暖背景下随着林龄增加, 林分会逐渐受到气温升高导致的水分亏缺的限制, 导致明显的生长下降趋势, 因而气候变暖对成熟落叶松人工林威胁更为严重, 所以要注重对成熟林的优先保护, 同时可以预测, 随着东北地区今后气候进一步变暖, 可能将逐步影响到林龄更小的林分的生长, 因此需要进一步研究如何在落叶松人工林经营中采取科学的措施来更好地应对未来气候变化。     

What tree rings can tell us about the competition between trees and lianas? A case study based on growth,anatomy, density,and carbon accumulation

In tropical forest, landscape fragmentation and the consequent degradation of disturbed forests increase the incidence of light and dry hot winds, causing a disturbance on natural regeneration. Under these conditions, lianas (woody vines) development is stimulated instead of other species, which are more suited to mature forest and under less influence of the edge effect. For this, lianas colonization is an important variable for assessing the disturbance level of a forest. In this context, it becomes important to understand the nature of the competitive relationships between hyper-abundant lianas and ring growth of the host trees. Here, we selected trees with occupation or absence of lianas from two tropical species – Pinus caribaea var. hondurensis (Caribbean pine) and Tectona grandis (teak) – localized in a semideciduous forest fragment in southeastern Brazil, aiming to compare growth, climatic response, anatomy (vessels and intra-annual density fluctuations), wood density and carbon, by tree-ring analysis. The results showed that the lianas caused a change in tree-ring anatomy of host trees in last 10 years, mainly. We observed that trees occupied by lianas had a decrease the radial growth and carbon in the two species, an increase of the vessels size in teak and a decrease of the IADF frequency in Caribbean pine. In teak, the climate-tree relationship indicated that trees with lianas had lower response to rainfall and higher response to temperature in the summer (rainy and hottest period); in Caribbean pine, we observed that trees with lianas had a 2-month delay in the radial growth response to rainfall in the dry season. In the teak group, we observed that host trees had higher wood density values than liana-free tree in the outer rings, and the opposite was showed for pine. These findings show that tree-ring growth of host trees are a strong bioindicator of forest disturbance caused by aggressive colonization of lianas. We believe that these methods are applicable to future studies relating to the effects of habitat fragmentation and forest degradation on biodiversity and ecosystem services, particularly in the context of global climate change.     

川西亚高山不同年龄紫果云杉径向生长对气候因子的响应

运用树木年轮气候学的基本方法,建立王朗自然保护区紫果云杉在集中分布上限区域的年轮宽度年表,选取差值年表分析不同年龄云杉的径向生长同逐月气候因子的相关及响应关系,结果显示:幼龄组云杉年表的敏感度高于中龄组和老龄组云杉,幼龄组云杉对生长季前及生长季的气温状况显著正相关;中龄组云杉年表仅与当年4月份和7月份的月平均最低气温显著正相关;老龄组云杉的年轮宽度指数同上年生长季(上年8月份)的月平均气温和月平均最低温显著负相关,上年生长季高温的"滞后效应"在老龄组云杉体现的更为突出;幼龄组与中龄组云杉对当年6月份降水持续增加显示出明显的负相关关系,上年12月份的降水会对幼龄组和老龄组云杉径向生长不利。研究表明幼龄组云杉包含的气候信息要优于中龄组和老龄组云杉,在该区域进行相关研究时应根据研究需要选取不同年龄跨度的云杉年表。     

长白山北坡不同林型内红松年表特征及其与气候因子的关系

运用树木年轮学的基本原理和方法,选取了长白山北坡保存完好的典型性植被阔叶红松林,探讨了杨桦红松林和椴树红松林内建群种红松(Pinus koraiensis)径向生长对气候要素的响应.结果表明,长白山北坡红松的径向生长对降水较为敏感,杨桦红松林和椴树红松林中,红松年轮宽度均与当年7月以及上一年9月的降水呈显著正相关关系.不同林型内红松的生长与气候因子的关系也有差异.椴树红松的年轮宽度还与上年7月的降水显著负相关,与当年3、4月份的平均气温呈显著正相关.而杨桦红松林内红松年轮宽度和平均温度没有显著的相关关系.特征年分析进一步验证了响应函数相关分析的结果,即当年生长季以及上年生长季末的降水充足促进了红松的径向生长;椴树红松林中,初春温度的升高有利于红松的生长.     

Reproductive and genetic characteristics of rare,disjunct pitch pine populations at the northern limits of its range in Canada

Pitch pine, Pinus rigida Mill., is a rare species in Canada, existing as a disjunct population in the St. Lawrence River Valley in eastern Ontario and two northern outlier stands in southern Quebec along Canada's southern border with the United States. Reproductive and genetic characteristics of these small, scattered stands were investigated to develop a foundation for management and restoration in the event of range expansion northwards under anticipated climate warming. Seed yields and seed quality appear to be comparable to other eastern conifers, and to pitch pine at the center of its geographic range. For seed and seedling growth traits, most of the variation was attributable to differences among trees within stands and, to a lesser extent, among stands within a population; whereas the population effect was non-significant. For reproductive traits, such as numbers of filled and empty seeds per cone, reproductive efficiency, and inbreeding estimates, high levels of variation (ranging from 26% to 33%) were found among stands, suggesting that stand structural features, such as stand size and tree density within stands, play an important role in pollination environment and overall reproductive success. Estimates of genetic diversity at 32 allozyme gene loci indicate that these small, isolated stands have maintained relatively high levels of genetic diversity compared with populations at the center of its geographic range, and also relative to other widely dispersed eastern conifers. The relatively high levels of viable seed production and genetic diversity in native pitch pine populations indicate that native Canadian populations may be suitable seed sources for species restoration and range expansion in Canada.     

长白山红松年轮碳同位素与净初级生产力的关系

解析树木年轮碳同位素变化是揭示气候与环境变化的有效途径,然而少有研究分析年轮碳同位素对森林净初级生产力(NPP)的指示效应.基于生长季气象因子,分析了长白山红松年轮δ¹³C记录序列与阔叶红松林红松种群NPP变化趋势以及两者之间的相关关系.结果表明: 1970年以前,研究区红松年轮δ¹³C与红松种群NPP变化同步,且两者间呈极显著线性正相关关系,年轮δ¹³C记录了气候变化对红松种群NPP的影响;1970年以后,红松年轮δ¹³C与红松种群NPP变化出现分异,两者呈负相关关系但相关性不显著,表明干旱等其他环境因子的影响使年轮δ¹³C对气候变化的敏感性和对NPP指示性降低.这一相关关系在红松年轮δ¹³C与当年和下一年红松种群NPP之间都存在,表明当年环境状况对来年红松生长的重要意义.说明长白山红松年轮δ¹³C对红松种群NPP有良好的指示性,树木年轮δ¹³C有重建森林历史NPP长期变化的潜力.     

红松种源材质性状研究

对帽儿山地区17 年生26 个红松种源试验林进行了材质性状研究, 主要包括材质性状在不同种源间遗传变异情况以及从髓心到树皮径向变异模式研究。结果表明:红松各项材质在不同种源间均存在着极显著差异。各项材质均存在着一定的径向变异模式, 主要表现在:红松各种源的管胞长度、管胞直径、木材密度和晚材率总体趋势是从髓心向外呈迅速增加趋势, 而年轮宽从髓心向外呈递减趋势。本项研究为红松材性早期选择和预测提供依据。     

Exploring wood anatomy,density and chemistry profiles to understand the tree-ring formation in Amazonian tree species

Long-term analysis of tree growth using annual tree rings is increasingly in demand for tropical tree species. The basis of these studies has traditionally been the anatomical identification of the annual ring boundary. However, the structure of these annual rings has been sparsely explored for complementary physical and chemical wood traits. Here, we explore the relationships among wood density features and chemical elements (S, K, Ca, Mn) involved in the annual tree ring formation of 12 tropical tree species from non-flooded forest in the southern Amazon basin. Transverse wood sections were used for each species to determine: 1) macroscopic distinction (radial growth and wood density), 2) microscopic analyse of vessels, axial and ray parenchyma (anatomy) and 3) X-ray densitometry (physical) and X-ray fluorescence (chemical). For some species, the profiles of wood density, and Ca and Mn content showed intra- and inter-annual patterns that allowed to define and characterize the growth boundary of tree rings. Ca, K and S were mainly distributed in axial parenchyma cells, and around vessels, whereas, Mn was mainly distributed in fibres. Our results showed significant species-specific correlations between tree-ring width, density and concentrations of Ca, K and Mn. The anatomical characterization and the complementary information provided by the density and chemical profiles in some Amazonian species can represent a valuable proxy to improve the definition of annual ring-boundaries and improve the understanding of long-term growth and physiological patterns.