Climate impacts on lodgepole pine (Pinus contorta) radial growth in a provenance experiment

A potentially confounding radial-growth interaction exists at the intersection of two well-known principles, one in the field of dendrochronology and the other in quantitative genetics. From a dendrochronology perspective, tree populations growing in climatically marginal environments are expected to be more sensitive to seasonal and annual climate than those growing in optimal climate zones. From a genetics perspective, marginal populations may be adapted to grow a small amount each year and then shut down to prevent climate-induced mortality, or they may be adapted to respond to favourable climate conditions when available. We examined the relative strength of these forces using data from 12 populations of 34-year-old lodgepole pine (Pinus contorta) trees growing in 16 provenance-trial sites in western Canada. Growth generally correlated positively with annual temperature and negatively with summer aridity. The sensitivity of radial-growth to interannual climate fluctuations was both site and provenance-related, with the highest sensitivities occurring among populations from warm, central provenances growing at cold, marginal sites, and among populations from cold, marginal provenances growing at warm, central sites. The correlations between climate and growth varied regionally; notably, populations from warm provenances growing at warm sites responded more strongly to summer aridity, while populations from cold provenances growing at warm sites responded more to annual temperature. Our finding that sensitivity varied among populations growing under similar climate conditions indicates that sensitivity is influenced by genetics as well as by site climate, but the regional specificity of the growth responses did not support a single hypothesis for the influence of genetics on growth among populations from marginal vs. central locations. Implications of our study for forest productivity under climate change are more positive for trees growing in cool locations, where overall warmer temperatures will lead to increased growth, than in warm locations, where the negative effects of arid summers may counteract the positive effects of warmer annual temperatures.     

卡特兰雌配子的发育

采用石蜡切片法观察了卡特兰雌配子的发育过程。结果表明:卡特兰在授粉前,胎座和胚珠并没有开始发育;在成功授粉后,胚珠原基开始发育;卡特兰的胚珠为倒生型,具薄珠心和2层珠被,成熟胚囊为8核;从授粉到胚囊成熟需要112 d左右;外珠被的过早发育可能是造成胚败育的原因之一。     

翅果油树大小孢子发生及雌雄配子体发育

利用常规石蜡切片技术,对濒危植物翅果油树大、小孢子发生及雌、雄配子体发育的过程进行研究,探讨在其发育过程中是否存在生殖障碍。结果表明:翅果油树花药四囊型,花药壁由表皮、药室内壁、2层中层、绒毡层5层细胞组成,绒毡层为腺质型;小孢子母细胞减数分裂过程中的胞质分裂方式为同时型,小孢子四分体排列方式为四面体型,有时为交叉型或左右对称型;成熟花粉粒为2-细胞型,有3个萌发孔;子房上位,基生胎座,倒生胚珠,双珠被,厚珠心;胚囊发育属于蓼型。其雌、雄配子体发育过程多数正常。     

Critical Factors Affecting Ex Vitro Performance of Somatic Embryo Plants of Picea abies

The potential to use somatic embryos for large-scale propagation of elite genotypes, for integration into breeding programmes and for connecting breeding and mass propagation, is receiving much attention. However, before the methods are applied it is important that the plants regenerated via somatic embryogenesis grow as expected, i.e. as seedlings or cuttings. Growth of somatic embryo plants is under a cumulative influence of a number of treatments given during the in vitro phase and during the ex vitro establishment phase. The aim of this study was to identify treatments with a negative influence on the subsequent growth of somatic embryo plants of Norway spruce (Picea abies L. Karst.). Based on the results, the time of contact with abscisic acid during somatic embryo maturation and the length of continuous light treatment (CLT) during the first growth period strongly affect the height growth during two successive growth periods. In both cases longer treatments exerted negative effects. Based on these results a new method was set up, which includes: (1) prematuration treatment of the suspension culture in a growth regulator-free medium, by which the maturation step is synchronized and contracted; and (2) a two-phase germination treatment, first on a solidified medium and then in a liquid medium. This treatment avoids extended CLT during the first growth period. Another advantage of the two-phase germination treatment is a better root-system development. Somatic embryo plants produced according to this method can be transferred directly from in vitro conditions to the greenhouse.     

雷竹大孢子发生与雌配子体发育

利用常规石蜡切片技术对雷竹大孢子发生与雌配子体发育的过程进行研究。结果表明:雷竹单子房,子房1室,内具1个倒生胚珠;双珠被,厚珠心;大孢子母细胞由1个雌性孢原细胞直接发育而成,大孢子四分体呈线型;蓼型胚囊,成熟胚囊包括1个卵细胞、2个助细胞、2个极核组成的中央细胞以及多个反足细胞,助细胞具明显丝状器。雌配子体发育多数正常,不是造成雷竹结实率低的主要原因。     

Tree-rings reflect the impact of climate change on Quercus ilex L. along a temperature gradient in Spain over the last 100 years

We analyzed tree rings over the past 100 years to understand the response of Quercus ilex L. to climate change at four different sites along a temperature gradient in a highly anthropogenically transformed ecosystem. To test the hypothesis of a climate change related decrease in productivity at warmer sites, we discuss the effect of historical management on the growth of forest stands and the spatio-temporal variability of growth in response to climate, analyzing departures from linearity in that relationship. We reconstructed stand history and investigated past growth trends using tree-rings. Then we used a dendroecological approach to study the regional, local and age-dependent response to climate, analyzing the relationship between precipitation and tree growth using non-linear mixed models. Tree rings reflected the origin of the studied landscape, mainly a simplification of an original closed forest and progressive canopy opening for agrosilvopastoral purposes after the mid 1800s. As expected, trees were principally responding to water availability, and regional growth (as expressed by the first principal component from the matrix of chronologies) was highly responsive to hydrological year precipitation (r = 0.7). In this water limited ecosystem, the response of growth to precipitation was asymptotic and independent of age, but variable in time. Maximum growth was variable at the different sites and the non-linear function of growth saturated (i.e. reached an asymptote) at temperature dependent site specific precipitation levels within the range considered in the region to lead a shift towards deciduous species dominated woodlands (around 600 mm, variable with mean temperature). Only trees at warmer sites showed symptoms of growth decline, most likely explained by water stress increase in the last decades affecting the highly transformed open (i.e. low competition) tree structure. Stands at colder locations did not show any negative growth trend and may benefit from the current increase in winter temperatures. Coinciding with the decrease in productivity, trees at warmer sites responded more to moisture availability, exhibited a slower response to precipitation and reached maximum growth at higher precipitation levels than trees at colder sites. This suggests that warmer stands are threatened by climate change. The non-linear response of growth to precipitation described is meaningful for different ecological applications and provides new insights in the way trees respond to climate.     

Forest responses to climate change in the northwestern United States: Ecophysiological foundations for adaptive management

Climate change resulting from increased concentrations of atmospheric carbon dioxide ([CO₂]) is expected to result in warmer temperatures and changed precipitation regimes during this century. In the northwestern U.S., these changes will likely decrease snowpack, cause earlier snowmelt, increase summer evapotranspiration, and increase the frequency and severity of droughts. Elevated [CO₂] and warmer temperatures may have positive effects on growth and productivity where there is adequate moisture or growth is currently limited by cold. However, the effects of climate change are generally expected to reduce growth and survival, predispose forests to disturbance by wildfire, insects, and disease; and ultimately change forest structure and composition at the landscape scale. Substantial warming will likely decrease winter chilling resulting in delayed bud burst, and adversely affect flowering and seed germination for some species. The extent of these effects will depend on the magnitude of climate change, the abilities of individual trees to acclimate, and for tree populations to adapt in situ, or to migrate to suitable habitats. These coping mechanisms may be insufficient to maintain optimal fitness of tree populations to rapidly changing climate. Physiological responses to climatic stresses are relatively well-understood at the organ or whole-plant scale but not at the stand or landscape scale. In particular, the interactive effects of multiple stressors is not well known. Genetic and silvicultural approaches to increase adaptive capacities and to decrease climate-related vulnerabilities of forests can be based on ecophysiological knowledge. Effective approaches to climate adaptation will likely include assisted migration of species and populations, and density management. Use of these approaches to increase forest resistance and resilience at the landscape scale requires a better understanding of species adaptations, within-species genetic variation, and the mitigating effects of silvicultural treatments.     

Environmental Effects on Norway Spruce Needle Wax

In spring, tubular wax structures were observed on the needle surfaces of new Norway spruce needles and also on previous-year needles. In young plants, as in mature trees, the fusing and eroding of the wax tubes on needles of increasing age had taken place in relation to their exposure, particularly to wind and precipitation. The quantity of needle wax and certain fractions of the extracted wax differed in mature trees at two field sites and in young clonal plants. No difference in wax quantity and only small differences in wax composition occurred among young plants which had undergone different treatments (fumigation with O³, SO², ambient air, filtered air, or planting at the sites). Environmental effects on wax structure, wax quantity and the presence of certain compounds in the wax were identified.     

Drought sensitivity of Norway spruce is higher than that of silver fir along an altitudinal gradient in southwestern Germany

Context

For Central Europe, climate projections foresee an increase in temperature combined with decreasing summer precipitation, resulting in drier conditions during the growing season. This might negatively affect forest growth, especially at sites that are already water-limited, i.e., at low elevation. At higher altitudes trees might profit from increasing temperatures.

Aims

We analyzed variations in radial growth of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) along an altitudinal gradient from 400 until 1,140 m a.s.l. in the Black Forest, to assess climate responses with increasing elevation.

Methods

Climate–growth relationships were analyzed retrospectively using tree-ring and climate data. In total, we sampled stem discs of 135 trees to build 27 species- and site-specific chronologies (n _fir?=?13, n _spruce?=?14).

Results

Our results indicate distinct differences in climate–growth relations between fir and spruce along the gradient. Growth of high-altitude fir was positively related to temperature from January till March. Growth of low-altitude fir and spruce at all elevations was positively related to precipitation and negatively to temperature during the growing season, particularly in July. A self-calibrating Palmer drought severity index underlined summer drought sensitivity of these trees.

Conclusion

Overall, we found that climatic control of tree growth changes over altitude for fir. For spruce, a remarkable synchrony in growth variation and climate response was shown, which indicates that this species is drought sensitive at all studied elevations. In a future warmer climate, the growth of low-altitude fir and spruce along the entire studied gradient may be negatively affected in the Black Forest, if an increased evaporative demand cannot be compensated by increased water supply.     

Effects of climate on radial growth of Norway spruce and interactions with attacks by the bark beetle Dendroctonus micans (Kug., Coleoptera: Scolytidae): a dendroecological study in the French Massif Central

Samples of Norway spruce (Picea abies [L.] Karst) were dendrochronologically investigated in order to detect infestations by Dendroctonus micans (Kug.), the great spruce bark beetle (Col. Scolytidae), a relatively recent introduction to France. Uninfested natural forests located in the north-eastern French Alps and heavily infested plantations in the Ardèche region (Massif Central) were compared. The penetration holes bored in trunks by the bark beetle induced visible marks on wood, such as extreme ring width reductions, locally missing rings and crescent-shaped resin patches between consecutive rings that make possible a post-infestation dating.The outbreak began in 1979, 5 years prior to first insect visual detection by foresters. In the infested forest, tree basal area growth was not as sustained as in uninfested natural stands, but showed an inflection point at an unusually young tree age (from 30 to 40 years). Ring widths showing extreme synchronous radial growth reductions were caused either by excessively cold periods (e.g. in 1948, 1980, 1984, 1992) or by summer drought (as in 1986). Most of these weak growth years were shared with uninfested sites. In healthy forests, the consequences of extremely cold years were usually recorded only in high elevation stands, especially near the timberline, whereas summer drought effects were mostly visible in low altitude forests. By contrast, both phenomena were recorded in the infected Ardèche plantation. An analysis of tree-rings and monthly climate confirmed that Norway spruce growth in Ardèche plantations was reduced by excessively low minimum temperature during most parts of the year prior to ring formation, by higher than average maximum temperature during current spring and summer, and by drought in winter, spring and summer. Thus, the regional Ardèche climate with both cold winters and dry summers (especially in July) seems to weaken spruce trees planted there. Moreover, tree sensitivity to climate was found to be greatly enhanced by insect infestation. Such interactions between climatic stress and insect outbreak led to forest dieback in a 15–20-year period, when trees were still young (less than 70 years), and without any tree recovery. Therefore, in that region spruce plantations should be replaced by non-host species of Dendroctonus micans, especially where soil conditions may exacerbate drought effects.     

Incidence of Root and Butt Rot in Consecutive Rotations of Picea abies

The incidence of butt rot in two consecutive rotations of Norway spruce [Picea abies (L.) Karst.] in 28 permanent sample plots at four different sites in Denmark was evaluated. Incidence of butt rot was estimated by visual examination of stumps at final felling of the previous rotation and by examination of bore cores taken at the butt from a random sample of trees before first thinning of the subsequent rotation. There was no correlation between the incidence of butt rot at final felling of the previous rotation of Norway spruce and the incidence of butt rot at first thinning of the subsequent rotation of Norway spruce. The incidence of butt rot at final felling was between 19 and 100%, and at first thinning between 0 and 20%. The S-form of Heterobasidion annosum (Fr.) Bref. was the most commonly found decay-causing organism at all sites. Root systems of 28 trees without decay at stump height in the present rotation were excavated to estimate the incidence of root rot. Heterobasidion annosum was found in only one root. Resinicium bicolor (Alb. & Schw. ex Fr.) Parm. was found in 25% of the excavated root systems. The result of the study shows that the incidence of butt rot at first thinning of Norway spruce is not necessarily higher on sites where the previous rotation was heavily infected than on sites where infection in the previous rotation was low.     

西藏林芝地区气候变化趋势与森林资源消退关系研究

分析西藏林芝地区近40年以来气候变化趋势,结果表明,西藏林芝地区的气温呈逐渐变暖、降水量逐年增多的趋势.进一步分析了造成气候变化的主要因素为森林资源消退、旅游人数剧增、全球温室效应等,提出控制温室气体的排放,加大植树造林和封山育林力度等建议.     

西藏林芝地区气候变化趋势与森林资源消退关系研究

分析西藏林芝地区近40年以来气候变化趋势，结果表明，西藏林芝地区的气温呈逐渐变暖、降水量逐年增多的趋势．进一步分析了造成气候变化的主要因素为森林资源消退、旅游人数剧增、全球温室效应等，提出控制温室气体的排放，加大植树造林和封山育林力度等建议．     

Impact of elevated carbon dioxide concentration and temperature on bud burst and shoot growth of boreal Norway spruce

Effects of elevated temperature and atmospheric CO2 concentration ([CO2]) on spring phenology of mature field-grown Norway spruce (Picea abies (L.) Karst.) trees were followed for three years. Twelve whole-tree chambers (WTC) were installed around individual trees and used to expose the trees to a predicted future climate. The predicted climate scenario for the site, in the year 2100, was 700 micromol mol-1 [CO2], and an air temperature 3 degrees C higher in summer and 5 degrees C higher in winter, compared with current conditions. Four WTC treatments were imposed using combinations of ambient and elevated [CO2] and temperature. Control trees outside the WTCs were also studied. Bud development and shoot extension were monitored from early spring until the termination of elongation growth. Elevated air temperature hastened both bud development and the initiation and termination of shoot growth by two to three weeks in each study year. Elevated [CO2] had no significant effect on bud development patterns or the length of the shoot growth period. There was a good correlation between temperature sum (day degrees>or=0 degrees C) and shoot elongation, but a precise timing of bud burst could not be derived by using an accumulation of temperature sums.     

Occurrence of the moth Cydia pactolana is associated with the spruce canker fungus Neonectria fuckeliana

Several young damaged Norway spruce stands in eastern and central Finland were observed from 2013 to 2016. The damage included trees with heavy resin flow, necrotic foliage, stem and branch cankers and dead trees. Pest identification resulted in the tortricid moth Cydia pactolana whose occurrence was always associated with the presence of the ascomycete pathogen Neonectria fuckeliana. Both the insect and the disease contributed to the extent of the damage, but it is not possible to say in which order they had attacked the trees. Apparently, changed climate has affected the increased occurrence of both the fungus and the moth. However, the characteristics of the insect–fungus interaction and the factors contributing to the coincidences are unknown. Emerging coexistence or potential symbiosis of the two damaging agents is a serious threat for Norway spruce cultivation. Understanding the biology of this fungus–insect interaction is important for controlling them.     

Aspen Populus tremula is a key habitat for tree-dwelling bats in boreonemoral and south boreal woodlands in Norway

This study aims to identify which tree species bats use as maternity roosts in woodland in southern Norway and, to assess the effectiveness of bat boxes as potential mitigating measures if roosts are lost to forestry or other factors. Radio telemetry was used to find roosts in summer during late pregnancy and through the lactation period by tagging two common species; Daubenton's bats Myotis daubentonii (Kuhl 1817) and soprano pipistrelles Pipistrellus pygmaeus (Leach 1825). In addition, tree roosts were searched using ultrasound detectors at three sites. A total of 35 roosts in natural crevices were found using radio telemetry and through searches with ultrasound detectors and 31 of these roosts were in hollow trees, the majority in aspens Populus tremula L. (n?=?30). Some tree roosts were even used by the northern bat Eptesicus nilssonii (Keyserling &; Blasius 1839) which is considered a house-dwelling species in Norway. Aspens have some qualities preferred by bats and were both warmer and safer than other hollow tree species in this study area. Bat boxes are used as maternity roosts mainly by soprano pipistrelles at these latitudes, thus at present, there is no suitable alternative to hollow trees that would satisfy all tree-dwelling bat species.     

Modelling above and below ground carbon dynamics in a mixed beech and spruce stand influenced by climate

Tree growth and carbon dynamics are important issues especially in the context of climate change. However, we essentially lack knowledge about the effects on carbon dynamics especially in mixed stands. Thus, the objective of this study was to test the effects of climatic changes on the above and below ground carbon dynamics of a mixed stand of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) by means of scenario simulations. To account for the typical tree interactions in a mixed-species stand a spatial explicit tree growth model based on eco-physiological processes was applied. Three different climate scenarios considering altered precipitation, temperature, and radiation were calculated for an unthinned and a thinned stand. The results showed significant changes of above and belowground biomass over time, especially when temperature and radiation were increased additionally to decreased precipitation. The reduction in biomass increments of Norway spruce were more attenuated above than below ground. In contrast, the results for beech were the opposite: The belowground increments were reduced more. These results suggest a shift in the species contribution to above and belowground biomass under dryer and warmer conditions. Distinct effects were also found when thinned and unthinned stands were compared. A reduced stand density changed the proportions of above and below ground carbon allocation. As a main reason for the changed growth reactions the water balance of trees was identified which lead to changed biomass allocation pattern. This article belongs to the special issue “Growth and defence of Norway spruce and European beech in pure and mixed stands”.     

Silvicultural alternatives in an uneven-sized forest dominated by Picea abies

In this article, we report on the increments in basal area and tree diameter as well as the structural development observed in variously thinned plots that underwent either uneven- or even-sized treatment. The experimental forest was originally an uneven-sized mixed stand dominated by Picea abies. Twenty-eight randomized sample plots underwent each treatment, and the trees were monitored for 15 growing seasons after thinning. The uneven-sized plots retained a reverse J-shaped diameter distribution, but this was changed into a bell shape by low thinning in the even-sized plots. Absolute basal area increment was positively correlated with basal area in the even-sized treatment but not in the uneven-sized treatment. In the latter, all of the plots grew almost equally well, and only the basal area of broadleaves explained slightly positively the increment variation. Relative basal area increment was negatively correlated with basal area in both treatments. Additionally, the basal area of Scots pine was a positive explanatory variable in the relative increment variation in the even-sized treatment. For the dominant Norway spruce trees, diameter increment was negatively correlated with basal area in both treatments and, conversely, heavy removal increased the diameter increment. Relative basal area increment averaged 5% annually in uneven-sized plots representing the “target selection.” This was more than double the increment observed for the even-sized plots that represented the “prevailing practice.” Likewise, the diameter increment of Norway spruce trees was 48% greater in the uneven-sized compared to the even-sized plots.     

Pre-commercial thinning,birch admixture and sprout management in planted Norway spruce stands in South Sweden

Early management of the regenerated seedlings shapes the future stand properties. To address these issues, pre-commercial thinning (PCT) and control treatments were applied to planted Norway spruce (Picea abies L. Karst) and naturally regenerated birch (Betula pendula Roth., Betula pubescens Ehrh.) stands in forest experiments in southern Sweden (lat. 56–57?N) containing 1.1–5.5?m tall saplings. The treatments were retention of 1000 or 2000?stems?ha^?1 of Norway spruce, with no birch or birch at 1000?stems?ha^?1. Treatments were replicated with and without annual removal of birch sprouts from stumps. The periodic annual increment (PAI) over five years was calculated for total stand volume and individual trees. The mean PAI of dominant trees was significantly higher both following all PCT treatments than controls, and following low rather than high-density PCT. Birch retention did not affect growth of the dominant trees but PAI was lower in plots with uncontrolled sprouting. The PAI of birch was significantly higher in low-density Norway spruce plots than in control plots and the high-density plots. The treatment response was significant even in stands with initial heights of only 1–2?m.     

Short-term effects of whole-tree harvesting on understory plant species diversity and cover in two Norway spruce sites in southern Norway

Effects of clear-cut harvesting on ground vegetation plant species diversity and their cover are investigated at two Norway spruce sites in southern Norway, differing in climate and topography. Experimental plots at these two sites were either harvested conventionally (stem-only harvesting) or whole trees including crowns, twigs and branches were removed (whole-tree harvesting), leaving residue piles on the ground for some months. We compare the number of plant species in different groups and their cover sums before and after harvesting, and between the different treatments, using non-parametric statistical tests. An overall loss of ground vegetation biodiversity is induced by harvesting and there is a shift in cover of dominant species, with negative effects for bryophytes and dwarf shrubs and an increase of graminoid cover. Differences between the two harvesting methods at both sites were mainly due to the residue piles assembled during whole-tree harvesting and the physical damage made during the harvesting of residues in these piles. The presence of the residue piles had a clear negative impact on both species numbers and cover. Pile residue harvesting on unfrozen and snow-free soil caused more damage to the forest floor in the steep terrain at the western site compared to the eastern site.