Climatic impacts and drought control of radial growth and seasonal wood formation in Pinus halepensis

Short- and long-term growth responses to drought and climatic influences still remain poorly understood. In this study, we investigated the impact of climatic drivers (temperature, precipitation) and drought, using the Standardized Precipitation Index (SPI) calculated at different time scales (1–48?months), on earlywood (EW) and latewood (LW) widths in Pinus halepensis. Nine forests subjected to dry summer conditions were sampled in Mediterranean semi-arid areas from north-eastern Spain. In addition, we explored the seasonal dynamics of cambial activity and wood formation in relation to short-term climate variability. We found two peaks of tracheid cell production corresponding to EW (May–June) and LW (mid-July–August) growth phases, associated with a sharp decrease in enlarging cells in early July in response to low water availability. In the period of analysis (1970–2005), EW growth was positively correlated with precipitation in previous December and current January, April, May and June, while it was negatively correlated with temperature in June and July. LW was correlated positively with minimum temperatures in January. Probably this was an indirect relationship as a consequence of increased EW width at higher January temperatures. Drought affected more negatively EW than LW formation as evidenced the higher SPI-EW correlation (r?=?0.72) than the SPI-LW one (r?=?0.54). The strongest EW response to drought was observed in July, whereas the highest LW response to drought occurred in September; and this seasonal pattern matched the phases of lowest EW and LW tracheid production. Under a future reduction of winter and spring precipitation, the studied forests may show a decrease in tracheid cell production, causing a decline of radial growth, a reduction in hydraulic conductivity and, indirectly, a hampered carbon uptake in such semi-arid woodlands.     

Reconstruction of pre-monsoon weather conditions in northwestern Thailand from the tree-ring widths of Pinus merkusii and Pinus kesiya

Two pine species, Pinus merkusii and Pinus kesiya, native in Thailand were studied for their potential to reconstruct past weather conditions from their tree rings. Altogether, cores from 209 Merkus pines and from 205 Khasi pines were sampled at 16 sites. Standard methods were applied to assemble tree-ring chronologies for each site and tree species. The longest site chronology of Merkus pine and Khasi pine covered 314 and 183 years, respectively. Principal component analysis (PCA) illustrated a close clustering of all, except two, site chronologies on the PC1 axis, indicating a strong signal common for both pine species throughout NW Thailand. However, along the PC2 axis, there was a distinct separation between the Merkus pine and the Khasi pine site chronologies, which was possibly due to the species-specific differences in site altitudes. Simple correlations between the PC1 time series and a regionalized climate data set showed a highly negative association with temperature during the pre-monsoon period from March to May. Since, pre-monsoon temperature and rainfall were negatively correlated with each other, we used the PC1 as a direct proxy for pre-monsoon weather conditions (warm/dry vs. cool/wet) back to 1834 AD.     

Relationship between wood anatomy,tree-ring widths and wood density of Pinus sylvestris L. and climate at high latitudes in northern Sweden

In this study, wood anatomy, tree-ring width and wood density of Pinus sylvestris at the northern timberline in Fennoscandia were used to identify relationships among the parameters and to screen them for their climatic signals. Furthermore we investigated the influence of the juvenile wood section for all parameters developed. The measurements of wood anatomy were conducted with confocal laser scanning microscopy (CLSM) while the density profiles were produced using an Itrax MultiScanner. We developed chronologies of ring width, wood density and anatomy for a period between 1940 and 2010. Correlations between wood density and wood anatomy were strong in the latewood part. For some wood anatomy and density chronologies youth trends were found in the juvenile part. Wood density decreased from the pith up to the 9th ring and stabilized afterwards, while cell lumen diameter and lumen area increased simultaneously up to the 15th ring. All chronologies contained strong summer temperature signals. The wood anatomical variables provided additional information about seasonal precipitation which could not be found in wood density and tree-ring widths. Our study confirmed previous results stating that the parameter maximum density contains the strongest climate signal, that is, summer temperatures at the northern timberline. Nevertheless, the intra-annual data on tracheid dimensions showed good potential to supply seasonal climatic information and improve our understanding of climatic effects on tree growth and wood formation.     

The environmental signals of stable carbon isotope in various tree-ring components of Pinus tabulaeformis

Chinese pine (Pinus tabulaeformis) trees were sampled in the Helan Mountain, northwest China. The stable carbon isotope (δ¹³C) values of whole wood, holocellulose and alpha-cellulose in tree rings over 30 years (1968–1997) were measured to study the δ¹³C response of different tree-ring components to past environmental change. There were obvious differences in the δ¹³C values of the three components. The Pearson correlation coefficient between the δ¹³C of alpha-cellulose and that of holocellulose was 0.547 (ρ < 0.01); between alpha-cellulose and whole wood, the coefficient was −0.126 (ρ > 0.10); between holocellulose and whole wood, the coefficient was −0.056. Correlation function analyses indicated that the δ¹³C content of tree-ring alpha-cellulose correlated strongly with the average temperature from June to August (r = 0.427, ρ < 0.05), more than that of holocellulose (0.324, ρ < 0.10) or total wood (−0.245, ρ > 0.10). Significant correlations were observed between δ¹³C of tree-ring alpha-cellulose and the precipitation from the current year’s February to July (r = −0.514, ρ < 0.01) that were much higher than that of holocellulose (−0.481, ρ < 0.05) or total wood (−0.249, ρ > 0.10). A significant correlation (−0.545, ρ < 0.01) was also found between the ring width and the δ¹³C residual chronologies. These results suggest that more past environmental information is retained in the δ¹³C of tree-ring alpha-cellulose. Thus, the δ¹³C of alpha-cellulose of tree rings is the most suitable among the studied parameters for reconstructing the past climatic conditions during the growing season. The δ¹³C values of other organic compounds in Pinus tabulaeformis xylem were affected by the external environment after carbon was fixed from the atmosphere.     

Climatic impact on tree-ring widths in Abies borisii-regis forests from South-East Albania

The long-term radial growth responses to drought and climatic variability of less-studied species such as Abies borisii-regis (Mattf.) remain poorly understood.We tested the hypothesis that severe short-term drought conditions during summer months will impact the radial growth of A. borisii-regis (Mattf.) trees and such impact will have a more pronounced effect on latewood (LW) than earlywood (EW) width.Correlation analysis was employed to investigate the impact of climatic drivers (temperature, precipitation) and drought, using the Standardized Precipitation Evapotranspiration Index (SPEI) calculated at cumulative time scales (1–12 months), on EW, LW and tree-ring width (TRW) in A. borisii-regis (Mattf.) trees from South-Eastern Albania.We found that EW width was positively correlated with precipitation in July and previous September, while the LW width and TRW was enhanced by the current June–July precipitation. Previous autumn and current summer high temperatures constrained the radial growth in A. borisii-regis (Mattf.) trees, particularly the LW and TRW. All the tree-ring widths components showed the highest significant response to drought at short cumulative time scales (<4 months) mainly during July, August and September. The highest impact of drought was observed for the LW width.Under a future reduction of summer precipitation and temperature increase, the A. borisii-regis (Mattf.) may show a decrease in EW formation, causing a decline of radial growth, leading to a reduction in hydraulic conductivity and carbon uptake in these forests.     

Isotopic and anatomical signals for interpreting fire-related responses in Pinus halepensis

Key message

Our study aims to define isotopic and anatomical responses to fires of P. halepensis . Main results: decrease in tree growth and relative conductivity and increase in water use efficiency.

Abstract

We investigated the ecophysiological responses of a Pinus halepensis Mill. stand surviving two wildfires in southern France. Basal area, isotope composition and anatomical traits were analysed before and after fires, using tree rings to assess the ecological responses of trees to heat-related damage. The years were determined based on the presence of fire scars. Stable isotopes (δ¹³C and δ¹⁸O) were measured in tree rings before and after the “fire years”. Anatomical observations allowed qualitative analysis of the scar region and quantification of tracheid size in tree rings before and after the fire years. Relative and percentage conductivity of earlywood and latewood far from the woundwood were estimated. Results showed a decrease in tree growth after the fire events accompanied by an increase in ¹³C-derived water use efficiency (WUE_i) and a decrease in relative conductivity. The positive relationship between δ¹³C and δ¹⁸O suggested that both isotopic variations are mostly driven by changes in stomatal conductance following fire events. P. halepensis proved to be a strong isohydric species, able to survive frequent fires with temporary ecophysiological modifications and anatomical adaptations. Our findings afford new insights into post-fire survival strategies of this species in an environment where fires are predicted to increase in frequency during the twenty-first century.     

Physiological response of Pinus halepensis needles under ozone and water stress conditions

The aim of this study was to evaluate how physiological processes of potted Pinus halepensis plants, grown under controlled conditions, were affected by ozone (O₃) and/or water stress, integrating the gas exchange and biochemical data with fluorescence OJIP polyphasic transient data. Plants submitted to only water stress (T₁) and with ozone (T₃) showed a strong decrease in stomatal conductance and gas exchange, coinciding with a reduction of maximum yield of photochemistry ( φ _po) and very negative values of leaf water potential. Simultaneously, a great increase of both PSII antenna size, indicated by absorption per reaction centre, and electron transport per reaction centre were found. The reduction of photosynthesis in the O₃-treated plants (T₂) by a slowing down of the Calvin cycle was supported by the increase of related fluorescence parameters such as relative variable fluorescence, heat de-excitation constant, energy de-excitation by spillover, and the decrease of φ _po. We suggest an antagonistic effect between the two stresses to explain the delayed ozone-induced decrease of stomatal conductance values for T₃ with respect to T₁ plants, by an alteration of the physiological mechanisms of stomatal opening, which involve the increase of intra-cellular free-calcium induced by ABA under co-occurring water shortage. We emphasise the importance of considering the intensity of the individual stress factor in studies concerning the interaction of stresses.     

Trophic plasticity of Antarctic echinoids under contrasted environmental conditions

Echinoids are common members of Antarctic zoobenthos, and different groups can show important trophic diversity. As part of the ANT-XXIX/3 cruise of RV Polarstern, trophic plasticity of sea urchins was studied in three neighbouring regions (Drake Passage, Bransfield Strait and Weddell Sea) featuring several depth-related habitats offering different trophic environments to benthic consumers. Three families with contrasting feeding habits (Cidaridae, Echinidae and Schizasteridae) were studied. Gut content examination and stable isotopes ratios of C and N suggest that each of the studied families showed a different response to variation in environmental and food conditions. Schizasteridae trophic plasticity was low, and these sea urchins were bulk sediment feeders relying on sediment-associated organic matter in all regions and/or depth-related habitats. Cidaridae consumed the most animal-derived material. Their diet varied according to the considered area, as sea urchins from Bransfield Strait relied mostly on living and/or dead animal material, while specimens from Weddell Sea fed on a mixture of dead animal material and other detritus. Echinidae also showed important trophic plasticity. They fed on various detrital items in Bransfield Strait, and selectivity of ingested material varied across depth-related habitats. In Weddell Sea, stable isotopes revealed that they mostly relied on highly ¹³C-enriched food items, presumably microbially reworked benthic detritus. The differences in adaptive strategies could lead to family-specific responses of Antarctic echinoids to environmental and food-related changes.     

油松(Pinus tabulaeformis)树轮宽度与气候因子统计相关的生理机制--以贺兰山地区为例

通过分析贺兰山地区气候因子的变化特征,并且参考北美、俄罗斯的研究结果,确定了Vaganov-Shashkin生理响应模型的参数,并根据研究区的实际情况对参数作了相应修正,得出两组不同的参数,两组参数拟合出相同的结果,说明模型内在的机理是稳定的,更能反映出它的生理学特性.对树轮宽度序列和气候因子进行相关分析,发现比较明显的相关有与5～8月份降水的正相关,与5～8月份温度的负相关,与9、10月份温度的正相关.模型分析结果认为,从5月中旬到8月底,降水不能满足树木生长的需要,故形成树轮宽度与该时段降水的正相关,而在这个时段,由于温度和降水呈负相关,表现为树轮宽度与温度的负相关,其实这在很大程度上是一种间接的相关.到9月份,由于温度下降比较明显,这时树木生长的主要控制因素已转化为温度,所以形成了与9、10月份温度的正相关.模型分析还发现,树轮宽度的形成与生长结束时间相关性强,而与生长开始时间的关系不明显.     

Relations between apical structure and growth patterns in Pinus halepensis Mill,seedlings

Abstract

The process of primary growth in 2-year-old seedlings of 11 populations of Pinus halepensis Mill. is described. At the end of the first growing season one type of apical structure was observed: type-1, a tuft of primary needles placed close together, surrounding and protecting a meristematic apex.

At the end of the 2nd growing season, three types of apical structure were observed: type-1; type-2, a terminal winter bud; and type-3; a «bud» with characteristics of both type-1 and type-2.

Morphological observation along with an anatomical examination of the winter bud led to the conclusion that the definitive growth pattern in juvenile P. halepensis is monocyclic with a variable number of summer shoots. This growth pattern is reached by some P. halepensis populations in 3–4 years, by contrast, in other pine species two years are usually needed.

The populations studied differed both in growth potential (differences in number of cycles, ratio of first cycle to total growth, growth rates) and in the developmental stages of the apical meristem.

Four groups could be identified: (i) Morocco and Spain, (a limited growth, few cycles, a high ratio of 1st cycle to total growth, and growth in 2nd season almost entirely due to free growth); (ii) Algeria and Greece, (moderate to low growth, a large number of cycles, a low ratio of 1st cycle to total growth, and very early formation of apical structure with preformed primordia); (iii) Israel and Central Italy (a high growth, a large number of cycles, a medium ratio of 1st cycle to total growth, and early formation of apical structure with preformed primordia); (iiii) Greece, France and Italy, which was intermediate between group (i) and the other 2 groups.     

Climatic control of intra-annual wood density fluctuations of Pinus pinaster in NW Spain

Tree rings of Pinus pinaster often contain intra-annual density fluctuations (IADFs), which have been attributed to the succession of dry and rainy periods typical of Mediterranean climate, but their formation has not been studied yet under Atlantic climate. We analyzed the occurrence and climatic significance of replicated IADFs in ten monospecific stands in NW Spain. The frequency of IADFs was higher than previously reported for this species under Mediterranean conditions and consistently decreased with increasing elevation. The formation of bands of latewood-like tracheids within the earlywood was favored by dry previous August, cold previous winter and dry April. Bands of earlywood-like tracheids within the early latewood were also favored by low winter temperatures. However, their occurrence was geographically heterogeneous, with two groups of stands being defined by their distances to the shoreline. In coastal stands, cold May–August triggered IADFs formation, while in inland stands their formation was favored by dry May–July. Regional winter temperatures and April water balance were strongly related to the East Atlantic (EA) pattern, which greatly conditioned the occurrence of IADFs in the earlywood and the early latewood. By contrast, the presence of bands of earlywood-like tracheids in the late latewood was independent of the EA pattern, being strongly related to warm conditions in spring and especially to a wet October. The link between regional climate and the EA pattern strongly controlled the physiological processes that determine intra-annual growth dynamics and short-term cell enlargement of P. pinaster in NW Spain.     

Influence of micro-site conditions on tree-ring climate signals and trends in central and northern Sweden

Tree-ring chronologies are important indicators of pre-instrumental, natural climate variability. Some of the longest chronologies are from northern Fennoscandia, where ring width measurement series from living trees are combined with series from sub-fossil trees, preserved in shallow lakes, to form millennial-length records. We here assess the recent ends of such timeseries by comparing climate signals and growth characteristics in central and northern Sweden, of (1) trees growing at lakeshore micro-sites (representing the source of sub-fossil material of supra-long chronologies), with (2) trees collected in dryer micro-sites several meters “inland”. Calibration trials reveal a predominating June–September temperature signal in N-Sweden and a weaker but significant May–September precipitation signal in C-Sweden. At the micro-site level, the temperature signal in N-Sweden is stronger in the lakeshore trees compared to the inland trees, whereas the precipitation signal in C-Sweden remains unchanged among the lakeshore and inland trees. Tree-rings at cambial ages >40 years are also substantially wider in the lakeshore micro-site in C-Sweden, and juvenile rings are more variable (and wider) in the dryer micro-site in N-Sweden (compared to the adjacent micro-sites). By combining the data of the various micro-sites with relict samples spanning the past 1,000 years, we demonstrate that growth rate differences at the micro-site scale can affect the low frequency trends of millennial-length chronologies. For the supra-long chronologies from northern Fennoscandia, that are derived from sub-fossil lake material, it is recommended to combine these data with measurement series from only lakeshore trees.     

中国祁连山西段青海云杉树轮密度记录的气候信号

树轮气候学是研究过去气候变化的重要手段之一。以往研究表明,树轮密度是生长季温度的重要代用资料。本文建立了祁连山西段青海云杉132～135年的树轮最大密度年表、树轮最小密度年表、树轮晚材平均密度年表和树轮早材平均密度年表,比较了不同密度年表指示的最优气候信号及其季节组合,评估了其作为气候代用资料的潜力与价值。结果表明:树轮晚材平均密度和树轮早材平均密度对于树木生长季气候信号的响应远低于树轮最大密度和树轮最小密度;树轮早材平均密度和树轮最小密度与帕尔默干旱指数有很强的联系,表明树轮早材平均密度和树轮最小密度有成为干旱代用指标的潜力。     

A 323-year long reconstruction of drought for SW Romania based on black pine (Pinus Nigra) tree-ring widths

Increase in temperature and decrease in precipitation pose a major future challenge for sustainable ecosystem management in Romania. To understand ecosystem response and the wider social consequences of environmental change, we constructed a 396-year long (1615–2010) drought sensitive tree-ring width chronology (TRW) of Pinus nigra var. banatica (Georg. et Ion.) growing on steep slopes and shallow organic soil. We established a statistical relationship between TRW and two meteorological parameters—monthly sum of precipitation (PP) and standardised precipitation index (SPI). PP and SPI correlate significantly with TRW (r?=?0.54 and 0.58) and are stable in time. Rigorous statistical tests, which measure the accuracy and prediction ability of the model, were all significant. SPI was eventually reconstructed back to 1688, with extreme dry and wet years identified using the percentile method. By means of reconstruction, we identified two so far unknown extremely dry years in Romania—1725 and 1782. Those 2 years are almost as dry as 1946, which was known as the “year of great famine.” Since no historical documents for these 2 years were available in local archives, we compared the results with those from neighbouring countries and discovered that both years were extremely dry in the wider region (Slovakia, Hungary, Anatolia, Syria, and Turkey). While the 1800–1900 period was relatively mild, with only two moderately extreme years as far as weather is concerned, the 1900–2009 period was highly salient owing to the very high number of wet and dry extremes—five extremely wet and three extremely dry events (one of them in 1946) were identified.     

Morphological and Chemical Differentiation between Tunisian Populations of Pinus halepensis,Pinus brutia,and Pinus pinaster

The lipid fraction of seeds from different pine species and populations was studied regarding total lipid content, fatty acid profile and vitamin E composition. The investigated seeds contained a high percentage of lipid (13.6 to 31.5 %). Lipid fractions were found to be rich in vitamin E, which varied significantly among species and populations. P. halepensis (Ph−Hn) showed the highest content of vitamin E (256.3 mg/kg of seeds) and the uppermost content of α-tocopherol (44 mg/kg). However, P. halepensis (Ph−Kas) was the richest in γ-tocopherol (204.9 mg/kg). Lipid fractions had a low content of δ-tocopherol (1.2 to 3.6 mg/kg. The highest content of γ-tocotrienol (∼18 %) was determined for P. halepensis (Ph−Dc and Ph−Hn). Thirteen fatty acids were identified by GC-FID with significant variation between the investigated species. The linoleic acid was the major fatty acid followed by oleic acid and palmitic acid. The chemical differentiation among species for the composition of fatty acids and vitamin E was confirmed by PCA. Significant correlations were observed between the content of vitamin E and fatty acids and ecological parameters of P. halepensis populations.     

Comparative root system structure of post-fire Pinus halepensis Mill. and Cistus monspeliensis L saplings.

To describe root system topology of Pinus halepensis and Cistus monspeliensis saplings co-inhabiting natural post-fire sites, 55 P. halepensis and 26 C. monspeliensis saplings were extracted by the total excavation method from a burnt pine stand. Seedlings were individually labelled when emerging after fire and extracted three years later, at the sapling phase. In order to evaluate the effect of inter-specific competition of C. monspeliensis on P. halepensis root system, a stratified sampling was carried out according to density and height of the saplings. Topological parameters considered in the analysis were magnitude, total external pathlength, and altitude of the root systems. Weight and length of roots were also measured in order to estimate the specific root length, an index commonly used in morphological studies. Results clearly evidenced greater variability in root system topology of P. halepensis than C. monspeliensis saplings. Herringbone architecture (i.e., the most ordered pattern possible, with branching confined to the main axis) characterised small pine saplings, regardless of competition from C. monspeliensis, which changed to random branching in large saplings. In medium sized saplings, the root system was affected by inter-specific competition, which delayed changes in root branching. In contrast, C. monspeliensis invariably adopted randomly branched architecture, regardless of intra-specific competition. It is concluded that such different topological patterns make C. monspeliensis more competitive during the early stages of post-fire succession, because its root system is much more transport-efficient in the nutrient-rich environment. Those pines which finally branch roots by random pattern will reach higher stem height and magnitude, a factor which allows them to successfully compete with C. monspeliensis for soil nutrients and water. The morphological analysis showed a significant increase in the specific root length with competition, both in P. halepensis and C. monspeliensis saplings, which could be interpreted as a consequence of the reduction of root diameter in response to nutrient depletion.