Adjustment Capacity of Maritime Pine Cambial Activity in Drought-Prone Environments

Intra-annual density fluctuations (IADFs) are anatomical features formed in response to changes in the environmental conditions within the growing season. These anatomical features are commonly observed in Mediterranean pines, being more frequent in younger and wider tree rings. However, the process behind IADF formation is still unknown. Weekly monitoring of cambial activity and wood formation would fill this void. Although studies describing cambial activity and wood formation have become frequent, this knowledge is still fragmentary in the Mediterranean region. Here we present data from the monitoring of cambial activity and wood formation in two diameter classes of maritime pine (Pinus pinaster Ait.), over two years, in order to test: (i) whether the differences in stem diameter in an even-aged stand were due to timings and/or rates of xylogenesis; (ii) if IADFs were more common in large trees; and (iii) if their formation is triggered by cambial resumption after the summer drought. Larger trees showed higher rates of cell production and longer growing seasons, due to an earlier start and later end of xylogenesis. When a drier winter occurs, larger trees were more affected, probably limiting xylogenesis in the summer months. In both diameter classes a latewood IADF was formed in 2012 in response to late-September precipitation, confirming that the timing of the precipitation event after the summer drought is crucial in determining the resumption of cambial activity and whether or not an IADF is formed. It was the first time that the formation of a latewood IADF was monitored at a weekly time scale in maritime pine. The capacity of maritime pine to adjust cambial activity to the current environmental conditions represents a valuable strategy under the future climate change conditions.     

Growth reactions of Pinus sylvestris L. and Quercus pubescens Willd. to drought years at a xeric site in Valais, Switzerland

In Valais, an inner-Alpine dry valley in Switzerland, low-elevation Scots pine (Pinus sylvestris L.) forests are changing. While pine shows high mortality rates, deciduous species, in particular pubescent oak (Quercus pubescens Willd.), are becoming more abundant. We hypothesise that increasing drought and the species-specific drought tolerance are key factors in these processes. In this study, the growth reaction to drought years of pine and oak growing at a xeric site in Valais was analysed using dendrochronological and wood anatomical methods. Congruent with theoretical expectations, the tree-ring widths of both species, the mean lumen area of earlywood vessels in oak and the number of tracheids in a radial row in pine decreased in response to dry conditions. However, both species also showed reactions deviating from those known from mesic sites: In oak, the mean lumen area of latewood vessels increased in drought years. In pine, in the driest year of the period (1976), the mean radial diameter increased in latewood and decreased only slightly in earlywood. These results emphasises that the process of wood formation and cell functionality at xeric sites is not completely understood yet. Both species seem to have difficulties to adapt the size of their water-conducting cells to strongly reduced water availability in drought years. Additionally, the cell number is strongly reduced. Thus it remains unclear if both species can maintain sufficient water transport under increasingly dry conditions.     

Wood borer performance and wood characteristics of drought‐stressed Scots pine seedlings

Four‐year‐old Scots pine [Pinus sylvestris L. (Pinaceae)] seedlings were exposed to medium and severe drought stress for two consecutive years. The anatomical properties of drought‐stressed Scots pine wood and their impact on the performance of destructive wood boring early instars of Hylotrupes bajulus L. (Coleoptera: Cerambycidae) were studied. Drought stress significantly decreased diameter of earlywood tracheids in both growing years and diameter of latewood tracheids after the second growing season only. Cell lumen area was significantly decreased by both medium and severe drought stress compared to well‐watered controls. In addition, area of cell lumen was significantly smaller in severe drought than in medium drought treatment. The drought stress marginally increased the number of resin canals in the wood, but did not affect the size of resin canals either in wood or bark. The relative growth rate of xylophagous H. bajulus neonatal larvae was not significantly affected by drought stress during the 106‐day feeding period on Scots pine wood blocks. The results show that although water availability was an important factor affecting the development and anatomy of wood cells, observed changes in wood characteristics did not affect the performance of early instars feeding on wood processed from drought‐stressed young Scots pine seedlings.     

Growth patterns in relation to drought-induced mortality at two Scots pine (Pinus sylvestris L.) sites in NE Iberian Peninsula

Drought-related tree mortality has become a widespread phenomenon. Scots pine (Pinus sylvestris L.) is a boreal species with high ecological amplitude that reaches its southwestern limit in the Iberian Peninsula. Thus, Iberian Scots pine populations are particularly good models to study the effects of the increase in aridity predicted by climate change models. A total of 78 living and 39 dead Scots pines trees were sampled at two sites located in the NE of the Iberian Peninsula, where recent mortality events have been recorded. Annual tree rings were used to (1) date dead trees; (2) investigate if there was an association between the occurrence of tree death and severe drought periods characterized by exceptionally low ratios of summer precipitation to potential evapotranspiration (P/PET); and (3) to compare the growth patterns of trees that died with those of surviving ones. Mixed models were used to describe the relationships between tree growth (in terms of basal area increment, BAI, and the percentage of latewood, LW%) and climate variables. Our results showed a direct association between Scots pine mortality and severe drought periods characterized by low summer water availability. At the two sites, the growth patterns of dead trees were clearly distinguishable from those of the trees that survived. In particular, the BAI of dead trees was more sensitive to climate dryness (low P/PET_summer, high temperatures) and started to decline below the values of surviving neighbors 15–40 years before the time of death, implying a slow process of growth decline preceding mortality.     

Living on the edge: Legacy of water availability on Tetraclinis articulata secondary growth under semiarid conditions in Morocco

Tetraclinis articulata is a xerothermic Mediterranean conifer native from NW Africa that grows under semiarid conditions in the Atlas Mountains. This species is particularly well adapted to water scarcity and forms xeric woodlands subjected to recurring drought, aridification and overexploitation. Wood of T. articulata has been considered of limited dendrochronological value due to abundant anatomical anomalies in their growth rings. We studied tree-ring growth and tree-ring features to check its dendrochronological potential and evaluate their climatic signals and drought legacy effects at two semiarid sites with contrasting elevation in Morocco. Tree-ring boundaries were properly identified on wood cores from 60 to 68% of the sampled trees. Intra-annual wood density fluctuations (IADF), microrings, missing rings and rings with undefined limits were most abundant at the low-elevation coastal site than at the mountain site. Microring and missing ring abundance was inversely related to tree-ring growth, independent of cambial age, and mostly dependent on drought occurrence. IADF frequency was independent of tree-ring growth but inversely related to cambial age. A complex response of IADF formation to cool March to June conditions was found at the low-elevation coastal site, while IADF formation in the mountain site was related to May water availability. Undefined ring limits were observed only at the low-elevation site, showing positive relationships with elevated maximum temperatures in previous winter. Previous winter rainfall favored radial growth, but a significant legacy effect of water availability was evident up to three years prior to growth. Tree-ring growth was mostly related to water availability 33 and 26 months prior to growth at low-elevation and mountain sites, respectively. The obtained results reveal a very plastic cambial activity and a strong ability of T. articulata to withstand drought even for long periods.     

How needle phenology indicates the changes of xylem cell formation during drought stress in Pinus sylvestris L.

Temperature, water availability and photoperiod are the primary drivers of tree phenological processes. However, there is lack of information how the spring temperature and summer drought affect phenology of both cambium and needles. We evaluated the interplay between xylem and needle development of mature Scots pine trees in three consecutive years differing in the spring temperatures and onset and duration of summer drought. Cambial activity began on the day of a year (DOY) 83–87, while the bud break occurred on DOY 113–119, when also the secondary cell wall of tracheids started its formation. While the timing of bud break correlated with the sum of effective temperatures, the beginning of cambial zone activity did not. The needles were fully unfolded around DOY 170, which corresponded to the transition between earlywood and latewood tracheids. Summer drought did not affect needle development, but it changed the rate of production and morphology of latewood tracheids. Latewood tracheids from the year 2015 when the drought was longest (i.e. until the second half of August in 2015 compared to the duration from June until the middle of July in the two other years) were 32% narrower and they had 34% thinner cell walls than in 2014 and 2016. The improvement of tree water status in July resulted in the formation of intra-annual density fluctuations (IADF). The interplay between needle and cambium phenology provided an insight into how the tree allocates the resources with varying temperature and soil water availability.     

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.     

Growth and response patterns of Picea crassifolia and Pinus tabuliformis to climate factors in the Qilian Mountains,northwest China

Different tree species exhibit different phenological and physiological characteristics, leading to complexity in inter-species comparison of stem radial growth response to climate change. This study explored the climate-growth responses of Qinghai spruce (Picea crassifolia) and Chinese pine (Pinus tabulaeformis) in the Qilian Mountains, Northwest China. Meanwhile, Vaganov-Shashkin model (VS-oscilloscope) was used to simulate the relationships between radial growth rates and phenology. The results showed that 1) in their radial growth patterns, Qinghai spruce showed a significant increasing trend, while Chinese pine showed a decreasing trend, and Qinghai spruce has a longer growing season than Chinese pine. 2) For the radial growth-climate dynamic response, Qinghai spruce was influenced in an unstable manner by the mean temperature in the mid-growing season of the current year and the late growing season of the previous year and by the mean minimum temperature in the mid-growing season of the current year, while Chinese pine was influenced in a stable manner by the mean temperature and mean maximum temperature during the growing season of the current year. 3) The radial growth rates of the two conifer species were limited by temperature at the initiation and cessation of growth and by soil moisture at the peak of growth. But Chinese pine was more severely affected by soil moisture than Qinghai spruce in the middle of growth. Therefore, different management and restoration measures should be taken based on the differences in ecological responses and physical and physiological properties of the two conifer species to climate change in the subalpine forest ecosystems in the semiarid and arid regions of Northwest China.     

Growth rings, growth ring formation and age determination in the mangrove Rhizophora mucronata

BACKGROUND AND AIMS: The mangrove Rhizophora mucronata has previously been reported to lack annual growth rings, thus barring it from dendrochronological studies. In this study the reported absence of the growth rings was reconsidered and the periodic nature of light and dark brown layers visible on polished stem discs investigated. In addition, the formation of these layers in relation to prevailing environmental conditions, as well as their potential for age determination of the trees, was studied. METHODS: Trees of known age were collected and a 2.5-year cambial marking experiment was conducted to determine the periodic nature of the visible growth layers. KEY RESULTS: Annual indistinct growth rings were detected in R. mucronata and are defined by a low vessel density earlywood and a high vessel density latewood. The formation of these growth rings and their periodic nature was independent from site-specific environmental conditions in two forests along the Kenyan coast. However, the periodic nature of the rings was seriously affected by slow growth rates, allowing accurate age determination only in trees with radial growth rates above 0.5 mm year(-1). The onset of the formation of the low vessel density wood coincided with the onset of the long rainy season (April-May) and continues until the end of the short rainy season (November). The high vessel density wood is formed during the dry season (December-March). Age determination of the largest trees collected in the two studied forests revealed the relatively young age of these trees (+/-100 years). CONCLUSIONS: This study reports, for the first time, the presence of annual growth rings in the mangrove R. mucronata, which offers further potential for dendrochronological and silvicultural applications.     

Climate effects on inter- and intra-annual larch stemwood anomalies in the Mongolian forest-steppe

Climate response of tree-ring width and intra-annual wood anomalies were studied in stands of Siberian larch (Larix sibirica) on Mt. Bogd Uul in the forest-steppe ecotone of Mongolia. Climate on Mt. Bogd Uul is characterized by an increase of the annual mean temperature by 1.5 K between 1965 and 2007, the lack of a long-term trend for annual precipitation and, with it, an increase in aridity. Tree-ring width increases with increasing June precipitation of the current year (June) and increasing late summer precipitation of the previous year. In >100-year old trees, also a negative correlation of tree-ring width with the July temperature of the year prior to tree-ring formation was found. Decreasing tree-ring width with increasing snowfall in December can be explained with the protection of the frost-sensitive eggs of gypsy moth by snow cover, which is a major herbivore of larch in Mongolia and causes reduction in the annual stem increment. The most significant change in wood anatomy was the decline of wide latewood, which is attributable to the increase of summer days with a mean temperature > 15 °C and drought periods in summer without precipitation. Increasing summer drought is also thought to have caused the repeated occurrence of missing rings since the 1960s, which were not observed in the late 19th and early 20th centuries.     

Climate–tree growth relationships of longleaf pine (Pinus palustris Mill.) in the Southeastern Coastal Plain, USA

Knowledge of tree growth/climate response relationships is important to dendroecological studies and dendroclimatic reconstructions, particularly in the Southeastern Coastal Plain where few such studies have been attempted. To this end, we developed tree-ring chronologies of total ring width, earlywood width, and latewood width from longleaf pine (Pinus palustris Mill.) at three sites in the Southeastern Coastal Plain to examine the climate–growth relationships for this tree species. The length of these chronologies is unprecedented for southern pine chronologies in the Southeast. We compared the tree-ring chronologies to monthly temperature, precipitation, Palmer drought severity index (PDSI), and Palmer hydrological drought index (PHDI) data from the pertinent climate divisions. We found that PDSI and PHDI have the highest correlation with longleaf pine growth, and the strongest relationships between longleaf pine growth and these variables occur between July and November. Precipitation in the spring and summer was also positively related to growth at all sites. The relationship between temperature and growth was the weakest among all climate variables, but warm summer temperatures had a consistent, negative relationship with longleaf pine growth. The climate signal in the latewood was generally more robust than for total ring width and earlywood width.     

High-resolution climatic analysis of wood anatomical features in Corsican pine from Corsica (France) using latewood tracheid profiles

Key message

We propose a new methodology to identify intra-annual density fluctuations in latewood using cell features and relative radial position within the latewood of pine trees growing on Corsica, France. Climatic forcing of latewood wood anatomical anomalies was analyzed.

Abstract

We analyzed latewood anatomical features from Corsican pine (Pinus nigra ssp. laricio) of high-elevation sites in Corsica (France) derived from digital images of the wood surface. Latewood of each ring during the period 1950–2008 was partitioned into ten equal parts P1–P10. Mean values of the cell parameters cell lumen area (CLA), radial cell width (RCW), radial cell wall thickness (CWT), and modeled latewood density (MLD) were calculated for P1–P10. The cellular profiles for each cell parameter were subjected to principal component analyses. It was possible to quantify macroscopically visible variations of wood anatomy like intra-annual density fluctuations (IADFs) by latewood profiles of different cell parameters. A combination of cell parameter characteristics including their relative radial position within latewood provides a quantification of the cell anatomical variations in an IADF. Individual cell parameter chronologies and principal components of cell parameter profiles were correlated with climate data to determine the climatic forcing on latewood formation. Average cell parameter profiles and deviations from the long-term means are able to describe “normal” and “anomalous” environmental conditions during latewood formation. Cell feature anomalies throughout the latewood during individual years allow the reconstruction of past weather conditions with a high temporal resolution.     

东亚地区夏季干旱、强台风事件与松树枯萎病的关系

松树枯萎病的发生和流行给东亚的日本、中国和韩国带来了不小的损失,引起许多国家的重视. 本文应用实地观测和气象数据分析等方法,研究了东亚地区松树枯萎病的发生和流行特点,以及与灾害气象事件之间的关系.结果表明: 在日本、中国和韩国,持续的夏季干旱少雨和强台风等极端气象事件能够诱发松树枯萎.在极端干热的环境中,松树常出现能量代谢失调, 以至于受胁迫的松树整株枯萎;而在低温多雨的年份,松树枯萎少,甚至没有发生枯死现象.在松材线虫及其媒介昆虫侵染之前,松树的活力业已下降.松树枯萎病似乎应该局限在台风频发和持续干热的地区.在自然环境优越、少有台风和干热事件出现且没有不当扩大松树栽培范围的地区,松树枯萎病大面积发生的可能性不大.     

Identification of QTLs influencing wood property traits in loblolly pine (Pinus taeda L.). I. Physical wood properties

Physical wood property traits for loblolly pine (Pinus taeda L.) were analyzed for the presence of quantitative trait loci (QTLs) in a three-generation outbred pedigree. These traits include wood specific gravity (wsg), volume percentage of latewood (vol%), and microfibril angle (mfa). Phenotypic data were collected for rings 2–11 for wsg and vol%, and rings 3, 5 and 7 for mfa. Both earlywood and latewood were analyzed for each trait. An interval mapping approach designed for an outbred pedigree was used to estimate the number of QTLs, the magnitude of QTL effects, and their genomic position. Nine unique QTLs were detected for composite traits (average of all rings) for wsg, five for vol%, and five for mfa. The majority of these QTLs were verified by analyses of individual-ring traits. Additional QTLs for each trait were also detected by these individual-ring analyses. Most QTLs for wsg were specific to either earlywood or latewood, whereas each QTL for mfa was detected for both earlywood and latewood. Before these QTLs are utilized in a breeding program, they should be verified in larger experiments and in different genetic and environmental backgrounds. Received: 15 February 2000 / Accepted: 31 March 2000     

Fast response of Scots pine to improved water availability reflected in tree‐ring width and δ13C

Drought‐induced forest decline, like the Scots pine mortality in inner‐Alpine valleys, will gain in importance as the frequency and severity of drought events are expected to increase. To understand how chronic drought affects tree growth and tree‐ring δ¹³C values, we studied mature Scots pine in an irrigation experiment in an inner‐Alpine valley. Tree growth and isotope analyses were carried out at the annual and seasonal scale. At the seasonal scale, maximum δ¹³C values were measured after the hottest and driest period of the year, and were associated with decreasing growth rates. Inter‐annual δ¹³C values in early‐ and latewood showed a strong correlation with annual climatic conditions and an immediate decrease as a response to irrigation. This indicates a tight coupling between wood formation and the freshly produced assimilates for trees exposed to chronic drought. This rapid appearance of the isotopic signal is a strong indication for an immediate and direct transfer of newly synthesized assimilates for biomass production. The fast appearance and the distinct isotopic signal suggest a low availability of old stored carbohydrates. If this was a sign for C‐storage depletion, an increasing mortality could be expected when stressors increase the need for carbohydrate for defence, repair or regeneration.     

Elemental and stable isotope composition of Pinus halepensis foliage along a metal(loid) polluted gradient: implications for phytomanagement of mine tailings in semiarid areas

Background and aims

Aleppo pine (Pinus halepensis Mill.) is a widely used species for restoring degraded semiarid areas, but its use for the revegetation of metal(loid) polluted soils has not been thoroughly investigated. The main goal of this research was to study the ecophysiological status and elemental composition of spontaneous populations of Pinus halepensis growing on mine tailings to assess their use in phytomanagement of mine spoils in semiarid climates.

Methods

Edaphic characteristics and the physiological (by stable isotopes) and nutritional status of pine trees were determined on mine tailings, in the metalloid-polluted surroundings and a non-polluted control area.

Results

Low soil phosphorus availability at the tailings was found to be a more important limiting factor for pine physiological performance than high soil metal(lloid)s concentrations. Foliar phosphorus concentrations showed a strong negative correlation with foliar sulphur concentrations along the studied transect. The carbon and oxygen isotopic composition (δ¹³C and δ¹⁸O) of pine needles indicated that trees at the tailings were less water stressed than those in surroundings or control areas. The low foliar δ¹⁵N of pines growing at the tailings was due to low soil fertility and/or a heavy reliance on symbiotic ectomycorrhizal fungi for nitrogen uptake.

Conclusions

The results of this study indicate that Pinus halepensis is a suitable tree species for the phytostablisation of neutral or slightly-alkaline mining wastes in semiarid environments, thanks to its drought hardiness and good adaptation to low soil fertility and salinity.     

False ring formation in eastern hemlock branches: impacts of hemlock woolly adelgid and elongate hemlock scale

Herbivores can alter plant physiology through the induction of abnormal wood formation. Feeding by some insects induces the formation of false rings, a band of thick-walled latewood cells within the earlywood portion of the tree ring that reduces water transport. Hemlock woolly adelgid (Adelges tsugae Annand) and elongate hemlock scale (Fiorinia externa Ferris) are invasive insects that both feed on eastern hemlock [Tsuga canadensis (L.) Carrière]. Adelges tsugae has a greater effect on tree health than F. externa, but the mechanism underlying their differential effect is unknown. We explored the effects of these herbivores by assessing growth ring formation in branches of trees that had been experimentally infested for 4 yr with A. tsugae, F. externa, or neither insect. We measured false ring density, ring growth, and earlywood: latewood ratios in the two most recently deposited growth rings. Branches from A. tsugae-infested trees had 30% more false rings than branches from F. externa-infested trees and 50% more than branches from uninfested trees. In contrast, branches from F. externa-infested trees and control trees did not differ in false ring formation. Radial growth and earlywood: latewood ratios did not differ among treatments. Our results show that two invasive herbivores with piercing-sucking mouth parts have differing effects on false ring formation in eastern hemlock. These false rings may be the product of a systemic plant hypersensitive response to feeding by A. tsugae on hemlock stems. If false rings are responsible for or symptomatic of hemlock water stress, this may provide a potential explanation for the relatively large effect of A. tsugae infestations on tree health.     

太行山南麓不同径级栓皮栎生长对气候要素及干旱事件的响应

为评估径级对树木的气候-生长关系的影响,建立太行山南麓低海拔地区栓皮栎全轮、早材、晚材宽度年表,对比两个径级栓皮栎人工林径向生长对气候响应的敏感性差异,使用叠加时代分析揭示干旱事件对不同径级栓皮栎的影响,为气候变化背景下研究区栓皮栎人工林可持续经营提供参考数据。结果表明: 大径级栓皮栎全轮、晚材年表的平均敏感度高于小径级栓皮栎年表,但小径级栓皮栎早材年表的平均敏感度高于大径级栓皮栎早材年表。晚材是树轮气候响应最敏感的组分。两径级栓皮栎标准年表指数对当年气候因子的响应模式相似。小径级栓皮栎全轮、晚材生长对当年6—8月的气候因子更敏感,其早材对生长季之前(1—2月)气候因子更敏感;而大径级栓皮栎早材对去年的气候因子更敏感,有更强的滞后效应。研究区栓皮栎对干旱非常敏感,干旱年份两径级栓皮栎全轮、晚材的径向生长量显著低于上一年,且大径级栓皮栎的减少幅度更高,但干旱后大径级栓皮栎生长恢复程度更高。     

Climatic signals in tree-ring widths and wood structure of Pinus halepensis in contrasted environmental conditions

Tree-ring widths (RW), earlywood (EW) and latewood (LW) widths, the transition from early to latewood (T) and the occurrence of intra-annual density fluctuations in EW (E-ring) and in LW (L-ring), as well as the presence of resin canals in EW and LW, were analyzed in Aleppo pine (Pinus halepensis Mill.) from three sites in Spain and one in Slovenia to find out if the anatomical characteristics can provide additional seasonal climate–growth information from contrasted environmental conditions. Principal component analysis was applied to elucidate the relationship between the measured parameters and climate. Principal component factor PC1 proved to be related to parameters of EW and the climatic variables of winter-spring; PC2 to parameters of LW and climatic variables of summer–autumn; PC3 to conditions during transitions from humid to dry periods. The three PCs vary between sites and are determined by the climatic conditions during their formation. The study demonstrates that wood anatomical features may provide complementary information to that contained in tree-ring widths. Since such results are obtained on contrasting sites, it is likely that it may be generalized over the wide range of P. halepensis distribution representing a useful proxy for studies on a regional scale.