Does the removal of snowpack and the consequent changes in soil frost affect the physiology of Norway spruce needles?

Climate change may cause a decrease in snow cover in northern latitudes. This, on the other hand, may result in more severe soil frost even in areas where it is not common at present, and may lead to increased stress on the tree canopy. We studied the effects of snow removal and consequent changes in soil frost and water content on the physiology of Norway spruce (Picea abies [L.] Karst.) needles and implications on root biomass. The study was conducted at a 47-year-old Norway spruce stand in eastern Finland during the two winters of 2005/06 and 2006/07. The treatments in three replicates were: (i) natural snow accumulation and melting (CTRL), (ii) artificial snow removal during the winter (OPEN), and (iii) the same as OPEN, but the ground was insulated in early spring to delay soil thawing (FROST). In spite of the deeper soil frost in the OPEN than in the CTRL treatment, soil warming in spring occurred at the same time, whereas soil warming in the FROST was delayed by 2 and 1.5 months in 2006 and 2007, respectively. The soil water content was affected by snow manipulations, being at a lower level in the OPEN and FROST than CTRL in spring and early summer. The physiological measurements of the needles (e.g. starch, carbon and nitrogen content and apoplastic electrical resistance) showed differences between soil frost treatments. The differences were mostly seen between the CTRL and FROST, but also in the case of the starch content in early spring 2007 between the CTRL and OPEN. The needle responses in the FROST were more evident after the colder winter of 2006. The physiological changes seemed to be related to the soil temperature and water content in the early growing season rather than to the wintertime soil temperature. No difference was found in the fine root (diameter < 2 mm) biomass between the treatments assessed in 2007. In the future, conditions similar to the OPEN treatment may be more common than at present in areas experiencing a thick snow cover. The present experiment took place over the course of two years. It is possible that whenever thin snow cover occurs yearly, the reduced starch content during the early spring may be reflected in the tree growth itself as a result of reduced energy reserves.     

Effects of soil frost on soil respiration and its radiocarbon signature in a Norway spruce forest soil

Apart from a general increase of mean annual air temperature, climate models predict a regional increase of the frequency and intensity of soil frost with possibly strong effects on C cycling of soils. In this study, we induced mild soil frost (up to −5 °C in a depth of 5 cm below surface) in a Norway spruce forest soil by removing the natural snow cover in the winter of 2005/2006. Soil frost lasted from January to April 2006 and was detected down to 15 cm depth. Soil frost effectively reduced soil respiration in the snow removal plots in comparison to undisturbed control plots. On an annual basis 6.2 t C ha⁻¹ a⁻¹ were emitted in the control plots compared with 5.1 t C ha⁻¹ a⁻¹ in the snow removal plots. Only 14% of this difference was attributed to reduced soil respiration during the soil frost period itself, whereas 63% of this difference originated from differences during the summer of 2006. Radiocarbon (Δ¹⁴C) signature of CO₂ revealed a considerable reduction of heterotrophic respiration on the snow removal plots, only partly compensated for by a slight increase of rhizosphere respiration. Similar CO₂ concentrations in the uppermost mineral horizons of both treatments indicate that differences between the treatments originated from the organic horizons. Extremely low water contents between June and October of 2006 may have inhibited the recovery of the heterotrophic organisms from the frost period, thereby enhancing the differences between the control and snow removal plots. We conclude that soil frost triggered a change in the composition of the microbial community, leading to an increased sensitivity of heterotrophic respiration to summer drought. A CO₂ pulse during thawing, such as described for arable soils several times throughout the literature, with the potential to partly compensate for reduced soil respiration during soil frost, appears to be lacking for this soil. Our results from this experiment indicate that soil frost reduces C emission from forest soils, whereas mild winters may enhance C losses from forest soils.     

N2O emission in a Norway spruce forest due to soil frost: concentration and isotope profiles shed a new light on an old story

In mountain regions of Central Europe an increase of soil frost periods is predicted for this century due to reduced snow fall. To investigate the effects of freezing and thawing on soil N₂O fluxes in a mature Norway spruce forest in the mountainous Fichtelgebirge, Germany, the natural snow cover on three experimental plots was removed to induce soil frost. Three plots with natural snow cover served as controls. Soil N₂O fluxes were recorded in biweekly to monthly intervals during the frost and subsequent thawing period of the below-average cold winter in 2005/2006 and in the above-average warm winter in 2006/2007. In addition, N₂O concentrations and isotope signatures in soil air were measured along soil profiles in six different depths (from 6 to 70 cm). The soil of the snow removal plots was frozen down to 15 cm depth from January to April 2006 while the soil of control plots remained unfrozen under snow cover. Both soil freezing and thawing resulted in almost tenfold enhanced N₂O fluxes on snow removal plots contributing 84% to annual N₂O emissions. In the subsequent winter without soil frost no effects were observed. Vertical gradients of N₂O concentrations together with isotope abundance suggest that the subsoil of all plots was a probably weak, but continuous N₂O source throughout the year. Isotope signatures and N₂O concentration gradients in the soil profile indicate that microbial N₂O production and reduction of N₂O to N₂ did not or just marginally occur in frozen soil layers of the snow removal plots. Consequently, elevated N₂O fluxes in the late winter were attributed to the release of accumulated N₂O originating from the subsoil. At unfrozen soil, however, N₂O emissions were reduced due to a shift of the N₂O production-consumption ratio towards more consumption in the topsoil of both the control and snow removal plots. These findings contradict the general assumption that N₂O production in the organic layer is responsible for bursts of N₂O due to soil frost.     

Intra-annual tracheid formation of Norway spruce provenances in southern Finland

We studied the intra-annual wood formation in a Norway spruce provenance experiment in southern Finland from 2004–2008. Two Finnish provenances, northern and southern, as well as German and Hungarian provenances were included. Timing of tracheid formation and differentiation, and tracheid dimensions were determined from periodically extracted microcores. The aim was to determine the differences between the years and provenances in the timing of the xylogenesis and in the xylem characteristics. Year-to-year variation was high both in timing of tracheid formation and xylem characteristics, while between-provenance differences were small. The onset of tracheid formation varied from early May to late June in different trees in different years. The onset of tracheid formation was not closely related to the annual variations of temperature sum. In all the years, daily temperatures exceeded the threshold +5°C for several weeks before the onset of tracheid formation. The highest tracheid formation rate occurred after the summer solstice in all years and generally coincided with the highest daily temperatures during the growing season. Tracheid production ceased early in 2006 due to a mid-summer drought. Cell differentiation continued late in autumn as non-mature tracheids were still observed around mid-September. No clear differences between the provenances in the timing of tracheid formation were observed, although the Finnish provenances tended to initiate tracheid formation slightly earlier than the other provenances. The tree-ring widths of the Finnish provenances were also wider, while tracheid diameter of the German provenance was slightly smaller. Our results indicate that between-tree variation in the timing of wood formation is high compared with the latitude effect of seed source.     

模拟降水减少对帽儿山地区兴安落叶松径向生长的影响

以2004年在小兴安岭帽儿山地区栽植的兴安落叶松(Larix gmelinii)为研究对象,在2012年,经过100%减雨、50%减雨、冬季除雪和对照4种处理,在控雨的前期(6月)和后期(8月)用微生长锥取样利用石蜡切片方法测量扩大细胞的径向长度和数量。在10月末用生长锥取样,打磨后测量成熟管胞的径向长度和数量,进而分析降水减少对兴安落叶松径向生长的影响。结果表明:在帽儿山地区,生长季温度与兴安落叶松径向生长显著正相关,降水在生长季后期为负相关。大树芯成熟管胞测定表明,3块样地综合而言,除雪组成熟管胞数量最多,其次是对照组,50%减雨组和100%减雨组成熟管胞数量最少;管胞径向长度在各个处理组之间差异不显著。但是,在3块样地中,成熟管胞数量和径向长度在相同处理间测定结果有差别,样地1的100%减雨组、50%减雨组、除雪组和对照组中成熟管胞数量随土壤湿度增加而增加,径向管胞长度差异不明显;样地2的除雪组中成熟管胞径向长度显著小于其他3个实验组,而50%减雨组、100%减雨组和对照组的成熟管胞径向长度和数量均差异不显著;样地3中4个实验组的成熟管胞数量均差异不显著,成熟管胞径向长度100%减雨组略小于其他3个实验组,这与样地3遮阴较大、土壤湿度较高有关。石蜡切片的形成层扩大细胞分析表明:在减雨处理的初期(6月份),样地3在扩大细胞的数量上,随着不同处理间土壤湿度的减小而减少。在减雨处理的后期(8月份),样地1扩大细胞的数量随着不同处理间土壤湿度的减小而减少。研究结果表明降水减少会对兴安落叶松径向生长产生影响,其影响程度取决于微环境条件的差异,且不同时期降水减少的影响也有差异。     

Effects of soil frost on nitrogen net mineralization, soil solution chemistry and seepage losses in a temperate forest soil

Freezing and thawing may alter element turnover and solute fluxes in soils by changing physical and biological soil properties. We simulated soil frost in replicated snow removal plots in a mountainous Norway spruce stand in the Fichtelgebirge area, Germany, and investigated N net mineralization, solute concentrations and fluxes of dissolved organic carbon (DOC) and of mineral ions (NH₄⁺, NO₃⁻, Na⁺, K⁺, Ca²⁺, Mg²⁺). At the snow removal plots the minimum soil temperature was −5 °C at 5 cm depth, while the control plots were covered by snow and experienced no soil frost. The soil frost lasted for about 3 months and penetrated the soil to about 15 cm depth. In the 3 months after thawing, the in situ N net mineralization in the forest floor and upper mineral soil was not affected by soil frost. In late summer, NO₃⁻ concentrations increased in forest floor percolates and soil solutions at 20 cm soil depth in the snow removal plots relative to the control. The increase lasted for about 2–4 months at a time of low seepage water fluxes. Soil frost did not affect DOC concentrations and radiocarbon signatures of DOC. No specific frost effect was observed for K⁺, Ca²⁺ and Mg²⁺ in soil solutions, however, the Na⁺ concentrations in the upper mineral soil increased. In the 12 months following snowmelt, the solute fluxes of N, DOC, and mineral ions were not influenced by the previous soil frost at any depth. Our experiment did not support the hypothesis that moderate soil frost triggers solute losses of N, DOC, and mineral ions from temperate forest soils.     

Phytochrome action and frost hardening in black spruce seedlings

Black spruce [ Picea mariana (Mill.) B.S.P.] development is sensitive to photoperiod. To date the implication of photoperiod, and especially phytochrome, in the frost hardening process of black spruce has not been fully tested. Two light fluence rates, night interruption of darkness, and red vs far-red radiation treatments were applied to black spruce seedlings, followed by freezing at –6°C. Parallel to the freezing test, growth measurements, bud formation and the xylem water potential estimates of the seedlings were done. While dry weight accumulation depends on the irradiation energy level, bud formation and freezing tolerance are photoperiodically sensitive. Furthermore, bud formation and frost hardening are dependent upon whether phytochrome is in the active form or inactive form, as demonstrated by the positive effect of short days, far-red radiation and the reversal of the red effect by far-red radiation. Also, xylem water potential appears to be influenced by short day and far-red conditioning.     

Tracheids in white spruce seedling's long lateral roots in response to nitrogen availability

This study examined how the availability of inorganic nitrogen (N) modified the anatomical characteristics of white spruce (Picea glauca (Moench) Voss) roots related to their hydraulic properties. Seedlings were grown for one growing season in 4 L capacity pots filled with sand under one of three N levels: low (10 ppm), medium (50 ppm) and high (125 ppm). First order lateral roots with intact tips were sampled from dormant seedlings in October. Root segments were collected from 4, 10, and 14 cm distances above the root tip for fixation and sectioning and for maceration. Additional specimens were collected from the 4 and 14 cm distances for maceration and scanning electron microscopy of xylem pits. Root diameter and surface area occupied by the xylem in root cross sections increased basipetally in all treatments but exceptions were found. Higher N-levels significantly increased root diameter and surface area occupied by the xylem. In the two higher N treatments secondary root development was more advanced near the root tip than in the low N treatment. There was a strong positive correlation between root diameter and cross-sectional root area occupied by the xylem (30–50% of the root cross section) but not in portions with little secondary development. Non-conducting space within the xylem occupied 10–13% of its cross-sectional surface. Tracheids of the primary xylem were larger, had larger lumens but thinner cell walls than those of the secondary xylem. Low N treatment seedling tracheids had smaller total cross-sectional area, less lumen, and less cell wall surface area than the two other N treatments. Tracheid diameter means were between 19–20 μm in the high and medium N treatments, and 15.2 μm in the low N treatment. The range was 4.5–51.3 μm. Tracheid length was not significantly affected by N. The average tracheid was about 1000 μm long, and the range was 110–3530 μm. Pit-border diameters ranged between 4.1–20.6 μm (average 10–11 μm) and were not affected by the N treatment. Pit aperture diameters were within 0.62–10.2 μm range (average between 3–4 μm) and were also not significantly affected by the N treatment, although tracheids from the medium N-treatment roots tended to have larger apertures. The pit border diameter equals that of the margo while the aperture size should be similar to that of the torus of the pit membrane. If the capacity for axial water transport in spruce roots is affected by N, it would be by its impact on conduit diameter and, possibly on the pit-membrane pore sizes but not by changes to conduit length and to the size of the pit membrane surface area. This revised version was published online in June 2006 with corrections to the Cover Date.     

Ascorbic acid and xylem development in trunks of the Siberian larch trees

The contents of ascorbic acid (AA) and its oxidized form, dehydroascorbic acid (DHA), were assessed as related to the tracheid differentiation in the course of early and late wood development in the Siberian larch (Larix sibirica Ldb.) trees. The samples of the cambium, cell enlargement zone and mature cells were collected at the successive developmental stages by scraping tissues off layer by layer from trunk segments of the 20-year-old trees according to anatomical and histochemical criteria. While cambium initials were rapidly dividing, the AA contents per dry weight and per cell considerably exceeded the corresponding values characteristic of the late xylem development; such difference corresponded to the higher number of early tracheids per annual ring, as compared to the late tracheids. The AA content decreased as cells enlarged. The radial growth of the early wood tracheids, as compared to the late wood tracheids, was accompanied with a threefold increase in the AA and a decline in the DHA contents. The AA/DHA ratio was in line with the early tracheid enlargement. The maximum AA content was observed at the early stage of the secondary cell wall thickening in the tracheids of early and late xylem preceding lignification. During this stage of early wood development, the DHA content exceeded sixfold the corresponding value in the late xylem; as a result, the initial rates of lignification were different in two tissues. The rate of lignification in a newly developing layer of the early xylem increased gradually and was the highest in the completely differentiated tracheids. In the late xylem, the lignification rate was at its highest at the very beginning and then declined in the course of tracheid maturation. The dissimilar patterns of lignification in the early and late xylem were primarily associated with the DHA content, which increased in the early xylem and decreased in the maturing late xylem. Thus, the AA content and its accessibility to oxidation in the growing and mature xylem cells exhibited the diverse developmental patterns in the early and late xylem: two tissues differed in the tracheid number and radial diameter as well as in the rate of lignification.Translated from Fiziologiya Rastenii, Vol. 52, No. 1, 2005, pp. 97–107.Original Russian Text Copyright © 2005 by Antonova, Chaplygina, Varaksina, Stasova.     

Cloning, characterization and localization of three novel class III peroxidases in lignifying xylem of Norway spruce (Picea abies)

Plant class III peroxidases (POXs) take part in the formation of lignin and maturation of plant cell walls. However, only a few examples of such peroxidases from gymnosperm tree species with highly lignified xylem tracheids have been implicated so far. We report here cDNA cloning of three xylem-expressed class III peroxidase encoding genes from Norway spruce (Picea abies). The translated proteins, PX1, PX2 and PX3, contain the conserved amino acids required for heme-binding and peroxidase catalysis. They all begin with putative secretion signal propeptide sequences but diverge substantially at phylogenetic level, grouping to two subclusters when aligned with other class III plant peroxidases. In situ hybridization analysis on expression of the three POXs in Norway spruce seedlings showed that mRNA coding for PX1 and PX2 accumulated in the cytoplasm of young, developing tracheids within the current growth ring where lignification is occurring. Function of the putative N-terminal secretion signal peptides for PX1, PX2 and PX3 was confirmed by constructing chimeric fusions with EGFP (enhanced green fluorescent protein) and expressing them in tobacco protoplasts. Full-length coding region of px1 was also heterologously expressed in Catharanthus roseus hairy root cultures. Thus, at least the spruce PX1 peroxidase is processed via the endoplasmic reticulum (ER) most likely for secretion to the cell wall. Thereby, PX1 displays correct spatiotemporal localization for participation in the maturation of the spruce tracheid secondary cell wall.     

Embolism formation during freezing in the wood of Picea abies

Freeze-thaw events can cause embolism in plant xylem. According to classical theory, gas bubbles are formed during freezing and expand during thawing. Conifers have proved to be very resistant to freeze-thaw induced embolism, because bubbles in tracheids are small and redissolve during thawing. In contrast, increasing embolism rates upon consecutive freeze-thaw events were observed that cannot be explained by the classical mechanism. In this study, embolism formation during freeze-thaw events was analyzed via ultrasonic and Cryo-scanning electron microscope techniques. Twigs of Picea abies L. Karst. were subjected to up to 120 freeze-thaw cycles during which ultrasonic acoustic emissions, xylem temperature, and diameter variations were registered. In addition, the extent and cross-sectional pattern of embolism were analyzed with staining experiments and Cryo-scanning electron microscope observations. Embolism increased with the number of freeze-thaw events in twigs previously dehydrated to a water potential of -2.8 MPa. In these twigs, acoustic emissions were registered, while saturated twigs showed low, and totally dehydrated twigs showed no, acoustic activity. Acoustic emissions were detected only during the freezing process. This means that embolism was formed during freezing, which is in contradiction to the classical theory of freeze-thaw induced embolism. The clustered pattern of embolized tracheids in cross sections indicates that air spread from a dysfunctional tracheid to adjacent functional ones. We hypothesize that the low water potential of the growing ice front led to a decrease of the potential in nearby tracheids. This may result in freezing-induced air seeding.     

Analysis of freeze-thaw embolism in conifers. The interaction between cavitation pressure and tracheid size

Ice formation in the xylem sap produces air bubbles that under negative xylem pressures may expand and cause embolism in the xylem conduits. We used the centrifuge method to evaluate the relationship between freeze-thaw embolism and conduit diameter across a range of xylem pressures (Px) in the conifers Pinus contorta and Juniperus scopulorum. Vulnerability curves showing loss of conductivity (embolism) with Px down to -8 MPa were generated with versus without superimposing a freeze-thaw treatment. In both species, the freeze-thaw plus water-stress treatment caused more embolism than water stress alone. We estimated the critical conduit diameter (Df) above which a tracheid will embolize due to freezing and thawing and found that it decreased from 35 microm at a Px of -0.5 MPa to 6 microm at -8 MPa. Further analysis showed that the proportionality between diameter of the air bubble nucleating the cavitation and the diameter of the conduit (kL) declined with increasingly negative Px. This suggests that the bubbles causing cavitation are smaller in proportion to tracheid diameter in narrow tracheids than in wider ones. A possible reason for this is that the rate of dissolving increases with bubble pressure, which is inversely proportional to bubble diameter (La Place's law). Hence, smaller bubbles shrink faster than bigger ones. Last, we used the empirical relationship between Px and Df to model the freeze-thaw response in conifer species.     

Temporal dynamic of wood formation in Pinus cembra along the alpine treeline ecotone and the effect of climate variables

We determined the temporal dynamic of cambial activity and xylem development of stone pine (Pinus cembra L.) throughout the treeline ecotone. Repeated micro-sampling of the developing tree ring was carried out during the growing seasons 2006 and 2007 at the timberline (1,950 m a.s.l.), treeline (2,110 m a.s.l.) and within the krummholz belt (2,180 m a.s.l.) and the influence of climate variables on intra-annual wood formation was determined. At the beginning of both growing seasons, highest numbers of cambial and enlarging cells were observed at the treeline. Soil temperatures at time of initiation of cambial activity were c. 1.5°C higher at treeline (open canopy) compared to timberline (closed canopy), suggesting that a threshold root-zone temperature is involved in triggering onset of above ground stem growth. The rate of xylem cell production determined in two weekly intervals during June through August 2006–2007 was significantly correlated with air temperature (temperature sums expressed as degree-days and mean daily maximum temperature) at the timberline only. Lack of significant relationships between tracheid production and temperature variables at the treeline and within the krummholz belt support past dendroclimatological studies that more extreme environmental conditions (e.g., wind exposure, frost desiccation, late frost) increasingly control tree growth above timberline. Results of this study revealed that spatial and temporal (i.e., year-to-year) variability in timing and dynamic of wood formation of P. cembra is strongly influenced by local site factors within the treeline ecotone and the dynamics of seasonal temperature variation, respectively.     

Morphological adaptations of tracheid structure to water stress gradients in the crown of Sequoiadendron giganteum

Summary Mean radial diameter of tracheids in young branches of a 90 m Sequoiadendron giganteum decreases linearly with height along a gradient correlated linearly with decreasing xylem pressure potential. These smaller tracheid diameters provide strength to resist strong mechanical tensions in the xylem column and hypothetically allow greater efficiencies of water conduction. Tracheid length is not significantly correlated with either water stress or tracheid diameter.     

Xylem wall collapse in water-stressed pine needles

Wall reinforcement in xylem conduits is thought to prevent wall implosion by negative pressures, but direct observations of xylem geometry during water stress are still largely lacking. In this study, we have analyzed the changes in xylem geometry during water stress in needles of four pine species (Pinus spp.). Dehydrated needles were frozen with liquid nitrogen, and xylem cross sections were observed, still frozen, with a cryo-scanning electron microscope and an epifluorescent microscope. Decrease in xylem pressure during drought provoked a progressive collapse of tracheids below a specific threshold pressure (P(collapse)) that correlates with the onset of cavitation in the stems. P(collapse) was more negative for species with smaller tracheid diameter and thicker walls, suggesting a tradeoff between xylem efficiency, xylem vulnerability to collapse, and the cost of wall stiffening. Upon severe dehydration, tracheid walls were completely collapsed, but lumens still appeared filled with sap. When dehydration proceeded further, tracheids embolized and walls relaxed. Wall collapse in dehydrated needles was rapidly reversed upon rehydration. We discuss the implications of this novel hydraulic trait on the xylem function and on the understanding of pine water relations.     

Daylength Effects on Growth, Tracheid Development, and Photosynthesis in Seedlings of Picea sitchensis and Pinus sylvestris

Effects of change in daylength on tracheid expansion and tracheidwall thickening are considered in relation to shoot and rootgrowth, assimilation rate, and starch content, in seedlingsof Picea sitchensis and Pinus sylvestris grown in controlledenviroments. Rates of wall thickening decreased in the first tracheids tobegin wall thickening after decrease in daylength. A sharp decreasein starch-grain size at the same time supports previous suggestionsthat wall thickness is related to available substrate; however,rates of assimilation were found to be far greater in shortdays than in long, suggesting that a surplus of carbohydratemay be produced in long days. It is pointed out that if availablesubstrate does limit wall thickening it is likely to determineto al accumulation of wall material; wall material per tracheidwould follow logistically from this depending on the numberof tracheids around the xylem, and wall thickness would dependalso on the radial tracheid diameter. Unexpectedly, rate of shoot growth accelerated after transferto short days before its final cessation with terminal-bud formation.Root growth declined in short days, but later increased againafter terminal bud formation. Rate of root growth did not appearto be associated with tracheid development in the shoot.     

Effects of environmental factors on wood formation in larch (Larix sibirica Ldb.) stems

 Effects of temperature and precipitation on xylem cell production by the cambium, radial cell expansion and secondary wall thickening in larch stems have been studied. The observations were carried out over two seasons on ten 50- to 60-year-old trees, growing in central Siberia and chosen according to growth rate (the number of cells in radial rows of each of two of the preceding seasons was equal). The data on the number of cells in differentiation zones and mature xylem along radial rows of tracheids, radial and tangential sizes of tracheids and their lumina were used for calculating cambial activity, the rates and durations of cell development in the zones, and both the thickness and cross-sectional areas of tracheid walls. The mean day air temperature, mean maximum diurnal and mean minimum nocturnal temperatures as well as precipitation have been shown by correlation and regression analyses to affect differentially separate stages of tracheid differentiation. Throughout all the seasons it was temperature that had the main influence on the initial divisions in the xylem, radial cell expansion and biomass accumulation. However, the levels of such an effect on separate stages of cytogenesis were different, especially the influence of nocturnal temperature on xylem cell production by cambium and primary wall growth. The optimum values of temperature and precipitation for cell production by cambium, for radial cell expansion and secondary wall thickening have been calculated. These optimum values of the first and second processes proved to be practically equal, while the last differs considerably in response to temperature. The data are discussed in connection with formation of early and late tracheids. Received: 3 July 1996 / Accepted: 7 February 1997     

Longer growing seasons lead to less carbon sequestration by a subalpine forest

As global temperatures increase, the potential for longer growing seasons to enhance the terrestrial carbon sink has been proposed as a mechanism to reduce the rate of further warming. At the Niwot Ridge AmeriFlux site, a subalpine forest in the Colorado Rocky Mountains, we used a 9‐year record (1999–2007) of continuous eddy flux observations to show that longer growing season length (GSL) actually resulted in less annual CO₂ uptake. Years with a longer GSL were correlated with a shallower snow pack, as measured using snow water equivalent (SWE). Furthermore, years with a lower SWE correlated with an earlier start of spring. For three years, 2005, 2006, and 2007, we used observations of stable hydrogen isotopes (δD) of snow vs. rain, and extracted xylem water from the three dominant tree species, lodgepole pine, Engelmann spruce, and subalpine fir, to show that the trees relied heavily on snow melt water even late into the growing season. By mid‐August, 57% to 68% of xylem water reflected the isotopic signature of snow melt. By coupling the isotopic water measurements with an ecosystem model, SIPNET, we found that annual forest carbon uptake was highly dependent on snow water, which decreases in abundance during years with longer growing seasons. Once again, for the 3 years 2005, 2006, and 2007, annual gross primary productivity, which was derived as an optimized parameter from the SIPNET model was estimated to be 67% 77%, and 71% dependent on snow melt water, respectively. Past studies have shown that the mean winter snow pack in mountain ecosystems of the Western US has been declining for decades and is correlated with positive winter temperature anomalies. Since climate change models predict continuation of winter warming and reduced snow in mountains of the Western US, the strength of the forest carbon sink is likely to decline further.     

Changes in phenolic acids during maturation and lignification of scots pine xylem

The content and fractional composition of alcohol soluble phenolic acids (PhA) in cells with different degree maturation and lignification in the course of early and late wood formation in the pine (Pinus sylvestris L.) stem during vegetation were studied. Phenolic compounds (PhC), extracted by 80% ethanol, were divided into free and bound fractions of PhA. In turn, the esters and ethers were isolated from bound PhA. The contents of all substances were calculated per dry weight and per cell. Considerable differences have been found to exist in both the contents and the composition of the fractions PhA on successive stages of tracheid maturation of early and late xylem. Early wood tracheids at all secondary wall thickening steps contained PhC less and free PhA more than late wood tracheids. Throughout earlywood tracheid maturation, the pool of free PhA per cell declined at the beginning of lignification and then increased gradually while that of bound PhA decreased. The maturation of late wood tracheids were accompanied by the rise of free PhA pool and the diminution of bound PhA pool. In the composition of bound PhA, the ethers were always dominant, and the amount of that in earlywood cells was less than in latewood cells. The cells of early xylem at all steps of maturation contained more of esters. The sum total of free hydroxycinnamic acids, precursors of monolignols, gradually decreased during early xylem lignification as the result of the reduction of the pools of p-coumaric, caffeic, ferulic and synapic acids, while that of their esters rised. In the course of late xylem lignification, the pools of free p-coumaric, ferulic and, especially, synapic acids increased. Simultaneously, the amount of ferulic acid ester and synapic acid ether increased too. According to the data, lignin biosynthesis in early xylem and late xylem occurs with different dynamics and the structure of lignins of two xylem types might be different too.     

Xylem traits of peatland Scots pines reveal a complex climatic signal: A study in the Eastern Italian Alps

Long-term climate reconstructions are frequently based on tree-ring high-resolution proxies extracted from subfossil peatland trees. Peatlands are peculiar ecosystems characterized by high moisture in the upper soil part which creates a harsh living environment for trees. The climate mostly indirectly influences tree growth determining seasonal variations in the water table level. Within this framework, the aim of this study was to investigate climate responses of trees (Pinus sylvestris L.) growing inside and outside a Southern Alpine peat bog, by using tree-ring and wood anatomical traits (e.g. tracheid number and dimension, cell-wall thickness). Our results showed differences in the xylem structure and climate signal recorded by peatland and mineral soil trees. Peatland trees were characterized by narrow rings and tracheids with thinner cell wall. Summer temperature and precipitation were the major drivers of xylem formation in peatland trees. At intra-annual level wood anatomical traits revealed a complex within-ring signal during the growing season. The multi-parameters approach together with the high-resolution gained by using tree-ring sectors allowed us to obtain new detailed information on the xylem development of peatland trees and climate drivers that influenced it.