Effects of the time of drought occurrence within the growing season on growth and survival of Pinus ponderosa seedlings

Key message

A drought event during spring produces a stronger and long lasting decrease in growth of ponderosa pine seedlings than a summer drought event. However, survival is not differentially affected.

Abstract

Although there is certainty about the increasing frequency of extreme climatic events, the consequences of changing patterns of drought events within the growing season on the growth and survival of different species are much less certain. In particular, little knowledge is available on the differential effect on tree seedlings of a drought event at different times within the growing season. The objective of this study was to quantify the effect of a drought event imposed at different times over the growing season on the growth, survival and some related morphological and physiological variables of Pinus ponderosa seedlings from two seed sources. Four treatments were applied: control conditions; spring drought; summer drought and spring plus summer drought (SpSuD). A drought event in spring reduced stem growth and biomass accumulation in ponderosa pine seedlings during the occurrence of the drought and afterwards, even when plant water status had recovered. The lack of growth recovery could not be associated with loss of stem hydraulic conductivity or reduction in stomatal conductance after drought. However, the spring drought did not differentially affect plant survival, as was the case with prolonged drought in the SpSuD treatment. The summer drought event had a significant but much smaller impact on plant growth. Our results suggest different consequences of a drought event in spring or in summer in ponderosa pine seedlings. This knowledge may be relevant to understand and predict tree seedlings responses to changing patterns of drought events within the growing season in the framework of climatic change.     

Impact of soil texture and water availability on the hydraulic control of plant and grape-berry development

Aims

All components of the soil-plant-atmosphere (s-p-a) continuum are known to control berry quality in grapevine (Vitis vinifera L.) via ecophysiological interactions between water uptake by roots and water loss by leaves. The scope of the present work was to explore how the main hydraulic components of grapevine influence fruit quality through changes in liquid- and gas-phase hydraulic conductance.

Methods

To reach our objectives, determinations of shoot growth, berry size and sugar content, leaf gas exchange, predawn leaf water potential (as a proxy of soil water potential), midday stem water potential and leaf water potential were performed in conjunction with anatomical measurements of shoot xylem. All measurements were conducted in two different cultivars (Cabernet franc and Merlot) and on three different soil types (clayey, gravelly, and sandy).

Results

Shoot xylem morphometric characteristics and whole-plant hydraulic conductance were influenced by cultivar and soil type. Differences in leaf gas exchange parameters and water potentials were determined by soil type significantly more than by cultivar. Between the two extremes (gravelly soil imposing drought conditions and sandy soil with easily accessible water) the clayey soil expressed an intermediate plant water consumption and highest sugar accumulation in berry.

Conclusions

Hydraulic and non hydraulic limitations to vine/berry interactions supported the conclusion that water availability in the soil overrides differences due to cultivar in determining the productive potential of the vineyard. Non hydraulic stomatal control was expected to be an important component on plants grown on the clayey soil, which experienced a moderate water stress. Possible links between hydraulic traits and berry development and quality are discussed.     

Time change of aluminium toxicity in the acid bulk soil and the rhizosphere in Norway spruce (Picea abies (L.) Karst.) and beech (Fagus sylvatica L.) stands

Context

In acidic forest soils, aluminium can alter tree health due to its potential toxicity. Aluminium phytotoxicity is mainly influenced by its chemical form and its availability.

Methods

As physical-chemical indicators of Al toxicity in soil, Al speciation in soil solutions and in the exchange complex was measured in the rhizosphere and the bulk soil of two tree species (Norway spruce (Picea abies (L.) Karst.) and European Beech (Fagus sylvatica L.) in an acidic soil and in 4 months (November, February, May and August) representing the four seasons in a year.

Results

In the bulk soil, Al toxicity was generally higher under Norway spruce than under beech. Furthermore, temporal changes in Al behaviour were identified under Norway spruce but not under beech. The monomeric Al in the soil solutions and the exchangeable Al in the solid soil increased significantly in February under Norway spruce and were positively correlated with nitrate concentration, suggesting that nitrate influence Al speciation and mobility under Norway spruce. In the rhizosphere, Al toxicity was restricted through Al complexation by organic compounds and by nutrient contents independently from the season. The ecological importance of the rhizosphere in Al detoxification is discussed.

Conclusions

This study suggests that plant specific differences as well as seasonal changes in plant physiology, microbial activity and microclimatology influence aluminum toxicity in acid forest soils.     

How important is woody tissue photosynthesis in poplar during drought stress?

Key message

Woody tissue photosynthesis might play a key role in maintaining plant carbon economy and hydraulic function under unfavourable conditions such as drought stress.

Abstract

Within trees, a portion of respired CO₂ is assimilated by bark and woody tissue photosynthesis, but its physiological role remains unclear, in particular under unfavour able conditions like drought stress. We hypothesised that woody tissue photosynthesis will contribute to overall tree carbon gain both under sufficient water supply and during drought, and plays a role in maintaining the hydraulic function. We subjected half of the trees to a stem and branch light-exclusion treatment to prevent bark and woody tissue photosynthesis. Then, we measured leaf gas exchange and stem growth in Populus deltoides x nigra ‘Monviso’ trees both under well-watered and dry conditions. We additionally measured cavitation using acoustic emission in detached control and light-excluded branches to illustrate the role of woody tissue photosynthesis in xylem embolism repair. Under well-watered conditions, light exclusion resulted in reduced stem growth relative to control trees by 30 %. In response to drought, stem shrinkage of light-excluded trees was more pronounced as compared to control trees. During drought stress also maximum photosynthesis and transpiration rate tended to decrease more rapidly in light-excluded trees compared to control trees. Leaf fall in light-excluded branches together with the larger number of acoustic emissions in control branches indicates that in the latter more xylem vessels were still hydraulically functional under drought. Therefore, our study highlights that photosynthesis at branch and stem level might be a key factor in the resilience of trees to drought stress by maintaining both the plant carbon economy and hydraulic function.

     

Growth trends and climate responses of Norway spruce along elevational gradients in East-Central Europe

Key message

Decadal growth variability of Norway spruce increases with elevation. Recent temperature sensitivity and growth enhancement are limited to trees growing in the zone adjacent to timberline.

Abstract

Growth trends and climate responses of forest trees along elevational gradients are not fully understood. A deeper insight is, however, fundamental for predicting ecosystem functioning and productivity under future climate change. Supplementary to the effects of elevation and regional provenance on tree growth are sample depth, uneven representation of sample age and varying site conditions. Furthermore, there is only a limited number of studies addressing growth changes along elevational gradients, while at the same time applying tree-ring standardization methods that are sensitive to trend preservation. Here, we introduce 12 novel tree-ring width chronologies of Norway spruce (Picea abies[L.] Karst.) from four elevational belts encompassing montane forests and the local timberline in three regions in East-Central Europe between 15° and 19°E. Each chronology is characterized by sufficient sample replication and a comparable age structure between 1906 and 2010. Tree growth near timberline revealed substantial medium-frequency variability and sharply increasing ring widths since the 1980s. Medium-frequency growth variability of lower elevation trees was, however, relatively small, and growth rates over the last decade were either stable or even decreased. During the last four decades, Norway spruce from higher elevations exhibited a reduced response to autumn temperatures preceding ring formation. In contrast, trees from the lower-montane zone increased their sensitivity to drought during the same time. Our results emphasize not only different but also instable growth trends and climate responses of forest trees along altitudinal gradients, which should be considered in future forest management strategies.

     

Water loss regulation to soil drought associated with xylem vulnerability to cavitation in temperate ring-porous and diffuse-porous tree seedlings

Key message

Sustainable stomatal opening despite xylem cavitation occurs in ring-porous species and stomatal closure prior to cavitation in diffuse-porous species during soil drought.

Abstract

To elucidate the relationship between water loss regulation and vulnerability to cavitation associated with xylem structure, stomatal conductance (g _s), defoliation, vulnerability curves, and vessel features were measured on seedlings of ring-porous Zelkova serrata and Melia azedarach, and diffuse-porous Betula platyphylla var. japonica, Cerasus jamasakura and Carpinus tschonoskii. Under prolonged drought conditions, the percentage loss of hydraulic conductivity (PLC) increased and g _s decreased gradually with decreasing predawn (Ψ_pd) or xylem water potential (Ψ_xylem) in Z. serrata. During the gentle increase of PLC in M. azedarach, g _s increased in the early stages of dehydration while leaves were partly shed. A sharp reduction in g _s was observed before the onset of an increase in the PLC for drying plants of the three diffuse-porous species, suggesting cavitation avoidance by stomatal regulation. In the ring-porous species, xylem-specific hydraulic conductivity (K _s) was higher, whereas the vessel multiple fractions, the ratio of the number of grouped vessels to total vessels, was lower than that in the diffuse-porous species, suggesting that many were distributed as solitary vessels. This may explain the gradual increase in the PLC with decreasing Ψ_xylem because isolated vessels provide less opportunity for air seeding. Different water loss regulation to soil drought was identified among the species, with potential mechanisms being sustainable gas exchange at the expense of xylem dysfunction or partial leaf shedding, and the avoidance of xylem cavitation by strict stomatal regulation. These were linked to vulnerability to cavitation that appears to be governed by xylem structural properties.     

Hydraulic limitation on maximum height of Pinus strobus trees in northern Minnesota,USA

Key message

Using comparisons within and between trees, the authors show evidence for hydraulic limitation of tree height in a humid-climate species that is far from the global maximum tree height.

Abstract

We measured water status and two indicators of drought stress as a function of height within the canopies of four tall (32–35 m) eastern white pines (Pinus strobus) at an old-growth site in northern Minnesota, USA. Pre-dawn and midday xylem pressure potential measured on terminal shoots (Ψ _shoot), needle length, and foliar carbon isotope discrimination (δ ¹³C) all showed within-canopy gradients consistent with increasing drought stress with height. Midday Ψ _shoot near tree tops was ?1.8 MPa, close to values associated with stomatal closure for other temperate conifers. Pre-dawn Ψ _shoot decreased with height at >2× the gradient in gravitational potential. δ ¹³C was strongly correlated with height and weakly correlated with light. Needles were 15–25 % shorter at canopy top compared to the bottom of the canopy. Midday Ψ _shoot and needle length showed significant differences in regression model coefficients from tree to tree. The patterns are consistent with hydraulic constraints on height growth of white pine at this site.     

Hydraulic properties of European elms: xylem safety-efficiency tradeoff and species distribution in the Iberian Peninsula

Key message

Ulmus minor and U. glabra show a trade-off between safety and efficiency in water transport, and U. laevis shows adaptations to waterlogged environments.

Abstract

Three native elm species grow in Europe: Ulmus minor Mill., U. glabra Huds. and U. laevis Pall., and within the Iberian Peninsula their habitats mainly differ in water availability. We evaluated firstly whether vulnerability to xylem embolism caused by water-stress has been a determinant factor affecting their distribution; secondly, if their xylem anatomy differs due to water availability dissimilarities; and thirdly, if these species present a trade-off between water transport safety and efficiency. Plants of the three species were grown in a common-garden in Madrid, Central Spain. The centrifuge method was used for constructing the vulnerability curves, and anatomical measurements were carried out with an optical microscope. We found clear differences in conductivity and cavitation vulnerability between the three species. Although all three elms were highly vulnerable to cavitation, U. minor was significantly more resistant to water stress cavitation. This species reached 50 % loss in conductivity at ?1.1 MPa, compared to U. glabra that did so at ?0.5 MPa, and U. laevis at ?0.4 MPa. Maximum xylem specific conductivity and maximum leaf specific conductivity were two to three times higher in U. glabra when compared to U. minor. A clear trade-off between safety against losses of conductivity and water transport efficiency was observed considering both U. minor and U. glabra samples. Ulmus minor’s hydraulic configuration was better adapted to overcome drought episodes. The expected aridification of the Iberian Peninsula could compromise Ulmus populations due to their high vulnerability to drought stress.     

Antioxidative response patterns of Norway spruce bark to low-density Ceratocystis polonica inoculation

Key message

Based on time courses of individual antioxidant compounds, bark phenolic metabolism has been recognised to integrate ascorbate–glutathione system as a redox hub in Norway spruce defence against Ceratocystis polonica infection.

Abstract

Temporal courses of individual phenolics, thiols and ascorbate were studied in Norway spruce phloem over a 5-month period after inoculation at low density with Ceratocystis polonica. The initial reaction of Norway spruce 3 days after inoculation was characterised by significantly increased isorhapontin and taxifolin concentrations, accompanied by significantly lowered catechin contents. On later sampling dates, catechin concentrations within infected bark increased until September. The slightly accumulated astringin contents in April and May diminished at later sampling dates in response to infection. The isorhapontin levels strongly raised in April and were slightly lowered from June onwards. Compared to the controls, taxifolin concentrations were higher in the infected samples showing a double peak with maxima in April and June. The taxifolin values eased later but remained above the control levels. The initial response of the ascorbate–glutathione system to fungal infection was characterised by a significantly more oxidised glutathione pool, slightly more reduced ascorbate system and by higher glutathione reductase activity. Three weeks later an accumulation of thiols was observed, whereas total ascorbate was significantly lowered and the ascorbate redox state shifted towards more oxidised values. Until the middle of July a gradual increase of total glutathione was determined within the infected bark, which was accompanied by significantly increased cysteine contents, higher glutathione reductase activity, but significantly lowered total ascorbate contents. The increased pressure on the ascorbate system reflects its interaction with phenolics, as ascorbate is needed for reducing the phenoxyl radicals formed during pathogen defence.     

Seasonal dynamics of phloem and xylem formation in silver fir and Norway spruce as affected by drought

The dynamics of phloem growth ring formation in silver fir (Abies alba Mill.) and Norway spruce (Picea abies Karst.) at different sites in Slovenia during the droughty growing season of 2003 was studied. We also determined the timing of cambial activity, xylem and phloem formation, and counted the number of cells in the completed phloem and xylem growth rings. Light microscopy of cross-sections revealed that cambial activity started on the phloem and xylem side simultaneously at all three plots. However, prior to this, 1–2 layers of phloem derivatives near the cambium were differentiated without previous divisions. The structure of the early phloem was similar in silver fir and Norway spruce. Differences in the number of late phloem cells were found among sites. Phloem growth rings were the widest in Norway spruce growing at the lowland site. In all investigated trees, the cambium produced 5–12 times more xylem cells than phloem ones. In addition, the variability in the number of cells in the 2003 growth ring around the stem circumference of the same tree and among different trees was higher on the xylem side than on the phloem side. Phloem formation is presumably less dependent on environmental factors but is more internally driven than xylem formation.     

Comparative axial widening of phloem and xylem conduits in small woody plants

Key message

Along the stem axis phloem’s sieve elements increase in diameter basally at rates comparable to those of xylem conduits and in agreement with principles of hydraulic optimization.

Abstract

Plant physiology relies on the efficiency of the two long-distance transport systems of xylem and phloem. Xylem architecture comprises conduits of small dimensions towards the stem apex, where transpiration-induced tensions are the highest along the root-to-leaves hydraulic pathway, and widen basally to minimize the path length resistance to water flow. Instead, information on phloem anatomy and allometry is extremely scarce, although potentially relevant for the efficiency of sugar transportation. We measured the hydraulic diameter (Dh) of both xylem conduits and phloem sieve elements in parallel at different heights along the stem of a small tree of Picea abies, Fraxinus excelsior and Salix eleagnos. Dh increased from the stem apex to base in both xylem and phloem, with a higher scaling exponent (b) of sieve elements than that of tracheids in the conifer (0.19 vs. 0.14) and lower than that of vessels in the angiosperms (0.14–0.22 vs. 0.19–0.40). In addition, sieve elements were larger than tracheids in P. abies and narrower than angiosperms vessels at any height along the stem. In conclusion, axial conduit widening would seem to be a key feature of both xylem and phloem long-distance transport architectures.     

Estimating the absorptive root area in Norway spruce by using the common direct and indirect earth impedance methods

Aims

Estimates of root absorption magnitude are needed for the balanced management of forest ecosystems, but no methods able to work on the whole tree and stand level were available. Modified earth impedance method was developed recently and here it was tested, by comparing the results with those obtained by combination of several classical methods.

Methods

We used direct (soil cores, scanning and microscopy) and indirect (sap flow patterns and modified earth impedance) methods in an attempt to estimate the absorptive root area indexes (RAI) at two sites of about 25 and 40-years-old Norway spruce. We considered the geometric surfaces of all scanned fine roots to be equal to the fine root absorptive area (RAI _scan ). To estimate the potentially physically permeable area of fine roots, we microscopically evaluated the point of secondary xylem appearance and calculated the geometric area of root portions with primary structure (RAI _micro ). We termed the area of electrically conductive root surface as the active (ion) absorptive area (RAI _mei ) and measured its extent by the modified earth impedance (MEI) method.

Results

The highest values for absorptive root areas at the two experimental sites we obtained with the scanning method (RAI _scan was considered to be 100%), followed by the RAI _micro (51%) and RAI _mei (32%). RAImei reached about 2/3 of RAImicro. The surface area of the ectomycorrhizal hyphae was an order of magnitude larger than that of all fine roots, but the MEI did not measure such increase.

Conclusions

We showed that the absorptive root area, indirectly estimated by the MEI, provides consistent results that approach the values obtained for fine roots with a primary structure estimated by traditional direct methods. The similar range of the values for the absorptive root surface area obtained by microscopy and by the MEI method indicates that this method is feasible and that it could be used to determine the extent of active absorptive root surface areas in forests.     

The influence of alkaline and non-alkaline parent material on Norway spruce tree chemical composition and growth rate

Background and aims

This study investigated the influence of contrasting parent materials on tree chemical composition and growth rate under field conditions. On the island of Alnö, Sweden, alkaline intrusions are interspersed into non-alkaline gneiss bedrock, which provides a unique opportunity to conduct this study with a minimum of confounding effects.

Methods

Three plots with alkaline and three plots with non-alkaline parent material were established in a homogenous Norway spruce stand. The chemical composition of soil and soil solution was determined throughout the soil profiles. The chemical composition of bark, wood and needles was determined for each plot, and the latest 5 year basal area growth increment calculated.

Results

Concentrations of Ca in needles were correlated with the soil exchangeable Ca levels. Tree growth rate was significantly higher on the alkaline plots and positively correlated with soil concentrations of Ca, Mg, P, and Zn. The tree growth rate also tended to correlate with soil N concentrations, but levelled out for the highest soil N concentrations.

Conclusions

Tree growth was enhanced on the alkaline plots and was correlated with several elements. However, none of these elements could be confirmed as the limiting one for tree growth at the current site.     

Association genetics of phenolic needle compounds in Norway spruce with variable susceptibility to needle bladder rust

Key message

Accumulation of phenolic needle metabolites in Norway spruce is regulated by many genes with small and additive effects and is correlated with the susceptibility against fungal attack.

Abstract

Norway spruce accumulates high foliar concentrations of secondary phenolic metabolites, with important functions for pathogen defence responses. However, the molecular genetic basis underlying the quantitative variation of phenolic compounds and their role in enhanced resistance of spruce to infection by needle bladder rust are unknown. To address these questions, a set of 1035 genome-wide single nucleotide polymorphisms (SNPs) was associated to the quantitative variation of four simple phenylpropanoids, eight stilbenes, nine flavonoids, six related arithmetic parameters and the susceptibility to infection by Chrysomyxa rhododendri in an unstructured natural population of Norway spruce. Thirty-one significant genetic associations for the flavonoids gallocatechin, kaempferol 3-glucoside and quercetin 3-glucoside and the stilbenes resveratrol, piceatannol, astringin and isorhapontin were discovered, explaining 22–59% of phenotypic variation, and indicating a regulation of phenolic accumulation by many genes with small and additive effects. The phenolics profile differed between trees with high and low susceptibility to the fungus, underlining the importance of phenolic compounds in the defence mechanisms of Norway spruce to C. rhododendri. Results highlight the utility of association studies in non-model tree species and may enable marker-assisted selection of Norway spruce adapted to severe pathogen attack.

     

Shoot hydraulic characteristics, plant water status and stomatal response in olive trees under different soil water conditions

Aims

To evaluate the impact of the amount and distribution of soil water on xylem anatomy and xylem hydraulics of current-year shoots, plant water status and stomatal conductance of mature ‘Manzanilla’ olive trees.

Methods

Measurements of water potential, stomatal conductance, hydraulic conductivity, vulnerability to embolism, vessel diameter distribution and vessel density were made in trees under full irrigation with non-limiting soil water conditions, localized irrigation, and rain-fed conditions.

Results

All trees showed lower stomatal conductance values in the afternoon than in the morning. The irrigated trees showed water potential values around ?1.4 and ?1.6 MPa whereas the rain-fed trees reached lower values. All trees showed similar specific hydraulic conductivity (K _s) and loss of conductivity values during the morning. In the afternoon, K _s of rain-fed trees tended to be lower than of irrigated trees. No differences in vulnerability to embolism, vessel-diameter distribution and vessel density were observed between treatments.

Conclusions

A tight control of stomatal conductance was observed in olive which allowed irrigated trees to avoid critical water potential values and keep them in a safe range to avoid embolism. The applied water treatments did not influence the xylem anatomy and vulnerability to embolism of current-year shoots of mature olive trees.     

Identification of three wheat globulin genes by screening a Triticum aestivum BAC genomic library with cDNA from a diabetes-associated globulin

Background

Plant systemic signaling characterized by the long distance transport of molecules across plant organs involves the xylem and phloem conduits. Root-microbe interactions generate systemic signals that are transported to aerial organs via the xylem sap. We analyzed the xylem sap proteome of soybean seedlings in response to pathogenic and symbiotic interactions to identify systemic signaling proteins and other differentially expressed proteins.

Results

We observed the increase of a serine protease and peroxidase in the xylem sap in response to Phytophthora sojae elicitor treatment. The high molecular weight fraction of soybean xylem sap was found to promote the growth of Neurospora crassa in vitro at lower concentrations and inhibit growth at higher concentrations. Sap from soybean plants treated with a P. sojae elicitor had a significantly higher inhibitory effect than sap from control soybean plants. When soybean seedlings were inoculated with the symbiont Bradyrhizobium japonicum, the abundance of a xyloglucan transendoglycosyl transferase protein increased in the xylem sap. However, RNAi-mediated silencing of the corresponding gene did not significantly affect nodulation in soybean hairy root composite plants.

Conclusion

Our study identified a number of sap proteins from soybean that are differentially induced in response to B. japonicum and P. sojae elicitor treatments and a majority of them were secreted proteins.     

The natural abundance of 15N in litter and soil profiles under six temperate tree species: N cycling depends on tree species traits and site fertility

Aims

We investigated the influence of tree species on the natural ¹⁵N abundance in forest stands under elevated ambient N deposition.

Methods

We analysed δ¹⁵N in litter, the forest floor and three mineral soil horizons along with ecosystem N status variables at six sites planted three decades ago with five European broadleaved tree species and Norway spruce.

Results

Litter δ¹⁵N and ¹⁵N enrichment factor (δ¹⁵N_litter–δ¹⁵N_soil) were positively correlated with N status based on soil and litter N pools, nitrification, subsoil nitrate concentration and forest growth. Tree species differences were also significant for these N variables and for the litter δ¹⁵N and enrichment factor. Litter from ash and sycamore maple with high N status and low fungal mycelia activity was enriched in ¹⁵N (+0.9 delta units) relative to other tree species (European beech, pedunculate oak, lime and Norway spruce) even though the latter species leached more nitrate.

Conclusions

The δ¹⁵N pattern reflected tree species related traits affecting the N cycling as well as site fertility and former land use, and possibly differences in N leaching. The tree species δ¹⁵N patterns reflected fractionation caused by uptake of N through mycorrhiza rather than due to nitrate leaching or other N transformation processes.     

Overstorey and juvenile response to thinning and drought in a jarrah (Eucalyptus marginata Donn ex Sm.) forest of southwestern Australia

Aims

Forest thinning is expected to affect tree water use and carbon assimilation, but the related influence from climate variability is little known. Recent forest thinning in the Wungong catchment coincided with a record dry year following the thinning, which provides a rare opportunity to understand the climate influence on the thinning effect.

Methods

A field experiment was conducted to examine changes before and after thinning, especially the rainfall, soil moisture, leaf water status, tissue isotope signature (¹³?C and ¹⁵?N) and N concentration of overstorey and understorey juvenile trees of Eucalyptus marginata (Donn ex Sm.).

Results

Despite the post-thinning drought, surface soil was moister and juvenile jarrah plants were less water stressed, attributable to reduced rain interception and transpiration as a result of less canopy cover. The overstorey was under stress but mainly due to drought rather than by thinning. The concentration of N declined in both tree stems and juvenile leaves along with available N in soil, suggesting a soil N limitation. No treatment effects were detected from leaf relative water content and tissue isotope signature (¹³?C and ¹⁵?N).

Conclusions

The drought effects were superimposed over the thinning effects on overstorey growth, with stemwood δ¹³C being a major indicator of water stress. The water relations and carbon assimilation of understorey juveniles were however dependent more on topsoil moisture, and the wetter soil during the year following thinning enhanced growth activity and hence the depletion of ¹³?C (more negative δ¹³C) in juvenile leaves.     

Xylem and phloem phenology in co-occurring conifers exposed to drought

Key message

Variability in xylem and phloem phenology among years and species is caused by contrasting temperatures prevailing at the start of the growing season and species-specific sensitivity to drought.

Abstract

The focus of this study was to determine temporal dynamics of xylem and phloem formation in co-occurring deciduous and evergreen coniferous species in a dry inner Alpine environment (750 m a.s.l., Tyrol, Austria). By repeated micro-sampling of the stem, timing of key phenological dates of xylem and phloem formation was compared among mature Pinus sylvestris, Larix decidua and Picea abies during two consecutive years. Xylem formation in P. sylvestris started in mid and late April 2011 and 2012, respectively, and in both years about 2 week later in P. abies and L. decidua. Phloem formation preceded xylem formation on average by 3 week in P. sylvestris, and c. 5 week in P. abies and L. decidua. Based on modeled cell number increase, tracheid production peaked between early through late May 2011 and late May through mid-June 2012. Phloem formation culminated between late April and mid-May in 2011 and in late May 2012. Production of xylem and phloem cells continued for about 4 and 5–6 months, respectively. High variability in xylem increment among years and species is related to exogenous control by climatic factors and species-specific sensitivity to drought, respectively. On the other hand, production of phloem cells was quite homogenous and showed asymptotic decrease with respect to xylem cells indicating endogenous control. Results indicate that onset and culmination of xylem and phloem formation are controlled by early spring temperature, whereby strikingly advanced production of phloem compared to xylem cells suggests lower temperature requirement for initiation of the former.