Effects of prolonged drought on the anatomy of sun and shade needles in young Norway spruce trees

Predicted increases in the frequency and duration of drought are expected to negatively affect tree vitality, but we know little about how water shortage will influence needle anatomy and thereby the trees’ photosynthetic and hydraulic capacity. In this study, we evaluated anatomical changes in sun and shade needles of 20‐year‐old Norway spruce trees exposed to artificial drought stress. Canopy position was found to be important for needle structure, as sun needles had significantly higher values than shade needles for all anatomical traits (i.e., cross‐sectional needle area, number of tracheids in needle, needle hydraulic conductivity, and tracheid lumen area), except proportion of xylem area per cross‐sectional needle area. In sun needles, drought reduced all trait values by 10–40%, whereas in shade needles, only tracheid maximum diameter was reduced by drought. Due to the relatively weaker response of shade needles than sun needles in drought‐stressed trees, the difference between the two needle types was reduced by 25% in the drought‐stressed trees compared to the control trees. The observed changes in needle anatomy provide new understanding of how Norway spruce adapts to drought stress and may improve predictions of how forests will respond to global climate change.     

Stem radius changes and their relation to stored water in stems of young Norway spruce trees

Changes in the stem radius of young Norway spruce [Picea abies (L.) Karst.] were related to changes in stem water content in order to investigate the relationship between diurnal stem size fluctuations and internally stored water. Experiments were performed on living trees and on cut stem segments. The defoliated stem segments were dried under room conditions and weight (W), volume (V), and xylem water potential (O_s) were continuously monitored for 95 h. Additionally, photos of cross-sections of fresh and air-dried stem segments were taken. For stem segments we found that the change in V was linearly correlated to the change in W as long as O_s was >-2.3ǂ.3 MPa (phase transition point). Stem contraction occurred almost solely in the elastic tissues of the bark (cambium, phloem, and parenchyma), and the stem radius changes were closely coupled to bark water content. For living trees, it is therefore possible to estimate the daily contribution of "bark water" to transpiration from knowledge of the stem size and continuous measurements of the stem radius fluctuations. When O_s reaches the phase-transition point, water is also withdrawn from the inelastic tissue of the stem (xylem), which - in the experiment with stem segments - was indicated by an increasing ratio between (V and (W. We assume that for O_s below the transition point, air is sucked into the tracheids (cavitation) and water is also withdrawn from the xylem. Due to the fact that in living P. abies O_s rarely falls below -2.3ǂ.3 MPa and the xylem size is almost unaffected by radius fluctuations, dendrometers are useful instruments with which to derive the diurnal changes in the bark water contents of Norway spruce trees.     

Triadimefon reduces transpiration and increases yield in water stressed plants

The total free amino acid pools in radicles of watermelon seeds, investigated during imbibition of water at 25°C, were higher under the most (darkness) than under the least (continuous broad spectrum far-red light) favourable light regime for germination. When seeds were imbibed in an appropriate osmotic solution of PEG-6000 (fully suppressing germination), in darkness or under continuous red or far-red light, the biochemical analyses of the radicles after 1,2,3 and 4 days from the onset of imbibition show that while the total soluble sugar content remains rather constant in all treatments, significant changes are observed in the total free amino acid pools. After the first day, a considerable increase characterizes the "darkness" pool in contrast to a moderate one under red, while the "far-red" pool remains constant. Ultimately, at 4 days, the three pools are 190,142 and 123% of the 0 day radicle one. The qualitative free amino acid determination of the 4 day darkness and far-red pools shows a considerably increased percentage contribution of glutamic acid, arginine and citrulline in the "darkness" pool. The free amino acid increase in non-illuminated radicles may be correlated to germinability; moreover, it is evidently a phytochrome-mediated, pre-germinatory event, probably due to the hydrolysis of proteins (known to be rich in glutamic acid and arginine), stored in the radicle.     

No relation between drought stress and ethylene production in Norway spruce

The relationship between water availability and ethylene production was studied in 24‐year‐old trees of Norway spruce [ Picea abies (L.) Karst.] growing on experimental plots with different water availability. Ethylene and oxygen were collected from the cambial/xylem region and heartwood in the stem using non‐destructive sampling methods. Xylem sap flow was measured in stems using a heat‐balance technique. Pre‐dawn water potential of shoots was used to assess the water status of the trees. Growth was calculated from increments in stem basal area. The highest ethylene concentrations were found in irrigated trees that also, as compared to the other treatments, showed the most rapid flow rate of sap, the highest pre‐dawn water potential, and the most rapid growth. By contrast, the lowest ethylene concentrations were measured in trees to which artificial drought was induced. Such trees also showed the lowest water transport, lowest water potential and relatively slow growth. Thus, no signs of drought‐induced ethylene production were found in this study, contrary to the general contention of a positive relation between drought stress and ethylene production.     

Effect of nitrogen on drought strain and nutrient uptake in Norway spruce Picea abies (L.) Karst.) trees

A field lysimeter study was established with the aim of investigating the effect of nitrogen availability upon drought strain in Norway spruce trees. Forest soil (Typic Udipsamment) was filled in lysimeters 1 m in diameter and 1 m deep. Small trees of Norway spruce from five different clones were planted in the lysimeters. Roofs under the canopy of the trees ensured full control of water and nutrient input. Three levels of nitrogen were given to the trees during five years; ambient rainwater, and five and fifteen times this N concentration, respectively. Additional N was given as NH₄NO₃ in irrigation water. Mean annual N-addition during the five years corresponded to 5, 27 and 82 kg per ha and year for the three treatments, respectively. During the third and fifth growth season drought was artificially induced. In addition to a watered control, two levels of drought were applied, representing water deprivation for 2 and 3 months, respectively, in 1990 and 3 and 4 months, respectively in 1992. A higher water consumption in the nitrogen fertilized trees during the droughts resulted in a significantly lower pre-dawn shoot water potential compared to the trees receiving ambient rain N. The interaction between drought and nitrogen fertilization was clear also for photosynthesis and transpiration. A decrease in height- and diameter increment caused by drought was most pronounced in the 82 kg N ha^–1 yr^–1 treatment. A water strain integral showed a strong positive correlation to the needle biomass of the trees. Foliar concentrations of several nutrients decreased significantly with increasing drought strain in the trees. Concentration of potassium and boron were especially low and visual symptoms of deficiency occurred.     

Redistribution of water via layering branches between connected parent and daughter trees in Norway spruce clonal groups

Key message

We measured sap flow and shoot water potentials in clonally connected parent and daughter trees. We found bidirectional flow patterns in branches mediating the connection between parent and daughter trees.

Abstract

Layering is an important mode of vegetative reproduction at treeline, in which clonal daughter trees are formed by the rooting of lower (“layering”) branches of the parent tree. These branches mediate the connection between parent (PT) and daughter tree (DT). Here, we measured quantity and direction of sap flow in layering branches as well as PT and DT, and measured shoot water potentials in the crowns of a connected PT and DT. We found bidirectional sap flow pattern in layering branches, with the bidirectionality of the flow resulting from water potential dynamics of the parent and daughter trees varying diurnally. We found that 4.3 % of the total water transpired by the DT was supplied by the PT root system, with up to 25 % of the instantaneous daughter tree sap flow coming from the parent tree. In contrast, water provided by the daughter’s root system to the parent tree comprised only a negligible amount, less than 1 % of the parent’s entire sap flow. Additionally, after experimental excavation of part of the DT roots, layering branch flow towards the DT increased, while flows in the opposite direction almost vanished. This study showed that aboveground clonal connections can facilitate a new type of hydraulic redistribution where water is transported bidirectionally through branches. This transfer of water and nutrients is vital especially in the first years of the daughter tree but supplies considerable amounts of water even several years after the establishment of a new clonal tree.

     

Photosynthesis and transpiration of healthy and diseased spruce trees in the course of three vegetation periods

Summary CO₂- and H₂O-gas exchange of 20- to 25-year-old spruce trees from a plantation in the Hunsrück mountains were investigated over a period of 3 years. All measurements were made as pair comparisons, i.e., in each case the gas exchange of a damaged tree and of a relatively healthy tree in its immediate vicinity was measured simultaneously. A second plantation in the Westerwald mountains consisted of 18-year-old apparently healthy spruce trees. Pair comparison at this location meant comparison of two healthylooking trees. The investigations at both locations included diurnal course measurements of photosynthesis and transpiration, and light saturation curves and CO₂-saturation curves of photosynthesis. The reduced photosynthesis parameters of the phenotypically damaged trees at the Hunsrück location indicates massive damage to the photosynthetic apparatus. Measurements of H₂O-gas exchange showed that there are disturbances in stomatal regulation of the needles of damaged trees. As a result, the water use efficiency of these needles proved to be significantly lower. In addition, apparent photorespiration of the damaged trees was decreased, whereas their light- and CO₂-compensation points and their dark respiration were increased. In contrast to the Hunsrück plantation, no such effects were detectable when the healthy-looking Westerwald trees were subjected to pair comparison of gas exchange. Reduced photosynthetic capacity and disturbances of the stomatal regulation of the phenotypically damaged Hunsrück trees may be due to damage in the cellular membranes. Furthermore, a comparison of three growing seasons led to the conclusion that the gas exchange of spruce trees in their natural habitat is markedly influenced by climatic conditions.     

Seasonal variation in CO2 efflux of stems and branches of Norway spruce trees

BACKGROUND AND AIMS: Stem and branch respiration, important components of total forest ecosystem respiration, were measured on Norway spruce (Picea abies) trees from May to October in four consecutive years in order (1) to evaluate the influence of temperature on woody tissue CO2 efflux with special focus on variation in Q10 (change in respiration rate resulting from a 10 degrees C increase in temperature) within and between seasons, and (2) to quantify the contribution of above-ground woody tissue (stem and branch) respiration to the carbon balance of the forest ecosystem. METHODS: Stem and branch CO2 efflux were measured, using an IRGA and a closed gas exchange system, 3-4 times per month on 22-year-old trees under natural conditions. Measurements of ecosystem CO2 fluxes were also determined during the whole experiment by using the eddy covariance system. Stem and branch temperatures were monitored at 10-min intervals during the whole experiment. KEY RESULTS: The temperature of the woody tissue of stems and branches explained up to 68% of their CO2 efflux. The mean annual Q10 values ranged from 2.20 to 2.32 for stems and from 2.03 to 2.25 for branches. The mean annual normalized respiration rate, R10, for stems and branches ranged from 1.71 to 2.12 micromol CO2 m(-2)s (-1) and from 0.24 to 0.31 micromol CO2 m(-2) s(-1), respectively. The annual contribution of stem and branch CO2 efflux to total ecosystem respiration were, respectively, 8.9 and 8.1% in 1999, 9.2 and 9.2% in 2000, 7.6 and 8.6% in 2001, and 8.6 and 7.9% in 2002. Standard deviation for both components ranged from 3 to 8% of the mean. CONCLUSIONS: Stem and branch CO2 efflux varied diurnally and seasonally, and were related to the temperature of the woody tissue and to growth. The proportion of CO2 efflux from stems and branches is a significant component of the total forest ecosystem respiration, approx. 8% over the 4 years, and predictive models must take their contribution into account.     

土壤干旱条件下不同施钾水平对烟草光合速率和蒸腾效率的影响

研究了土壤干旱条件下,不同的施钾水平对烟草光合速率,胞间CO2浓度,气孔导度,蒸腾速率,蒸腾效率及生物量的影响。结果表明：在土壤干旱条件下适量施钾可以减少叶肉细胞光合活性的下降,消弱非气孔因素对光合的限制,增强气孔调节能力,提高蒸腾效率,并获得较高的生物量。     

Effects of drought and waterlogging on ultrastructure of Scots pine and Norway spruce needles

Effects of water stress on needle ultrastructure of 2-year-old Scots pine (Pinus sylvestris L.) and 5-year-old Norway spruce [Picea abies (L.) Karst.] seedlings were studied in greenhouse experiments. Drought stress was induced by leaving seedlings without watering, and waterlogging stress was produced by submerging the seedling containers in water. Needle samples for ultrastructural analyses were collected several times during the experiments, and samples for nutrient analyses at the end of the experiments. In drought stress, plasmolysis of mesophyll and transfusion parenchyma tissues, aggregation of chloroplast stroma and its separation from thylakoids and decreased size and abundance of starch grains in needles of both species were observed. The concentration of lipid bodies around the chloroplasts were detected in pine needles. Calcium and water concentrations in spruce needles were lower by the end of the experiments compared to controls. In waterlogging treatment, swelling of phloem cells in pine needles and large starch grains, slight swelling of thylakoids and increased translucency of plastoglobuli in chloroplasts of both species studied were observed. The phosphorus concentration in pine needles was higher while phosphorus, calcium and magnesium concentrations in spruce needles were lower in the waterlogging treatments compared to controls. Typical symptoms induced by drought stress, e. g. aggregation of chloroplast stroma and its separation from thylakoids, were detected, but, in waterlogging stress, ultrastructural symptoms appeared to be related to the developing nutrient imbalance of needles.     

Size-dependent responses to summer drought in Scots pine, Norway spruce and common oak

In this study, we provide a detailed analysis of tree growth and water status in relation to climate of three major species of forest trees in lower regions of Bavaria, Southern Germany: Scots pine (Pinus sylvestris), Norway spruce (Picea abies) and common oak (Quercus robur). Tree-ring chronologies and latewood δ¹³C were used to derive measures for drought reaction across trees of different dimensions: growth reduction associated with drought years, long-term growth/climate relations and stomatal control on photosynthesis. For Scots pine, growth/climate relations indicated a stronger limitation of radial growth by high summer temperatures and low summer precipitation in smaller trees in contrast to larger trees. This is corroborated by a stronger stomatal control on photosynthesis for smaller pine trees under average conditions. In dry years, however, larger pine trees exhibited stronger growth reductions. For Norway spruce, a significantly stronger correlation of tree-ring width with summer temperatures and summer precipitation was found for larger trees. Additionally, for Norway spruce there is evidence for a change in competition mode from size-asymmetric competition under conditions with sufficient soil water supply to a more size-symmetric competition under dry conditions. Smaller oak trees showed a weaker stomatal control on photosynthesis under both dry and average conditions, which is also reflected by a significantly faster recovery of tree-ring growth after extreme drought events in smaller oak trees. The observed patterns are discussed in the context of the limitation-caused matter partitioning hypothesis and possible species-specific ontogenetic modifications.     

Water relations in Norway spruce trees growing at ambient and elevated CO2 concentrations

Water relations were studied in Norway spruce [Picea abies (L.) Karst.] trees grown at ambient (AC, 350 μmol mol⁻¹) and elevated (EC, 700 μmol mol⁻¹) CO₂ concentrations under temperate water stress. The results suggested that both crown position and variability in atmospheric CO₂ concentration are responsible for different patterns of crown water relations. Mean hourly sap flux density (FSA) showed higher values in upper crown position in comparison with the whole crown in both AC and EC treatments. Mean soil-to-leaf hydraulic conductance (G_Tsa) was 1.4 times higher for the upper crown than that calculated across the whole crown for the trees in AC. However, G_Tsa did not vary significantly with crown position in EC trees, suggesting that elevated CO₂ may mitigate differences in hydraulic supply for different canopy layers. The trees in EC treatment exhibited significantly higher values of F_SA measured on the whole crown level and slightly higher soil water content compared to AC treatment, suggesting more economical use of soil water and therefore an advantage under water-limited conditions.     

Cell structural changes in the needles of Norway spruce exposed to long-term ozone and drought

Effects of ozone and/or drought on Norway spruce needles werestudied using light microscopy and electron microscopy. Saplingswere exposed to ozone in open-top chambers during 1992–1995and also to drought in the late summers of 1993–1995.Samples from current and previous year needles were collectedfive times during 1995. Ozone increased the numbers of peroxisomesand mitochondria, which suggests that defence mechanisms againstoxidative stress were active. The results from peroxisomes suggestthat the oxidative stress was more pronounced in the upper sideof the needles, and those from mitochondria that defence wasmore active in the younger needle generation. Possibly due tothe good nitrogen status and the active defence, no ozone-specificchloroplast alterations were seen. At the end of the season,older needles from ozone treatments had smaller central vacuolescompared with other needles. Cytoplasmic vacuoles around thenucleus were increased by ozone in the beginning of the experiment,and did not increase towards the end of the season as in thecontrols. These results from vacuoles may indicate that ozoneaffected the osmotic properties of the cells. Decreased numberand underdevelopment of sclerenchyma cells and proliferationof tonoplast were related to nutrient imbalance, which was enhancedby drought. Larger vascular cylinders and more effective starchaccumulation before and after the drought periods compensatedfor the reduced water status. Numbers of peroxisomes and mitochondriawere increased in the drought-exposed needles before the onsetof drought treatments of the study year, i.e. these changeswere memory effects. Interactions between ozone and droughtwere few.     

Assessing the impact of elevated O3 and CO2 on gas exchange characteristics of differently K supplied clonal Norway spruce trees during exposure and the following season

 Well-supplied and K-deficient 4-year-old clonal Norway spruce trees were exposed to combinations of two levels of ozone (20 and 80 nl l^{–  1} O₃) and carbon dioxide (350 and 750 μl l^{–  1} CO₂) to study the effects of possible future climate factors on gas exchange characteristics. The fumigation was performed in environmental chambers for a complete growing season. After the exposure, plants were cultivated outdoors to investigate possible recovery and delayed effects. During the exposure 1-year-old needles responded to the 80 nl l^{–  1} O₃ treatment by a sharp but transient decrease of both apparent carboxylation efficiency (CE) and maximum photosynthetic capacity (A₂₅₀₀). Elevated CO₂ also reduced CE and A₂₅₀₀. The effect became stronger in the course of the exposure and was accompanied by decreases of N and P as well as chlorophyll contents. In case of K deficiency, the acclimation response of current-year needles was even more pronounced reflecting lower sink capacities for carbon metabolites. The joint application of elevated O₃ and CO₂ resulted in the lowest values of gas exchange parameters and chlorophyll contents. At the beginning of the growing season after the exposure and under outdoor conditions, all these treatment effects disappeared in the needles which had developed during the fumigation. In the course of the development of the new flush, however, the well-supplied 1-year-old needles which had been treated with 80 nl l^{–  1} O₃ and 350 μl l^{–  1} CO₂ in the year before, exhibited a sharp decline of CE and A₂₅₀₀. Simultaneously, chlorotic mottle and bands developed. These delayed symptoms are discussed in the context of the previously published “memory” effect for O₃ (Sandermann et al. 1989). Additionally, evidence is presented that shoot development is altered in plants which had been exposed to elevated O₃. Accepted: 5 August 1996     

Effect of nitrates supplied with the transpiration flow on assimilate translocation

Solutions of nitrates (0.5% KNO₃, 0.2% NH₄NO₃) or urea (0.15%) were fed under the pressure of 10⁴ Pa to 50–60-cm-long detached shoots of common flax (Linum usitatissimum L.). One hour after the start of supplying the solutions, an assimilation clip chamber was fastened to the middle part of the shoot (¹⁴C source area), and ¹⁴CO₂ was blown through in the light for 2.5 min. The analysis of distribution of ¹⁴C among the labeled products of photosynthesis produced by source leaves showed that nitrates reduced the incorporation of the label into sucrose. At the same time, the ratio of labeled sucrose to labeled hexoses decreased, and the incorporation of the label into serine greatly increased. Urea did not produce such effects. The pattern of distribution of ¹⁴C within the plant 3 h after the assimilation of ¹⁴CO₂ points to the suppression of assimilate efflux from the leaves of plants fed with nitrates. In plants supplied with water or urea, 17–20% of labeled carbon was found below the ¹⁴C source area of the shoot, in nitrate type of treatment, only 3–5% was found there. In plants supplied with nitrates, the cortex tissue below the source leaf contained more ¹⁴C in proteins and less in low-molecular substances. In the wood tissue, such a correlation was not observed. When the shoot was supplied with water or urea, the content of ¹⁴C in sucrose in the source leaves in 3 h declined from 55–60% to 38–42%. When the shoot was fed with nitrates, the share of label in sucrose increased from 50 to 62–73%. Autoradiography of the source leaves showed that, in plants supplied with water or urea, the label was predominantly accumulated in large vascular bundles, and in nitrate type of treatment, it was accumulated outside large bundles. Electron microscopy showed that, in nitrate plants, the companion cells of phloem endings were very much vacuolated.     

Patterns and mechanisms of transpiration in a large subalpine Norway spruce (Picea abies (L.) Karst.)

In situ water relations of a large subalpine Norway spruce (Picea abies) were analyzed by simultaneous measurements of sap flow at different crown positions. In the diurnal scale, transpiration varied greatly, both spatially and temporally. Over longer periods, however, different parts of the crown transpired in fairly constant proportions. The average estimated transpiration was about 3.5 times greater in the upper than in the lower half and decreased 1.6-fold from south to north. Water intercepted from rain, fog and dew buffered and significantly decreased the transpiration. The effect was strongest in those parts which were least coupled to the free atmosphere. The top of the crown seemed to experience a regular shortage of water shortly after starting transpiration, when it was forced to switch from internal reserves to sources in the soil. Further, lower branches then started transpiring, which may have led them to compete for the water. An enhanced nocturnal sap flow during warm and dry winds (Foehn) indicated that the tree also transpired at night. Shaded twigs had more capacity to intercept water externally than twigs in the sun. The significance of the crown structure for interaction with water in both liquid and vapour phases is discussed.     

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.     

Stem respiration of Norway spruce trees under elevated CO2 concentration

Measurements of stem respiration were conducted for a period of four years (1999–2002) in 14-year old Norway spruce (Picea abies [L.] Karst) trees exposed to ambient (CA) and elevated CO₂ concentration (CE; ambient plus 350 μmol mol⁻¹). Stem respiration measurements of six trees per treatment were carried out 2–3 times per month during the growing season. Stem respiration in CE treatment was higher (up to 16 %) than in CA treatment. Temperature response of stem respiration (Q₁₀) for the whole experimental period ranged between 1.65–2.57 in CA treatment and 2.24–2.56 in CE treatment. The mean stem respiration rate normalized to 10 °C (R₁₀) in CA and CE treatments ranged between 1.67–1.95 and 2.19–2.72 μmol(CO₂) m⁻² s⁻¹, respectively. Seasonal variations in stem respiration were related to temperature and tree growth.     

Effect of drought stress on chlorophyll a fluorescence and electrical admittance of shoots in Norway spruce seedlings

Effects of mild and severe soil drought on the water status of needles, chlorophyll a fluorescence, shoot electrical admittance, and concentrations of photosynthetic pigments in needles of seedlings of Picea abies (L.) Karst. were examined under controlled greenhouse conditions. Drought stress reduced shoot admittance linearly with a decrease in shoot water potential (_w) and increase in water deficit (WD) and led to a decrease in concentrations of chlorophyll a, b and carotenoids. Severe water stress (shoot _w=–2.4 MPa) had a negative effect on chlorophyll a fluorescence parameters including PSII activity (F_v/F_m), and the vitality index (R_fd). Variations in these parameters suggest an inhibition of the photosynthetic electron transport in spruce needles. Water stress led to a decrease in the mobility of electrolytes in tissues, which was reflected by decreased shoot electrical admittance. After re-watering for 21 days the WD in needles decreased and the shoot water potential increased. In the re-watered plants, the chloroplast function was restored and chlorophyll a fluorescence returned to a similar level as in the control plants. This improved hydraulic adjustment in the seedlings triggered a positive effect on ion flow in the tissues and increased shoot electrical admittance. We conclude that the shoot electrical admittance and photosynthetic electron transport in leaves are closely linked to changes in water status and their decrease is among the initial responses of seedlings to water stress.