Seasonal dynamics of major cations in xylem sap and needles of Norway spruce (Picea abies L. Karst.) in stands with different soil solution chemistry

Soil solution, xylem sap and needles of mature trees were sampled in three spruce stands over one vegetation period and analysed for major cations. Investigations of nutrient distribution between these three pools and evaluations of seasonal dynamics give the following results: The highest nutrient concentrations in the xylem sap occur at the time of bud break and become gradually lower during the vegetation period. The trees show similar trends of xylem sap concentrations with time for potassium, calcium and magnesium regardless of the nutritional status of the plots. No coupling of xylem sap to soil solution composition can be observed in spite of a high variability of soil solution chemistry in time. The major cations in the current needles exhibit a significantly different trend with time. No time-based correlations for nutrient contents could be found for the needles from the previous year.Despite mobilisation of storage pools in the deficient stand, trees are not able to increase the Ca and Mg contents in the needles up to the level of the other stands. Potassium could be retranslocated in sufficient extent for nutrition of current needles. Due to seasonal variability and dependence upon internal processes, such as retranslocation and mobilisation of nutrients, xylem sap does not seem to be a good tool for the estimation of the nutritional status of forest sites.It was concluded that only minor transport into new foliage via xylem sap will proceed after nutrient flush during the bud break and the nutrient content in the new biomass will be governed by dilution due to biomass growth or by nutrient transport by other means than xylem sap.     

Scarification,fertilization and herbicide treatment effects on planted conifers and soil fertility

The influences of soil surface modification (blade scarification and plastic mulching), fertilization and herbicide application on soil nutrient and organic carbon content and tree growth and foliar nutrient status were examined after seven years in a study located within the Great Lakes-St. Lawrence forest region of Canada. Plots had been planted with white pine (Pinus strobus L.) and white spruce (Picea glauca [Moench] Voss) seedlings. Light (PAR), soil moisture and temperature were monitored and recorded throughout the growing season. Forest floor and soil mineral (0–20 cm layer) samples were collected from all experimental plots, except those which had plastic mulching. Foliar samples were collected in autumn and analysed for N, P and K and storage compounds. Seedling mortality was 20% higher in unscarified plots. Combined silvicultural treatments increased productivity as much as 14 times, but scarification reduced soil carbon and nutrient capital 2–3 fold. Herbicide application reduced soil carbon by at least 20 %. Foliar nutrient, protein, starch and lipid contents in autumn were little affected by treatment. The future management of such stands in Canada probably will include more shelterwood harvesting and crop rotations, silvicultural systems that are more closely aligned with natural forest succession.     

Seasonal development of the photosynthetic performance of Norway spruce (Picea abies [L.] Karst.) under magnesium deficiency

In order to investigate the influence of different magnesium nutrition on photosynthesis, one hundred 6-year-old spruce trees derived from one clone were planted in October 1990 into a special out-door experimental construction, where they were cultivated in sand culture with an optimal supply of nutrients, except magnesium, via circulating nutrient solutions. Magnesium was added to the nutrient solutions in three different concentrations, varying from optimal to severe deficient supplies. During the first vegetative period in 1991, photosynthetic performance and carboxylation efficiency were measured under saturating light, controlled CO₂ conditions, optimal temperature and humidity, using a minicuvette system.During summer, the trees under moderate magnesium deficiency developed tip yellowing symptoms on older needles, while the youngest needles remained green with unchanged chlorophyll contents. Trees under severe magnesium deficiency showed yellowing symptoms on all needle age classes combined with decreased chlorophyll contents in the youngest needles as well. In comparison with the controls, the photosynthetic performance of the 1-year-old needles was significantly lower in both deficiency treatments. The same was observed in the youngest needles of the trees under severe deficiency. Trees under moderate deficiency treatment decreased in photosynthetic performance during the summer without reduction of chlorophyll contents. The reduction of photosynthetic rates corresponded to a decrease in carboxylation efficiency, which is taken as a measure of the activity of the enzyme ribulose-1,5-bisphosphate carboxylase. This reduction, together with the observed increase of carbohydrate contents in needles of trees growing under magnesium deficiency, led to the assumption that the photosynthetic carbonfixation is reduced as a consequence of the accumulation of carbohydrates.     

Effect of Soil pH on Growth and Cation Deposition in the Root Tip of <Emphasis Type="BoldItalic">Zea mays</Emphasis> L.

Abstract The effects of sandy soil pH on the distribution of growth velocities and on cation concentrations and deposition rates in root growth zones of Zea mays L. seedlings were investigated. The pH values of the rooting medium varied between 4.2 and 8.6 in sand culture (70% saturated) without external supply of nutrients. At all pH values, densities (in 7moles per g fresh weight) of potassium, magnesium, and calcium increased toward the root tip. Lower pH in the medium increased calcium tissue density fivefold and magnesium density 1.7-fold, whereas the density of potassium, the overall elongation rate, and the growth velocity distribution did not show any significant pH dependence. Throughout the growth zone the deposition rates of the divalent cations, as calculated on the basis of the continuity equation, increased with lower pH. The data are consistent with the hypothesis that the effects of pH on the cation deposition rates are due to the increase in the divalent cation concentration of the soil solution at low pH and that the abundant uronic acid residues of the young walls of the meristem provide a reservoir of storage capacity for Ca and Mg under conditions of low nutrient availability.     

Responses of Pinus clausa, Pinus serotina and Pinus taeda seedlings to anaerobic solution culture. II. Changes in tissue nutrient concentration and net acquisition

Fifteen or 30 days of anaerobic growth conditions significantly reduced shoot and root nitrogen, potassium, phosphorus, iron and manganese concentrations in seedlings of pond pine ( Pinus serotina Michx.), sand pine [ P. clausa (Engelm.) Sarg.] and drought-hardy and wet-site loblolly pine ( P. taeda L.) grown in a culture system using non-circulating, continuously flowing solution. Calcium and shoot magnesium levels were least affected by anaerobic growth conditions – largely reflecting the passive nature of their uptake. Shoot and root nutrient content (mg nutrient pot^-1) followed similar trends, with wet-site loblolly and pond pine seedlings least affected by anaerobic solution culture. Shoot biomass of wet-site loblolly and pond pine seedlings was not affected by anaerobiosis, suggesting an increase in shoot nutrient utilization efficiency. Root biomass was significantly reduced by 15 or 30 days of anaerobiosis, with sand pine exhibiting the largest reduction in root dry weight (57%).
These results suggest that anaerobiosis interferes with net nutrient acquisition, even under the high nutrient conditions provided by solution culture. Sand pine suffered the largest reductions in shoot and root biomass and nutrient concentrations, showing earlier symptoms of waterlogging injury and nutrient stress than drought-hardy loblolly pine seedlings. Whether net nutrient acquisition decreased because of the reduction in root surface area available for absorption and/or reduced uptake efficiency cannot be ascertained from these data.     

Competition and facilitation between tree species change with stand development

Processes governing tree interspecific interactions, such as facilitation and competition, may vary in strength over time. This study tried to unveil them by performing dendrometrical analyses on black spruce Picea mariana, trembling aspen Populus tremuloides and jack pine Pinus banksiana trees from pure and mixed mature boreal forest stands in the Clay Belt of northwestern Quebec and on the tills of northwestern Ontario. We cored 1430 trees and cut 120 for stem analysis across all stand composition types, tree species and study regions. Aspen annual growth rate was initially higher when mixed with conifers, but then progressively decreased over time compared to pure aspen stands, while jack pine growth rate did not differ with black spruce presence throughout all stages of stand development. When mixed with aspen, black spruce showed a contrary response to aspen, i.e. an initial loss in growth but a positive gain later. On the richer clay soil of the Quebec Clay Belt region, however, both aspen and spruce responses in mixed stands reversed between 37 and 54 years. Overall, our results demonstrate that interspecific interactions were present and tended to change with stand development and among species. Our results also suggest that the nature of interspecific interactions may differ with soil nutrient availability.     

Fractionated extraction and speciation of macronutrients from Scots pine and Norway spruce needles of different ages

Summary The relationship between proteins and the macroelements potassium, magnesium, calcium, sulphur and phosphorus was studied in homogenates of needles of different ages from Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.]. Complete extractions by acid digestion, protein extractions by a buffer-detergent and non-protein extractions by a buffer alone showed that most of the potassium and magnesium of the needles was soluble independent of the proteins. Only a minor part (50–60 ppm, dry weight) of the magnesium could be referred to the chlorophyll content of the needles. Both potassium and magnesium appeared independent of the protein gel filtration. This was also valid for the minor fraction of the total calcium content, which was extractable in a buffer. Heterogeneous calcium deposits are also suggested by extraction using a chelator (EGTA). Part of the buffer-soluble sulphur and phosphorus compounds occurred independently of solubilized proteins, and had complex distribution patterns after gel filtration, even outside the high molecular separation range. It is suggested that further analyses along the present lines, compared with conventional analyses on total extracts, may extend the usefulness of mineral nutrient analyses in plants.     

Mineral Nutrient Requirements of Pinus silvestris and Picea abies Seedlings

The mineral nutrient requirements of Pinus silvestris L. and Picea abies Karst. were studied according to previously published methods applied to a series of various plant species. The optimum nutrient proportions are similar to those of Vaccinium, with a lower relative potassium requirement than birch and other broad-leaf species. Various ratios between ammonium and nitrate nitrogen were about equally efficient except for a minor growth reduction with pure nitrate, which gave a comparatively low nitrogen content and a high cation/nitrogen ratio. The rate of ammonium uptake was much higher than that of nitrate when both sources were supplied. The required total concentration in the nutrient solution for maximum growth is lower in pine than in spruce, but both fall within the low salt range. Both species, especially pine, are sensitive to high salt concentrations. Although pine and spruce grow on the same type of soils as Vaccinium— leached soils with low base saturation – accumulation of calcium or other cations is not as pronounced as in Vaccinium, especially not in pine. The results are compared with results from similar experiments with a series of other conifers. All the conifers have more flexible cation uptake mechanisms than Vaccinium but the results indicate tendencies to accumulation of anions, nitrate and phosphate. Recommended fertilizer compositions for forest fertilization and nurseries are discussed.     

Variation in tissue element concentrations in Quercus ilex L. over a range of different soils

In order to study the variability in nutrient concentrations in four tissues of Q. ilex in relation to soil properties, we selected fifteen stands in both Quercus ilex forests and Q. ilex-Pinus halepensis mixed forests. These stands had developed on soils derived from eight different parent materials. Three soil groups were differentiated according to their chemical properties: calcareous soils, siliceous soils, and volcanic soils. Across sites, nutrient concentrations were generally less variable in current-year tissues than in older tissues. Nitrogen and potassium showed the lowest variability among sites, their concentrations in current-year leaves ranging from 1.17% to 1.39% for N and from 0.53% to 0.68% for K. There were few statistically significant correlations between tissue element concentrations, the most frequent being the antagonistic relationship between calcium and magnesium. Nitrogen concentration in current-year leaves was negatively correlated with soil chemical fertility (nitrogen, phosphorus and potassium). This may reflect a nutritional imbalance between nitrogen and other nutrients, some of which may be more limiting than nitrogen to Q. ilex growth in Catalonia forests. Negative correlations were also found between plant magnesium and soil calcium, and positive correlations between plant calcium and soil calcium.     

Studies on chemical composition ofChenopodium album L.

Summary Field studies conducted at the Haryana Agricultural University, Hissar, India for two years revealed thatChenopodium album L. contained very high degree of nitrogen, phosphorus, potassium, calcium, magnesium, iron and manganese. Its nutrient content declined with advancement in age of the plant.     

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

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

Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary Photosynthetic rates and nutrient contents of spruce needles were measured in a region with high levels of air pollution in NE Bavaria, Germany (FRG), and compared to spruce grown under clean air conditions at Craigieburn, in the South Island of New Zealand (NZ). The absolute rates of CO₂ uptake, the slope of the CO₂ response curve at 240 l l^–1 internal CO₂ concentration, and the change of photosynthetic rates with needle age at ambient and saturated CO₂ concentrations were virtually identical at both measuring sites. These results confirm an earlier conclusion, that there is no long-term effect of atmospheric pollutants directly on photosynthetic CO₂ uptake rates with persistent exposure at the FRG site to high levels of anthropogenic air pollution. Photosynthetic capacity at saturating CO₂ concentration was three times higher in the NZ spruce. Needles with high photosynthetic capacity in NZ had lower nitrogen and higher calcium concentrations per unit dry weight but higher concentrations of nitrogen, phosphorus, potassium, magnesium and calcium per unit leaf area, and twice the specific leaf weight.     

Variation in fine root biomass of three European tree species: Beech (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.), and Scots pine (Pinus sylvestris L.)

Abstract

Fine roots (<2 mm) are very dynamic and play a key role in forest ecosystem carbon and nutrient cycling and accumulation. We reviewed root biomass data of three main European tree species European beech, (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.) and Scots pine (Pinus sylvestris L.), in order to identify the differences between species, and within and between vegetation zones, and to show the relationships between root biomass and the climatic, site and stand factors. The collected literature consisted of data from 36 beech, 71 spruce and 43 pine stands. The mean fine root biomass of beech was 389 g m^?2, and that of spruce and pine 297 g m^?2 and 277 g m^?2, respectively. Data from pine stands supported the hypothesis that root biomass is higher in the temperate than in the boreal zone. The results indicated that the root biomass of deciduous trees is higher than that of conifers. The correlations between root biomass and site fertility characteristics seemed to be species specific. There was no correlation between soil acidity and root biomass. Beech fine root biomass decreased with stand age whereas pine root biomass increased with stand age. Fine root biomass at tree level correlated better than stand level root biomass with stand characteristics. The results showed that there exists a strong relationship between the fine root biomass and the above-ground biomass.     

Effects of aluminium on growth and nutrient uptake of small Picea abies and Pinus sylvestris plants

Summary The effects of aluminium concentrations between 0.2 and 30 mM at pH 3.8 ±0.2 on small plants of Norway spruce [(Picea abies (L.) Karst], Scots pine (Pinus sylvestris L.), and Scots pine infected with the ectomycorrhizal fungus Suillus bovinus (L. ex Fr.) O. Kuntze were investigated. The plants were grown at maximum relative growth rate (R_G % day^–1) with free access but very low external concentrations of nutrients. Steady-state conditions with respect to relative growth rate (R_G) and internal nutrient concentrations were achieved before addition of aluminium, which was added as AlCl₃ and/or Al(NO₃)₃. There were reductions in rg at aluminium concentrations of 0.3 mM in spruce, 6 mM in pine and 10 mM in ectomycorrhizal pine, i. e. at aluminium concentrations considerably higher than those normally occurring in the top layer of the mineral soil where most fine roots are found. Nutrient uptake rate per unit root growth rate was calculated for different nutrient elements. The uptake rate of calcium and magnesium was reduced at aluminium concentrations of 0.2 mM (spruce), 1 mM (pine) and 3 mM (ectomycorrhizal pine), without influencing R_g. The results question the validity of the hypothesis of aluminium toxicity to forest tree species at low external concentrations.     

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.     

Quantitative study and modelling of the litter decomposition in a European alluvial forest. Is there an influence of overstorey tree species on the decomposition of ivy litter (Hedera helix L.)?

The influence on the decomposition rate of ivy litter (Hedera helix L.) of three ligneous overstorey species (oak, Quercus robur L., white poplar, Populus alba and ash, Fraxinus excelsior L.) that support ivy was studied in an alluvial hardwood forest. The ivy provides an abundant litter at the end of spring. The decomposition of ivy litter and the nutrient release rate were analyzed over four months during the growing season of the canopy trees, the hypothesis being that throughfall could slow down the mass loss rate of ivy depending on the support species. Mathematical models for mass loss and nutrient (P, N, K and Mg) release rates were developed. Mass loss rate and the release rate of magnesium, nitrogen and phosphorus show significant differences depending on species whereas no influence of species was observed on the release rate of potassium. The results illustrate the significant effect of oak compared to ash and poplar in slowing down mass loss rate and nitrogen, phosphorus and magnesium release rates. The mass loss rate over time under the three species followed the proposed exponential model, whereas the release rates of phosphorus and magnesium did not follow this model. This model is fitted to the data in two cases out of three for potassium. Magnesium and potassium are released more rapidly when nitrogen accumulates and the phosphorus content does not change significantly. No species × date interaction was observed, except in the case of magnesium. The species effect is interpreted as an effect of composition of throughfalls and presence or absence of inhibitory substances such as phenolic compounds.     

Production of Seeds and Cones and Consequences for Wood Radial Increment in Norway Spruce (Picea Abies (L.) Karst.)

Abstract

The presence of a trade–off between growth and reproduction was tested in four sites in a subalpine Norway spruce (Picea abies (L.) Karst.) forest by measuring annual stem diameter increments at breast height and seed and cone productions during the periods 1962–1985 and 1983–1990, respectively. Trees growing in forest stands near the timber line (about 1900 m above sea level) had the greatest reduction in annual stem diameter increment during mast years; while trees growing at about 1300–1500 m above sea level did not show any reduction. Trees growing at about 1700 m showed only a limited reduction. At the same elevation, trees growing within closed forest stands suffered a greater reduction in stem growth when compared with trees growing at the edge of a cutting.     

Parameter-specific hydroclimatic sensitivity of a low-elevation network of living and historical tree-ring series from north-eastern Austria

Compared to the alpine regions of Austria, the eastern part of the country is overall warmer and drier with some tree species growing at the dry limit within their ecological range. This suggests that tree rings may be a valuable proxy for hydroclimatic variability. In this study, we develop ring-width, earlywood-width and latewood-width chronologies obtained from drought sensitive living trees and historical timber from one of the driest and warmest regions of Austria, the Weinviertel. For this, samples of four different tree species − fir (Abies alba Mill.), spruce (Picea abies (L.) Karst.), oak (Quercus petraea Liebl., Q. robur L., Q. cerris L.), and pine (Pinus sylvestris L., P. nigra Arnold) were collected from 88 sites (23 forest stands and 65 historical constructions). 1589 samples (oak: 592 samples, pine: 577, spruce: 212, fir: 208) were dated. Dendro-provenancing was required to ensure that only regional material has entered the chronologies. According to the analyses, historical pine and oak wood were nearly 100% regional, while spruce and fir wood were imported to a great extent with only 44% spruce and 35% fir confirmed to be regional. Because site conditions of historical wood samples are unknown but can have a significant influence on tree growth, changes to site replication over time were important in the assessment of chronology reliability. We also controlled for potential artificial increases in sample depth when more than one construction element is made from a single trunk.We assessed the pine (1584-2011 AD) and oak (1244-2011 AD) chronologies’ potential for reconstructing past hydroclimatic variability by means of response functions in a 17-months window from previous June to current October with climate data from three weather stations (Vienna, Retz, Brno) from 1897 to 2010 AD. The highest seasonal response coefficients are for oak and pine ring width (∼0.43) with respect to aggregated March to July precipitation totals and Thornthwaite climatic water balance. These chronologies reveal a high potential for estimating past changes in regional-scale moisture availability during the earlier growing season.     

Climate‐diameter growth relationships of black spruce and jack pine trees in boreal Ontario,Canada

To predict the long‐term effects of climate change – global warming and changes in precipitation – on the diameter (radial) growth of jack pine (Pinus banksiana Lamb.) and black spruce (Picea mariana [Mill.] B.S.P.) trees in boreal Ontario, we modified an existing diameter growth model to include climate variables. Diameter chronologies of 927 jack pine and 1173 black spruce trees, growing in the area from 47°N to 50°N and 80°W to 92°W, were used to develop diameter growth models in a nonlinear mixed‐effects approach. Our results showed that the variables long‐term average of mean growing season temperature, precipitation during wettest quarter, and total precipitation during growing season were significant (alpha = 0.05) in explaining variation in diameter growth of the sample trees. Model results indicated that higher temperatures during the growing season would increase the diameter growth of jack pine trees, but decrease that of black spruce trees. More precipitation during the wettest quarter would favor the diameter growth of both species. On the other hand, a wetter growing season, which may decrease radiation inputs, increase nutrient leaching, and reduce the decomposition rate, would reduce the diameter growth of both species. Moreover, our results indicated that future (2041–2070) diameter growth rate may differ from current (1971–2000) growth rates for both species, with conditions being more favorable for jack pine than black spruce trees. Expected future changes in the growth rate of boreal trees need to be considered in forest management decisions. We recommend that knowledge of climate–growth relationships, as represented by models, be combined with learning from adaptive management to reduce the risks and uncertainties associated with forest management decisions.     

Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary A declining Picea abies (L.) Karst. stand produced as much foliage and branches as a healthy stand but less stemwood at a similar leaf area index and climate. Nutrient analyses revealed that most biomass components at the declining site had lower concentrations of calcium and magnesium, but similar nitrogen and potassium (except for lower potassium in younger needles) and higher phosphorus, manganese and aluminum than the respective components at the healthy site. Comparison of these data with the results from studies on the nutrition and growth of P. abies seedlings (Ingestad 1959) led to the conclusion that the healthy stand is in a balanced nutritional state, while trees at the declining stand have only 56% of the foliar magnesium concentration required to permit growth at a rate which could be achieved at their nitrogen status. It appears that acidic deposition, which involves an input of nitrogen and a leaching of cations from the soil, causes an imbalance in the availability of nitrogen and magnesium. Growth is eventually reduced as magnesium becomes limiting.