Photosynthesis,nutrient, growth and soil investigations of a declining Norway spruce (Picea abies) stand in the coastal region of Northern Germany

The research site, Wingst Compartment 123B, is a 68-year-old Norway spruce (Picea abies (Karst.)) stand located in the coastal area of northern Germany. This area receives high atmospheric inputs of ammonium and also has relatively high ozone concentrations (0.061 mg m⁻³).Ten trees were categorized as healthy to slightly damaged (3–29% needle loss) or severely damaged (49–71% needle loss). Apparent net photosynthetic rates were measured on detached branches at light saturation (1000 μE m⁻² s⁻¹). Needles were analyzed for chlorophyll, N, C, Ca, Mg, K, Zn, Mn and Fe.When compared to the healthy-to-slightly-damaged trees, the severely damaged trees tended to have higher rates of net apparent photosynthesis in the 1 and 2-year-old needles and similar rates in the current-year needles. All three needle ages from the severely damaged trees had higher average stomatal conductances to water vapor (g_s. Although the damaged trees had significantly less total chlorophyll in all needle ages sampled, there was no statistically significant difference in the chlorophyll a:b ratio between the healthy and severely damaged trees.Nitrogen contents of the current-year needles were slightly lower in the severely damaged trees. Carbon and calcium levels did not significantly differ between the damage classes, although the average Ca content of all younger needles was generally under the recommended sufficiency levels. The current-year needles of the severely damaged trees had significantly less magnesium (0.42–0.46 mg Mg g⁻¹) than those of the healthy trees, and all trees had Mg values in the deficiency range (< 0.7 mg g⁻¹). The severely damaged trees also had lower average potassium levels in the older needles.The annual volume increment per unit crown surface area declined with increasing crown damage. Trees with a 50% needle loss showed a 62% loss of volume increment.Soil investigations revealed conditions of high soil acidity and poor nutrient capacity. The low pH values (pH < 3.8) in 64% of all samples indicated a high risk of acid toxicity for plant roots in the investigated area.The significance of these results relative to the current ideas concerning forest decline is discussed.     

Crown architecture and stemwood production in Norway spruce (Picea abies (L.) Karst.)

The morphological data for Norway spruce (Picea abies (L.) Karst.) trees published by Burger (1953) were reanalyzed to examine the relationships between crown shape, needle and branch mass density of crown volume, needle efficiency in stemwood production, harvest index and stemwood production efficiency per unit of crown projected area. Production of stemwood per unit of crown projected area was higher in narrow-crowned trees than in broad-crowned trees because the narrow-crowned trees had (i) smaller horizontal space requirements for equal crown volumes, (ii) higher leaf area index due to both the geometrical crown shape and higher needle density per unit of crown volume, and (iii) higher harvest index indicating high allocation of dry matter production to stem.     

Vitality rating of picea abies by defoliation class and other vigour indicators

The study was made to assess the relationship between visually estimated defoliation class and a number of other variables depicting the vitality of Norway spruce (n=50). Variables characterizing crown size and condition were determined on the standing trees. Electrical impedance (an indication of the physiological status of a tree) was measured in the inner bark tissue of the living trees. Shoot growth and needle variables were determined from the detached sample branches. Negative correlations were found between defoliation class and the growth parameters (5‐year height, radial and volume growth of the trunk). Positive correlations were observed between the needle loss class and the abundance of branch damage, secondary shoots and impedance values. 18 variables (defoliation excluded) describing tree size and vitality were summarized in a factor analysis incorporating 5 factors. These were interpreted as 1) vitality, 2) foliage discoloration, 3) tree size, 4) needle number and 5) needle size factors. The tree‐specific score values for the first factor were interpreted as “vitality indices”;. The rank correlation between these score values and defoliation degree was 0.835. This result suggests that the defoliation class and vitality parameters contributing to the first factor give a consistent estimate of the tree condition.     

Relationships between crown projected area and components of above-ground biomass in Norway spruce trees in even-aged stands: Empirical results and their interpretation

Sample tree material was reanalyzed in order to study the relationships between horizontal crown projected area and components of above-ground biomass in Norway spruce (Picea abies (L.) Karst.) trees growing in even-aged stands. The needle mass of dominant trees increased linearly with the increase in crown projected area, but in co-dominant and dominanted trees the increase in needle mass levelled off toward larger crown projected areas. The branch mass of dominant and co-dominant trees accumulated faster than linearly with increasing crown projected area, whereas in dominated trees an approximately linear relationship existed between these two variables. The increase in needle and branch mass per unit increase in crown projected area was highest in dominant trees and decreased to co-dominant and dominated trees, i.e. with tree position in the canopy. The stem mass accumulated obviously faster than linearly and similarly in all tree classes with the increase in crown projected area. The narrow crown shape indicated a high density of all components of above-ground biomass per unit of crown projected area.     

Treatment of Severely Checked Sitka Spruce Trees with Injected Nutrients

Matched pairs of nutrient deficient Sitka spruce (Picea sitchensis(Bong.) Carr.) of mean height 3.0 m on a drained acid infertiledeep peat were injected in early summer by infusion with anaqueous solution of 40, 20, 66 and 0.2 g l^–1 N, P, K,Mg respectively, pH 7.2 and high conductivity. Three growingseasons later there were no responses in injected trees in treeheight increment, foliage nutrient levels of N, P, K, Ca, needlecolour, needle weight or crown density. Foliage Mg levels wereraised significantly. The injection procedure caused severetissue damage around the Infusion point and 84 per cent of thetrees had stain in stem cross sections 50 cm above the treatmentlocation.     

Variation in the ratio of shoot silhouette area to needle area in fertilized and unfertilized Norway spruce trees

We compared the range and variation in shoot silhouette area to projected leaf area ratio (SPAR) in fertilized and unfertilized (control) Norway spruce (Picea abies (L.) Karst.) trees. We measured SPAR for several view directions of 169 shoots at different depths in the crown of fertilized and control trees. There was an increase in SPAR with depth in the crown in both control and fertilized trees. In the fertilized trees, however, mean SPAR was larger overall, the increase with depth in the crown was steeper, and there was a larger variation in SPAR with inclination and rotation angle of the shoot (relative to the view direction). In particular, shoots in the lower crown of fertilized trees were rotationally asymmetrical ("flat") and had high values of the maximum ratio of shoot silhouette area to projected leaf area (SPAR(max)). Differences in SPAR between fertilized and control trees were explained by changes in shoot structure in response to fertilization and shading. Shoots of fertilized trees were larger and had more needle area than shoots of control trees. However, the ratio of needle area to shoot size was smaller in fertilized trees than in control trees, implying less within-shoot shading and, consequently, a larger SPAR. Also, the increase in SPAR with increased shading (depth in the crown) could be explained by a decrease in the ratio of needle area to shoot size. In addition, because fertilized trees had more needle area than control trees, the effect of shading at a given depth in the crown was more pronounced in fertilized trees than in control trees.     

Effects of thinning on growth of six tree species in north-temperate forests of Lithuania

This paper summarises the results from 35 years-observed thinning experiments on 256 permanent sample plots in 10–60 year-old stands of ash, aspen, birch, oak, pine and spruce in Lithuania. Thinning enhanced crown projection area increment of residual trees. The largest effect was observed in stands of aspen and birch (growth increase by 200%), followed by ash and oak (over 100%), and spruce and pine (about 80%). Thinning also promoted dbh increment, especially in younger stands, and the increase of dbh increment was positively correlated with the thinning intensity. The strongest reaction was exhibited by oak and aspen, while ash, birch and conifers reacted to a lower extent. Low and moderate intensities of thinning stimulated volume production in younger stands while the opposite was observed in older stands with increasing removals. Spruce stands exhibited relatively strongest increase of volume increment and pine, –the weakest, while the effect on deciduous species was intermediate. The results demonstrate that significant increase in volume increment is achievable with thinning of only young forest stands, e.g. 10–20 year-old pine, birch and ash, or 10–30 year-old oak, aspen and spruce.     

The effects of Arceuthobium sichuanense infection on needles and current‐year shoots of mature and young Qinghai spruce (Picea crassifolia) trees

Arceuthobium sichuanense is a hemiparasitic angiosperm that infects Qinghai spruce (Picea crassifolia Kom.) in Qinghai province, China, and causes severe damage to spruce forests in Qinghai‐Tibet Plateau. In this study, the impact of A. sichuanense infection on mature and young trees of Qinghai spruce was evaluated by examining needle and current‐year shoot morphology, needle water and nitrogen‐use efficiency (NUE) and needle nitrogen concentration. The most apparent effect of A. sichuanense infection was a significant reduction in both needle size distal to infection and current‐year shoot length in the infected branches (p < 0.001). Per cent reductions in needle and current‐year shoot length were similar between mature and young trees (58.9 vs. 56.3%; 59.7 vs. 62.9%). There was a high degree of correlation in foliar δ¹⁵N values between the dwarf mistletoe and its host trees (R² = 0.9017, p < 0.001), while the foliar δ¹³C values of A. sichuanense were similar to those of infected mature and young spruce trees. The dwarf mistletoe infection also resulted in a significant decrease in host needle N concentration and δ¹³C values (p < 0.001). The per cent reduction in needle N concentration in young trees was nearly twice as much as that in mature trees (20.49 vs. 11.54%), while the per cent reduction in needle δ¹³C values was similar between young and mature trees (?0.98 vs.?1.1‰). The NUE in mature trees was not affected by A. sichuanense infection, but the NUE in young trees was increased by the infection.     

Rapid freezing induces winter injury symptomatology in red spruce foliage

Red spruce (Picea rubens Sarg.) suffers frequent and extensive injury to current-year foliage during the winter. Experimental freezing of red spruce foliage at cooling rates > 10 degrees C min(-1) induced visible symptomatology similar to natural winter injury at the branch, needle and cellular levels. Such damage was associated with a low-temperature exotherm near -10 to -12 degrees C, a loss in needle fluorescence, massive cellular disruption, foliar discoloration, and low needle survival. Susceptibility of individual trees to rapid freezing injury was associated with historical winter injury patterns and alterations in foliar nutrition. We conclude that anthropogenic deposition may alter the sensitivity of trees to winter injury caused by rapid temperature changes.     

Resistance to the green spruce aphid (Elatobium abietinum Walker) in progenies of Sitka spruce (Picea sitchensis (Bong) Carr.)

Damage caused by Elatobium abietinum was estimated as needle loss in a progeny trial of Sitka spruce in Denmark in 1990 and 1992. The parents of the progeny had been selected for putative aphid resistance during a massive attack of aphids in 1957. Needle loss was significantly less in the progeny than in a standard provenance. Family heritability was estimated as 0.72 (P>0.0001). Aphid damage had an impact on growth and vigour of trees following warm winters in 1989 and since, but no correlation could be found between growth parameters and damage of different families. Due to high heritability and large variation, high gains are expected by breeding. Practical and theoretical consequences of including aphid resistance in breeding programmes of Sitka spruce are discussed.     

Inherited narrow crown form, harvest index and stem biomass production in Norway spruce, Picea abies

The effect of crown form on stem biomass production was investigated in an 18-19-year-old Norway spruce stand (Picea abies (L.) Karst.). The harvest index was 0.271 in pendula trees, which have a heritable narrow crown form, and 0.235 in normal-crowned trees and the dry weights of stem biomass were 2.57 kg and 3.37 kg, respectively. However, the production of stem biomass per crown projected area was more than twice as much in pendula trees as in normal-crowned trees. Results indicate that the crown form of pendula spruce is less plastic than that of normal-crowned spruce suggesting that such trees might maintain a high stemwood yield per unit ground area with increasing stand density. Because pendula spruce has a higher harvest index and a less flexible crown than normal-crowned spruce, it might be useful for crop tree ideotype breeding.     

Crown architecture of understory and open-grown white pine (Pinus strobus L.) saplings

Crown architecture and growth allocation were studied in saplings of eastern white pine (Pinus strobus L.), a species classified as intermediate in shade tolerance. A comparison was made of 15 understory saplings and 15 open-grown saplings that were selected to have comparable heights (mean of 211 cm, range of 180-250 cm). Mean ages of understory and open-grown trees were 25 and 8 years, respectively. Understory trees had a lower degree of apical control, shorter crown length, and more horizontal branch angle, resulting in a broader crown shape than that of open-grown trees. Total leaf area was greater in open-grown saplings than in understory saplings, but the ratio of whole-crown silhouette (projected) leaf area to total leaf area was significantly greater in understory pine (0.154) than in open-grown pine (0.128), indicating that the crown and shoot structure of understory trees exposed a greater percentage of leaf area to direct overhead light. Current-year production of understory white pine was significantly less than that of open-grown white pine, but a higher percentage of current-year production was allocated to foliage in shoots of understory saplings. These modifications in crown structure and allocation between open-grown and understory white pine saplings are similar to those reported for more shade-tolerant fir (Abies) and spruce (Picea) species, but the modifications were generally smaller in white pine. As a result, white pine did not develop the flat-topped "umbrella" crown structure observed in understory fir and spruce, which approaches the idealized monolayer form that maximizes light interception. The overall change to a broader crown shape in understory white pine was qualitatively similar, but much more limited than the changes that occurred in fir and spruce. This may prevent white pine from persisting in understory shade as long as fir and spruce saplings.     

Mechanisms of Douglas-fir resistance to western spruce budworm defoliation: bud burst phenology, photosynthetic compensation and growth rate

We compared growth rates among mature interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) trees showing resistance or susceptibility to defoliation caused by western spruce budworm (Choristoneura occidentalis Freeman), and among clones and half-sib seedling progeny of these trees in a greenhouse. We also investigated bud burst phenology and photosynthetic responses of clones to budworm defoliation in greenhouse experiments. Resistant mature trees had a higher radial growth rate than susceptible trees, especially during periods of budworm defoliation. Clones from resistant trees grew larger crowns than clones from susceptible trees, whereas stem base diameter at the ground line and height did not differ. Half-sib seedling progeny from resistant trees had larger stem diameter, height, and total biomass than progeny from susceptible trees. Mean 5-year radial growth increment of mature trees was more strongly correlated with growth of seedlings than with growth of clones. Clones from resistant trees had later bud burst than clones from susceptible trees, and budworm defoliation of clones depended on the degree of synchrony between bud burst phenology and budworm larval feeding. Clones of resistant and susceptible mature trees showed similar responses of net photosynthetic rate to 2 years of budworm defoliation. We conclude that phenotypic differences in crown condition of Douglas-fir trees following western spruce budworm defoliation are influenced by tree genotype and that high growth rate and late bud burst phenology promote tree resistance to budworm defoliation.     

Radial,Height and Volume Increment Variation in Picea abies (L.) Karst. Stands with Varying Thinning Intensities

Variation in radial increment at different stem heights was compared with stem volume increment and height increment using thinning experiments of Picea abies (L.) Karst. in southern Finland. The magnitude of growth variation was similar on trees from different crown classes and stand densities. However, there was higher autocorrelation in the codominant and intermediate trees and trees on dense plots. The relationship between short-term volume increment and radial increment variation at breast height was not affected by thinnings. While short-term growth variation was similar at different stem heights, some evidence was found for differences in medium-term variation between radial increment at breast height and volume increment. Height increment reacted to unfavourable climatic events later than radial and volume increment. Radial and volume increments were negatively correlated to temperature and positively correlated to precipitation of the current summer. Height increment was positively related to current early summer temperature, but negatively to temperature of the previous summer.     

Autumn frost damage: clonal variation in Sitka spruce

Current-year needles of Sitka spruce (Picea sitchensis) can be injured by frost after hardening of stem and buds in the autumn, and in Scotland trees of southern provenances are most susceptible to damage. In October 1993 a frost of −5°C caused needle damage to clonal plants of Alaskan, Queen Charlotte Islands, and Washington provenances of Sitka spruce that had been selected for variation in the onset of root dormancy. Visual damage assessment revealed significant differences among clones in the severity of damage, and this damage was positively correlated with the lateness of root dormancy. Measurement of electrolyte leakage from shoot samples confirmed the observed differences between clones. The large clonal differences in frost hardiness found in this study demonstrate a potential for selecting frost tolerant trees from within southern provenances of Sitka spruce.     

Examining age- and altitude-related variation in tree architecture and needle efficiency in Norway spruce using trend surface analysis

The aim of this study was to examine the variation in tree architecture and needle efficiency in stemwood production in Norway spruce in relation to tree age and altitude of growing site. The data, which were obtained from the literature, described individual Norway spruce (Picea abies (L.) Karst.) trees from even-aged stands in Switzerland. Second-order trend surface models, with tree age and altitude as independent variables, were used in the analysis. The fitted models for stem, branch and needle dry masses explained 95%, 75% and 64% of the variation, respectively. The model for the estimated mean branch density in the crown (kg m⁻³) explained 64% of the variation and the model for mean needle density in the crown (kg m⁻³) only 28% of the variation. Crown structural characteristics, which showed age- and altitude-related variations, included live crown ratio (59% of variation explained), number of living whorls (43% of variation explained), mean weight of single needle (40% of variation explained) and specific needle area (27% of variation explained). Tree age had a strong effect on needle efficiency in stemwood production, so that needle efficiency increased up to the age of 50–70 years, depending on altitude.     

Fine root condition relates to visible crown damage in Norway spruce in acidified soils

Allochthonous Norway spruce stands in the Kysucké Beskydy Mts. (north‐western Slovakia) have been exposed to substantial acid deposition in the recent past and grow in acidified soil conditions with mean pH of about 4.0 in the topsoil. We selected 90 spruce trees representing 30 triplets of different crown status: healthy, stressed and declining to assess the relationship between crown and fine root status. Sequential coring and in‐growth bags were applied to each triplet to investigate fine root mass and growth in the soil depths of 0–10 and 10–20 cm. Fine root quantity (biomass and necromass), turnover (production over standing stock), morphological features (specific root length, root tip density) and chemical properties (Ca:Al molar ratio) were compared among the abovementioned health status categories. Living fine root biomass decreased with increasing stress, while the ratio of dead to living biomass increased. Annual fine root production decreased and specific root length increased in stressed trees when compared to healthy or declining trees, a situation which may be related to the position of trees in the canopy (healthy and declining – dominant, stressed – co‐dominant). The Ca:Al ratio decreased with increasing crown damage, indicating a decreased ability to filter out aluminium. In conclusion, fine root status appears to be linked to visible crown damage and can be used as a tree health indicator.     

Evidence that longer needle retention of spruce and pine populations at high elevations and high latitudes is largely a phenotypic response

There is abundant evidence that evergreen conifers living at high elevations or at high latitudes have longer-lived needles than trees of the same species living elsewhere. This pattern is likely caused by the influence of low temperature in combination with related factors such as a short growing season and low nutrient availability. Because it is not known to what degree such patterns result from phenotypic versus genotypic variation, we evaluated needle longevity for common-garden-grown lowland populations of European Scots pine (Pinus sylvestris L.) of wide latitudinal origin and Norway spruce (Picea abies L.) of wide elevational origin. Nine-year-old trees of 16 Scots pine populations ranging in origin from 47 degrees to 60 degrees N were studied in Kórnik, Poland (52 degrees N) and 18-year-old trees of 18 Norway spruce populations ranging in origin from 670 to 1235 m elevation in southwestern Poland were studied near Morawina, Poland (51 degrees N, 180 m elevation). There was no tendency in either species for populations from northern or high elevation origins to retain needles longer than other populations. All of the Scots pine populations had between 2.5 to 3.0 needle age cohorts and all of the Norway spruce populations had between 6.4 and 7.2 needle age cohorts. Thus, extended needle retention in Scots pine and Norway spruce populations in low-temperature habitats at high elevations and high latitudes appears to be largely an environmentally regulated phenotypic acclimation.     

Snow damage in lodgepole pine stands brought into thinning and fertilization regimes

Several heavy wet snowfalls occurred during 2007-2009 across a broad-scale thinning and fertilization experiment to bring overstocked juvenile lodgepole pine (Pinus contorta var. latifolia) in the foothills of Alberta, Canada into an intensive management regime. We examined the bending and breakage of trees in relation to thinning and fertilization and used a multimodel information-theoretic approach to model stand and tree level predictors of snow damage. Fertilized stands suffered the greatest amount of snow damage, and this was most noteworthy when stands were also thinned; here 22% (17% broken stems) of trees were damaged compared to 8% (4% broken stems) in the thinned and unfertilized stands. At the stand level, needle weight and crown cover were reliable predictors of snow damage. At the tree level, separate models were developed for each combination of thinning and fertilization. All models used total tree volume; usually the smaller trees in the stands were more susceptible to damage but in the thinned and fertilized stands larger but slender trees with large asymmetrical crowns tended to be damaged. Also, trees with lower total stem volume were more susceptible to damage. Only in the thinned and fertilized stands were variables related to crown shape and asymmetry important predictors of snow damage. We conclude that snow damage is an important agent for self-thinning in unthinned stands and fertilization tends to exacerbate damage because of increase in foliage size. In areas with regular occurrence of heavy snow, we do not recommend fertilization at the same time as thinning, as the larger and more economically important trees in the stand are at risk.     

Effect of soil chemical properties on growth,foliation and nutrition of Norway spruce stand affected by yellowing in the Bohemian Forest Mts., Czech Republic

In 1998, a fertilizer experiment aiming to investigate the effects of slow-release N, P, K and Mg fertilizer (SILVAMIX Mg NPK^®) on a 60-year-old spruce stand with symptoms of yellowing was established. In this paper, trees were selected to investigate the relation between annual diameter increment, yellowing, foliation, needle and soil chemical properties: ten from the fertilized treatment (F), ten green trees from the control (CG) and ten yellow trees from the control (CY). CG and CY trees were growing in close proximity at a distance of only several meters apart under the same soil conditions. In treatment F, increased annual diameter increment, improved foliation, needle Mg concentration, plant-available Mg and P concentrations in the soil and absence of yellow trees were recorded 7 years after a single application of the fertilizer. During the last 15 years, annual growth increment and foliation of CY trees have continuously decreased while relatively stable values were recorded for CG trees and increased for F trees. In 2006, CG and CY trees differed significantly in Mg concentration in needles, foliation, yellowing and annual diameter increment. Although differences in soil chemical properties between CG and CY treatments were not significant, lower concentrations of plant-available Mg²⁺ and higher concentrations of H⁺ and Al³⁺ were found in soils under CY trees. There was a negative correlation between soil concentration of Mg and yellowing, but this correlation was relatively weak, indicating that there is no simple relation between soil and needle concentrations of Mg. In the investigated locality, the “new type” of yellow tree decline has been a long-term gradual process.