Effects of Alkalization of the Environment on the Anatomy of Scots Pine (Pinus sylvestris) Needles

The effects of alkaline dust emitted from a cement plant for over 40 years on the anatomy of needles of Scots pine and lignin accumulation were analysed. Comparative analytical studies were conducted in stands similar as to their silvicultural indicators, climate and age in alkalised and in a relatively unpolluted area. Cross-section of needles were stained, photographed under microscope and measured. It was found that, due to the alkalisation of the environment, the total area of the needle cross-section, needle width and thickness and the area of mesophyll had decreased. At the same time, the vascular bundles and epidermis had increased. The greatest anatomical and biochemical differences between the needles from trees growing under optimum conditions and in the alkalised area were observed in the oldest needles. Visual analysis of cross-sections and biochemical analysis showed accumulation of lignin in older needles but more intensively in alkalised areas than in control.     

Relationships between Lignin and Nutrients in Picea Abies L. under Alkaline Air Pollution

The influence of alkaline dust pollution (pH of water solution12.3–12.6) from a cement plant (Kunda, Estonia) and alkalizedgrowth substrate on lignin and mineral composition of the needles of 75–90-yr-old Norway spruce was studied on sample plots established at different distances from the emission source. Disbalanced content of nutrients and an increase in thelignin content were found under the high level of air pollution.The contents of K, Ca, S and B in needles were higher, while those of Mn and N were lower in heavily polluted plots. A strongcorrelation was found between lignin and K, N and B concentrations as well as Ca/K in needles and shoots. Stimulation of lignification of needles correlated with the pollution load. Reduction of needle growth was observed and a negative correlation was established between the lignin concentration in tissues and the length of needles under the impact of dust pollution.     

The influence of storm-induced microsites to tree regeneration patterns in boreal and hemiboreal forest

We reviewed studies dealing with regeneration under variable conditions in boreal and hemiboreal forests as affected by different microsite types by tree species functional groups. Generally, the importance of storm-induced microsites for regeneration dynamics in boreal forests depends on several factors: (1) distribution and type of microsites (generated by storm characteristics and stand conditions); (2) viable seed supply (stand history, species dispersal traits and status of surviving trees) and their species’ life history strategy; (3) climatic and site conditions (pre-storm conditions and storm-induced changes); and (4) delayed storm effects, such as retarded falling of trees, favoured vegetation growth, etc. Studies acknowledging the significance of microsites were mostly related to intermediate or severe events, causing sufficient changes in resource levels and growth conditions, and influencing extrinsic factors such as frost heaving, erosion and browsing. Also, the dispersal traits of available tree species, including sprouting and response of surviving trees, such as canopy expansion, should be considered in evaluating microsite importance in individual cases. In intermediate to severe windstorm events, pioneer species are generally profiting most from the additional offer in microsites, requiring bare mineral soil and elevated locations for their establishment and growth. Under gap dynamics, shade-tolerant species benefit from dead wood and elevated locations as these offer safe sites in stands with abundant understorey vegetation.     

The Quality of Stemwood of Pinus sylvestris in an Alkalised Environment

The impact of long-term dust pollution emitted from a cement plant on soil chemistry, and the concentrations of plant nutrients, lignin, cellulose and hemicellulose in the stemwood of 80–85-year-old Pinus sylvestris was investigated at different distances from the emission source. It was found that alkaline cement dust (pH 12.3–12.6) emissions for over 40 years resulted in an alkalisation (pH 6.7–7.9) of the polluted soil compared to a pH value of 3.8 in unpolluted soil. There were also nutrient imbalances in the soil, as well as certain disturbances in mineral nutrition processes and accumulation of nutrients in the tree stems. The average concentrations of K, Ca and Mg in stems were higher and those of N and P lower than in the unpolluted area. The lignin (L) content in stemwood increased, hemicellulose (Hc) decreased, while cellulose (Ce) did not change. A variation in the partitioning of L, Ce, Hc and nutrients between different sections of stems and between trees from different sample plots was found. L, Ce and Hc were not related to the internal K, Ca and Mg concentrations. Correlations were established between L, Ce, or Hc content and C content, and between L and Hc content in stem tissues. The contents of wood components were not related to N or P in the alkalised areas, but seemed to be more associated with P than with N. Alterations in the arbitrarily chosen ratio L/(Ce + Hc) indicated changes in wood quality, and a negative correlation with N/P was found in stem tissue in the polluted area, while positive correlations with N/Mg and Ca/Mg were found in the control area.     

Aboveground biomass and nutrient accumulation dynamics in young black alder, silver birch and Scots pine plantations on reclaimed oil shale mining areas in Estonia

The growth, aboveground biomass production and nutrient accumulation in black alder (Alnus glutinosa (L.) Gaertn.), silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) plantations during 7 years after planting were investigated on reclaimed oil shale mining areas in Northeast Estonia with the aim to assess the suitability of the studied species for the reclamation of post-mining areas. The present study revealed changes in soil properties with increasing stand age. Soil pH and P concentration decreased and soil N concentration increased with stand age. The largest height and diameter of trees, aboveground biomass and current annual production occurred in the black alder stands. In the 7-year-old stands the aboveground biomass of black alder (2100 trees ha⁻¹) was 2563 kg ha⁻¹, in silver birch (1017 trees ha⁻¹) and Scots pine (3042 trees ha⁻¹) stands respective figures were 161 and 1899 kg ha⁻¹. The largest amounts of N, P, K accumulated in the aboveground part were in black alder stands. In the 7th year, the amount of N accumulated in the aboveground biomass of black alder stand was 36.1 kg ha⁻¹, the amounts of P and K were 3.0 and 8.8 kg ha⁻¹, respectively. The larger amounts of nutrients in black alder plantations are related to the larger biomass of stands. The studied species used N and P with different efficiency for the production of a unit of biomass. Black alder and silver birch needed more N and P for biomass production, and Scots pine used nutrients most efficiently. The present study showed that during 7 years after planting, the survival and productivity of black alder were high. Therefore black alder is a promising tree species for the reclamation of oil shale post-mining areas.     

Plasticity in mesophyll volume fraction modulates light-acclimation in needle photosynthesis in two pines

Acclimation potential of needle photosynthetic capacity varies greatly among pine species, but the underlying chemical, anatomical and morphological controls are not entirely understood. We investigated the light-dependent variation in needle characteristics in individuals of Pinus patula Schlect. & Cham., which has 19-31-cm long pendulous needles, and individuals of P. radiata D. Don., which has shorter (8-17-cm-long) stiffer needles. Needle nitrogen and carbon contents, mesophyll and structural tissue volume fractions, needle dry mass per unit total area (M(A)) and its components, volume to total area ratio (V/A(T)) and needle density (D = M(A)/(V/A(T))), and maximum carboxylase activity of Rubisco (V(cmax)) and capacity of photosynthetic electron transport (J(max)) were investigated in relation to seasonal mean integrated irradiance (Q(int)). Increases in Q(int) from canopy bottom to top resulted in proportional increases in both needle thickness and width such that needle total to projected surface area ratio, characterizing the efficiency of light interception, was independent of Q(int). Increased light availability also led to larger M(A) and nitrogen content per unit area (N(A)). Light-dependent modifications in M(A) resulted from increases in both V/A(T) and D, whereas N(A) changed because of increases in both M(A) and mass-based nitrogen content (N(M)) (N(A) = N(M)M(A)). Overall, the volume fraction of mesophyll cells increased with increasing irradiance and V/A(T) as the fraction of hypodermis and epidermis decreased with increasing needle thickness. Increases in M(A) and N(A) resulted in enhanced J(max) and V(cmax) per unit area in both species, but mass-based photosynthetic capacity increased only in P. patula. In addition, J(max) and V(cmax) showed greater plasticity in response to light in P. patula. Species differences in mesophyll volume fraction explained most of the variation in mass-based needle photosynthetic capacity between species, demonstrating that differences in plastic adjustments in mass-based photosynthetic activities among these representative individuals were mainly associated with contrasting investments in mesophyll cells. Greater area-based photosynthetic plasticity in P. patula relative to P. radiata was associated with larger increases in M(A) and mesophyll volume fraction with increasing irradiance. These data collectively demonstrate that light-dependent increases in mass-based nitrogen contents and photosynthetic activities were associated with an increased mesophyll volume fraction in needles at higher irradiances. They also emphasize the importance of light-dependent anatomical modifications in determining needle photosynthetic capacity.     

Growth and Biomass Partitioning of 6-Year-Old Spruces under Alkaline Dust Impact

Two-year-old seedlings of Picea abies L., P. glauca L. and P. mariana L. were planted in a sample plot subjected to high concentrations of cement dust and in an unpolluted (control) area in 1990. In 1994, the six-year-old trees were dug up in the pre-bud-break period for morphological assessment. Comparison of biomass formation and the lengths of different organs of the trees showed that alkalization of the growth substrate (pH 8.1) and the high level of dust pollution load (600–2400 g m^-2yr^-1) are serious factors inhibiting height growth by 61%, the length of shoots and needles by 75 and 28%, respectively, and the dry weights of roots by 88%, stems by 90%, shoots by 91% and needles by 55%. The ratio of roots to shoots dry weights increased in all species of spruces under stress. P. mariana was the most sensitive to dust pollution impact and alkalization of the environment.     

Ground vegetation under natural stress conditions in Scots pine forests on fixed sand dunes in southwest Estonia

Ecosystems on dunes are influenced by critical environmental factors (mineral nutrients, water deficiency, etc.) considered decisive for their existence. The present paper is based on studies carried out on dunes on the coastal area of the Baltic Sea, southwest Estonia. The nature of forest ecosystems on dunes was studied from the aspects of chemical characteristics of soil, vascular plant species richness and diversity. Sampling sites on the dunes with different heights were selected in Cladina and in Vaccinium vitis-idaea site-type Scots pine forests. Vascular plant species richness and diversity were related to edaphic gradients. On the dune with a height of 32.1 m a.s.l., significant relationships were revealed between the number of species of ground vegetation, pH, volumetric water content in soil and the position of the sample plots. No relationships were revealed between the number of vascular plant species, soil pH, volumetric water content and mineral nutrients on the dune with a height of 9 m a.s.l. The most frequent and abundant plant species on the higher dune were Deschampsia flexuosa, Vaccinium vitis-idaea and V. myrtillus; the highest number of species were found at the bottom of the dune, while on the top only some xeromorphic species such as Festuca ovina, Sedum acre and Crepis tectorum occurred. On the lower dune, the most frequent were Vaccinium vitis-idaea, V. myrtillus and Melampyrum pratense, while V. uliginosum was found only on the bottom and slope and Empetrum nigrum on the top of the dune.     

Dendrochronological analysis of the growth and growth-climate relationships of conifers in the region of alkaline dust deposition

Relationships between climate and radial growth of Oxalis-Myrtillus-site type Norway spruce and Scots pine stands under different cement dust loads were investigated. Dendrochronological methods were used. Long-term alkaline (pH 13.2-12.7) dust pollution emitted over 40 years from a cement plant was the reason of alkalisation (pH 7.8-8.1) and high concentrations of K, Ca and Mg in soil of affected territories. Two study sites, Kunda and Malla (2.5 and 5.0 km E from the emission source), were influenced by the dust emissions of the cement plant. Two other study sites, Eru (38 km W) and Revoja (34 km W), were situated on a relatively unpolluted area and served as control sites. The relationships between the radial growth and climate were almost similar on the control and dust-polluted sites. The climatic variables that had a significant effect on the radial growth of Norway spruce at both control and polluted sites were the temperature and precipitation of summer months of the current and preceding years. Specific to the Scots pine stands growing under dust pollution was a significant positive impact of the current spring temperature on the radial growth. A positive effect of precipitation during winter months on the radial growth was found at all sites. A significant negative effect of cement dust on the radial growth of Scots pine stands during the period of large amounts of dust emission (1966-1991) was detected. The effect of cement dust emissions on the radial growth of Norway spruce stands was also negative but weak. The decrease in the dust emissions since 1992 improved the growth conditions at pine stand sites.     

Dependence of needle architecture and chemical composition on canopy light availability in three North American Pinus species with contrasting needle length

Morphology and chemical composition of needles of shade-intolerant southern conifers (Pinus palustris Mill. (mean needle length +/- SD = 29.1 +/- 4.1 cm), P. taeda L. (12.3 +/- 2.9 cm) and P. virginiana Mill. (5.1 +/- 0.8 cm)) were studied to test the hypothesis that foliage acclimation potential to canopy light gradients is generally low for shade-intolerant species, and in particular, because of mechanical limitations, in species with longer needles. Plasticity for each needle variable was defined as the slope of the foliar characteristic versus irradiance relationship. A novel geometrical model for needle area and volume calculation was employed for the three-needled species P. palustris and P. taeda. Needle thickness (T) strongly increased, but width (W) was less variable with increasing daily integrated quantum flux density averaged over the season (Q(int)), resulting in changes in cross-sectional needle shape that were manifested in a positive relationship between the total to projected needle area ratio (A(T)/A(P)) and Q(int) in the three-needled species. In contrast, cross-sectional needle geometry was only slightly modified by irradiance in the two-needled conifer P. virginiana. Needle dry mass per unit total needle area (M(T)) was positively related to Q(int) in all species, leading to greater foliar nitrogen contents per unit area at higher irradiances. Separate examination of the components of M(T) (density (D) and the volume (V) to A(T) ratio; M(T) = DV/A(T)) indicated that the positive effect of light on M(T) resulted solely from increases in V/A(T), i.e., from increases in the thickness of foliage elements. Foliar chlorophyll content per unit mass increased with increasing Q(int), allowing an improvement in light-harvesting efficiency in low light. The variables characterizing needle material properties (D, the dry to fresh mass ratio, and needle carbon content per unit mass) were generally independent of Q(int), suggesting that needles were less stiff and had greater tip deflections under their own weight at lower irradiances because of smaller W and T. Comparisons with the literature revealed that plasticity in foliar characteristics tended to be lower in the studied shade- intolerant species than in shade-tolerant conifers, but plasticity among the investigated species was unaffected by needle length. However, we argue that, because of mechanical limitations, plastic changes in needle cross section in response to low irradiance may decrease rather than increase light-interception efficiency in long-needled species.