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.     

Changes in the canopies of Pinus sylvestris and Picea abies under alkaline dust impact in the industrial region of Northeast Estonia

Studies were carried out in 1999, 2005 and 2007 in the area of Kunda cement plant in Northeast Estonia on sample plots 3 km W and 2.5 and 5 km E of Kunda. As control stands, two plots for pine and spruce were established in Lahemaa National Park (34-38 km W of Kunda). The selected pine and spruce stands were 75-85-year-old Myrtillus site type, of 0.7-0.8 density and II quality class, with moderately dense or sparse understorey. The values concerning needle density and number of needle scars were higher for shoots formed in the period of higher pollution than for the shoots grown under a considerably lower pollution load. Although the cement dust pollution has notably decreased from year to year, the number of needle pairs per 1 cm of the shoot was 1.8-2.1 times greater in the shoots formed in 1998 than in those formed in 2003, whereas the changes were statistically reliable. Possibly the low temperatures at the time of shoot and needle formation affected the density of needles on all sample plots, and thus the number of needles on shoots formed in 2003 was many times smaller. After the significant fall in the pollution load since 1996 the length growth of needles intensified around Kunda cement plant, at the same time no changes occurred in the length growth of needles in the control area. As compared to the data from 1998, the length growth of pine needles had improved, especially 2.5 and 5 km E from the cement plant, needles being respectively 1.5 and 1.1 cm longer than 6 years ago. The stimulation of the growth of pine and spruce needles 2.5 and 5 km E of the cement plant may be a sign of a positive effect of reduced doses of cement dust in soil. The greater length of pine and spruce needles is the reason for the larger biomass of the needles.     

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.     

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.     

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.