Wood properties of Populus and Betula in long‐term exposure to elevated CO2 and O3

We studied the interactive effects of elevated concentrations of CO₂ and O₃ on radial growth and wood properties of four trembling aspen (Populus tremuloides Michx.) clones and paper birch (Betula papyrifera Marsh.) saplings. The material for the study was collected from the Aspen FACE (free‐air CO₂ enrichment) experiment in Rhinelander (WI, USA). Trees had been exposed to four treatments [control, elevated CO₂ (560 ppm), elevated O₃ (1.5 times ambient) and combined CO₂ + O₃] during growing seasons 1998–2008. Most treatment responses were observed in the early phase of experiment. Our results show that the CO₂‐ and O₃‐exposed aspen trees displayed a differential balance between efficiency and safety of water transport. Under elevated CO₂, radial growth was enhanced and the trees had fewer but hydraulically more efficient larger diameter vessels. In contrast, elevated O₃ decreased radial growth and the diameters of vessels and fibres. Clone‐specific decrease in wood density and cell wall thickness was observed under elevated CO₂. In birch, the treatments had no major impacts on wood anatomy or wood density. Our study indicates that short‐term impact studies conducted with young seedlings may not give a realistic view of long‐term ecosystem responses.     

Severe photobiont injuries of lichens are strongly associated with air pollution

The photobiont ultrastructure of the epiphytic lichens Bryoria fuscescens and Bryoria fremontii was studied along the pollution gradient from two Cu-Ni smelters in Nikel and Monchegorsk in northern Finland and north-western Russia. The relationship between ultrastructural characteristics of B. fuscescens and environmental factors (i.e. climate, atmospheric SO₂ and bark element concentrations) was studied by using a principal component analysis (PCA) aiming to assess the air quality in a northern environment. Based on PCA, increased plasmolysis and mitochondrial changes in the Trebouxia photobiont were significantly correlated with elevated pollutant concentrations. Degenerated cells, showing altered chloroplasts and electron-translucent pyrenoglobuli, occurred in lichens growing 35–50 km from the Monchegorsk smelter. Cell wall and cytoplasmic lipid volumes, and size of pyrenoglobuli, positively correlated with the distance from the Monchegorsk smelter. Vacuoles and electron-opaque vacuolar deposits were significantly increased at the Finnish site in the vicinity of a pulp mill. Swelling of mitochondrial cristae and thylakoids showed little correlation with environmental factors, but indicated of initial stage of injuries and were observed at several slightly polluted sites in northern Finland and north-western Russia. The results suggest that the severe photobiont injuries of lichens are strongly associated with poor air quality.     

Herbivores inhibit climate-driven shrub expansion on the tundra

Recent Pan-Arctic shrub expansion has been interpreted as a response to a warmer climate. However, herbivores can also influence the abundance of shrubs in arctic ecosystems. We addressed these alternative explanations by following the changes in plant community composition during the last 10 years in permanent plots inside and outside exclosures with different mesh sizes that exclude either only reindeer or all mammalian herbivores including voles and lemmings. The exclosures were replicated at three forest and tundra sites at four different locations along a climatic gradient (oceanic to continental) in northern Fennoscandia. Since the last 10 years have been exceptionally warm, we could study how warming has influenced the vegetation in different grazing treatments. Our results show that the abundance of the dominant shrub, Betula nana , has increased during the last decade, but that the increase was more pronounced when herbivores were excluded. Reindeer have the largest effect on shrubs in tundra, while voles and lemmings have a larger effect in the forest. The positive relationship between annual mean temperature and shrub growth in the absence of herbivores and the lack of relationships in grazed controls is another indication that shrub abundance is controlled by an interaction between herbivores and climate. In addition to their effects on taller shrubs (>0.3 m), reindeer reduced the abundance of lichens, whereas microtine rodents reduced the abundance of dwarf shrubs (<0.3 m) and mosses. In contrast to short-term responses, competitive interactions between dwarf shrubs and lichens were evident in the long term. These results show that herbivores have to be considered in order to understand how a changing climate will influence tundra ecosystems.     

Stem wood properties of mature Norway spruce after 3 years of continuous exposure to elevated [CO2] and temperature

The objective of the study was to investigate the interactive effects of elevated atmospheric carbon dioxide concentration, [CO₂], and temperature on the wood properties of mature field-grown Norway spruce ( Picea abies (L.) Karst.) trees. Material for the study was obtained from an experiment in Flakaliden, northern Sweden, where trees were grown for 3 years in whole-tree chambers at ambient (365 μmol mol⁻¹) or elevated [CO₂] (700 μmol mol⁻¹) and ambient or elevated air temperature (ambient +5.6 °C in winter and ambient +2.8 °C in summer). Elevated temperature affected both wood chemical composition and structure, but had no effect on stem radial growth. Elevated temperature decreased the concentrations of acetone-soluble extractives and soluble sugars, while mean and earlywood (EW) cell wall thickness and wood density were increased. Elevated [CO₂] had no effect on stem wood chemistry or radial growth. In wood structure, elevated [CO₂] decreased EW cell wall thickness and increased tracheid radial diameter in latewood (LW). Some significant interactions between elevated [CO₂] and temperature were found in the anatomical and physical properties of stem wood (e.g. microfibril angle, and LW cell wall thickness and density). Our results show that the wood material properties of mature Norway spruce were altered under exposure to elevated [CO₂] and temperature, although stem radial growth was not affected by the treatments.     

Sex‐specific selection on energy metabolism – selection coefficients for winter survival

Selection for different fitness optima between sexes is supposed to operate on several traits. As fitness‐related traits are often energetically costly, selection should also act directly on the energetics of individuals. However, efforts to examine the relationship between fitness and components of the energy budget are surprisingly scarce. We investigated the effects of basal metabolic rate (BMR, the minimum energy required for basic life functions) and body condition on long‐term survival (8 winter months) with manipulated densities in enclosed populations of bank voles (Myodes glareolus). Here, we show that survival selection on BMR was clearly sex‐specific but density‐independent. Both the linear selection gradient and selection differential for BMR were positive in females, whereas survival did not correlate with male characteristics. Our findings emphasize the relative importance of individual physiology over ecological factors (e.g. intra‐specific competition). Most current models of the origin of endothermy underline the importance of metabolic optima in females, whose physiology evolved to fulfil demands of parental provisioning in mammals. Our novel findings of sex‐specific selection could be related to these life history differences between sexes.     

Winter herbivory by voles during a population peak: the relative importance of local factors and landscape pattern

1. We studied the relative role of local habitat variables and landscape pattern on vole–plant interactions in a system with grey-sided voles ( Clethrionomys rufocanus (Sund.)) and their favourite winter food plant, bilberry ( Vaccinium myrtillus L.). The study was conducted during a vole peak year (1992–93) in a tundra area in northern Norway.
2. Using Mantel statistics we were able to separate the direct effects of the spatial patterning of habitats and the indirect effects due to spatial aggregations of similar habitats.
3. Results indicate that knowledge about the explicit spatial patterning of patches does not improve our understanding of the system. Instead, two local factors, vegetation height and bilberry biomass, explained more than 50% of the variation in cutting intensity in winter (defined as the proportion of above-ground shoots cut). Increasing vegetation height increased, and increasing bilberry biomass decreased, the cutting intensity.
4. The conclusion that grey-sided voles are able to distribute themselves relative to habitat quality was also partially supported by our estimated over-winter persistence by voles in the various habitats. Vole persistence was uncorrelated with vegetation height, the important predictor of autumn vole density, but tended to correlate with the deviation from the relation between vegetation height and autumn vole density. This conforms to the expectations from the theory of ideal-free habitat distribution.
5. The cue for vole habitat choice, i.e. vegetation height, indicates that either predation or freezing risk is important for voles when selecting over-wintering habitat.     

Stomatal density, anatomy and nutrient concentrations of Scots pine needles are affected by elevated CO2 and temperature

Stomatal density, anatomy and nutrient concentrations of Scots pine (Pinus sylvestris L.) needles were studied during 3 years of growth at elevated CO₂ (693 ± 30 µmol mol⁻¹), at elevated temperature (ambient +2·8–6·2 °C depending on the time of the year) and in a combination of elevated CO₂ and temperature in closed-top chambers. The treatments were started in August 1996. At elevated temperature, the needles that were grown in the first year (i.e. the 1997 cohort) were thinner, had thinner mesophyll in the abaxial side, thinner vascular cylinder and lower stomatal density than those grown at ambient temperature. The proportion of mesophyll area occupied by vascular cylinder or intercellular spaces were not changed. Lower stomatal density apparently did not lead to decreased use of water, as these needles had higher concentrations of less mobile nutrients (Ca, Mg, B, Zn and Mn), which could indicate increased total transpiration. In the 1997 and 1998 cohorts, elevation of temperature decreased concentrations of N, P, K, S and Cu. In the 1999 cohort, contradictory, higher concentrations of N and S at elevated temperature may be related to increased nutrient mineralization in the soil. Elevation of CO₂ did not affect stomatal density, needle thickness, thickness of epidermis or hypodermis, vascular cylinder or intercellular spaces. Concentrations of N, P, S and Cu decreased at elevated CO₂. Reductions were transient and most distinct in the 1997 cohort. The effects of CO₂ and temperature were in some cases interactive, which meant that in the combined treatment stomatal density decreased less than at elevated temperature, and concentrations of nutrients decreased less than expected on the basis of separate treatments, whereas the thickness of the epidermis and hypodermis decreased more than in the separate treatments. In conclusion, alterations in the anatomy and stomatal density of Scots pine needles were more distinct at elevated temperature than at elevated CO₂. Both elevated CO₂ and temperature-induced changes in nutrient concentrations that partly corresponded to the biochemical and photosynthetic alterations in the same cohorts ( Luomala et al. Plant, Cell and Environment 26, 645–660, 2003 ) Reductions in nutrient concentrations and alterations in the anatomy were transient and more evident in the needle cohort that was grown in the first treatment year.     

The stability–diversity relationship in stream macroinvertebrates: influences of sampling effects and habitat complexity

1. Theory predicts that the stability of a community should increase with diversity. However, despite increasing interest in the topic, most studies have focused on aggregate community properties (e.g. biomass, productivity) in small‐scale experiments, while studies using observational field data on realistic scales to examine the relationship between diversity and compositional stability are surprisingly rare. 2. We examined the diversity–stability relationship of stream invertebrate communities based on a 4‐year data set from boreal headwater streams, using among‐year similarity in community composition (Bray–Curtis coefficient) as our measure of compositional stability. We related stability to species richness and key environmental factors that may affect the diversity–stability relationship (stream size, habitat complexity, productivity and flow variability) using simple and partial regressions. 3. In simple regressions, compositional stability was positively related to species richness, stream size, productivity and habitat complexity, but only species richness and habitat complexity were significantly related to stability in partial regressions. There was, however, a strong relationship between species richness and abundance. When abundance was controlled for through re‐sampling, stability was unrelated to species richness, indicating that sampling effects were the predominant mechanism producing the positive stability–diversity relationship. By contrast, the relationship between stability and habitat complexity (macrophyte cover) became even stronger when the influence of community abundance was controlled for. Habitat complexity is thus a key factor enhancing community stability in headwater streams.     

Effects of elevated O3, alone and in combination with elevated CO2, on tree leaf chemistry and insect herbivore performance: a meta-analysis

We reviewed the effects of elevated ozone (O₃), alone and in combination with elevated carbon dioxide (CO₂) on primary and secondary metabolites of trees and performance of insect herbivores by means of meta‐analysis. Our database consisted of 63 studies conducted on 22 species of trees and published between 1990 and 2005. Ozone alone had no overall effect on concentrations of carbohydrates or nutrients, whereas in combination with CO₂, elevated O₃ reduced nutrient concentrations and increased carbohydrate concentrations. In contrast to primary metabolites, concentrations of phenolics and terpenes were significantly increased by 16% and 8%, respectively, in response to elevated O₃. Effects of ozone in combination with elevated CO₂ were weaker than those of ozone alone on phenolics, but stronger than those of ozone alone on terpenes. The magnitude of secondary metabolite responses depended on the type of ozone exposure facility and increased in the following order: indoor growth chamber 3 than gymnosperms, as shifts in concentrations of carbohydrate and phenolics were observed in the former, but not in the latter. Elevated O₃ had positive effects on some indices of insect performance: pupal mass increased and larval development time shortened, but these effects were counteracted by elevated CO₂. Therefore, despite the observed increase in secondary metabolites, elevated O₃ tends to increase tree foliage quality for herbivores, but elevated CO₂ may alleviate these effects. Our meta‐analysis clearly demonstrated that effects of elevated O₃ alone on leaf chemistry and some indices of insect performance differed from those of O₃+CO₂, and therefore, it is important to study effects of several factors of global climate change simultaneously.     

Meteorological drivers of the dynamics of autotrophic picoplankton

1. The tiny non‐motile autotrophic picoplankton (APP; size range 0.2–2 μm) occur in all types of aquatic habitats and are comprised of prokaryotic as well as eukaryotic taxa. In the Boreal Zone, the majority of lakes have high concentrations of coloured humic substances that can adversely affect lake light climate and cause steep summertime stratification resulting in epilimnetic nutrient depletion. APP are more effective in nutrient and light acquisition than larger phytoplankton and thus should be competitive in humic lakes. 2. Most lacustrine APP studies have been based on short sampling periods, and thus, interannual variation and its drivers are still unclear. We studied APP in the small, boreal, humic Lake Valkea‐Kotinen during five open‐water periods in 2002–06 to determine interannual variation and the importance of meteorological drivers for APP dynamics. 3. Total APP showed a bimodal annual pattern, but the timing and vertical location of the two maxima varied during the study. In general, APP thrived in warm water and the most important abiotic factor controlling APP was stability of the water column (N_s). On average, 82% of APP were found in the epilimnion or metalimnion during summertime stratification. 4. There was niche separation of APP and larger phytoplankton in the lake because, with only one exception, APP maxima occurred separately from the maxima of larger phytoplankton. 5. Two groups, solitary eukaryotic APP and colonial picocyanobacteria (Merismopedia warmingiana), responded differently to the abiotic factors. Solitary APP preferred high water colour and low pH, both of which occurred after heavy rain, whereas colonial APP did not fare well when water colour was high. Our findings suggest that when future climate change‐related processes increase incoming allocthonous organic matter load from the catchment, solitary eukaryotic APP will be favoured.