Pollution resistance of mountain birch, Betula pubescens subsp. czerepanovii, near the copper-nickel smelter: natural selection or phenotypic acclimation?

Mountain birch, Betula pubescens subsp. czerepanovii (Orlova) Hamet-Ahti, is one of the very few woody plants surviving in the heavily polluted industrial barrens of the Kola Peninsula, north-western Russia. The present study was designed to find out whether long-lasting pollution impact caused selection of resistant birch genotypes, and to investigate possible trade-offs between pollution resistance and performance by exploring growth of birch seedlings in the different environments. In the reciprocal planting experiments, progenies of birches from polluted sites performed slightly better in the polluted environment (in terms of survival, leaf number, and leaf size) than progenies of birches from clean sites; and vice versa, in the clean environment progenies of birches from polluted sites performed slightly worse than progenies of birches from clean sites. However, this pattern was detected only in some of statistical tests, while the majority of comparisons showed no genotype or genotype by environment effects. Although some of the results indicate that long-lasting pollution impact may have increased pollution resistance of the affected populations by elimination of sensitive genotypes (survival selection), this conclusion should be considered tentative. The resistance did not reach the level sufficient to assure re-colonisation of polluted habitats: progenies of all trees completely died on the contaminated soils by the end of the first growth season. Therefore persistence of mountain birch populations near the strong polluters may be transient, being explained not only by higher resistance of the survivors, but also by phenotypic acclimatisation of mature trees to gradual increase in pollution.     

Diet-mediated effects of heavy metal pollution on growth and immune response in the geometrid moth Epirrita autumnata

The potential capacity of larval growth and immune response traits of the autumnal moth to adapt to heavy metal polluted environment was tested experimentally. Both the relative growth rate (RGR) and pupal weight were significantly higher in control trees than on polluted trees, indicating that metal pollution prevented the insect from achieving maximal growth on birch leaves. Larval growth rates of different broods differed significantly between metal contaminated and control birches. However, pupal weight of broods, which is considered more important for fitness than growth rate, in response to pollution did not differ. Immune response was significantly higher in moths exposed to pollution than in moths that were exposed to control environment suggesting that pollution enhances the immune defense of defoliators. Encapsulation rate tended to differ between broods indicating that the immune function has potential to respond to selection.     

Consequences of elevated carbon dioxide and ozone for foliar chemical composition and dynamics in trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera).

Atmospheric chemical composition affects foliar chemical composition, which in turn influences the dynamics of both herbivory and decomposition in ecosystems. We assessed the independent and interactive effects of CO2 and O3 fumigation on foliar chemistry of quaking aspen (Populus tremuloides) and paper birch (Betula papyrifera) at a Free-Air CO2 Enrichment (FACE) facility in northern Wisconsin. Leaf samples were collected at five time periods during a single growing season, and analyzed for nitrogen. starch and condensed tannin concentrations, nitrogen resorption efficiencies (NREs), and C:N ratios. Enriched CO2 reduced foliar nitrogen concentrations in aspen and birch; O3 only marginally reduced nitrogen concentrations. NREs were unaffected by pollution treatment in aspen, declined with 03 exposure in birch, and this decline was ameliorated by enriched CO2. C:N ratios of abscised leaves increased in response to enriched CO2 in both tree species. O3 did not significantly alter C:N ratios in aspen, although values tended to be higher in + CO2 + O3 leaves. For birch, O3 decreased C:N ratios under ambient CO2 and increased C:N ratios under elevated CO2. Thus, under the combined pollutants, the C:N ratios of both aspen and birch leaves were elevated above the averaged responses to the individual and independent trace gas treatments. Starch concentrations were largely unresponsive to CO2 and O3 treatments in aspen. but increased in response to elevated CO2 in birch. Levels of condensed tannins were negligibly affected by CO2 and O3 treatments in aspen, but increased in response to enriched CO2 in birch. Results from this work suggest that changes in foliar chemical composition elicited by enriched CO2 are likely to impact herbivory and decomposition, whereas the effects of O3 are likely to be minor, except in cases where they influence plant response to CO2.     

Budburst phenology of white birch in industrially polluted areas

Effects of environmental contamination on plant seasonal development have only rarely been properly documented. Monitoring of leaf growth in mountain birch, Betula pubescens subsp. czerepanovii, around a nickel-copper smelter at Monchegorsk hinted advanced budburst phenology in most polluted sites. However, under laboratory conditions budburst of birch twigs cut in late winter from trees naturally growing around three point polluters (nickel-copper smelter at Monchegorsk, aluminium factory at Kandalaksha, and iron pellet plant at Kostomuksha) showed no relationship with distance from the emission source. In a greenhouse experiment, budburst phenology of mountain birch seedlings grown in unpolluted soil did not depend on seedling origin (from heavily polluted vs. clean sites), whereas seedlings in metal-contaminated soil demonstrated delayed budburst. These results allow to attribute advanced budburst phenology of white birch in severely polluted sites to modified microclimate, rather than to pollution impact on plant physiology or genetics.     

Exposure to moderate concentrations of tropospheric ozone impairs tree stomatal response to carbon dioxide

With rising concentrations of both atmospheric carbon dioxide (CO₂) and tropospheric ozone (O₃), it is important to better understand the interacting effects of these two trace gases on plant physiology affecting land-atmosphere gas exchange. We investigated the effect of growth under elevated CO₂ and O₃, singly and in combination, on the primary short-term stomatal response to CO₂ concentration in paper birch at the Aspen FACE experiment. Leaves from trees grown in elevated CO₂ and/or O₃ exhibited weaker short-term responses of stomatal conductance to both an increase and a decrease in CO₂ concentration from current ambient level. The impairement of the stomatal CO₂ response by O₃ most likely developed progressively over the growing season as assessed by sap flux measurements. Our results suggest that expectations of plant water-savings and reduced stomatal air pollution uptake under rising atmospheric CO₂ may not hold for northern hardwood forests under concurrently rising tropospheric O₃.     

Slow growth of Empetrum nigrum in industrial barrens: combined effect of pollution and age of extant plants

We studied the impact of industrial pollution on population demography (age structure), growth and reproduction of crowberry, Empetrum nigrum L. Crowberry growing in severely polluted sites near non-ferrous smelters (at Harjavalta, Monchegorsk and Nikel) was on average twice as old as in unpolluted habitats, as indicated by the number of annual rings at root collar. Shoot length decreased both with plant ageing and due to pollution impact, while neither the proportion of generative plants nor berry production was affected by pollution or plant age. Our results suggest that death of the extant individuals of E. nigrum near the non-ferrous smelters is to a large extent explained by age-related damage of the main stem accelerated by pollution. Since vegetative propagation, seed germination and seedling establishment are hampered by soil toxicity, E. nigrum populations near the smelters continue to decline with ageing in spite of the gradual decline of emissions.     

Density patterns of gall mites (Acarina:Eriophyidae) in a polluted area

Trends in the densities of six species of gall mites on European aspen (Populus tremula) and on two birch species (Betula pubescens and B. pendula) were compared in an air pollution gradient from the Harjavalta copper-nickel smelter, SW Finland. The densities of gall mites on both birch species decreased towards the smelter and were negatively correlated with the levels of copper and nickel in the birch leaves. In contrast, the densities of aspen mites correlated neither with distance from the pollution source nor with the content of heavy metals in aspen leaves. Both birch and aspen trees tended to produce smaller leaves near the smelter, but a significant correlation between gall mite densities and site-specific leaf areas was only found for one species of mite.     

Effects of rural roadside levels of nitrogen dioxide on Polytrichum formosum Hedw

Polytrichum formosum Hedw. was exposed to 60 nl litre(-1) (122.4 microm(-3)) NO2 for 37 weeks in a closed chamber fumigation system. This concentration was chosen to simulate roadside levels in rural areas. Over an initial winter period (October-January) growth of existing shoots was stimulated by NO2. When new growth was recorded in April and May, NO2 pollution over winter and spring had resulted in a 36% reduction in new (< 1 cm) shoot production, and a 46% reduction in old shoots showing new growth. It is concluded that plants of Polytrichum formosum Hedw. growing near to roads may be adversely affected by NO2 pollution. Adverse effects of NO2 on plants and possible synergistic effects with other pollutants could cause growth reductions in sensitive species, thus affecting species composition of roadside vegetation.     

Growth responses of birch and Sitka spruce exposed to acidified rain

Seedlings of birch and Sitka spruce were grown on a range of British soils for 2 years and exposed to simulated acid rainfall treatments of pHs 5.6, 4.5, 3.5 and 2.5. Both species developed visible leaf injury patterns when exposed to the pH 2.5 treatment. In Sitka spruce this leaf injury was followed by high needle loss during the first winter and greater mortality. Generally, height growth of Sitka spruce was unaffected by treatments, but acid rainfall at pH 2.5 increased the height of birch. Mean height of both species was strongly affected by soil type. Significant soils x treatment effects on the heights of both species indicated that on some soils plant growth responses to the treatments did not fit the general pattern. Hence, while the results indicate that generally ambient acidities of rainfall in the UK are unlikely to adversely affect the growth of birch or Sitka spruce, plants growing on some soils may be susceptible to injury.     

Rape (Brassica chinensis L.) seed germination, seedling growth, and physiology in soil polluted with di-n-butyl phthalate and bis(2-ethylhexyl) phthalate

Phthalic acid esters (PAEs) pollution in agricultural soils caused by widely employed plastic products is becoming more and more widespread in China. PAEs polluted soil can lead to phytotoxicity in higher plants and potential health risks to human being. We evaluated the individual toxicity of di-n-butyl phthalate (DnBP) and bis(2-ethylhexyl) phthalate (DEHP), two representative PAEs, to sown rape (Brassica chinensis L.) seeds within 72 h (as germination stage) and seedlings after germination for 14 days by monitoring responses and trends of different biological parameters. No significant effects of six concentrations of PAE ranging from 0 (not treated/NT) to 500 mg?kg^?1 on germination rate in soil were observed. However, root length, shoot length, and biomass (fresh weight) were inhibited by both pollutants (except root length and biomass under DEHP). Stimulatory effects of both target pollutants on malondialdehyde (MDA) content, superoxide dismutase (SODase) activity, ascorbate peroxidase (APXase) content, and polyphenoloxidase (PPOase) activity in shoots and roots (SODase activity in shoots excluded) were in the same trend with the promotion of proline (Pro) but differed with acetylcholinesterase activity (except in shoots under DnBP) for analyzed samples treated for 72 h and 14 days. Responses of representative storage compounds free amino acids (FAA) and total soluble sugar (TSS) under both PAEs were raised. Sensitivity of APXase and Pro in roots demonstrates their possibility in estimation of PAE phytotoxicity and the higher toxicity of DnBP, which has also been approved by the morphological photos of seedlings at day 14. Higher sensitivity of the roots was also observed. The recommended soil allowable concentration is 5 mg DnBP?kg^?1 soil for the development of rape. We still need to know the phytotoxicity of DEHP at whole seedling stage for both the growing and development; on the other hand, soil criteria for PAE compounds are urgently required in China.     

Effects of acid rain on growth and nutrient concentrations in Scots pine and Norway spruce seedlings grown in a nutrient-rich soil

The effects of artificially applied acid precipitation on growth and nutrient concentrations of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies [L.] Karst.) seedlings were investigated in a long-term acid irrigation experiment in field conditions. Seedlings of northern and southern origin were planted in boxes containing peat and composted soil rich in nutrients, and sprinkler irrigated with water acidified with nitric and sulphuric acids to pH 3 or pH 4 for periods varying from two to three and a half growing seasons during 1986-1989. Water irrigated (pH 5.4-7.6) and non-irrigated groups of seedlings were also included in the experiment. At the end of the experiment needles, main and lateral shoots and roots were collected from the seedlings for the determination of height growth and biomass partitioning, and for the analysis of S, N, Mg, P, K, Ca, Mn and Fe concentrations. The treatment effects compared to the irrigated control were studied using multivariate analyses of variance and covariance. In the pine seedlings the total dry matter production increased by 25-70% compared with the irrigated controls when the total wet deposition to the seedlings exceeded 67 kg S ha(-1) and 36 kg N ha(-1) (e.g. after two growing seasons' exposure of the pH 3 treatment). The increase was mainly due to an increase in needle dry weight (54-72% greater at pH 3) and root weight (20-65% greater at pH 3), whereas the height growth or shoot weight growth were less affected. The northern provenance pine seedlings responded more clearly to the pH 3 irrigation than the southern ones. The treatments had no consistent effects on any of the growth variables studied in the spruce seedlings, however. The pines had higher root and foliage Ca concentrations as a result of the acid irrigation, whereas in spruce, acid rain decreased the Ca concentration in needles and shoots. Root Mn and Fe concentrations were higher in both species as a result of the pH 3 treatment. A higher soil conductivity and Ca concentration resulted from the prolonged pH 3 treatment. The results strongly support the hypothesis that the long-term growth and nutrient allocation response of conifers to acid precipitation is dependent both on the tree species and on the nutritional status of the soil.     

The effect of nitrogen additions on oak foliage and herbivore communities at sites with high and low atmospheric pollution

To evaluate plant and herbivore responses to nitrogen we conducted a fertilization study at a low and high pollution site in the mixed conifer forests surrounding Los Angeles, California. Contrary to expectations, discriminant function analysis of oak herbivore communities showed significant response to N fertilization when atmospheric deposition was high, but not when atmospheric deposition was low. We hypothesize that longer-term fertilization treatments are needed at the low pollution site before foliar N nutrition increases sufficiently to affect herbivore communities. At the high pollution site, fertilization was also associated with increased catkin production and higher densities of a byturid beetle that feeds on the catkins of oak. Leaf nitrogen and nitrate were significantly higher at the high pollution site compared to the low pollution site. Foliar nitrate concentrations were positively correlated with abundance of sucking insects, leafrollers and plutellids in all three years of the study.     

Nitrilotriacetate- and citric acid-assisted phytoextraction of cadmium by Indian mustard (Brassica juncea (L.) Czernj, Brassicaceae)

In a pot experiment the effects of nitrilotriacetate (NTA) and citric acid applications on Cd extractibility from soil as well as on its uptake and accumulation by Indian mustard (Brassica juncea) were investigated. Plants were grown in a sandy soil with added CdS at four levels ranging from 50 to 200 mg Cd kg(-1) soil. After 30 days of growth, pots were amended with NTA or citric acid at 10 and 20 mmol kg(-1). Control pots were not treated with chelates. Harvest of plants was performed immediately before and one week after chelate addition. Soil water-, NH(4)NO(3)- and EDTA-extractable Cd fractions increased constantly with both increasing soil metal application and chelate concentration. Shoot dry weights did not suffer significant reductions with increasing Cd addition to the soil except for both NTA treatments in which at 200 mg Cd kg(-1) a 30% decrease in dry matter was observed. Generally, following NTA and citric acid amendments, Cd concentration in shoots increased with soil Cd level. However, due to Cd toxicity, at the highest metal application rate both NTA treatments lowered Cd concentration in the above-ground parts. Compared to the control, at 10 mmol kg(-1) citric acid did not change Cd concentration in shoots, whereas NTA-treated plants showed an about 2-fold increase. The addition of chelates at 20 mmol kg(-1) further enhanced Cd concentration in shoots up to 718 and 560 microg g(-1) dry weight in the NTA and citrate treatments, respectively.     

Variations in chemical composition of birch foliage under air pollution stress and their consequences for Eriocrania miners

The relationships between chemical composition of birch foliage, individual performance and population density of both solitary and gregarious species of Eriocrania leafminers (Lepidoptera: Eriocraniidae) were studied in the vicinity of the Harjavalta copper-nickel smelter, southwestern Finland. The contents of heavy metals (Cu, Ni, Fe and Zn) in birch foliage exponentially decreased with the distance from the factory whereas the content of manganese showed the opposite trend. Performance parameters of Eriocrania miners were correlated neither with the distance from the pollution source nor with the foliar content of heavy metals. However, larval masses of both solitary and gregarious species were highest at the zone of moderate pollution. The levels of total nitrogen and total carbon and the carbon: nitrogen ratio in birch leaves showed no clear patterns in relation to distance from the factory. Feeding efficiency of solitary larvae increased with higher foliar nitrogen content but decreased when the carbon: nitrogen ratio was high. Survival of solitary Eriocrania species was negatively correlated with total leaf carbon content. However, the population density of solitary miners showed a negative correlation with total foliar nitrogen and a positive correlation with the carbon:nitrogen ratio. Performance of the gregarious E. haworthi was not correlated with the chemical composition of birch leaves, whereas population density was highest close to the factory complex and correlated positively with the levels of copper and nickel in birch leaves. Consequently, pollution-induced changes in measured host plant chemicals were unlikely to affect population densities of Eriocrania miners via altered larval performance.     

Sources of variation in concentrations of nickel and copper in mountain birch foliage near a nickel-copper smelter at Monchegorsk, north-western Russia: results of long-term monitoring

Concentrations of nickel and copper, two principal metal pollutants of the 'Severonikel' smelter at Monchegorsk, NW Russia, were measured in unwashed leaves of mountain birch, Betula pubescens subsp. czerepanovii, collected in eight study sites along the pollution gradient during 1991-2003. In spite of significant decline in metal emissions, concentrations of foliar metals in most of the study sites did not decrease, indicating that soil contamination remains extremely high. Multiyear mean values peaked at 6.6 km S of the smelter, where they were 20-25 times higher than in the most distant study site. Concentrations of both metals demonstrated pronounced annual variation, which was explained by the meteorological conditions of early summer: higher precipitation in May increased foliar concentrations of both metals, whereas higher precipitation in June resulted in lower foliar concentrations of nickel. These data suggest that ecotoxicological situation in metal-contaminated areas can be modified by the expected climate change. In heavily polluted sites individual birch trees generally retained their ranks in terms of metal contamination during 1995-2003, demonstrating that the use of the same set of trees can significantly increase the accuracy of the monitoring data.     

Indication of airborne pollution by birch and spruce in the vicinity of copper smelter

The aim of the study was determination of air pollution impact of the copper smelter in Bor and its surroundings (Serbia) by assessing the suitability of birch (Betula pendula Roth.) and spruce (Picea abies L.) for the purposes of biomonitoring and comparing it with previously published data from the same study area. The concentrations of Cu, Zn, Pb and Mn in leaves/needles, branches, roots and soil were determined. Sampling was performed during 2009 in two zones with high load of air pollution due to copper mining and smelting activities, and one background zone. Metal accumulation and translocation was evaluated in terms of biological factors. In addition, plant enrichment factor was calculated. According to the results, plant foliage was not enriched through soil, which indicates absorption from the air, with both species acting as excluders of Cu, Pb, Zn and Mn. Leaves were more enriched with all the metals than needles, indicating a better response of birch to airborne pollution than spruce. Cluster analysis showed different level of pollution at the sites, while correlations between Cu and Pb obtained by Principal Component Analysis indicated their anthropogenic origin. Regarding previously published results, beside birch leaves, pine needles (which showed higher level of response to pollution compared to linden leaves) could be applied in air biomonitoring surveys near copper smelters.     

The sensitivity of early 20th century cultivars of wheat to air pollution

The effects of a mixture of SO(2) and NO(2) at a concentration of 0.08 to 0.10 ppm (in microg m(-3): 164-205 for NO(2); 229-286 for SO(2)) were tested for four weeks on two old varieties of winter wheat, Little Joss and Holdfast, introduced in 1908 and 1938, respectively, grown in simulated autumn conditions. After two weeks, root dry weights of both varieties were significantly decreased but, although there was some leaf necrosis, shoot dry weights were unaltered. After a further two weeks, the dry weights of the shoots as well as the roots were significantly decreased. These effects, amounting to a combined 40% decrease in total dry weight, were greater than those in identical experiments carried out with the present day variety Avalon, in which the decrease was 20%. The results suggest that the modern cultivar Avalon is more tolerant of SO(2) and NO(2) than Little Joss and Holdfast, which were grown many years ago.     

Effect of birch (Betula spp.) and associated rhizoidal bacteria on the degradation of soil polyaromatic hydrocarbons, PAH-induced changes in birch proteome and bacterial community

Two birch clones originating from metal-contaminated sites were exposed for 3 months to soils (sand-peat ratio 1:1 or 4:1) spiked with a mixture of polyaromatic hydrocarbons (PAHs; anthracene, fluoranthene, phenanthrene, pyrene). PAH degradation differed between the two birch clones and also by the soil type. The statistically most significant elimination (p ≤ 0.01), i.e. 88% of total PAHs, was observed in the more sandy soil planted with birch, the clearest positive effect being found with Betula pubescens clone on phenanthrene. PAHs and soil composition had rather small effects on birch protein complement. Three proteins with clonal differences were identified: ferritin-like protein, auxin-induced protein and peroxidase. Differences in planted and non-planted soils were detected in bacterial communities by 16S rRNA T-RFLP, and the overall bacterial community structures were diverse. Even though both represent complex systems, trees and rhizoidal microbes in combination can provide interesting possibilities for bioremediation of PAH-polluted soils.     

Cadmium accumulation and its effect on the in vitro growth of woody fleabane and mycorrhized white birch

The effect of Cd on woody fleabane (Dittrichia viscosa (L.) Greuter) and white birch (Betula celtiberica Rothm. & Vasc.) was examined. Woody fleabane and white birch were grown in vitro in Murashige, T., Skoog, F., [1962. A revised medium for rapid growth and bioassays with tobacco tissue cultures. Physiol. Plant. 15, 473-479] (MS) plus Cd (10 mg Cd kg(-1)) and except for root length in white birch, plant development was inhibited when Cd was added. Cd accumulation in above-ground tissues showed differences among clones, reaching 1300 and 463 mg Cd kg(-1) dry wt. in selected clones of woody fleabane and white birch, respectively. Tolerance of Paxillus filamentosus (Scop) Fr. to Cd was also examined before mycorrhization. Plants of mycorrhized white birch grown in the presence of Cd had a better development and accumulated more Cd in their shoots than the non-mycorrhized ones. The use of selected clones of woody fleabane and the mycorrhization of white birch enhance extraction efficiency from contaminated soils in phytoremediation programs.     

Accumulation of Cd,Cu and Zn in shoots of maize (<Emphasis Type="Italic">Zea mays</Emphasis> L.) exposed to 0.8 or 20 nM Cd during vegetative growth and the relation with xylem sap composition

This work focuses on the exposure of maize plants to nanomolar concentrations of Cd, which is relevant for agricultural soils cropped with food and feed plants. Maize plants were cultivated in nutrient solution at 0.8 or 20 nM Cd during the vegetative growth stages. No significant hormesis or toxic effects of Cd were observed on maize growth, but a decrease in the allocation of Cd to shoots between the 0.8 and 20 nM Cd exposures revealed that the plants already responded to these low concentrations of Cd according to a shoot Cd excluder strategy. The Cd, Cu and Zn concentrations in shoots decreased with time as the result of an early decrease in the root/shoot ratio and of a decrease in the coefficient of allocation to aboveground for Zn and Cd at 20 nM. As a consequence, shoots of young plants were richer in micronutrients Cu and Zn but also in toxic Cd. The rate of delivery of Cd, Cu and Zn from xylem sap was successfully used to predict the time course of concentrations of Cd, Cu and Zn in the shoot. However, it overestimated the actual concentrations of Cd in the shoot, presumably because the reallocation of this trace element from shoots back to roots was not taken into account.