Impact of artificial, ammonium-enriched rainwater on soils and young coniferous trees in a greenhouse. Part II--Effects on the trees

To obtain an insight into the effects of a high ammonium deposition on trees young, coniferous trees were planted in pots in a greenhouse and treated with artificial ammonium-enriched rainwater. Application of 480 kg ammonium N ha(-1) year(-1) resulted in an increase of the shoot/root ratio. The biomass of fine roots strongly declined, as did the numbers of mycorrhizae. The fructification of the mycorrhizal fungi was totally inhibited. The nitrogen content of the needles was enhanced, but the levels of potassium, magnesium and calcium decreased sharply. The phosphorus content remained almost unaffected. Much of the nitrogen was stored as arginine. The levels of leaf pigments also increased. Within one year of treatment many of the trees died. The trees that were treated with 48 kg ammonium-N ha(-1) year(-1) did not show any signs of deterioration.     

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.     

Stress responses of Calluna vulgaris to reduced and oxidised N applied under 'real world conditions'

Effects and implications of reduced and oxidised N, applied under 'real world' conditions, since May 2002, are reported for Calluna growing on an ombrotrophic bog. Ammonia has been released from a 10 m line source generating monthly concentrations of 180-6 microg m(-3), while ammonium chloride and sodium nitrate are applied in rainwater at nitrate and ammonium concentrations below 4mM and providing up to 56 kg N ha(-1) year(-1) above a background deposition of 10 kg N ha(-1) year(-1). Ammonia concentrations, >8 microg m(-3) have significantly enhanced foliar N concentrations, increased sensitivity to drought, frost and winter desiccation, spring frost damage and increased the incidence of pathogen outbreaks. The mature Calluna bushes nearest the NH3 source have turned bleached and moribund. By comparison the Calluna receiving reduced and oxidised N in rain has shown no significant visible or stress related effects with no significant increase in N status.     

Effects of simultaneous ozone exposure and nitrogen loads on carbohydrate concentrations, biomass, growth, and nutrient concentrations of young beech trees (Fagus sylvatica)

Beech seedlings were grown under different nitrogen fertilisation regimes (0, 20, 40, and 80 kg Nha(-1)yr(-1)) for three years and were fumigated with either charcoal-filtered (F) or ambient air (O3). Nitrogen fertilisation increased leaf necroses, aphid infestations, and nutrient ratios in the leaves (N:P and N:K), as a result of decreased phosphorus and potassium concentrations. For plant growth, biomass accumulation, and starch concentrations, a positive nitrogen effect was found, but only for fertilisations of up to 40 kg Nha(-1) yr(-1). The highest nitrogen load, however, reduced leaf area, leaf water content, growth, biomass accumulation, and starch concentrations, whereas soluble carbohydrate concentrations were enhanced. The ozone fumigation resulted in reduced leaf area, leaf water content, shoot growth, root biomass accumulation, and decreased starch, phosphorus, and potassium concentrations, increasing the N:P and N:K ratios. A combined effect of the two pollutants was detected for the leaf area and the shoot elongation, where ozone fumigation amplified the nitrogen effects.     

Arbuscular mycorrhizal fungi can decrease the uptake of uranium by subterranean clover grown at high levels of uranium in soil

Subterranean clover inoculated or not with the arbuscular mycorrhizal (AM) fungus Glomus intraradices was grown on soil containing six levels of 238U in the range 0-87 mg kg(-1). Increasing U concentration in soil enhanced the U concentration in roots and shoots of both mycorrhizal and nonmycorrhizal plants but had no significant effects on plant dry matter production or root AM colonization. Mycorrhizas increased the shoot dry matter and P concentration in roots and shoots, while in most cases, it decreased the Ca, Mg and K concentrations in plants. The AM fungus influenced U concentration in plants only in the treatment receiving 87 mg U kg(-1) soil. In this case, U concentration in shoots of nonmycorrhizal plants was 1.7 times that of shoots of mycorrhizal plants. These results suggested that mycorrhizal fungi can limit U accumulation by plants exposed to high levels of U in soil.     

The interactions between plant growth,vegetation structure and soil processes in semi-natural acidic and calcareous grasslands receiving long-term inputs of simulated pollutant nitrogen deposition

Regular applications of ammonium nitrate (35-140 kg N ha(-1) year(-1)) and ammonium sulphate (140 kg N ha(-1) year(-1)) to areas of acidic and calcareous grassland in the Derbyshire Peak District over a period of 6 years, have resulted in significant losses in both overall plant cover, and the abundance of individual species, associated with clear and dose-related increases in shoot nitrogen content. No overall growth response to nitrogen treatment was seen at any stage in the experiment. Phosphorus additions to the calcareous plots did however lead to significant increases in plant cover and total biomass, indicative of phosphorus limitation in this system. Clear and dose-related increases in soil nitrogen mineralization rates were also obtained, consistent with marked effects of the nitrogen additions on soil processes. High nitrification rates were seen on the calcareous plots, and this process was associated with significant acidification of the 140 kg N ha(-1) year(-1) treatments.     

Detection and localization of aluminum and heavy metals in ectomycorrhizal Norway spruce seedlings

Norway spruce seedlings colonized with Hebeloma crustuliniforme were grown in growth pouches. After formation of ectomycorrhizas, the seedlings were exposed to Al or the heavy metals Cd, Cu, Ni, or Zn at various concentrations for 5 weeks to estimate the detection limits of metals with X-ray microanalysis in the cryo-scanning electron microscope. When the lowest metal concentrations (1 mM Al(3+), 0.1 mM Cd(2+), 0.2 mM Cu(2+), 0.5 mM Ni(2+), 2 mM Zn(2+)) were applied, only Al and Zn were detected at low X-ray counts in the ectomycorrhizas. After application of 10-fold higher metal concentrations, distinct metal accumulation patterns were observed. Cd was detected predominantly in the Hartig net, Al and Ni in the Hartig net and in the cell walls of the cortex, and Zn in the Hartig net, the cortical cell walls and the fungal mantle. Cu was not detected at all. By combining X-ray microanalysis with absolute metal concentrations found in the roots, the estimated detection limits of X-ray microanalysis were: Al> or =0.86 mg g(-1), Cd> or =0.26 mg g(-1), Ni> or =1.30 mg g(-1), and Zn> or =0.54 mg g(-1), whereas Cu was not detectable even at root concentrations of 0.47 mg g(-1). Treatments with the highest metal concentrations showed high X-ray counts of metals in cells of the stele but reduced concentrations of the macronutrients K, Mg, and P in roots, indicating a possible disturbance of root and ectomycorrhizal function.     

The effects of excess nitrogen deposition on young Norway spruce trees. Part I the soil

The effects of wet-deposited nitrogen on soil acidification and the health of Norway spruce were investigated in a pot experiment using an open-air spray/drip system. Nitrogen was applied as ammonium ((NH(4))(2)SO(4)) or nitrate (HNO(3)/NaNO(3)) in simulated rain to either the soil or the foliage for a period of two years five months. Symptoms of forest decline were not reproduced. Adverse effects relating to soil acidification and N saturation were observed and depended on the chemical form of N. The plant-soil system absorbed most of the soil-applied NH(+)(4) at doses of up to 65 kgN ha(-1) year(-1) but only 54% at a dose of 125 kgN ha(-1) year(-1). About 60% of soil-applied NO(-)(3) was absorbed in all treatments. Ammonium treatments acidified the soil, increased base cation leaching, and mobilised acidic cations. Nitrification was not the major source of acidity, however. Nitrate inputs increased soil pH. Critical loads calculated using current criteria were 60-120 and 30-60 kgN ha(-1) year(-1) for ammonium and nitrate, respectively. Ammonium is more likely to damage forest ecosystems, however, illustrating the need for care in the definition of critical loads.     

Scots pine responses to CO2 enrichment--I. Ectomycorrhizal fungi and soil fauna

Ectomycorrhizal Scots pine seedlings were grown in unfertilized forest soil at ambient and double (ca 700 ppm) atmospheric concentrations of CO2. The biomass of seedlings and fungal biomass both in the roots and in the soil and the numbers of certain groups of soil animals were measured under summer conditions and after an artificial winter acclimation period. No biomass parameter showed any significant change due to CO2 elevation. Increases were found during the winter acclimation period in total and fine root biomasses, fungal biomass in the soil and total fungal biomass both in the roots and in the soil, while the ratio of needle biomass: fungal biomass and the shoot: root ratio decreased. The N concentration in previous-year needles was lower in the double CO2 environment than with ambient CO2. Enchytraeids almost disappeared in the double CO2 environment during winter acclimation, while the numbers of nematodes increased at the same time in both treatments.     

The arbuscular mycorrhizal fungus Glomus mosseae induces growth and metal accumulation changes in Cannabis sativa L

The effect of arbuscular mycorrhiza on heavy metal uptake and translocation was investigated in Cannabis sativa. Hemp was grown in the presence and absence of 100 microg g-1 Cd and Ni and 300 microg g-1 Cr(VI), and inoculated or not with the arbuscular mycorrhizal fungus Glomus mosseae. In our experimental condition, hemp growth was reduced in inoculated plants and the reduction was related to the degree of mycorrhization. The percentage of mycorrhizal colonisation was 42% and 9% in plants grown in non-contaminated and contaminated soil, suggesting a significant negative effect of high metal concentrations on plant infection by G. mosseae. Soil pH, metal bioavailability and plant metal uptake were not influenced by mycorrhization. The organ metal concentrations were not statistically different between inoculated and non-inoculated plants, apart from Ni which concentration was significantly higher in stem and leaf of inoculated plants grown in contaminated soil. The distribution of absorbed metals inside plant was related to the soil heavy metal concentrations: in plant grown in non-contaminated soil the greater part of absorbed Cr and Ni was found in shoots and no significant difference was determined between inoculated and non-inoculated plants. On the contrary, plants grown in artificially contaminated soil accumulated most metal in root organ. In this soil, mycorrhization significantly enhanced the translocation of all the three metals from root to shoot. The possibility to increase metal accumulation in shoot is very interesting for phytoextraction purpose, since most high producing biomass plants, such as non-mycorrhized hemp, retain most heavy metals in roots, limiting their application.     

Biomass and nutrient concentration of sweet corn roots and shoots under organic amendments application

Two field experiments were conducted at the Waimanalo research station on the island of O'ahu, Hawaii to study the effect of chicken (CM) and dairy (DM) manures on biomass and nutrient concentration in sweet corn roots and shoots. Sweet corn (super sweet 10, Zea Mays L. subsp. mays) was grown for two consecutive growing seasons under four rates of application (0, 168, 337, and 672 kg ha^{? 1} total N equivalent) and one time (OTA) or two time (TTA) applications of organic manure types and rates. There were significant effects of types, rates, and number of manure applications on dry biomass and macro- and micro-nutrient concentration in roots and shoots tissues. Results of root tissue indicated a significant accumulation of N and C under CM and DM treatments compared with the control treatment. Manure application rates significantly increased the accumulation of N and C in root tissue. Dry weight of roots and shoots and both macro- and micro-nutrient contents in the plant tissues significantly increased under TTA treatment compared with OTA treatment. There was a significant correlation (r² = 0.46 to 0.81) between root biomass, macro-, and micro-nutrient contents during both growing seasons. The results of the study indicates that amending soils with CM at the highest application rate provided the best crop performance in terms of root and shoot biomass, crop N, C, and other macro- and micro-nutrients.     

Clonal variation in heavy metal accumulation and biomass production in a poplar coppice culture. II. Vertical distribution and phytoextraction potential

Short rotation coppice cultures (SRC) are intensively managed, high-density plantations of multi-shoot trees. In April 1996, an SRC field trial with 17 different poplar clones was established in Boom (Belgium) on a former waste disposal site. In December 1996 and January 2001, all shoots were cut back to a height of 5 cm to create a coppice culture. For six clones, wood and bark were sampled at the bottom, middle and top of a shoot in August and November 2002. No significant height effect of metal concentration was found, but for wood, metal concentrations generally increased toward the top of the shoot in August, and decreased toward the top of the shoot in November. Phytoextraction potential of a clone was primarily determined by metal concentration and by biomass production. Shoot size and number of shoots per stool were less important, as a high biomass production could be achieved by producing a few large shoots or many smaller shoots. Clone Fritzi Pauley accumulated 1.4 kg ha(-1) of Al over two years; Wolterson and Balsam Spire showed a relatively high accumulation of Cd and Zn, i.e. averaging, respectively 47 and 57 g ha(-1) for Cd and 2.4 and 2.0 kg ha(-1) for Zn over two years.     

Mercury toxicity induces oxidative stress in growing cucumber seedlings

In this study, the effects of exogenous mercury (HgCl(2)) on time-dependent changes in the activities of antioxidant enzymes (catalase and ascorbate peroxidase), lipid peroxidation, chlorophyll content and protein oxidation in cucumber seedlings (Cucumis sativus L.) were investigated. Cucumber seedlings were exposed to from 0 to 500microM of HgCl(2) during 10 and 15 days. Hg was readily absorbed by growing seedlings, and its content was greater in the roots than the in shoot. Time and concentration-dependent reduction in root and shoot length was observed at all concentrations tested, equally in the roots and shoot, at both 10 and 15 days. At 50microM HgCl(2), root fresh weight of 15-day-old seedlings increased, and at other concentrations, it reduced. For 10-day-old seedlings, reduction in root and shoot fresh biomass was observed. At 15 days, only at 50microM HgCl(2) was there no observed reduction in shoot fresh biomass. Dry weight of roots increased at 500microM both at 10 and 15 days, though at 250microM HgCl(2) there was only an increase at 15 days. There was a significant effect on shoot dry weight at all concentrations tested. Hg-treated seedlings showed elevated levels of lipid peroxides with a concomitant increase in protein oxidation levels, and decreased chlorophyll content when exposed to between 250 and 500microM of HgCl(2). At 10 days, catalase activity increased in seedlings at a moderately toxic level of Hg, whereas at the higher concentration (500microM), there was a marked inhibition. Taken together, our results suggest that Hg induces oxidative stress in cucumber, resulting in plant injury.     

Genotypic Differences in Effects of Cadmium Exposure on Plant Growth and Contents of Cadmium and Elements in 14 Cultivars of Bai Cai

Abstract

Fourteen cultivars of bai cai (Brassica campestris L. ssp. chinensis var. communis) were grown in the nutrient solutions containing 0–0.5 μg mL^?1 of cadmium (Cd) to investigate genotypic differences in the effects of Cd exposure on the plant growth and uptake and distribution of Cd in bai cai plants. The Cd exposure significantly reduced the dry and fresh weights of roots and shoots, the dry weight ratio of shoot/root (S/R), total biomass, and chlorophyll content (SPAD value). Cd concentrations in bai cai ranged from 13.3 to 74.9 μg g^?1 DW in shoots and from 163.1 to 574.7 μg g^?1 DW in roots under Cd exposure, respectively. The considerable genotypic differences of Cd concentrations and accumulations in both shoots and roots were observed among 14 bai cai cultivars. Moreover, Cd mainly accumulated in the roots. Cd also caused the changes of uptake and distribution of nutrients in bai cai and under the influence of cadmium, the concentration of potassium (K) decreased in shoot and increased in root. However, the concentrations of magnesium (Mg), phosphorus (P), manganese (Mn), boron (B), and iron (Fe) increased in shoots and decreased in roots. In addition, Cd exposure resulted in an increase in calcium (Ca), sulphur (S), and zinc (Zn) concentrations in both shoots and roots but had no significant effects on the whole uptake of the examined mineral nutrients except for S.     

Possible evidence for transport of an iron cyanide complex by plants

Barley (Hordeum vulgare L.), oat (Avena sativa L.), and wild cane (Sorghum bicolor L.), were exposed to 15N-labeled ferrocyanide to determine whether these plant species can transport this iron cyanide complex. Plants were treated with ferrocyanide in a nutrient solution that simulated iron cyanide contaminated groundwater and soil solutions. This nutrient solution has been shown to maintain ferrocyanide speciation with minimal dissociation to free cyanide. Following treatment, all three plants showed dramatic enrichments in roots (delta 15N per thousand =1000-1500) and shoots (delta 15N per thousand =500). Barley and oat showed enrichment primarily in roots while wild cane showed a near equal enrichment in root and shoot tissues. Nitrogen-deficient barley plants treated with ferrocyanide showed a significantly greater 15N enrichment as compared to nitrogen-sufficient plants. While the results are suggestive of ferrocyanide transport by these plant species, additional study will be required to verify these results.     

Effect of ectomycorrhizae and ammonium on 134Cs and 85Sr uptake into Picea abies seedlings

Microorganisms play an important role in the fixation of radionuclides in forest soils. In particular, fungi have the capacity to absorb and translocate radionuclides. The role of the ectomycorrhizal fungus Hebeloma crustuliniforme in the uptake of radiocaesium (134Cs) and radiostrontium (85Sr) into seedlings of Norway spruce (Picea abies) was investigated in a pouch test system. Inoculated and non-inoculated seedlings; seedlings inoculated during 8 and 15 weeks; seedlings exposed during 2 and 3 weeks to the radioactive solution; and seedlings grown under low and high ammonium conditions prior to the application of the radionuclides were compared. The final 134Cs and 85Sr activity was determined in fine-roots, main-roots, stems and needles. The results showed that ectomycorrhizae reduced the uptake of 134Cs and 85Sr. The degree of ectomycorrhization was of crucial importance and seemed to be governed by the period during which ectomycorrhizae were allowed to develop and by the ammonium concentration in the nutrient solution. The radionuclide uptake increased with increasing exposure time. Both radionuclides were predominantly accumulated in fine-roots. However, needles proved to describe best the result of net root uptake and translocation to the shoot. The uptake-and and translocation-rates of 85Sr were smaller than those of 134Cs. It is assumed that the translocation is coupled with the intensity of water fluxes through the xylem and that 85Sr is more readily adsorbed into mycelium or plant tissue relative to 134Cs. The effect of high ammonium growth conditions was overcome by the effect of ectomycorrhization, except in needles with a very large biomass which behaved as a strong sink and led to a high accumulation of 134Cs.     

Empirical and simulated critical loads for nitrogen deposition in California mixed conifer forests

Empirical critical loads (CL) for N deposition were determined from changes in epiphytic lichen communities, elevated NO(3)(-) leaching in streamwater, and reduced fine root biomass in ponderosa pine (Pinus ponderosa Dougl. ex Laws.) at sites with varying N deposition. The CL for lichen community impacts of 3.1kg ha(-1) year(-1) is expected to protect all components of the forest ecosystem from the adverse effects of N deposition. Much of the western Sierra Nevada is above the lichen-based CL, showing significant changes in lichen indicator groups. The empirical N deposition threshold and that simulated by the DayCent model for enhanced NO(3)(-)leaching were 17kg N ha(-1) year(-1). DayCent estimated that elevated NO(3)(-) leaching in the San Bernardino Mountains began in the late 1950s. Critical values for litter C:N (34.1), ponderosa pine foliar N (1.1%), and N concentrations (1.0%) in the lichen Letharia vulpina ((L.) Hue) are indicative of CL exceedance.     

Urban polluted forest soils induce elevated root peroxidase activity in Scots pine (Pinus sylvestris L.) seedlings.

Plant biomass. mycorrhizal status and root peroxidase activity were measured in ectomycorrhizal Scots pine (Pinus sylvestris L.) seedlings grown in urban polluted and native, non-polluted forest soils with added ammonium or potassium sulphates simulating N and S deposition of urban areas. Peroxidase activity in the fine roots of seedlings planted in polluted forest soils was higher than in those planted in non-polluted soils and correlated positively with the activities measured in an earlier study in the roots of mature Scots pines growing at the sites from where the soils were collected. Growth of seedlings and mycorrhizal status were not affected by the origin of soil. Exposing the seedlings to winter acclimation conditions for 6 weeks elevated peroxidase activity in the roots. The addition of ammonium or potassium sulphate to non-polluted soils did not induce elevated root peroxidase activity, although at the levels of 0.5 and 1.0 g of ammonium sulphate a slight increasing trend was observed. We suggest, that indirect biotic factors, i.e. changes in the community structure of soil fungi, early stages of recognition, and defence reactions of pine roots against saprophytic and pathogenic fungi may be participating in the elicitation of peroxidase (POD) activity, although the possible role of heavy metals cannot be excluded.     

Effects of ammonium and aluminium on the development and nutrition of Pinus nigra in hydroculture

Application of ammonium and aluminium to young Pinus nigra var. maritima (Ait.) Melville trees resulted in a variety of negative effects. Excess ammonium led to an increase in shoot/root ratio. The biomass of the fine roots declined, resulting in an increase of the coarse/fine root ratio. The degree of mycorrhizal infection of the roots decreased. The nitrogen content of the trees increased considerably, whereas particularly the levels of calcium magnesium, manganese and zinc decreased sharply. Excess aluminium resulted in a simultaneous reduction of root and shoot biomass, a decline of the fine root system, an increase in the coarse/fine root ratio and a decrease in the degree of mycorrhizal infection. Uptake of the divalent cations calcium, magnesium, iron, manganese and zinc was restricted substantially, The nitrogen and phosphorus contents of the trees were hardly affected, whilst the potassium content of the shoot increased and of the roots decreased. This implicates that a deteriorating fine root system has to supply water and nutrients to a more demanding shoot. In the long term, high ammonium inputs and aluminium dissolution in forest ecosystems will lead to substantial nutrient deficiencies, just as has been found in the field.     

Arsenate (As) uptake by and distribution in two cultivars of winter wheat (Triticum aestivum L.)

Two cultivars of winter wheat (Triticum aestivum L.) (Jing 411 and Lovrin 10) were used to investigate arsenate (As) uptake and distribution in plants grown in hydroponic culture and in the soil. Results showed that without As addition, Lovrin 10 had higher biomass than Jing 411 in the soil pot experiment; in the hydroponic experiment Lovrin 10 had similar root biomass to and lower shoot biomass than Jing 411. Increasing P supply from 32 to 161 microM resulted in lower tissue As concentrations, and increasing As supply from 0 to 2,000 microM resulted in lower tissue P concentrations. Increasing P supply tended to increase shoot-to-root ratios of As concentrations, and increasing As supply tended to decrease shoot-to-root ratios of As concentrations. Both cultivars invested more in root production under P deficient conditions than under P sufficient conditions. Lovrin 10 invested more biomass production to roots than Jing 411, which might be partly responsible for higher shoot P and As concentrations and higher shoot-to-root ratios of As concentrations. Moreover, Lovrin 10 allocated less As to roots than Jing 411 and the difference disappeared with decreasing P supply.