Symbiotic N2 fixation of Alnus incana ssp. rugosa in shrub wetlands of the Adirondack Mountains, New York, USA

Surface waters in forested watersheds in the Adirondack Mountains and northern New York State are susceptible to nitrogen (N) saturation. Atmospheric deposition of N to watersheds in this region has been measured but the extent of internal N inputs from symbiotic N₂ fixation in alder-dominated wetlands is not known. We estimated N₂ fixation by speckled alder in these wetlands by the ¹⁵N natural abundance method and by acetylene reduction using a flow-through system. Foliar N derived from fixation (%N_dfa) was estimated for five wetlands. The '¹⁵N of speckled alder foliage from four of the five sites did not differ significantly (PА.05) from that of nodulated speckled alders grown in N-free water culture (-1.2ǂ.1‰). Estimates from the ¹⁵N natural abundance method indicated that alders at these sites derive 85-100% of their foliar N from N₂ fixation. At one of the sites, we also measured biomass and N content and estimated that the alder foliage contained 43 kg N ha^-1 of fixed N in 1997. This estimate was based on a foliar N content of 55.4lj kg N ha^-1 (mean-SE), 86dž%N_dfa, and an assumption that 10% of foliar N was derived from reserves in woody tissues. At this site, we further estimated via acetylene reduction that 37ᆞ kg N ha^-1 was fixed by speckled alders in 1998. This estimate used the theoretical 4:1 C₂H₂ reduction to N₂ fixation ratio and assumed no night-time fixation late in the season. Nitrogen inputs in wet and dry deposition at this site are approximately 8 kg N ha^-1 year^-1. We conclude that speckled alder in wetlands of northern New York State relies heavily on N₂ fixation to meet N demands, and symbiotic N₂ fixation in speckled alders adds substantial amounts of N to alder-dominated wetlands in the Adirondack Mountains. These additions may be important for watershed N budgets, where alder-dominated wetlands occupy a large proportion of watershed area.     

Seasonal flooding regimes influence survival,nitrogen fixation,and the partitioning of nitrogen and biomass in Alnus incana ssp. rugosa

Alteration of natural flooding regimes can expose lowlands to waterlogged soil conditions during any month of the year. The seasonality of flooding may have profound effects on the carbon and nitrogen budgets of N-fixing alders (Alnus spp.), and in turn, may impact the C and N economy of extensive alder-dominated, wetland ecosystems, including those dominated by speckled alder (Alnus incana ssp. rugosa). To better understand this process, two-year-old, nodulated seedlings of speckled alder were subjected to late spring (May 10 – July 10), summer (July 10 – September 8), and fall (September 8 – November 8) flooding treatments. Alders were root-flooded outdoors in tanks containing an N-free nutrient solution and compared with unflooded alders at the experimental site. Flooding arrested N fixation, photosynthesis, and growth of alders without recovery in all flooding treatments for the remainder of the growing season. Late spring and summer flooding resulted in complete mortality of alders while all seedlings survived flooding in the fall. Fall flooding increased foliar N resorption by 140% over unflooded seedlings. Eighty-seven percent of the total N fixed and 89% of biomass accumulation for the entire growing season occurred in unflooded alders after July 10. In unflooded alders, nitrogen fixation rates per unit mass declined by 63% for nodules, 28% for leaves, and 48% for whole seedlings during the fall, while total N fixed per plant in the fall was similar to that fixed in the summer. The majority of newly fixed N in unflooded alders was allocated to leaves before September 8 and to roots/nodules combined after September 8. In unflooded plants, the greatest proportion of new biomass was partitioned to leaves before July 10, to stems between July 10 and September 8, and equally to stems and roots/nodules after September 8. Fall-flooded alders did not increase root or nodule biomass. Proportional allocation of plant resources were such that the ratio of N fixed to seedling growth of unflooded alders decreased by 19% during summer before rebounding by 6% in fall. Seasonality of flooding alters seedling survival, growth, and resource allocation, and may be a critical determinant of speckled alder recruitment and occurrence in wetlands.     

Nitrogen cycling in dense plantings of hybrid poplar and black alder

Summary Nitrogen cycling was studied during the third growing season in pure and mixed plantings (33×33 cm spacing) of hybrid poplar and black alder in southeastern Canada. After 3 years, hybrid poplar growth and N content of living tissues in a plot and of individual hybrid poplar plants increased with the proportion of black alder in a planting. No differences were detected among N contents of individual alder plants regardless of plot treatment. Black alder allocated a larger portion of its N to roots than hybrid poplar. Symbiotic nitrogen fixation was estimated to account for 80% of the nitrogen in aboveground alder tissues in the pure treatment using natural¹⁵N dilution. N return in leaf litter was estimated to be 70kg ha^–1 in the pure alder treatment and decreased to a minimum of 20 kg ha^–1 in the pure hybrid poplar plots. No difference was detected among treatments for throughfall N content. Nitrogen concentration in roots and leaf litterfall of black alder was higher than hybrid poplar. Significant soil N accretion occurred in mixed plantings containing two alders to one poplar and pure black alder plantings. Nitrogen availability (NO₃–N) increased with the amount of black alder in a plot. Results suggest that the early increase in nitrogen accumulation of hybrid poplar in mixed treatments can be attributed to an increase of total soil N availability resulting from the input of large amounts of N from easily mineralizable alder tissue.     

Autumnal changes in tissue nitrogen of autumn olive,black alder and eastern cottonwood

Two experiments were conducted to determine patterns of N change in tissues of autumn olive (Elaeagnus umbellata Thunb.) and black alder (Alnus glutinosa [L.] Gaertn.) during autumn in central Illinois, U.S.A. In the first study leaf nitrogen concentrations of autumn olive decreased 40% at an infertile minespoil site and 39% at a fertile prairie site throughout autumn whereas nitrogen concentrations in respective bark samples increased by 39% and 37%. Salt-extractable protein concentrations increased in bark and decreased in leaves over the sampling period. Free amino acid concentrations of autumn olive leaves decreased over the course of the experiment from peak concentrations in August. Asparagine, glutamic acid and proline were major constituents of the free amino acid pools in leaves. Total phosphorus concentrations of autumn olive leaves declined by 40–46% during autumn while bark concentrations of P did not significantly change.In the second experiment non-nodulated seedlings of alder receiving a low level of N-fertilization did not exhibit net resorption of leaf N during autumn whereas foliar N concentration of contrasting nonactinorhizal cottonwood plants (Populus deltoides Bartr. ex. Marsh) under the same fertilization regime decreased by 27% after the first frost. A gradual but significant decrease of 38% in foliar N concentration of nodulated alder seedlings grown under a low N-fertilization regime was associated with the cessation of nitrogenase activity during autumn in nodules. Compared with the low N fertilization regime, the higher level of N-fertilization resulted in smaller autumnal decreases of foliar N concentration in nodulated alder (17%) and in cottonwood (20%); but there was no decrease in foliar N concentration in non-nodulated alder. The higher level of N-fertilization promoted a greater accumulation of N in the roots than in the bark of both tree species after the first frost.Our results suggest that black alder lackingFrankia symbionts does not exhibit net leaf N resorption and that autumnal decreases in leaf N ofFrankia-nodulated black alder result primarily from declining foliar N import relative to export due to low temperature inhibition of N₂ fixation. In contrast, autumn olive exhibited greater and more precipitous autumnal declines in foliar N concentration than those of alder, and the pattern of N decline was unaffected by site fertility.     

Growth,nitrogen accumulation,and symbiotic dinitrogen fixation in pure and mixed plantings of hybrid poplar and black alder

Summary Growth and N accumulation were assessed in pure and mixed plantings (2 years old) of hybrid poplar and black alder in southern Québec. Symbiotic dinitrogen fixation was evaluated by natural¹⁵N dilution. Growth of hybrid poplar plants and N accumulation in their tissues increased with their decreasing contribution to species ratio whereas no differences among treatments were measured for black alder. Yield and N content per hectare of aboveground components increased with the proportion of black alder in the plantation. Symbiotic dinitrogen fixation was estimated at 68% of alder nitrogen in both pure and mixed treatments. The maximum rate of N-fixation was 53kg ha^–1 yr^–1 in pure alder plots. The amount of nitrogen accumulated in entire plants of black alder from symbiotic fixation could be sufficient to balance the N export in harvested stems and branches of short-rotation plantations containing at least 33% of alder.     

Assessing the adaptability of the actinorhizal symbiosis in the face of environmental change

Human activity, and in particular industrial activity, has altered natural environments. Here we present an experimental approach adapted to study the actinorhizal symbiosis in alder trees and shrubs submitted to abiotic stress. We measured the impact of exogenous nitrogen on the establishment of the alder symbiosis with Frankia sp., and its primary function; nitrogen fixation. Results showed our version of the growth pouch method was functional, and corroborated the gradual inhibition of symbiosis in the presence of increasing exogenous nitrogen concentrations. In mountain alder (Alnus viridis ssp. crispa) there was a gradual and suppressive effect of nitrogen on the relative number or root nodules, while in black alder (Alnus glutinosa) results suggested a threshold effect at 45 ppm N. Shoot to root biomass ratios were increased in the presence of the microsymbiont, and this effect was generally maintained even in the presence of heavy metals (As, Se or V). Alders and the actinorhizal symbiosis were not heavily affected by the presence of heavy metals, confirming potential applications in soil rehabilitation, however the distribution of metals in plant tissues sometimes changed when high levels of metals were present. A. glutinosa plants exposed to high levels of As significantly increased the allocation of As to roots (≈90%), while those exposed to high levels of Se rose their aerial tissue Se allocation to roughly 86%. A. glutinosa plants exposed to high V levels did not change behavior: V was in all cases preferentially accumulated in underground tissues (≥90%). Our results detail the use of a high-throughput approach to study the plasticity of the actinorhizal symbiosis in the presence of fluctuating nitrogen and metal conditions. These methods are transposable to numerous actinorhizal studies in both fundamental and applied research.     

Root symbioses ofAlnus glutinosa (L.) gaertn. and their possible role in alder decline: A preliminary study

During the last few years alder has declined in South Bohemia. The possible role of mycorrhizal and actinorhizal symbioses is reviewed and some of the preliminary results from experiments testing the influence of these symbioses on alder growth and the influence of eutrophication on the development of these symbioses are reported. Seedlings ofAlnus glutinosa were inoculated with arbuscular mycorrhizal (AM) fungi and the actinomyceteFrankia in experiment 1, and with rhizosphere soil collected from field sites with different degrees of alder damage in experiment 2. In both experiments, a solution containing nitrate, ammonia and phosphorus in concentrations simulating eutrophic waters, was applied. Both symbioses markedly promoted the growth of the seedlings in experiment 1. The plants inoculated with the rhizosphere soil microflora in experiment 2 were larger than the control plants. Response of the seedlings to the inoculation with the soil from the rhizosphere of damaged alder trees from six field sites differs, even though no correlation was found relating growth to the health status of the trees. Nutrient treatment did not have any effect on the growth of seedlings in either experiment. The dry weight ofFrankia was greater in mycorrhizal plants compared to nonmycorrhizal plants and mycorrhizal colonization is reduced inFrankia inoculated plants supplemented with phosphorus in experiment 1. Nitrogen enhanced mycorrhizal colonization in nodulated plants which were not supplemented with phosphorus no effect of nitrogen on actinorhiza was observed.     

Physiological and morphological responses of red alder and sitka alder to flooding

Red alder (Alnus rubra Bong.) and sitka alder (A. viridis ssp. sinuata [Regel] Löve & Löve) are nitrogen-fixing woody species that grow sympatrically along the Pacific coast of North America. Red alder is found in poorly drained lowlands, as well as in soils of moist upland slopes, whereas sitka alder generally colonizes well-drained soils. To identify factors that contribute to flood tolerance, we conducted greenhouse experiments subjecting both species to a 20-day flood and 10-day recovery and red alder to a 50-day flood and 20-day recovery. We determined the effect of this stress on nitrogenase activity, root and nodule alcohol dehydrogenase (ADH) activity, lenticel and adventitious root development, relative growth rate (RGR), and leaf gas exchange. After 24 h of flooding, nitrogenase activity could not be detected in either species. Limited nitrogenase activity did return in red alder at the end of a 10-day recovery following the 20-day flood, but sitka alder showed no recovery of nitrogenase activity. After 50 days of continuous flooding, red alder nitrogenase activity returned to pretreatment levels. Red alder root and nodule ADH activity was more than twice that of sitka alder under flooded conditions. Sitka alder showed extensive root mortality and leaf abscission over the same 20-day flooding period. Flooded red alder exhibited an initial decline in root RGR, but recovered between days 10 and 20 with the formation of adventitious roots. Furthermore, initiation of adventitious roots in red alder coincided with an increase in stomatal conductance without a similar recovery of carbon dioxide exchange rate. Sitka alder formed few adventitious roots, lost much of its root and leaf biomass, and showed no restoration of growth during flooding or recovery. Different responses of red and sitka alder to flooding serve as a partial explanation for the different patterns of distribution of these species and suggest some adaptations of red alder that permit flood tolerance.     

Low Frankia inoculation potentials in primary successional sites at Mount St. Helens, Washington, USA

The ability of 23 year old volcanic soils from Mount St. Helens, USA, to nodulate actinorhizal Sitka alder (Alnus viridis ssp. sinuata) was studied by estimating Frankia Inoculation Unit density (FIU) in four types of primary successional communities using a host-plant bioassay. Rhizospheric Mycelial Inoculation Unit (MIU) density and the effects of inoculation with alder soil and phosphorus (P) on seedling growth were also examined. FIU and MIU were highest in alder thickets, lower in lupin patch and riparian zone sites, and lowest, with no FIUs, in bare areas. Seedling dry mass was higher in treatments inoculated with alder soil but not with P addition, while root:shoot mass ratios decreased with both inoculation and P. The extremely low densities of suitable symbionts in sparsely vegetated primary successional areas at Mount St. Helens may explain the surprisingly slow rate of spread of alder from nearby seed sources.     

N2 fixing alder (Alnus viridis spp. fruticosa) effects on soil properties across a secondary successional chronosequence in interior Alaska

Green alder (Alnus viridis ssp. fruticosa) is a dominant understory shrub during secondary successional development of upland forests throughout interior Alaska, where it contributes substantially to the nitrogen (N) economy through atmospheric N₂ fixation. Across a replicated 200+ year old vegetation chronosequence, we tested the hypotheses that green alder has strong effects on soil chemical properties, and that ecosystem-level N inputs via N₂ fixation decrease with secondary successional stand development. Across early-, mid-, and late-successional stands, alder created islands of elevated soil N and carbon (C), depleted soil phosphorus (P), and more acidic soils. These effects translated to the stand-level in response to alder stem density. Although neither N₂ fixation nor nodule biomass differed among stand types, increases in alder densities with successional time translated to increasing N inputs. Estimates of annual N inputs by A. viridis averaged across the upland chronosequence (6.6 ± 1.2 kg N ha^?1 year^?1) are substantially less than inputs during early succession by Alnus tenuifolia growing along Alaskan floodplains. However, late-succession upland forests, where densities of A. viridis are highest, may persist for centuries, depending on fire return interval. This pattern of prolonged N inputs to late successional forests contradicts established theory predicting declines in N₂-fixation rates and N₂-fixer abundance as stands age.     

Nitrate reductase activity, nitrogenase activity and photosynthesis of black alder exposed to chilling temperatures

Actinorhizal ( Frankia -nodulated) black alder [ Alnus glutinosa (L.) Gaertn.] seedlings fertilized with 0.36 m M nitrate (low nitrate fertilizer treatment) or 7.14 m M nitrate (high nitrate fertilizer treatment) and acclimated in a growth chamber for 2 weeks were exposed to 2.5 h of night-time chilling temperatures of −1 to 4°C. Cold treatment decreased nitrogenase activity (acetylene reduction activity) 33% for low nitrate fertilized plants and 41% for high nitrate fertilized plants. Recovery of nitrogenase activity occurred within 7 days after chilling treatment. In contrast, in vivo nitrate reductase (NR) activities of leaves and fine roots increased immediately after chilling then decreased as nitrogenase activities recovered. Fine roots of alder seedlings exhibited NR activities proportional to the amounts of nitrate in the rooting medium. In contrast, the NR activities of leaves were independent of substrate and tissue nitrate levels and corresponded to nitrogenase activity in the root nodules. In a separate experiment, net photosynthesis (PS) of similarly treated black alder seedlings was measured before and after chilling treatments. Net PS declined in response to chilling by 17% for plants receiving low nitrate fertilizer and 19% for plants receiving high nitrate fertilizer. After chilling, stomatal conductance (g_s) decreased by 39% and internal CO₂ concentration (c_i) decreased by 5% in plants receiving the high nitrate fertilizer, whereas plants receiving the low nitrate fertilizer showed no change in g_s and a 13% increase in c_i. Results indicate that chilling stimulates stomatal closure only at the high nitrate level and that interference with biochemical functions is probably the major impact of chilling on PS.     

Heterotrophic microbial activities and nutritional status of microbial communities in tropical marsh sediments of different salinities: the effects of phosphorus addition and plant species

Oligotrophic, phosphorus (P) limited herbaceous wetlands of northern Belize are being impacted by P loading from fertilizer runoff. P enrichment causes a shift in autotroph communities from a microphyte (cyanobacterial mats, CBM) to macrophyte (Eleocharis spp., Typha domingensis) dominated system. To document potential effects of P, salinity, and macrophyte species on the heterotrophic microbial community nutritional status (represented especially by specific phospholipids fatty acids and specific respiration rate), biomass and activities, we took soil samples from established P enrichment plots in replicated marshes of two salinity levels. P addition increased microbial biomass carbon (C), nitrogen (N) and P, as well as soil nutrient transformation rates (nitrogenase activity, N mineralization and immobilization, methanogenesis). The effect of plant species (Eleocharis vs Typha sites) was generally lower than the effect of P addition (CBM vs Eleocharis sites) and was most evident at the low salinity sites, where Eleocharis dominated plots had enhanced nitrogenase activity and P microbial immobilization. Salinity reduced the overall rates of microbial processes; it also weakened the positive effect of both P addition and plant species on microbial activities. Lastly, the amount of N stored in microbial cells, likely in form of osmoprotective compounds, was enhanced by salinity.     

Short-term influence of nitrate on acetylene reduction, net photosynthesis and nodule respiration in black alder (alnus glutinosa L. gaertn.) seedlings

The use of nitrogen-fixing trees such as black alder (Alnus glutinosa L. Gaertn.) as forest silvicultural tools has recently been recognized. The potential benefit of black alder in silvicultural practices may be reduced by nitrate fertilization. Fifteen-month-old, nodulated, black alder rooted cuttings were fertilized for 6 days with 0, 7.5 or 15 mM NO₃ to determine the influence of nitrate on acetylene reduction, nodule respiration and net photosynthesis. Acetylene reduction, net photosynthesis and nodule respiration were measured on the second, fourth and sixth days of nitrate application. Nitrate treatment significantly reduced acetylene reduction and nodule respiration by day 4. Acetylene reduction was 75% lower and nodule respiration 36% lower for the 15 mM NO₃ treatment when compared to that of the control treatment. By day 6, net photosynthesis and nodule respiration were significantly reduced by 29 and 59%, respectively, for seedlings treated with 15 mM NO₃. This study suggests that nitrate fertilization has a profound influence on nitrogenase activity and that nitrogen-fixing tree species may respond to nitrate fertilization by shifting photosynthetic rates.     

Growth, nutrition, and nitrogen fixation in grey alder at varied rate of nitrogen addition

Seedlings of grey alder (Alnus incana Moench), nodulated or unnodulated, were investigated at varied relative addition rate of nitrogen. Nitrogen fixation alone, without addition of mineral nitrogen, resulted in an almost optimum nitrogen status but only about half the maximum relative growth rate, probably mainly because of energy costs of nodulation and fixation. The growth deficit due to nodulation was much more than can be explained by the theoretical energy requirement for the amount of nitrogen fixed. Thus, the nitrogen fixation process was not very efficiently used. The nitrogen fixation rate was strongly stimulated by increasing nitrogen addition rate up to high levels. The fixation rate decreased rapidly close to optimum (maximum relative growth rate) and was negligible at maximum growth. A feed-back of mineral nitrogen on photosynthesis increased fixation rate with time, and the relative importance of fixation over mineral nitrogen nutrition increased. However, nitrogen fixation, also at maximum rate, supplied only a small proportion of the nitrogen amount required for maximum growth. The optimum nutrient solutions contained comparatively high nitrogen concentrations to secure free access to nitrogen. The nodules were damaged by this treatment, and it is concluded that the nitrogen additions must be adjusted to the current consumption of the plants to avoid an increased external nitrogen concentration. Strong linear regressions were found between relative growth rate, nitrogen status expressed as percentage content of fresh weight, and relative growth rate in unnodulated seedlings. There was a greater variability in nodulated seedlings than in unnodulated ones, because of the nitrogen fixation. The reactions of unnodulated grey alder were largely the same as previously reported for birch seedlings, but the maximum growth capacity was lower in grey alder. During an initial period of change in the internal nitrogen status, deficiency symptoms appeared, especially in unnodulated seedlings. As in birch, the leaves turned green again at stable nitrogen status, independent of level. The results are in sharp contrast to data from the literature where the external nitrogen concentration was used as the driving variable for the internal nitrogen status. The measured fixation rates for grey alder are much higher than those previously reported. Still, the maximum fixation rate observed is small compared to the total nitrogen uptake rate required for maximum growth, in contrast to reported relationships. These comparisons indicate that increased external nitrogen concentration obscures the real relations between mineral and fixed nitrogen, on one hand because of rapid inhibition of nitrogen fixation and, on the other hand, because of failure to obtain stable optimum nutrition and maximum growth by means of this treatment variable.     

Physiological indicators of nitrogen response in a short rotation sycamore plantation. I. CO2 assimilation, photosynthetic pigments and soluble carbohydrates

American sycamore ( Platanus occidentalis L.) seedlings were grown in the field under different urea-nitrogen fertilization regimes to identify physiological variables that characterize the growth response. Treatments included fertilization at the beginning of the growing season with 50, 150, 450 kg N ha⁻¹, fertilization 3 times each at 37.5 kg N ha⁻¹ and unfertilized control. The greatest aboveground biomass accumulation (3× that of control) occurred in plots fertilized with 450 kg N ha⁻¹, but nearly as much growth occurred when 37.5 kg N ha⁻¹ was added periodically. Photosynthesis, chlorophyll concentrations and growth increased rapidly after the midseason application of 37.5 kg N ha⁻¹ but not after the late-season application. Although nitrogen fertilization increased leaf area per plant, leaf nitrogen concentration did not differ between treatments. There was no evidence to indicate that fertilization extended the physiologically active season or increased susceptibility to drought or cold. Sycamore leaves accumulated sucrose and mannose in response to water stress in all treatments. Photosynthetic pigment concentrations and net photosynthetic rate were the most sensitive indicators of growth response to nitrogen fertilization in the first growing season. Careful timing (based on physiological indicators) of low level applications of nitrogen fertilizer can optimize growth.     

Density of red alder (Alnus rubra) in headwaters influences invertebrate and detritus subsidies to downstream fish habitats in Alaska

We investigated the influence of red alder (Alnus rubra) stand density in upland, riparian forests on invertebrate and detritus transport from fishless headwater streams to downstream, salmonid habitats in southeastern Alaska. Red alder commonly regenerates after soil disturbance (such as from natural landsliding or timber harvesting), and is common along streams in varying densities, but its effect on food delivery from headwater channels to downstream salmonid habitats is not clear. Fluvial transport of invertebrates and detritus was measured at 13 sites in spring, summer and fall during two years (2000–2001). The 13 streams encompassed a riparian red alder density gradient (1–82% canopy cover or 0–53% basal area) growing amongst young-growth conifer (45-yr-old stands that regenerated after forest clearcutting). Sites with more riparian red alder exported significantly more invertebrates than did sites with little alder (mean range across 1–82% alder gradient was about 1–4 invertebrates m^?3 water, and 0.1–1 mg invertebrates m^?3 water, respectively). Three-quarters of the invertebrates were of aquatic origin; the remainder was of terrestrial origin. Aquatic taxa were positively related to the alder density gradient, while terrestrially-derived taxa were not. Streams with more riparian alder also exported significantly more detritus than streams with less alder (mean range across 1–82% alder gradient was 0.01–0.06 g detritus m^?3 water). Based on these data, we predict that headwater streams with more riparian alder will provide more invertebrates and support more downstream fish biomass than those basins with little or no riparian alder, provided these downstream food webs fully utilize this resource subsidy.     

Comparison of fatty acids and elemental nutrients in periphyton, invertebrates, and cutthroat trout (Oncorhynchus clarki) in conifer and alder streams of western Washington state

Organism growth and reproduction are often limited by nutrient availability in freshwater ecosystems where, in some cases, food webs are primarily supported by allochthonous organic matter. Therefore, we hypothesized that the composition of riparian vegetation would influence the variability of N, P, and fatty acid content of in-stream consumers. Specifically, we predicted that organisms living in alder streams would have higher levels of N, P, and polyunsaturated fatty acids than organisms in coniferous streams. To determine this, we sampled fresh and aged leaf litter, periphyton, invertebrates, and cutthroat trout (Oncorhynchus clarki) from 6 streams in western Washington state: 3 streams had high densities of nitrogen-fixing red alder (Alnus rubra) in the riparian zone, whereas 3 had high densities of conifers. We found fresh alder litter had twice the total polyunsaturated fatty acid concentrations of hemlock vegetation while there were few statistical differences among aged alder and aged hemlock vegetation. Multidimensional plots showed fatty acid profiles were unique to vegetation and fish while periphyton and invertebrates shared the same multidimensional space. We used a mixed model to determine the relative importance of vegetation type (fixed factor: conifer or alder), trophic levels (fixed factor: periphyton, primary consumer, or fish), and streams (random factor) on individual fatty acid concentrations. Total polyunsaturated fatty acids, 16:0, 20:1, 20:3n6 and total n3 were the only fatty acids influenced by stream vegetation (vegetation + stream model or full model). 67% of the fatty acids were best supported by the trophic + stream model. Nitrogen, P, Ca, Fe, C:N, N:P, and C:N:P were all best supported by the trophic level + stream model, and Zn was the only nutrient supported best by the full model. Correlations of n3 and n6 fatty acid concentrations between periphyton and primary consumers, and primary consumers with trout indicated several fatty acid metrics, such as n3:n6, showed food resources may affect relative fatty acid abundances of consumers. Although vegetation type did not influence relative fatty acids of stream organisms, the importance of trophic level likely indicates organisms have different physical requirements for fatty acids. The significance of a random factor, ‘stream,’ suggests that the relative abundances of fatty acids in periphyton, invertebrates, and trout are stream-specific and are responding to local environmental or communal variables.     

Local and systemic effects of phosphorus and nitrogen on nodulation and nodule function in Alnus incana

Phosphorus (P) and nitrogen (N) effects on nodulation, nitrogenase activity and plant growth were studied in the root-hair-infected actinorhizal plant Alnus incana (L.) Moench. A split-root experiment, as well as a short-term experiment with entire root systems and a broader range of P concentrations, showed that P effects were specific on nodulation and not a general stimulation via a plant growth effect. These results indicate that nodule initiation and nodule growth have a high P demand. The split-root assay, comprising seven combinations of two N and two P levels, showed that P could counteract systemic N inhibition of nodulation, but did not counteract N inhibition of nitrogenase activity.     

大兴安岭北部多年冻土区河岸森林湿地土壤性质和微生物呼吸活性特征

多年冻土区河岸森林湿地是水文、生态和生物化学过程的关键区域。本研究以河岸森林湿地及其与泥炭地的交错带土壤为对象,分析了腐殖质层和不同深度土壤理化性质、生态化学计量和微生物呼吸活性( 微生物生物量碳、基础呼吸、微生物熵和代谢熵)特征。结果表明: 与大兴安岭多年冻土区泥炭地和河岸森林湿地的交错带相比,河岸森林湿地土壤理化性质主要分异在20 cm土层以下,其总碳、总氮含量和碳磷比、氮磷比显著降低,生态化学计量特征的变化主要是由于氮含量变化引起的,说明河岸森林湿地土壤氮转移相对较快,存在氮限制;交错带湿地土壤中钠、镁、钾和钙含量主要在30 cm土层发生分异,而河岸森林湿地土壤中钠、镁、钾和钙含量主要在20 cm土层发生分异,其镁含量与土壤总碳、总氮和总磷含量显著相关,说明土壤镁含量是大兴安岭河岸森林湿地的重要营养元素;河岸森林湿地和交错带腐殖质层微生物呼吸活性高于其他层土壤,说明其易分解的碳组分含量高;河岸森林湿地和交错带土壤微生物呼吸活性与土壤理化性质、生态化学计量特征及营养元素的相关性存在差异,而河岸森林湿地土壤总氮含量与微生物呼吸活性显著相关,说明大兴安岭河岸带湿地土壤微生物活性受氮的限制。     

Differential recovery of above‐ and below‐ground rich fen vegetation following fertilization

Abstract. For seven years we studied the recovery of vegetation in a Belgian P limited rich fen (Caricion davallianae), which had been fertilized with nitrogen (200 g.m^?2) and phosphorus (50 g.m^?2) in 1992. The vegetation in this fen has low above‐ground biomass production (< 100 g m^?2) due to the strong P limitation. Above‐ground biomass was harvested from 1992 to 1998 and P and N concentrations measured. In 1998, below‐ground biomass was also harvested. The response to fertilization differed markedly between below‐ and above‐ground compartments. Above‐ground, P was the single most important factor that continued to stimulate growth 7 yr after fertilization. Below‐ground, N tended to decrease live root biomass and increase dead root biomass and seemed to have a toxic effect on the roots. In the combined NP treatment the stimulating effect of P (an increase of live root biomass) was countered by N. The 1998 soil analysis showed no difference in soil P in the plots. Thus, Fe hydroxides are not capable of retaining P in competition with fen vegetation uptake. However, higher capture of P in root Fe coatings from N plots may partially explain this negative N effect. The results suggest that N root toxicity will be amplified in strongly P limited habitats but that its persistence will be less influenced by P availability. This mechanism may be a competitive advantage for N₂ fixing species that grow in strongly P limited wetlands.