The Estimated Impact of Fungi on Nutrient Dynamics During Decomposition of Phragmites australis Leaf Sheaths and Stems

Decomposition of culms (sheaths and stems) of the emergent macrophyte Phragmites australis (common reed) was followed for 16 months in the litter layer of a brackish tidal marsh along the river Scheldt (the Netherlands). Stems and leaf sheaths were separately analyzed for mass loss, litter-associated fungal biomass (ergosterol), nutrient (N and P), and cell wall polymer concentrations (cellulose and lignin). The role of fungal biomass in litter nutrient dynamics was evaluated by estimating nutrient incorporation within the living fungal mass. After 1 year of standing stem decay, substantial fungal colonization was found. This corresponded to an overall fungal biomass of 49 ± 8.7 mg g⁻¹ dry mass. A vertical pattern of fungal colonization on stems in the canopy is suggested. The litter bag experiment showed that mass loss of stems was negligible during the first 6 months, whereas leaf sheaths lost almost 50% of their initial mass during that time. Exponential breakdown rates were −0.0039 ± 0.0004 and −0.0026 ± 0.0003 day⁻¹ for leaf sheaths and stems, respectively (excluding the initial lag period). In contrast to the stem tissue—which had no fungal colonization—leaf sheaths were heavily colonized by fungi (93 ± 10 mg fungal biomass g⁻¹ dry mass) prior to placement in the litter layer. Once being on the sediment surface, 30% of leaf sheath's associated fungal biomass was lost, but ergosterol concentrations recovered the following months. In the stems, fungal biomass increased steadily after an initial lag period to reach a maximal biomass of about 120 mg fungal biomass g⁻¹ dry mass for both plant parts at the end of the experiment. Fungal colonizers are considered to contain an important fraction of nutrients within the decaying plant matter. Fungal N incorporation was estimated to be 64 ± 13 and 102 ± 15% of total available N pool during decomposition for leaf sheaths and stems, respectively. Fungal P incorporation was estimated to be 37 ± 9 and 52 ± 15% of total available P during decomposition for leaf sheaths and stems, respectively. Furthermore, within the stem tissue, fungi are suggested to be active immobilizers of nutrients from the external environment because fungi were often estimated to contain more than 100% of the original nutrient stock.     

Fungi on Leaf Blades of <Emphasis Type="Italic">Phragmites australis</Emphasis> in a Brackish Tidal Marsh: Diversity,Succession, and Leaf Decomposition

Although fungi are known to colonize and decompose plant tissues in various environments, there is scanty information on fungal communities on wetland plants, their relation to microhabitat conditions, and their link to plant litter decomposition. We examined fungal diversity and succession on Phragmites australis leaves both attached to standing shoots and decaying in the litter layer of a brackish tidal marsh. Additionally, we followed changes in fungal biomass (ergosterol), leaf nitrogen dynamics, and litter mass loss on the sediment surface of the marsh. Thirty-five fungal taxa were recorded by direct observation of sporulation structures. Detrended correspondence analysis and cluster analysis revealed distinct communities of fungi sporulating in the three microhabitats examined (middle canopy, top canopy, and litter layer), and indicator species analysis identified a total of seven taxa characteristic of the identified subcommunities. High fungal biomass developed in decaying leaf blades attached to standing shoots, with a maximum ergosterol concentration of 548 ± 83 μg g^–1 ash-free dry mass (AFDM; mean ± SD). When dead leaves were incorporated in the litter layer on the marsh surface, fungi experienced a sharp decline in biomass (to 191 ± 60 μg ergosterol g^–1 AFDM) and in the number of sporulation structures. Following a lag phase, species not previously detected began to sporulate. Leaves placed in litter bags on the sediment surface lost 50% of their initial AFDM within 7 months (k = −0.0035 day^–1) and only 21% of the original AFDM was left after 11 months. Fungal biomass accounted for up to 34 ± 7% of the total N in dead leaf blades on standing shoots, but to only 10 ± 4% in the litter layer. These data suggest that fungi are instrumental in N retention and leaf mass loss during leaf senescence and early aerial decay. However, during decomposition on the marsh surface, the importance of living fungal mass appears to diminish, particularly in N retention, although a significant fraction of total detrital N may remain associated with dead hyphae.     

Decomposition and CO2 Evolution from Standing Litter of the Emergent Macrophyte Erianthus giganteus

Abstract Decomposition of standing litter of the emergent macrophyte Erianthus giganteus (plumegrass) was quantified in a small freshwater wetland in Alabama, USA. Living green shoots of E. giganteus were tagged and periodically retrieved for determination of leaf and culm mass loss, litter-associated fungal biomass (ergosterol), and nitrogen and phosphorus concentrations. Laboratory studies were also conducted to examine the effects of plant litter moisture content and temperature on rates of CO₂ evolution from plant litter. Culm and leaf material lost 25 and 32% AFDM, respectively, during plant senescence and early litter decay. Fungal biomass, as determined by ergosterol concentrations, increased significantly in both leaf and culm litter during decomposition, with maximum biomass accounting for 3.7 and 6.7% of the total detrital weight in culm and leaf litter, respectively. Spatial differences in fungal biomass were observed along the culm axis, with upper regions of the culm accumulating significantly greater amounts of fungal mass than basal regions (p < 0.01, ANOVA). Rates of CO₂ evolution from both leaf and culm litter increased rapidly after wetting (0 to 76 μg CO₂−C g⁻¹ AFDM h⁻¹ within 5 min). In addition, rates of CO₂ evolution from water saturated culms increased exponentially as the temperature was increased from 10 to 30°C. These results provide evidence that considerable microbial colonization and mineralization of standing emergent macrophyte litter can occur before collapse of senescent shoot material to the water and sediment surface. Received: 5 December 1998; Accepted: 31 March 1999     

Estimation of fungal biomass in the decaying cones ofPinus densiflora

Fungal biomass in the decaying cones ofPinus densiflora was investigated. Leaching, immobilization and mobilization phases were recognized in the decomposition process of cones. Fungal biomass was estimated by the agar-film technique, using a conversion factor of 0.62 mg dry wt. mm⁻³ of hyphal volume to biomass and a factor of 2.5 for in-efficiencency of homogenization. The fungal biomass was 4.9±2.1 (mean±S.D.) mg dry wt. g⁻¹ dry matter in the cones on the tree, 11±6 mg g⁻¹ in the leaching phase, 19±7 mg g⁻¹ in the immobilization phase and 30±15 mg g⁻¹ in the mobilization phase. It significantly increased after cones had lain on the forest floor, and also in the immobilization phase. The latter result suggests that the fungal biomass contributed to the immobilization of nitrogen in the decomposition process. The ratio of ergosterol content to fungal biomass in the cones was 2.9–8.8 μg mg⁻¹ dry wt., lying in the range of 2–16 μg mg⁻¹ reported for mycelia. This suggested that the estimate of fungal biomass was reasonable. Reduction in this ratio with the dry weight loss in the cones suggested that the proportion of relatively active fungal biomass decreased with the progress of decomposition.     

Estimates of biomass and primary productivity in a high-altitude maple forest of the west central Himalayas

The paper describes the biomass and productivity of maple (Acer cappadocicum) forest occurring at an altitude of 2,750 m in the west central Himalayas. Total vegetation biomass was 308.3 t ha⁻¹, of which the tree layer contributed the most, followed by herbs and shrubs. The seasonal forest-floor litter mass varied between 5.4 t ha⁻¹ (in rainy season) and 6.6 t ha⁻¹ (in winter season). The annual litter fall was 6.2 t ha⁻¹, of which leaf litter contributed the largest part (59% of the total litter fall). Net primary productivity of total vegetation was 19.5 t ha⁻¹ year⁻¹. The production efficiency of leaves (net primary productivity/leaf mass) was markedly higher (2.9 g g⁻¹ foliage mass year⁻¹) than those of the low-altitude forests of the region.     

Litter N:P ratios indicate whether N or P limits the decomposability of graminoid leaf litter

The N:P ratio of leaf litter may determine if decomposability is N-limited (litter with low N:P ratio) or P-limited (litter with high N:P ratio). To test this hypothesis and to determine the threshold between N and P limitation, we studied relationships between litter N and P concentrations, litter mass loss and effects of fertilisation on litter mass loss in laboratory experiments. Leaf litter of 11 graminoid species was collected in Swiss and Dutch wetlands, yielding 84 litter samples with a broad range of N and P concentrations (3.2–15.1 mg N g⁻¹, 0.04–1.93 mg P g⁻¹) and with N:P mass ratios ranging from 5 to 100. On nutrient-free sand, dry mass loss after five or ten weeks (5.5–53% of initial mass) correlated positively with the N and P concentrations of the litter. Within species, mass loss correlated mainly with N for litter with low N:P ratio, and with P for litter with high N:P ratio, in agreement with our hypothesis. Among species, however, these relationships did not exist, and decomposition rather correlated with the specific leaf area. When the litter was incubated on fertilised sand, 35 out of 50 litter samples decomposed faster than on nutrient-free sand. Decomposition was generally accelerated by P fertilisation (i.e. P-limited) when the N:P ratio of the litter was above 25 and the P concentration below 0.22 mg g⁻¹, supporting our hypothesis. N-limited decomposition was not significantly related to the litter N:P ratio but occurred rarely for litter with N:P ratio greater than 25, and only for litter with N concentration below 11.3 mg g⁻¹. We conclude that the N:P ratio of leaf litter indicates whether its decomposability is more likely to be N- or P-limited. The critical N:P ratio (threshold between N and P limitation) appeared to be 25 for graminoid leaf litter.     

Annual production of leaf-decaying fungi in a woodland stream

1. Fungi are thought to be important mediators of energy flow in the detritus-based food webs of woodland streams. However, until recently, quantitative methods to assess their contribution have been lacking. Growth rates of leaf-decaying fungi can be estimated from rates of acetate incorporation into ergosterol which, together with estimates of fungal biomass from ergosterol concentrations, enables calculation of fungal production. In this study, I used this method to estimate total production of leaf-decaying fungi over an annual cycle in a small woodland stream, Walker Branch, Tennessee, U.S.A. To calculate fungal biomass and production on an areal basis, I determined the amount of leaf litter occurring in the stream by sampling transects randomly selected in each of ten 10-m sections every 20–50 days. Subsamples of leaves chosen from five of the transects were used to determine ergosterol concentrations and in situ rates of acetate incorporation into ergosterol. 2. Leaf litter, fungal biomass m^–2, and fungal production m^–2 were highly seasonal. Leaf litter ranged from 249 g m^–2 in November to less than 5 g m^–2 during the summer. Fungal biomass as percentage of leaf litter ranged from 4.4 to 8.8% during the year, but on an areal basis ranged from 11 to 13 g m^–2 during November to January to 0.25 g m^–2 in June, primarily due to the seasonal variation in amount of leaf litter present. Fungal growth rates averaged 2.6% day^–1 (0.9–7.0% day^–1) during the year. Daily production of leaf-decaying fungi ranged from 0.49 g m^–2 in November, when the amount of leaf litter was at its maximum, to 0.006 g m^–2 during the summer when the amount of leaf litter was low. Annual production of leaf-decaying fungi was 34 g m^–2, with an annual production to biomass ratio (P/B) of 8.2. 3. Fungal spore concentrations in the stream were also seasonal and were correlated with amount of leaf litter m^–2 and fungal biomass m^–2. Spore concentrations varied between one and four spores ml^–1 throughout most of the year, but increased to eighteen spores ml^–1 shortly after the greatest amount of leaf litter was present in the stream during November.     

Chemical composition of litter affects the growth and enzyme production by the saprotrophic basidiomycete Hypholoma fasciculare

Chemical composition of litter has previously been reported to affect in situ decomposition. To identify its effects on a single species level, the saprotrophic basidiomycete Hypholoma fasciculare was grown on 11 types of litter with variable chemical composition (N content of 3.4–28.9 mg g⁻¹), and the mass loss of litter and lignin, production of extracellular enzymes and fungal biomass were followed. After 12 weeks, mass loss ranged from 16 % to 34 %. During early decomposition stages, litter mass loss, fungal biomass production (estimated by ergosterol content) as well as fungal substrate use efficiency all increased with increasing initial N content of the litter. The initial litter decomposition rate was significantly positively correlated with the activities of arylsulfatase, cellobiohydrolase, endoxylanase and phosphatase. Contrary to expectations, the lignin content did not affect litter mass loss, when covariation with N content was accounted for. The ratio of lignin loss to total mass loss depended on the litter type and did not reflect the activities of ligninolytic enzymes.     

Fungal biomass and decomposition in Spartina maritima leaves in the Mondego salt marsh (Portugal)

Spartina maritima (Curtis) Fernald is a dominant species in the Mondego salt marsh on the western coast of Portugal, and it plays a significant role in estuarine productivity. In this work, leaf litter production dynamics and fungal importance for leaf decomposition processes in Spartina maritima were studied. Leaf fall was highly seasonal, being significantly higher during dry months. It ranged from 42 g m^-2 in June to less than 6 g m^-2 during the winter. Fungal biomass, measured as ergosterol content, did not differ significantly between standing-decaying leaves and naturally detached leaves. Fungal biomass increased in wet months, with a maximum of 614 g g^-1 of ergosterol in January in standing-decaying leaves, and 1077 g g^-1 in December, in naturally detached leaves, decreasing greatly in summer. Seasonal pattern of fungal colonization was similar in leaves placed in litterbags on the marsh-sediment surface. However, ergosterol concentrations associated with standing-decaying and naturally detached leaves were always much higher than in litterbagged leaves, suggesting that fungal activity was more important before leaf fall. Dry mass of litterbagged leaves declined rapidly after 1 month (about 50%), mostly due to leaching of soluble organic compounds. After 13 months, Spartina leaves had lost 88% of their original dry weight. The decomposition rate constant (k) for Spartina maritima leaves was 0.151 month^-1.     

Effect of Culture Conditions on Ergosterol as an Indicator of Biomass in the Aquatic Hyphomycetes

Ergosterol is a membrane component specific to fungi that can be used to estimate fungal biomass using appropriate factors of conversion. Our objectives were to determine the limits of use of ergosterol content as a measure of biomass for aquatic hyphomycetes, and to evaluate a previously established ergosterol-to-biomass conversion factor. We varied inoculum quality, growth medium, and degree of shaking of four aquatic hyphomycete species. In cultures inoculated with homogenized mycelium, we found a significant effect of shaking condition and culture age on ergosterol content. In liquid cultures with defined medium, ergosterol content reached 10 to 11 μg/mg of mycelium (dry mass) and varied by factors of 2.2 during exponential growth and 1.3 during stationary phase. The increase in ergosterol content during exponential phase could be attributed, at least in part, to rapid depletion of glucose. Oxygen availability to internal hyphae within the mycelial mass is also responsible for the differences found between culture conditions. Ergosterol concentration ranged from 0.8 to 1.6 μg/mg in static cultures inoculated with agar plugs. Ergosterol content varied by a factor of 4 in two media of different richnesses. For different combinations of these parameters, strong (r² = 0.83 to 0.98) and highly significant (P 0.001) linear relationships between ergosterol and mycelial dry mass (up to 110 mg) were observed. Overall, the ergosterol content varied by a factor of 14 (0.8 to 11 mg/g). These results suggest that care must be taken when the ergosterol content is used to compare data generated in different field environments.     

Total and Free Ergosterol in Mycelia of Saltmarsh Ascomycetes with Access to Whole Leaves or Aqueous Extracts of Leaves

Three species of saltmarsh ascomycetes were grown in the presence of all of the constituents of their natural substrate (leaves of cordgrass) or were presented only with aqueous extracts of the leaves. These two growth-condition treatments had no significant effect on total ergosterol content of the fungal mycelia, contrary to an earlier hypothesis that availability of plant lipids would lower fungal ergosterol contents. Mycelial content of free ergosterol was about twice as variable as that for total (free plus esterified) ergosterol. Total ergosterol (data pooled for all species) was strongly correlated to organic mycelial mass (r² = 0.43, P < 0.00001, and slope = 4.59 μg of ergosterol mg of organic mass^-1).     

Leaf traits variation during leaf expansion in <Emphasis Type="Italic">Quercus ilex</Emphasis> L.

The morphological, anatomical and physiological variations of leaf traits were analysed during Quercus ilex L. leaf expansion. The leaf water content (LWC), leaf area relative growth rate (RGR_l) and leaf dry mass relative growth rate (RGR_m) were the highest (76±2 %, 0.413 cm² cm⁻² d⁻¹, 0.709 mg mg⁻¹ d⁻¹, respectively) at the beginning of the leaf expansion process (7 days after bud break). Leaf expansion lasted 84±2 days when air temperature ranged from 13.3±0.8 to 27.6±0.9 °C. The net photosynthetic rate (P _N), stomatal conductance (g _s), and chlorophyll content per fresh mass (Chl) increased during leaf expansion, having the highest values [12.62±1.64 μmol (CO₂) m⁻² s⁻¹, 0.090 mol (H₂O) m⁻² s⁻¹, and 1.03±0.08 mg g⁻¹, respectively] 56 days after bud break. Chl was directly correlated with leaf dry mass (DM) and P _N. The thickness of palisade parenchyma contributed to the total leaf thickness (263.1±1.5 μm) by 47 %, spongy layer thickness 38 %, adaxial epidermis and cuticle thickness 9 %, and abaxial epidermis and cuticle thickness 6 %. Variation in leaf size during leaf expansion might be attributed to a combination of cells density and length, and it is confirmed by the significant (p<0.001) correlations among these traits. Q. ilex leaves reached 90 % of their definitive structure before the most severe drought period (beginning of June — end of August). The high leaf mass area (LMA, 15.1±0.6 mg cm⁻²) at full leaf expansion was indicative of compact leaves (2028±100 cells mm⁻²). Air temperature increasing might shorten the favourable period for leaf expansion, thus changing the final amount of biomass per unit leaf area of Q. ilex.     

Effects of water quality on habitat use by lesser scaup (Aythya affinis) broods in the boreal Northwest Territories, Canada

The dynamics of Rhizophora mangle litter production and decomposition were studied in a tropical coastal lagoon on the Gulf of Mexico in Veracruz, Mexico over a year (October 2002–October 2003). This region is characterized by three seasons: northerly winds (called ‘nortes’), dry, and rainy. Annual litter production (1116 g m⁻²) followed a seasonal pattern with leaf litter as the main fraction (70%) with two peaks in the dry and one in the rainy season. Leaf decomposition was evaluated with two types of litter bag in each season: fine mesh (1×1 mm) and coarse mesh (3×7 mm). Decomposition data were adjusted to a single negative exponential model. The results indicated faster decomposition rates in the coarse litter bag and significant differences among seasons. However these differences occurred after the 60th day of decomposition, indicating that leaching and microbial action were responsible for more than 50% of mass loss. After this period, the effects of aquatic invertebrates were evident but depended on climatic conditions. In the rainy season, the gastropod Neritina reclivata was associated with increasing leaf decomposition rate. In the ‘nortes’ season, the effect of aquatic invertebrates was smaller, and there were no differences in the decay constants calculated for the two litter bag types. High litter production represents an important input of organic matter which, through decomposition, may represent an important source of C, N, and P in this aquatic system.     

Productivity and mineral cycling in an oak coppice forest 2. Annual net production of the forest

Annual net production was estimated in the secondary coppice forest near Tokyo, which was dominated by a deciduous oak,Quercus serrata Thunb. Lateral growth of stems and old branches was directly estimated by examining the annual rings for 35 shoots in a clear-cut quadrat of 10m×10m. Phytomasses of current organs were also weighed in the quadrat. Preharvest losses of current organs were determined by twelve 0.5 m² litter traps for fine litter and twelve 6 m² quadrats for woody litter. Branch production was also assessed indirectly by use of the stem-branch allometry and death of branches. The results of the indirect method were in sufficient agreement with the result of the direct one. Grazing loss of leaves from the canopy was estimated directly from the loss in leaf area and indirectly from the animal faeces caught by the litter traps. Net production of the canopy trees was 149 kg a⁻¹ year⁻¹, in which leaf production was 36.9 kg. Animals grazed about 14% of the leaf area by the end of the growing season. True consumption of leaves by animals was 7.6% of leaf production or 10% of leaf mass. Production of undergrowth, mainly a dwarf bamboo,Pleioblastus chino Makino, was 28 kg a⁻¹ year⁻¹, being 15% of the total stand production. Productivity of this forest was significantly higher than that of cool-temperate deciduous broadleaf forests.     

Culm recruitment,dry matter dynamics and carbon flux in recently harvested and mature bamboo savannas in the Indian dry tropics

Culm recruitment, standing crop biomass, net production and carbon flux were estimated in mature (5 years after last harvest) and recently harvested bamboo (Dendrocalamus strictus (Roxb.) Nees) savanna sites in the dry tropics. During the 2 study years bamboo shoot recruitment was 1711–3182 and 1432–1510 shoots ha⁻¹ in harvested and mature sites, respectively. Corresponding shoot mortality was 66–93% and 62–69%, respectively. Total biomass was 34.9 t ha⁻¹ at the harvested site and 47.4 t ha⁻¹ at the mature site. Harvesting increased the relative contribution of belowground bamboo biomass. Annual litter input to soil was 2.7 and 5.9 t ha⁻¹ year⁻¹ at the harvested and mature sites, respectively. The bulk of the annual litterfall (78–88%) occurred in the cool dry season (November to February). The mean litter mass on the savanna floor ranged from 3.1 to 3.3 t ha⁻¹; at the harvested site wood litter contributed 70% of the litter mass and at the mature site leaves formed 77% of the litter mass. The mean total net production (TNP) for the two annual cycles was 15.8 t ha⁻¹ year⁻¹ at the harvested site and 19.3 t ha⁻¹ year⁻¹ at the mature site. Nearly half (46–57%) of the TNP was allocated to the belowground parts. Short lived components (leaves and fine roots) contributed about four-fifths of the net production of bamboo. Total carbon storage in the system was 64.4 t ha⁻¹ at the harvested site and 75.4 t ha⁻¹ at the mature site, of which 23–28% was distributed in vegetation, 2% in litter and 70–75% in soil. Annual net carbon deposition was 6.3 and 8.7 t ha⁻¹ year⁻¹ at harvested and mature sites, respectively.     

Litter production, leaf litter decomposition and nutrient return in Cunninghamia lanceolata plantations in south China: effect of planting conifers with broadleaved species

This study compared litter production, litter decomposition and nutrient return in pure and mixed species plantations. Dry weight and N, P, K, Ca, Mg quantities in the litterfall were measured in one pure Cunninghamia lanceolata plantation (PC) and two mixed-species plantations of C. lanceolata with Alnus cremastogyne (MCA) and Kalopanax septemlobus (MCK) in subtropical China. Covering 6 years of observations, mean annual litter production of MCA (4.97 Mg·ha⁻¹) and MCK (3.97 Mg·ha⁻¹) was significantly higher than that of PC (3.46 Mg·ha⁻¹). Broadleaved trees contributed 42% of the total litter production in MCA and 31% in MCK. Introduction of broadleaved tree species had no significant effect on litterfall pattern. Total litterfall was greatest in the dry season from November to March. Nutrient returns to the forest floor through leaf litter were significantly higher in both MCA and MCK than in PC (P < 0.05). The amounts of N, K, and Mg returned to the forest floor through leaf litter were highest in the MCA, and P and Ca returns were highest in the MCK. Percent contribution of broadleaf litter to total nutrient returns ranged from 41.7% to 86.9% in MCA and from 49.3% to 74.8% in MCK. The decomposition rate of individual leaf litter increased in the order: C. lanceolata < K. septemlobus < A. cremastogyne. Litter mixing had a positive effect on decomposition rate of the more recalcitrant litter and promoted nutrient return. Relative to mass loss of A. cremastogyne decomposing alone, higher mass loss of the mixture of C. lanceolata and A. cremastogyne was observed after 330 days of decomposition. These results indicate that mixed plantations of different tree species have advantages over monospecific plantations with regards to nutrient fluxes and these advantages have relevance to restoration of degraded sites. Responsible Editor: Alfonso Escudero.     

Response of Oxidative Enzyme Activities to Nitrogen Deposition Affects Soil Concentrations of Dissolved Organic Carbon

Recent evidence suggests that atmospheric nitrate (NO ₃ ⁻ ) deposition can alter soil carbon (C) storage by directly affecting the activity of lignin-degrading soil fungi. In a laboratory experiment, we studied the direct influence of increasing soil NO ₃ ⁻ concentration on microbial C cycling in three different ecosystems: black oak–white oak (BOWO), sugar maple–red oak (SMRO), and sugar maple–basswood (SMBW). These ecosystems span a broad range of litter biochemistry and recalcitrance; the BOWO ecosystem contains the highest litter lignin content, SMRO had intermediate lignin content, and SMBW leaf litter has the lowest lignin content. We hypothesized that increasing soil solution NO ₃ ⁻ would reduce lignolytic activity in the BOWO ecosystem, due to a high abundance of white-rot fungi and lignin-rich leaf litter. Due to the low lignin content of litter in the SMBW, we further reasoned that the NO ₃ ⁻ repression of lignolytic activity would be less dramatic due to a lower relative abundance of white-rot basidiomycetes; the response in the SMRO ecosystem should be intermediate. We increased soil solution NO ₃ ⁻ concentrations in a 73-day laboratory incubation and measured microbial respiration and soil solution dissolved organic carbon (DOC) and phenolics concentrations. At the end of the incubation, we measured the activity of β-glucosidase, N-acetyl-glucosaminidase, phenol oxidase, and peroxidase, which are extracellular enzymes involved with cellulose and lignin degradation. We quantified the fungal biomass, and we also used fungal ribosomal intergenic spacer analysis (RISA) to gain insight into fungal community composition. In the BOWO ecosystem, increasing NO ₃ ⁻ significantly decreased oxidative enzyme activities (−30% to −54%) and increased DOC (+32% upper limit) and phenolic (+77% upper limit) concentrations. In the SMRO ecosystem, we observed a significant decrease in phenol oxidase activity (−73% lower limit) and an increase in soluble phenolic concentrations (+57% upper limit) in response to increasing NO ₃ ⁻ in soil solution, but there was no significant change in DOC concentration. In contrast to these patterns, increasing soil solution NO ₃ ⁻ in the SMBW soil resulted in significantly greater phenol oxidase activity (+700% upper limit) and a trend toward lower DOC production (−52% lower limit). Nitrate concentration had no effect on microbial respiration or β-glucosidase or N-acetyl-glucosaminidase activities. Fungal abundance and basidiomycete diversity tended to be highest in the BOWO soil and lowest in the SMBW, but neither displayed a consistent response to NO ₃ ⁻ additions. Taken together, our results demonstrate that oxidative enzyme production by microbial communities responds directly to NO ₃ ⁻ deposition, controlling extracellular enzyme activity and DOC flux. The regulation of oxidative enzymes by different microbial communities in response to NO ₃ ⁻ deposition highlights the fact that the composition and function of soil microbial communities directly control ecosystem-level responses to environmental change.     

Primary production dynamics of dominant hydrophytes in Lake Provala (Serbia)

The objective of this investigation was to analyze the primary production of the dominant hydrophytes by monitoring levels of organic matter and organic carbon and estimating photosynthetic potential via the total chlorophyll content. The survey was conducted in Lake Provala (Serbia) throughout the peak vegetation period of the year 2000. The contents of organic matter and organic carbon for Myriophyllum spicatum L. were 105.11 g m⁻² and 73.66 g m⁻², Nymphoides peltata (Gmel.) Kunt. were 95.51 g m⁻² and 45.26 g m⁻² and Ceratophyllum demersum L. were 52.17 g m⁻² and 29.75 g m⁻². Chlorophyll A (Chl a) and chlorophyll A+B (Chl a+b) pigments ranged from 1.54 mg g⁻¹(Chl a) and 2.1 mg g⁻¹(Chl a+b) in M. spicatum to 5.27 mg g⁻¹(Chl a) and 7.53 mg g⁻¹(Chl a+b) in C. demersum. At full leaf out, the latter aquatic plants exceeded 50% cover of the open water surface. All species achieved maximum growth in June, but significant differences in growth dynamics were observed. At the end of the vegetation period, these plants sink to the bottom and decompose     

Seasonal growth and biomass ofTrapa japonica Flerov in Ojaga-Ike Pond,Chiba, Japan

In order to determine the seasonal growth and biomass ofTrapa japonica Flerov, field observations were carried out at Ojaga-ike Pond, Chiba, Japan, during 1979 and 1980. In spring, the plant showed exponential growth (c. 0.080 g g⁻¹ day⁻¹) and shoot elongation was as rapid as 10 cm day⁻¹. The plant attained its maximum biomass (380.5±35.1 g m⁻²) in late August, and about 50% of this was concentrated in the topmost 30-cm stratum (645.7±33.1 g m⁻³); maximum total stem length exceeded 6m. The plant produced large (500–800 mg per fruit), but small numbers of nut-like fruit (maximum, 5 fruits per rosette). Defoliation occurred almost linearly with time at a rate of 30.6 leaves m⁻² day⁻¹; annual net leaf production was estimated to be about twice as large as the seasonal maximum leaf biomass. While the number of leaves per rosette showed moderate seasonal change, rosette density, rosette area and leaf dry weight changed considerably during the year. From the negative log-log correlation between mean total leaf dry weight per rosette and rosette density, density-dependent rosette growth was assumed. The cause of the wide spread of this species in aquatic habitats is briefly discussed in terms of its seed size and morphology.     

Activity of surface-casting earthworms in a calcareous grassland under elevated atmospheric CO2

Earthworms make up the dominant fraction of the biomass of soil animals in most temperate grasslands and have important effects on the structure and function of these ecosystems. We hypothesized that the effects of elevated atmospheric CO₂ on soil moisture and plant biomass production would increase earthworm activity, expressed as surface cast production. Using a screen-aided CO₂ control facility (open top and open bottom rings), eight 1.2-m² grassland plots in Switzerland have been maintained since March 1994 at ambient CO₂ concentrations (350 μl CO₂ l⁻¹) and eight at elevated CO₂ (610 μl CO₂ l⁻¹). Cumulative earthworm surface cast production measured 40 times over 1 year (April 1995–April 1996) in plots treated with elevated CO₂ (2206 g dry mass m⁻² year⁻¹) was 35% greater (P<0.05) than that measured in plant communities maintained at ambient CO₂ (1633 g dry mass m⁻² year⁻¹). At these rates of surface cast production, worms would require about 100 years to egest the equivalent of the amount of soil now found in the Ah horizon (top 15 cm) under current ambient CO₂ concentrations, and 75 years under elevated CO₂. Elevated atmospheric CO₂ had no influence on the seasonality of earthworm activity. Cumulative surface cast production measured over the 7-week period immediately following the 6-week summer dry period in 1995 (no surface casting) was positively correlated (P<0.05) with the mean soil water content calculated over this dry and subsequent wetter period, when viewed across all treatments. However, no correlations were observed with soil temperature or with annual aboveground plant biomass productivity. No CO₂-related differences were observed in total nitrogen (N_tot) and organic carbon (C_org) concentration of surface casts, although concentrations of both elements varied seasonally. The CO₂-induced increase in earthworm surface casting activity corresponded to a 30% increase of the amount of N_tot (8.9 mg N m⁻² vs. 6.9 mg N m⁻²) and C_org (126 mg C m⁻² vs. 94 mg C m⁻²) egested by the worms in one year. Thus, our results demonstrate an important indirect stimulatory effect of elevated atmospheric CO₂ on earthworm activity which may have profound effects on ecosystem function and plant community structure in the long term. Received: 3 November 1996 / Accepted: 11 January 1997