Comparison of leaf processing rates under different temperature regimes in three headwater streams

1. We studied the effects of different temperature regimes on leaf litter processing in three forested Appalachian headwater streams of different pH (mean pH = 4.2, 6.5, 7.5).
2. We compared leaf breakdown rates, microbial biomass and macroinvertebrate shredder density and biomass between two 12-week processing periods (October–January and November–February) in each stream. Leaf processing rates were calculated both as k (day^–1) and k _d (degree day^–1).
3. There were no significant differences in processing rates ( k day^–1) between the two study periods for any leaf species in any stream. The average difference in temperature between the two study periods was 175 degree days. Shredder density was significantly higher during the earlier study period on 40% of the sample dates, but shredder biomass was not significantly different between the two study periods. ATP concentration was significantly higher during the early study period for 60% of the sample dates.
4. More significant differences in these variables (shredder density and biomass, ATP concentration) were seen among the three study streams than between the two study periods. This indicates that in this study other factors, particularly stream pH, contributed more to processing rate variation than did differences in thermal regime.     

Leaf Decay Processes during and after a Supra‐Seasonal Hydrological Drought in a Temperate Lowland Stream

Climate change models for Central Europe predict hydrological drought with fragmentation into pools during periods of high litter input in numerous lowland streams, presumably affecting in‐stream leaf decay processes. To investigate this assumption, we measured physicochemical parameters, macro‐invertebrate colonization, microbial activity, and decay rates of exposed leaves during and after a supra‐seasonal drought in a German lowland stream. Microbial activity, shredder colonization and leaf decay rates during fragmentation were low, presumably caused by drought‐related environmental conditions. Microbial activity and temperature‐corrected decay rates increased after the flow resumption but not leaf mass loss and shredder colonization. During both periods, exposed leaves appeared physically unaffected suggesting strongly reduced shredder‐mediated leaf decay despite shredder presence. Our results indicate that hydrological drought can affect organisms and processes in temperate lowland streams even after flow resumption, and should be considered in climate change scenarios. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Decomposition of leaf litter exposed to simulated acid rain in a buffered lotic system

SUMMARY. 1. Exposure to simulated acid rain resulted in changes in the chemical content of riparian vegetation and terrestrial leaf litter and had significant effects on leaf litter decay rates in a well-buffered lotic ecosystem.
2. Foliar nitrogen and phosphorus decreased with decreasing pH of the simulated acid rain and microbial activity was greater on leaf litter exposed to rain of pi I 5.4 than on leaf litter exposed to pH 3.0 or pH 4.0.
3. Detritivore numbers and biomass were significantly higher on leaf litter exposed to pH 5.4, probably due to the greater palatability of the leaf substrate.
4. Decay rates of leaf litter processed in the summer months were significantly lower than decay rates of leaf litter processed during the autumn/winter due to reduction in nitrogen content and microbial respiration.     

Decay rates of autumn and spring leaf litter in a stream and effects on growth of a detritivore

SUMMARY. 1. Differences in decay rates of autumn and spring balsam poplar (Populus balsamifera L.) leaf litter input to a stream and their effects on a lotic detritivore Tipula commiscibilis Diane were investigated.
2. Autumnal leaf litter decay rates were significantly greater than spring decay rates despite higher initial quality of spring leaves. Reduced spring/summer decomposition rates were the result of decreased microbial activity and biomass, and significantly lower numbers, kinds and biomass of macroinvertebrate detritivores.
3. Growth of the detritivore Tipula commiscibilis was significantly lower when fed spring leaves indicating that they were a poorer quality food source than autumn leaves.
4. Lower numbers of detritivores coupled with reduced leaf quality resulted in lower leaf litter decay rates characteristic of spring/summer.     

Ecology of leptocerid caddisfly larvae in an intermittent South Australian stream receiving Eucalyptus litter

SUMMARY. 1. Brown Hill Creck, a small intermittent stream in dry sclernphyll forest in South Australia, flows for about 6 months during winter and spring. When flow ceases the stream dries to isolated pools which receive high summer inputs of Eucalyptus obliqua litter. Decomposition of this material in remnant pools causes extremely dark waters and depressed oxygen concentrations for up to 90 days.
2. Only two fully aquatic insect species, Lepntorussa darlingtoni and Lectrides varians (Trichoptera: Leptoceridae), persist under these conditions. Both utilize litter as food, but have different feeding strategies. The more abundant L. darlingtoni ingests organic and microbial material from leaf surfaces, whereas L. varians is a leaf shredder.
3. Larval processing budgets based on laboratory and fleid experiments indicated that L. variant has a greater impact on litter decomposition rates than L. darlingtoni . However, because of low density, low oxygen concentrations in pools during summer, and rapid flushing of litter from pools once flow recommences. L. variaus larvae process a small proportion of the total litter input.     

Leaf litter input mediates tadpole performance across forest canopy treatments

Understanding the mechanisms limiting the distributions of organisms is necessary for predicting changes in community composition along habitat gradients. In many areas of the USA, land originally cleared for agriculture has been undergoing a process of reforestation, creating a gradient of canopy cover. For small temporary wetlands, this gradient can alter abiotic conditions and influence the resource base of wetland food webs by affecting litter inputs. As distributions of amphibians and many other temporary wetland taxa correlate with canopy cover, we experimentally manipulated shade levels and litter types in pond mesocosms to explore mechanisms limiting species performance in wetlands with canopy cover. Most differences between ponds were mediated by litter type rather than direct effects of shading. Although all three amphibian species tested are open-canopy specialists, spring peepers were the only species to show decreased survival in shaded ponds. Pond litter type generally had strong effects on growth and development rates, with tadpoles of two species in grass litter ponds growing to twice the size of, and metamorphosing 7 days earlier than, those in leaf litter ponds. Contrary to our initial hypothesis, shade level and litter type showed very few significant interactions. Our results indicate that the effects of shading cannot be considered in isolation of vegetation changes in pond basins when evaluating the effects of forest succession on temporary pond communities.     

Leaf litter processing rates in a Kenyan highland stream,the Njoro River

Mass loss of leaf litter from five common riparian trees in Kenya was determined using mesh bags over a 70-day period. The study was carried out in a river which previous research has shown to be apparently devoid of invertebrate shredders. There was a five fold difference between the most rapid and slowest decay rates, but all were within the fast range identified from temperate zone trees, demonstrating high rates of microbial activity. We conclude, therefore, that the low biomass of invertebrate shredders in Kenyan highland streams cannot be explained by poor quality of the leaf litter that enters them.     

Comparison of leaf processing rates under different temperature regimes in three headwater streams

1. We studied the effects of different temperature regimes on leaf litter processing in three forested Appalachian headwater streams of different pH (mean pH = 4.2, 6.5, 7.5).
2. We compared leaf breakdown rates, microbial biomass and macroinvertebrate shredder density and biomass between two 12-week processing periods (October–January and November–February) in each stream. Leaf processing rates were calculated both as k (day^–1) and k _d (degree day^–1).
3. There were no significant differences in processing rates ( k day^–1) between the two study periods for any leaf species in any stream. The average difference in temperature between the two study periods was 175 degree days. Shredder density was significantly higher during the earlier study period on 40% of the sample dates, but shredder biomass was not significantly different between the two study periods. ATP concentration was significantly higher during the early study period for 60% of the sample dates.
4. More significant differences in these variables (shredder density and biomass, ATP concentration) were seen among the three study streams than between the two study periods. This indicates that in this study other factors, particularly stream pH, contributed more to processing rate variation than did differences in thermal regime.     

Palatability of Leaves Conditioned in Streams Affected by Mine Drainage: A Feeding Experiment with Gammarus Pulex (L.)

Both the absence of leaf shredding macroinvertebrates and low microbial activity are of major importance in determining slow and incomplete leaf decay in extremely acidic (pH<3.5) mining streams. These streams are affected by a heavy ochre deposition causing the formation of massive iron plaques on leaf surfaces that hinder microbial exploitation. An investigation was carried out to determine whether iron plaques and leaf conditioning status (acid conditioned with and without iron plaques, neutral conditioned, unconditioned) affect the feeding preference of the shredder Gammarus pulex (L.). Leaf respiration rates and fungal biomass (ergosterol contents) were measured to determine microbial colonization. Neutral conditioned leaves had significantly higher microbial colonization than acid conditioned leaves with iron plaques. Notwithstanding, leaves of both conditioning types were consumed at high rates by G. pulex. The microbial colonization had no influence on feeding preference in the experiment. It is presumed that iron adsorbed organic material caused the high palatability of leaves with iron plaques. The results indicate that the large deposits of leaves coated with iron plaques will be available to the stream food web when water quality will be restored to neutral as planed in scenarios for the future development of mining streams.     

Catchment land-use, macroinvertebrates and detritus processing in headwater streams: taxonomic richness versus function

1. We used a litter bag technique to assess the effect of catchment land-use (forest, wetland, agriculture, urban) on the processing of red maple ( Acer rubrum L.) litter in 17 streams in Maine, U.S.A. Litter processing by fungi was predicted to increase with nutrient concentrations along a gradient of land use, from relatively unmodified to highly modified. Litter processing by litter-shredding macroinvertebrates was predicted to decline along this gradient because of a decline in their taxonomic richness and biomass.
2. Land use was associated with the anticipated gradient in nutrient and macroinvertebrate attributes, and a significant relationship was found between land use and nitrate concentration. There was, however, no significant relationship between land use and soluble reactive phosphorus (SRP) concentration. Similarly, shredder taxonomic richness was significantly related to land use type, whereas shredder biomass showed no significant relationship to land use.
3. Attributes of the shredder assemblage structure and nutrient concentrations were both strong determinants of litter processing. Increasing biomass and taxonomic richness of shredders was significantly related to increasing rates of litter mass loss. Increasing concentrations of nitrate and SRP were significantly related to increasing rates of litter softening below threshold concentrations (approximately 0.20 mg NO₃-N L^–1 and 5 μg SRP L^–1).
4. The potentially additive effects of nitrate and SRP concentrations or shredder richness and biomass on litter processing rates were confounded by the lack of significant correlation between these pairs of variables. Consequently, rates of litter processing (as rates of softening or mass loss) did not vary systematically among different land use regimes.     

Copper and zinc mixtures induce shifts in microbial communities and reduce leaf litter decomposition in streams

1. To assess the impact of metal mixtures on microbial decomposition of leaf litter, we exposed leaves previously immersed in a stream to environmentally realistic concentrations of copper (Cu) and zinc (Zn) (three levels), alone and in all possible combinations. The response of the microbial community was monitored after 10, 25 and 40 days of metal exposure by examining leaf mass loss, fungal and bacterial biomass, fungal reproduction and fungal and bacterial diversity.
2. Analysis of microbial diversity, assessed by denaturing gradient gel electrophoresis and identification of fungal spores, indicated that metal exposure altered the structure of fungal and bacterial communities on decomposing leaves.
3. Exposure to metal mixtures or to the highest Cu concentration significantly reduced leaf decomposition rates and fungal reproduction, but not fungal biomass. Bacterial biomass was strongly inhibited by all metal treatments.
4. The effects of Cu and Zn mixtures on microbial decomposition of leaf litter were mostly additive, because observed effects did not differ from those expected as the sum of single metal effects. However, antagonistic effects on bacterial biomass were found in all metal combinations and on fungal reproduction in metal combinations with the highest Cu concentrations, particularly at longer exposure times.     

Fungi are involved in the effects of litter mixtures on consumption by shredders

1. Decomposition of litter mixtures in both terrestrial and aquatic ecosystems often shows non‐additive diversity effects on decomposition rate, generally interpreted in streams as a result of the feeding activity of macroinvertebrates. The extent to which fungal assemblages on mixed litter may influence consumption by macroinvertebrates remains unknown. 2. We assessed the effect of litter mixing on all possible three‐species combinations drawn from four tree species (Alnus glutinosa, Betula pendula, Juglans regia and Quercus robur) on both fungal assemblages and the rate of litter consumption by a common shredder, Gammarus fossarum. After a 9‐week inoculation in a stream, batches of leaf discs were taken from all leaf species within litter mixture combinations. Ergosterol, an indicator of fungal biomass, and the composition of fungal assemblages, assessed from the conidia released, were determined, and incubated litter offered to G. fossarum in a laboratory‐feeding experiment. 3. Mixing leaf litter species enhanced both the Simpson’s index of the fungal assemblage and the consumption of litter by G. fossarum, but had no clear effect on mycelial biomass. Specifically, consumption rates of J. regia were consistently higher for mixed‐species litter packs than for single‐species litter. In contrast, the consumption rates of B. pendula were not affected by litter mixing, because of the occurrence of both positive and negative litter‐mixing effects in different litter species combinations that counteracted each other. 4. In some litter combinations, the greater development of some fungal species (e.g. Clavariopsis aquatica) as shown by higher sporulation rates coincided with increased leaf consumption, which may have resulted from feeding preferences by G. fossarum for these fungi. 5. Where litter mixture effects on decomposition rate are mediated via shredder feeding, this could be due to indirect effects of the fungal assemblage.     

Leaf Litter Fuels Methanogenesis Throughout Decomposition in a Forested Peatland

Decomposing leaf litter is a large supply of energy and nutrients for soil microorganisms. How long decaying leaves continue to fuel anaerobic microbial activity in wetland ecosystems is poorly understood. Here, we compare leaf litter from 15 tree species with different growth forms (angiosperms and gymnosperms, deciduous, and longer life span), using litterbags positioned for up to 4 years in a forested peatland in New York State. Periodically, we incubated partially decayed residue per species with fresh soil to assess its ability to fuel microbial methane (CH₄) production and concomitant anaerobic carbon dioxide (CO₂) production. Decay rates varied by leaf type: deciduous angiosperm > evergreen gymnosperm > deciduous gymnosperm. Decay rates were slower in leaf litter with a large concentration of lignin. Soil with residue of leaves decomposed for 338 days had greater rates of CH₄ production (5.8 µmol g^?1 dry mass d^?1) than less decomposed (<0.42 µmol g^?1 dry mass d^?1) or more decomposed (2.1 µmol g^?1 dry mass d^?1) leaf residue. Species-driven differences in their ability to fuel CH₄ production were evident throughout the study, whereas concomitant rates of CO₂ production were more similar among species and declined with degree of decomposition. Methane production rates exhibited a positive correlation with pectin and the rate of pectin decomposition. This link between leaf litter decay rates, biochemical components in leaves, and microorganisms producing greenhouse gases should improve predictions of CH₄ production in wetlands.     

Shredder–collector interactions in temporary streams of western Oregon

1. Field data and results from laboratory rearing are combined to describe shredder–collector interactions in western Oregon temporary streams.
2. The composition of functional feeding groups in the temporary streams conforms to that predicted for permanent headwaters by the River Continuum Concept (RCC).
3. A distinct functional feeding group phenology was found, with shredders emerging first and collectors emerging last in the season. Scrapers were intermediate.
4. Presence of shredders enhanced collector growth in laboratory experiments ( P < 0.01; n = 12). The species reared in the study trials accounted for 67% of the shredder biomass and 76% of the collector biomass in the natural systems.
5. Field data and laboratory rearing trials both support the hypothesis of a close shredder–collector interaction in the processing of allochthonous material in the summer-dry headwaters studied.     

Leaf litter breakdown rates in boreal streams: does shredder species richness matter?

1. Leaf litter breakdown rates were assessed in 23 boreal streams of varying size (first–seventh order) in central and northern Sweden. 2. Shredders were most abundant in small streams, while shredder species richness showed a hump-shaped relationship with stream order, with most species in fourth order streams. 3. In a partial least-squares regression analysis, year, water temperature, shredder species richness and shredder abundance were those factors correlating most strongly with leaf breakdown rates. Shredder species richness was more strongly correlated with leaf litter breakdown rates than shredder abundance, and shredder biomass showed no such correlation. 4. These data suggest that shredder species richness is an important variable in terms of leaf litter dynamics in streams.     

Effects of burial on leaf litter quality,microbial conditioning and palatability to three shredder taxa

Summary 1. Heterotrophic microorganisms are crucial for mineralising leaf litter and rendering it more palatable to leaf‐shredding invertebrates. A substantial part of leaf litter entering running waters may be buried in the streambed and thus be exposed to the constraining conditions prevailing in the hyporheic zone. The fate of this buried organic matter and particularly the role of microbial conditioning in this habitat remain largely unexplored. 2. The aim of this study was to determine how the location of leaf litter within the streambed (i.e. at the surface or buried), as well as the leaf litter burial history, may affect the leaf‐associated aquatic hyphomycete communities and therefore leaf consumption by invertebrate detritivores. We tested the hypotheses that (i) burial of leaf litter would result in lower decomposition rates associated with changes in microbial assemblages compared with leaf litter at the surface and (ii) altered microbial conditioning of buried leaf litter would lead to decreased quality and palatability to their consumers, translating into lower growth rates of detritivores. 3. These hypotheses were tested experimentally in a second‐order stream where leaf‐associated microbial communities, as well as leaf litter decomposition rates, elemental composition and toughness, were compared across controlled treatments differing by their location within the streambed. We examined the effects of the diverse conditioning treatments on decaying leaf palatability to consumers through feeding trials on three shredder taxa including a freshwater amphipod, of which we also determined the growth rate. 4. Microbial leaf litter decomposition, fungal biomass and sporulation rates were reduced when leaf litter was buried in the hyporheic zone. While the total species richness of fungal assemblages was similar among treatments, the composition of fungal assemblages was affected by leaf litter burial in sediment. 5. Leaf litter burial markedly affected the food quality (especially P content) of leaf material, probably due to the changes in microbial conditioning. Leaf litter palatability to shredders was highest for leaves exposed at the sediment surface and tended to be negatively related to leaf litter toughness and C/P ratio. In addition, burial of leaf litter led to lower amphipod growth rates, which were positively correlated with leaf litter P content. 6. These results emphasise the importance of leaf colonisation by aquatic fungi in the hyporheic zone of headwater streams, where fungal conditioning of leaf litter appears particularly critical for nutrient and energy transfer to higher trophic levels.     

Assessing the influence of wildfire on leaf decomposition and macroinvertebrate communities in boreal streams using mixed-species leaf packs

1. We investigated how compositional differences in riparian leaf litter derived from burned and undisturbed forests influenced leaf breakdown and macroinvertebrate communities using experimental mixed-species leaf packs in boreal headwater streams. Leaf pack mixtures simulating leaf litter from dominant riparian woody-stem species in burned and undisturbed riparian zones were incubated in two references and two fire-disturbed streams for 5 weeks prior to measuring temperature-corrected breakdown rates and macroinvertebrate community composition, richness, and functional metrics associated with decomposers such as shredder abundance and % shredders.
2. Leaf litter breakdown rates were higher and had greater variability in streams bordered by reference riparian forests than in streams where riparian forests had been burned during a wildfire. Streams bordered by fire disturbance showed significant effects of litter mixture on decomposition rates, observed as significantly higher decomposition rates of a fire-simulated leaf mixture compared to all other mixtures.
3. Variation among sites was higher than variation among litter mixtures, especially for macroinvertebrate community composition. In general, fire-simulated leaf mixtures had greater shredder abundances and proportions, but lower overall macroinvertebrate abundance; however, the shredder abundance trend was not consistent across all leaf mixtures at each stream.
4. These results show that disturbance-driven riparian forest condition and resulting composition of leaf subsidies to streams can influence aquatic invertebrate community composition and their function as decomposers. Therefore, if one of the primary goals of modern forest management is to emulate natural disturbance patterns, boreal forest managers should adapt silvicultural practices to promote leaf litter input that would arise post-fire, thereby supporting stream invertebrate communities and their function.

     

Periphytic Photosynthetic Stimulation of Extracellular Enzyme Activity in Aquatic Microbial Communities Associated with Decaying Typha Litter

We examined the effect of light on extracellular enzyme activities of periphytic/endogenous microbial assemblages associated with decomposing litter of an emergent macrophyte Typha angustifolia within a small inland wetland in southeastern Michigan. Standing-dead Typha leaf litter was collected, placed into floating wire mesh litter baskets, and submerged in a wetland pool. Enzyme saturation assays were conducted on three occasions following litter submergence (days 9, 28, and 44) to generate saturation curves for the individual enzymes tested and to examine potential differences in enzyme saturation kinetics during microbial colonization and development. Experimental light manipulations were conducted on two occasions during microbial development (days 10 and 29). Short-term (30 min) light exposure significantly increased extracellular β-glucosidase activity of litter-associated microbial communities. Activities of β-xylosidase and leucine-aminopeptidase were not stimulated, and stimulation of phosphatase activity was variable. The exact mechanism for increased enzyme activity remains unknown, but it may have been increased pH arising from periphytic algal photosynthesis. These results suggest that extracellular enzyme activity in microbial communities colonizing natural organic substrata may be influenced by light/photosynthesis, as has previously been demonstrated for periphyton communities grown on artificial, inert substrata. Thus, light/photosynthetic mediated stimulation of extracellular enzyme activities may be a common occurrence in microbial communities associated with natural decaying plant litter in wetlands and might engender diurnal patterns in other microbial decay processes (e.g., production, organic matter decomposition, and mineralization).     

Riparian plant species loss alters trophic dynamics in detritus-based stream ecosystems

Riparian vegetation is closely connected to stream food webs through input of leaf detritus as a primary energy supply, and therefore, any alteration of plant diversity may influence aquatic ecosystem functioning. We measured leaf litter breakdown rate and associated biological parameters in mesh bags in eight headwater streams bordered either with mixed deciduous forest or with beech forest. The variety of leaf litter types in mixed forest results in higher food quality for large-particle invertebrate detritivores (‘shredders’) than in beech forest, which is dominated by a single leaf species of low quality. Breakdown rate of low quality (oak) leaf litter in coarse mesh bags was lower in beech forest streams than in mixed forest streams, a consequence of lower shredder biomass. In contrast, high quality (alder) leaf litter broke down at similar rates in both stream categories as a result of similar shredder biomass in coarse mesh bags. Microbial breakdown rate of oak and alder leaves, determined in fine mesh bags, did not differ between the stream categories. We found however aquatic hyphomycete species richness on leaf litter to positively co-vary with riparian plant species richness. Fungal species richness may enhance leaf litter breakdown rate through positive effects on resource quality for shredders. A feeding experiment established a positive relationship between fungal species richness per se and leaf litter consumption rate by an amphipod shredder (Gammarus fossarum). Our results show therefore that plant species richness may indirectly govern ecosystem functioning through complex trophic interactions. Integrating microbial diversity and trophic dynamics would considerably improve the prediction of the consequences of species loss.     

Stronger effects of litter origin on the processing of conifer than broadleaf leaves: A test of home‐field advantage of stream litter breakdown

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