The effect of sunlight on leaf litter quality reduces growth of the shredder Klapopteryx kuscheli

• 1 The direct effect of sunlight on the conditioning, breakdown and incorporation of leaf litter in stream food webs has not yet been considered. The aim here was to evaluate the effects of light intensity on the colonization of leaf litter by microorganisms and its resulting quality as food for the stonefly shredder Klapopteryx kuscheli.
• 2 Leaf litter was conditioned for 2 months in an open reach of a second‐order stream in litter bags either exposed to or shaded from direct sunlight. Subsequently, we performed laboratory experiments to test larval consumption, growth, growth efficiency and feeding preference fed on both leaf litter treatments.
• 3 Leaf litter in the unshaded treatment had three times more chlorophyll‐a (Chl‐a) than that in the shaded treatment, 50% lower fungal biomass and similar bacterial abundance. Although larvae did not prefer either food and fed at the same rate on both leaf litter treatments, they grew twice as fast on the shade‐conditioned leaves and attained a two‐fold higher growth efficiency.
• 4 Sunlight can have significant effects on detritus‐based food webs. Riparian modification induced by human activities in forested catchments increases the potential for sunlight to influence detritus dynamics.

     

Fungal endophyte‐infected leaf litter alters in‐stream microbial communities and negatively influences aquatic fungal sporulation

Endophytes are ubiquitous plant‐associated microbes and although they have the potential to alter the decomposition of infected leaf litter, this has not been well‐studied. The endophyte Rhytisma punctatum infects the leaves of Acer macrophyllum (bigleaf maple), causing the appearance of black ‘tar spots’ that persist in senesced leaves. Other foliar fungi also cause visible damage in healthy tissues of this host plant system including an unidentified bullseye‐shaped lesion, common in western Washington. Using three treatments of endophyte infection status in leaf tissue (R. punctatum‐infected, bullseye‐infected, lesion‐free), leaf litter discs were submerged in a third‐order temperate stream using mesh litter bags and harvested periodically over two months to determine the effects of litter treatment and incubation time on litter mass loss, fungal sporulation, and microbial community colonization. Litter containing symptomatic endophyte infections (Rhytisma or bullseye) had reduced sporulation of aquatic hyphomycetes, but decomposed significantly faster than lesion‐free or bullseye‐infected litter. Using amplicon‐based sequencing, we found a significant difference in bacterial communities colonizing Rhytisma‐infected and bullseye‐infected leaf litter, a significant difference in fungal communities colonizing Rhytisma‐infected leaf litter compared to the two other treatments, and a change in both community structure and relative abundances of bacterial and fungal taxa throughout the study period. Indicator Species Analysis clarified the drivers of these community shifts at the genus level. Our results show that endophyte‐associated, in‐stream sporulation and microbial community effects are observable within one species of leaf litter.     

Effects of land use on the structure and function of leaf‐litter microbial communities in boreal streams

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Key role of streambed moisture and flash storms for microbial resistance and resilience to long‐term drought

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Resource–consumer diversity: testing the effects of leaf litter species diversity on stream macroinvertebrate communities

1. Understanding relationships between resource and consumer diversity is essential to predicting how changes in resource diversity might affect several trophic levels and overall ecosystem functioning. 2. We tested for the effects of leaf litter species diversity (i.e. litter mixing) on litter mass remaining and macroinvertebrate communities (taxon diversity, abundance and biomass) during breakdown in a detritus‐based headwater stream (North Carolina, U.S.A.). We used full‐factorial analyses of single‐ and mixed‐species litter from dominant riparian tree species with distinct leaf chemistries [red maple (Acer rubrum), tulip poplar (Liriodendron tulipifera), chestnut oak (Quercus prinus) and rhododendron (Rhododendron maximum)] to test for additivity (single‐species litter presence/absence effects) and non‐additivity (emergent effects of litter species interactions). 3. Significant non‐additive effects of litter mixing on litter mass remaining were explained by species composition, but not richness, and litter‐mixing effects were variable throughout breakdown. Specifically, small differences in observed versus expected litter mass remaining were measured on day 14; whereas observed litter mass remaining in mixed‐species leaf packs was significantly higher on day 70 and lower on day 118 than expected from data for single‐species leaf packs. 4. Litter mixing had non‐additive effects on macroinvertebrate community structure. The number of species in litter mixtures (two to four), but not litter species composition, was a significant predictor of the dominance of particular macroinvertebrates (i.e. indicator taxa) within mixed‐species packs. 5. In addition, the presence/absence of high‐ (L. tulipifera) and low‐quality (R. maximum) litter had additive effects on macroinvertebrate taxon richness, abundance and biomass. The presence of L. tulipifera litter had both positive (synergistic) and negative (antagonistic) effects on invertebrate taxon richness, that varied during breakdown but were not related to litter chemistry. In contrast, the presence/absence of L. tulipifera had a negative relationship with total macroinvertebrate biomass (due to low leaf mass remaining when L. tulipifera was present and higher condensed and hydrolysable tannins associated with leaf packs lacking L. tulipifera). Macroinvertebrate abundance was consistently lower when R. maximum was present, which was partially explained by litter chemistry [e.g., high concentrations of lignin, condensed tannins, hydrolysable tannins and total phenolics and high carbon to nutrient (N and P) ratios]. 6. The bottom‐up effects of litter species diversity on stream macroinvertebrates and litter breakdown are different, which suggests that structural attributes of macroinvertebrate communities may only partially explain the effects of litter‐mixing on organic matter processing in streams. In addition, stream macroinvertebrates colonising decomposing litter are influenced by resource diversity as well as resource availability. Broad‐scale shifts in riparian tree species composition will alter litter inputs to streams, and our results suggest that changes in the diversity and availability of terrestrial litter may alter stream food webs and organic matter processing.     

Effects of nutrient enrichment and leaf quality on the breakdown of leaves in a hardwater stream

1. The breakdown of leaf litter in streams is influenced strongly by leaf quality and the concentration of dissolved nutrients, primarily inorganic nitrogen (N) and phosphorus (P) in the water. We examined the effect of nutrient enrichment on the breakdown of three species of leaves in a hardwater, nutrient‐rich stream. The rate of microbial respiration was also measured on the decomposing leaves. 2. The breakdown rates of dogwood (Cornus stolonifera), aspen (Populus tremuloides) and birch (Betula occidentalis), k‐values of 0.0461, 0.0307 and 0.0186 day^–1, respectively, were unaffected by nutrient enrichment and generally faster than reported previously. Microbial respiration on the leaves was greater than reported previously for leaves of congeneric species. It appears that leaf breakdown in the study stream was not nutrient limited. 3. Nitrogen‐based measures of leaf quality, such as percentage N and carbon (C)/nitrogen ratio, did not correspond to measured breakdown rates among the three leaf types. The best predictors of relative breakdown rates were percentage lignin and the percentage of the total carbon that occurred as lignin. We suggest that, when leaf breakdown is not nutrient limited, measures of carbon quality (i.e. lignin‐based measures) are a better assessment of overall leaf quality than are N‐based measures. 4. Previous studies have indicated that the enzymes produced by aquatic hyphomycetes (microfungi) operate most efficiently at a basic pH and in the presence of calcium ions. The hardwater conditions (pH=8.6, total hardness > 300 mg CaCO₃ L^–1) and abundance of dissolved NO₃ and soluble reactive phosphorous (SRP) (approximately 50 μg L^–1, each) in the study stream appear to have provided conditions that resulted in a high respiration rate and rapid breakdown of leaf litter.     

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)     

Leaf‐Litter Mixtures Affect Breakdown and Macroinvertebrate Colonization Rates in a Stream Ecosystem

Previous work in terrestrial and aquatic ecosystems has suggested that the relationship between breakdown rates of leaf litter and plant species richness may change unpredictability due to non‐additive effects mediated by the presence of key‐species. By using single‐ and mixed‐species leaf bags (7 possible combinations of three litter species differing in toughness; common alder [Alnus glutinosa ], sweet chestnut [Castanea sativa ], and Spanish oak [Quercus ilex ilex ]), I tested whether leaf species diversity, measured as richness and composition, affects breakdown dynamics and macroinvertebrate colonization (abundance, richness and composition) during 90 days incubation in a stream. Decomposition rates were additive, i.e., observed decomposition rates were not different from expected ones. However, decomposition rates of individual leaf species were affected by the mixture, i.e., there were species‐specific responses to mixing litter. The invertebrate communities colonizing the mixtures were not richer and more diverse in mixtures than in single‐species leaf bags. On the opposite, mixing leaf species had a negative, non‐additive effect on rates of shredder and taxa colonization and on macroinvertebrate diversity. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Temporal and stoichiometric patterns of algal stimulation of litter-associated heterotrophic microbial activity

1. Periphyton communities associated with submerged plant detritus contain interacting autotrophic and heterotrophic microbes, and are sites of extracellular enzymatic activity. The strength and nature of these interactions might be expected to change over time as microbial communities develop on plant litter. Microbial interactions and enzymatic activity can be altered by nutrient availability, suggesting that litter stoichiometry could also affect these phenomena.
2. We grew wetland plants under ambient and nutrient-enriched conditions to generate plant litter of differing nutrient content. In two experiments, we investigated: (1) the influence of algal photosynthesis on fungal and bacterial production and the activities of four extracellular enzymes throughout a 54-day period of microbial colonisation and growth; and (2) the influence of litter stoichiometry on these relationships.
3. Ambient and nutrient-enriched standing-dead plant litter was collected and then submerged in wetland pools to allow for natural microbial colonisation and growth. Litter samples were periodically retrieved and transported to the laboratory for experiments manipulating photosynthesis using the photosystem II inhibitor DCMU (which effectively prevents algal photosynthetic activity). Algal (¹⁴C-bicarbonate), bacterial (³H-leucine), and fungal (¹⁴C-acetate) production, and β-glucosidase, β-xylosidase, leucine aminopeptidase, and phosphatase activities (MUF- or AMC-labelled fluorogenic substrates) were measured under conditions of active and inhibited algal photosynthesis.
4. Photosynthesis stimulated overall fungal and bacterial production in both experiments, although the strength of stimulation varied amongst sampling dates. Phosphatase activity was stimulated by photosynthesis during the first, but not the second, experiment. No other enzymatic responses to short-term photosynthesis manipulations were observed.
5. Microbial communities on high-nutrient litter occasionally showed increased extracellular enzyme activity, fungal growth rates, and bacterial production compared to communities on non-enriched litter, but algal and fungal production were not affected. Litter stoichiometry had no effects on fungal, bacterial, or enzymatic responses to photosynthesis, but the mean enzyme vector analysis angle (a measure of P- versus N-acquiring enzyme activity) was positively correlated to litter N:P, suggesting that elevated litter N:P led to an increase in the relative activity of P-acquiring enzymes.
6. These results supported the hypothesis that algal photosynthesis strongly influences heterotrophic microbial activity on macrophyte leaf litter, especially that of fungi, throughout microbial community development. However, the strength of this photosynthetic stimulation does not generally depend on small differences in litter nutrient content.
7. Stimulation of microbial heterotrophs by algal photosynthesis could drive diurnal shifts in periphyton community and aquatic ecosystem function, as well as linking green (photoautotroph-based) and brown (detrital-based) food webs.

     

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.     

Leaf breakdown in an Atlantic Rain Forest stream

The hypothesis of this study was that colonizers in decaying leaf litter prefer native species (Erythrina verna) to exotic ones (Eucalyptus camaldulensis and Protium heptaphyllum). Therefore, native species are expected to show higher breakdown rates, increased biomass, richness and density of invertebrate species, and increased biomass of decomposer fungi. Breakdown of leaf litter from these three species was assessed in an Atlantic Rain Forest stream. Four samples were collected during a period of 90 days and washed on a sieve to separate the invertebrates. Then, a series of leaf disks were cut to determine ash‐free dry mass and fungal biomass, and the remaining material was oven‐dried to determine the dry weight. Eucalyptus camaldulensis and E. verna showed higher breakdown rates than P. heptaphyllum, due to differences in leaf physical and chemical characteristics. The harder detritus (P. heptaphyllum) broke down more slowly than detritus with high concentrations of labile compounds (E. camaldulensis). The density of the invertebrates associated with detritus increased with time. There were no differences in density, taxonomic richness or biomass of invertebrates among the leaf types, which indicated that the invertebrates did not distinguish between exotic and native detritus. Fungal colonization varied among samples; E. camaldulensis showed the lowest ergosterol concentrations, mainly due to a high concentration of total phenolics. The detritus with the highest hardness value was colonized most slowly by fungi. These results showed that leaf breakdown in Atlantic Rain Forest streams could be affected either by changes in riparian vegetation, or by becoming more savanna‐like process due to climate change.     

Experimental shading alters leaf litter breakdown in streams of contrasting riparian canopy cover

1. Headwater stream ecosystems are primarily heterotrophic, with allochthonous organic matter being the dominant energy. However, sunlight indirectly influences ecosystem structure and functioning, affecting microbial and invertebrate consumers and, ultimately, leaf litter breakdown. We tested the effects of artificial shading on litter breakdown rates in an open‐canopy stream (high ambient light) and a closed‐canopy stream (low ambient light). We further examined the responses of invertebrate shredders and aquatic hyphomycetes to shading to disentangle the underlying effects of light availability on litter breakdown. 2. Litter breakdown was substantially slower for both fast‐decomposing (alder, Alnus glutinosa) and slow‐decomposing (beech, Fagus sylvatica) leaf litters in artificially shaded stream reaches relative to control (no artificial shading) reaches, regardless of stream type (open or closed canopy). 3. Shredder densities were higher on A. glutinosa than on F. sylvatica litter, and shading had a greater effect on reducing shredder densities associated with A. glutinosa than those associated with F. sylvatica litter in both stream types. Fungal biomass was also negatively affected by shading. Results suggest that the effects of light availability on litter breakdown rates are mediated by resource quality and consumer density. 4. Results from feeding experiments, where A. glutinosa litter incubated under ambient light or artificial shade was offered to the shredder Gammarus fossarum, suggest that experimental shading and riparian canopy openness influenced litter palatability interactively. Rates of litter consumption by G. fossarum were decreased by experimental shading in the open‐canopy stream only. 5. The results suggest that even small variations in light availability in streams can mediate substantial within‐stream heterogeneity in litter breakdown. This study provides further evidence that changes in riparian vegetation, and thus light availability, influence organic matter processing in heterotrophic stream ecosystems through multiple trophic levels.     

Small stonefly predators affect microbenthic and meiobenthic communities in stream leaf packs

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Decomposition of Eucalyptus globulus leaves and three native leaf species (Alnus glutinosa, Castanea sativa and Quercus faginea) in a Portuguese low order stream

Leaf decomposition of the exotic evergreen Eucalyptus globulus (eucalyptus), and three native deciduous tree species, Alnus glutinosa (alder), Castanea sativa (chestnut) and Quercus faginea (oak), was compared in a second order stream in Central Portugal. Changes in dry weight, nitrogen and polyphenolic compounds and microbial colonization were periodically assessed for three months.Negative exponential curves fit the leaf weight loss with time for all leaf species. Mass loss rate was in the order alder (K = 0.0161) > chestnut (K = 0.0079) > eucalyptus (K = 0.0068) > oak (K = 0.0037). Microbial colonization followed the same pattern as breakdown rates. Evidence of fungal colonization was observed in alder after 3 days in the stream, whereas it took 21 days in oak leaves to have fungal colonization. Fungal diversity was leaf species-dependent and increased with time. In all cases, percent nitrogen per unit leaf weight increased, at least, at the initial stages of decay while soluble polyphenolics (expressed as percentage per unit leaf weight) decreased rapidly in the first month of leaves immersion.Intrinsic factors such as nitrogen and polyphenolic content may explain differences in leaf decomposition. The possible incorporation of eucalyptus litter into secondary production in a reasonable time span is suggested, although community balance and structure might be affected by differences in allochthonous patterns determined by eucalyptus monocultures.     

Phagotrophic protists are a key component of microbial communities processing leaf litter under contrasting oxic conditions

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Shifts in leaf litter breakdown along a forest–pasture–urban gradient in Andean streams

Tropical montane ecosystems of the Andes are critically threatened by a rapid land‐use change which can potentially affect stream variables, aquatic communities, and ecosystem processes such as leaf litter breakdown. However, these effects have not been sufficiently investigated in the Andean region and at high altitude locations in general. Here, we studied the influence of land use (forest–pasture–urban) on stream physico‐chemical variables (e.g., water temperature, nutrient concentration, and pH), aquatic communities (macroinvertebrates and aquatic fungi) and leaf litter breakdown rates in Andean streams (southern Ecuador), and how variation in those stream physico‐chemical variables affect macroinvertebrates and fungi related to leaf litter breakdown. We found that pH, water temperature, and nutrient concentration increased along the land‐use gradient. Macroinvertebrate communities were significantly different between land uses. Shredder richness and abundance were lower in pasture than forest sites and totally absent in urban sites, and fungal richness and biomass were higher in forest sites than in pasture and urban sites. Leaf litter breakdown rates became slower as riparian land use changed from natural to anthropogenically disturbed conditions and were largely determined by pH, water temperature, phosphate concentration, fungal activity, and single species of leaf‐shredding invertebrates. Our findings provide evidence that leaf litter breakdown in Andean streams is sensitive to riparian land‐use change, with urban streams being the most affected. In addition, this study highlights the role of fungal biomass and shredder species (Phylloicus; Trichoptera and Anchytarsus; Coleoptera) on leaf litter breakdown in Andean streams and the contribution of aquatic fungi in supporting this ecosystem process when shredders are absent or present low abundance in streams affected by urbanization. Finally, we summarize important implications in terms of managing of native vegetation and riparian buffers to promote ecological integrity and functioning of tropical Andean stream ecosystems.     

天竺桂凋落叶添加对凤仙花生理特性的影响

采用盆栽试验,通过向土壤(每盆8kg)中添加0(CK)、20(L20)、40(L40)和80g·pot-1(L80)天竺桂(Cinnamomum japonicum)凋落叶,模拟其自然分解对凤仙花(Impatiens balsamina)生理特性的影响。结果显示:(1)天竺桂凋落叶处理显著增加了凤仙花现蕾前期和现蕾期叶片H2O2含量,同时显著抑制了凤仙花现蕾前期、现蕾期和盛花期超氧化物歧化酶(SOD)、过氧化物酶(POD)和过氧化氢酶(CAT)活性。(2)凋落叶添加处理显著增加了凤仙花现蕾前期抗坏血酸(ASA)含量,但现蕾期和盛花期ASA含量却显著下降。(3)添加凋落物处理显著降低了现蕾前凤仙花叶片MDA含量,但显著增加了现蕾期MDA含量,而盛花期MDA含量在L20、L80处理下显著降低,在L40处理下则显著增加。(4)天竺桂凋落叶添加对可溶性蛋白和可溶性糖两种渗透调节物质含量影响不显著。(5)凋落叶处理显著抑制了凤仙花地径生长和地上生物量积累。(6)天竺桂凋落叶中主要化感物质可能有香豆素、桉叶油醇、肉桂醛、反式石竹烯和松油醇等。研究认为,非酶促活性氧清除系统可能在凤仙花抵御化感物质氧化胁迫过程中发挥着比酶促系统更为重要的作用,其中的ASA在氧化胁迫的早期过程扮演着重要角色;天竺桂凋落叶添加处理所造成的氧化胁迫可能超过了两类活性氧清除系统的清除能力,对凤仙花膜系统造成明显伤害,导致其地径和生物量积累显著降低。     

An invasive species may be better than none: invasive signal and native noble crayfish have similar community effects

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Characterisation and reactivity continuum of dissolved organic matter in forested headwater catchments of Andean Patagonia

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