Microbial growth and detritus transformations during decomposition of leaf litter in a stream

SUMMARY. 1. Despite the widely accepted importance of bacteria and fungi in degrading detritus in aquatic ecosystems there is still very little quantitative information on the abundance and dynamics of these microorganisms. Using epifluorescent microscopy, we measured the biomass of bacteria and fungi during decomposition of three types of leaf detritus. Bacterial production was determined from the rate of incorporation of ³H-thymidine into DNA.
2. The transformation of leaf carbon into dissolved organic carbon and fine particulate organic carbon was followed in order to compare the amounts of leaf material that were converted into these 'end-products' of decomposition versus the amount converted into microbial biomass.
3. The amount of microbial carbon in the leaf-detritus complex never exceeded 5.2% of the total carbon, and fungal biomass was always much greater than bacterial biomass. Despite the greater standing stock of fungi, the rapid turnover of bacteria (doubling about once per day) implies that their role in degrading leaf litter or as a food source for detritivores might be as great as for fungi.
4. Removal of microbial biomass from leaf litter may occur as release of fungal spores and consumption or shedding of bacterial biomass. Fungal spores can be a significant part of the fine particulate organic carbon released from leaf detritus and potentially represent an important food resource for filter-feeding organisms.     

Effect of primary producers on the heterotrophic metabolism of a stream biofilm

1. ,The influence of benthic algae on heterotrophic metabolism in a forested Mediterranean stream was investigated. Bacterial density and ectoenzymatic activities, as well as algal biomass (chlorophyll- a ) and metabolism (the rate of ¹⁴C incorporation), were measured during colonization over 60 days of artificial substrata (clay tiles) under light and dark conditions.
2. ,Chlorophyll- a and the rate of ¹⁴C incorporation were significantly higher in light-grown than in dark-grown biofilms. Bacterial density and ectoenzymatic activity (especially β-glucosidase) were also significantly higher in light-grown biofilms.
3. ,Regressions of chlorophyll- a and ¹⁴C incorporation values on the ectoenzymatic activities were significant. The slopes of regression lines obtained for dark-grown biofilms were significantly higher than those obtained for light -grown biofilms.
4. ,The differences in the slope (of the regression lines) between dark and light-grown biofilms suggest that the response of the heterotrophs is faster in biofilms with low algal biomass accrual and slows down when algal biomass is increased.
5. ,It is concluded that algal accumulation in the epilithic biofilm influences the use of organic matter by the heterotrophic community by increasing the amount of organic substrate available for bacteria.     

Towards a budget of leaf litter decomposition in a first-order woodland stream

To construct a budget of carbon transformations occurring during leaf decomposition, alder leaves were placed in a woodland stream, later retrieved at weekly intervals, and rates of fungal and bacterial production, microbial respiration, and release of dissolved organic matter (DOM) and fine particulate organic matter (FPOM) were determined during short laboratory incubations. Carbon dioxide was the major decomposition product, explaining 17% of the microbially mediated leaf mass loss. DOM and FPOM were also important products (5 and 3% of total mass loss, respectively), whereas carbon flow to microbial biomass was low (2%). Fungal biomass in leaves always exceeded bacterial biomass (95–99% of total microbial biomass), but production of bacteria and fungi was similar, indicating that both types of microorganisms need to be considered when examining leaf decomposition in streams. Comparison of leaf mass loss in coarse and fine mesh bags revealed, in addition, that the shredder, Gammarus pulex, had a major impact on leaf decomposition in this study.     

A Cross-System Comparison of Bacterial and Fungal Biomass in Detritus Pools of Headwater Streams

The absolute amount of microbial biomass and relative contribution of fungi and bacteria are expected to vary among types of organic matter (OM) within a stream and will vary among streams because of differences in organic matter quality and quantity. Common types of benthic detritus [leaves, small wood, and fine benthic organic matter (FBOM)] were sampled in 9 small (1st-3rd order) streams selected to represent a range of important controlling factors such as surrounding vegetation, detritus standing stocks, and water chemistry. Direct counts of bacteria and measurements of ergosterol (a fungal sterol) were used to describe variation in bacterial and fungal biomass. There were significant differences in bacterial abundance among types of organic matter with higher densities per unit mass of organic matter on fine particles relative to either leaves or wood surfaces. In contrast, ergosterol concentrations were significantly greater on leaves and wood, confirming the predominance of fungal biomass in these larger size classes. In general, bacterial abundance per unit organic matter was less variable than fungal biomass, suggesting bacteria will be a more predictable component of stream microbial communities. For 7 of the 9 streams, the standing stock of fine benthic organic matter was large enough that habitat-weighted reach-scale bacterial biomass was equal to or greater than fungal biomass. The quantities of leaves and small wood varied among streams such that the relative contribution of reach-scale fungal biomass ranged from 10% to as much as 90% of microbial biomass. Ergosterol concentrations were positively associated with substrate C:N ratio while bacterial abundance was negatively correlated with C:N. Both these relationships are confounded by particle size, i.e., leaves and wood had higher C:N than fine benthic organic matter. There was a weak positive relationship between bacterial abundance and streamwater soluble reactive phosphorus concentration, but no apparent pattern between either bacteria or fungi and streamwater dissolved inorganic nitrogen. The variation in microbial biomass per unit organic matter and the relative abundance of different types of organic matter contributed equally to driving differences in total microbial biomass at the reach scale.     

Fungal and Bacterial Colonization of Submerged Leaf Litter in a Mediterranean Stream

Microbial colonization dynamics of fungi and bacteria were analyzed in an intermittent Mediterranean forested stream using two different leaf substrata (Platanus acerifolia and Populus nigra). Results showed that fungal and bacterial biomass accumulation was stimulated on both leaves due to a flooding episode that increased dissolved inorganic nitrogen (DIN) and dissolved oxygen (DO) availability in the stream water. Leaf mass loss coincided with the parallel increase in microbial biomass and extracellular enzymatic activities after the flood event. Differences in litter quality favoured bacterial biomass accumulation and β‐glucosidase and cellobiohydrolase enzymatic activities in the soft Populus species. Microbial heterotrophs colonization of submerged leaf litter and organic matter use in Mediterranean‐type streams are modulated by environmental conditions, especially the hydrological variability. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Influence of Algal Biomass on Extracellular Enzyme Activity in River Biofilms

Abstract Relationships between biofilm structural components (algal and bacterial biomass) and the activities of some extracellular enzymes that contribute to the ability to degrade organic matter) were explored for six Atlantic and three Mediterranean streams and rivers. The biofilms in these fluvial systems accounted for a wide range of bacterial and algal biomass and colonized the most common benthic habitats. Ratio of bacteria/algae biomass was lower in Atlantic than in Mediterranean streams, but enzymatic activities (β-glucosidase, β-xylosidase, phosphatase) were in general greater in the Mediterranean stream biofilms. Climatic characteristics (especially temperature) may explain the differences in enzymatic activities between biofilms of similar structure but different flow regime. The ratio β-xylosidase: β-glucosidase was similar (around 0.5) for all streams and substrata considered, showing that there is a general higher utilization of cellobiosic than xylobiosic molecules in fluvial systems. In general, highly heterotrophic biofilms showed lower extracellular enzymatic activities than more autotrophic biofilms. Maximum enzymatic activity is achieved when the algal biomass is two- to threefold higher than the bacterial biomass. The relevance of algal biomass on the heterotrophic ability of biofilms may be related to the physical proximity between the two, but also to the high proportion of polymeric carbohydrates included in algal exudates and lysis products, whose use is enzyme-mediated. Received: January 2000; Accepted: April 2000; Online Publication: 18 July 2000     

Effects of light and nutrients on the net accumulation and elemental composition of epilithon in boreal lakes

1. Two experiments in the Experimental Lakes Area (ELA) in north-western Ontario, Canada examined the effects of light and two key elements on the net accumulation and elemental composition of epilithon. In Lake (L) 224, benthic algae were grown under different light intensity and phosphorus supply, while in L302S we provided three levels of two different carbon sources (bicarbonate and glucose) to algae colonizing nutrient-diffusing substrata. After 1 month of accumulation, we sampled biofilms for chlorophyll (chl), carbon (C), phosphorus (P) and algal C.
2. Increased C supply did not significantly affect algal growth (C or chl) or elemental composition (C/P ratios) in L302S. However, P enrichment increased chl and algal C, dramatically reduced the C/P ratio of epilithon, and did not affect total organic C in L224. Phosphorus enrichment also increased the proportion of algal material in the total particulate organic matter and altered the taxonomic composition of algae in L224 biofilms. Shading had no significant effect on the C/P ratio and total organic C in epilithon from the L224 experiment.
3. Our results demonstrate that P supply affects the elemental composition of organic matter that collects on rock substrata. It thus appears that low availability of P relative to C and light drives the formation and retention of high C/P organic matter on rock surfaces in oligotrophic boreal lakes.     

Scales of observation of riparian forests and distributions of suspended detritus in a prairie river

1. Detrital inputs from riparian forests can provide the main source of energy to aquatic consumers in stream communities. However, the supply of coarse organic detritus to stream communities is difficult to predict. Patchy riparian inputs and connectivity between reaches have complicated studies and disrupted patterns of the distribution of suspended coarse particulate organic matter within streams and rivers.
2. In this study we emphasize the importance of spatial and temporal scales in determining potential distribution of instream leaf litter. Although large pulses of detritus are transported by streams during storm flows, the main supply of benthic leaf litter used by shredders and of suspended particulate organic matter used by filter feeders is transported during prolonged periods of baseflow. The local, fine-scale distribution of this organic matter is determined by the location and continuity of leaf litter sources (riparian vegetation) and specific features of channel roughness (such as woody debris, roots and rocks).
3. Viewing riparian vegetation at several scales results in variable conclusions regarding the amount of potential source area of leaf detritus. The percentage of suspended whole leaves at sites in the Little Washita River, Oklahoma, U.S.A. was best explained by the percentage of riparian forest cover in 500 m and 1000 m reaches upstream of the sites, as viewed by remote sensing imagery. The amount of leaf fragments was best explained by distance downstream along the longitudinal gradient. Ash-free dry mass of suspended coarse particulate matter did not correlate with any measures of riparian cover.
4. Our results suggest that leaves originate over longer reach lengths than those generally considered as source areas. Scale is an important consideration in studies of riparian patterns and related instream processes because of the need to integrate point dynamics as well as upstream influences.     

Scales of observation of riparian forests and distributions of suspended detritus in a prairie river

1. Detrital inputs from riparian forests can provide the main source of energy to aquatic consumers in stream communities. However, the supply of coarse organic detritus to stream communities is difficult to predict. Patchy riparian inputs and connectivity between reaches have complicated studies and disrupted patterns of the distribution of suspended coarse particulate organic matter within streams and rivers.
2. In this study we emphasize the importance of spatial and temporal scales in determining potential distribution of instream leaf litter. Although large pulses of detritus are transported by streams during storm flows, the main supply of benthic leaf litter used by shredders and of suspended particulate organic matter used by filter feeders is transported during prolonged periods of baseflow. The local, fine-scale distribution of this organic matter is determined by the location and continuity of leaf litter sources (riparian vegetation) and specific features of channel roughness (such as woody debris, roots and rocks).
3. Viewing riparian vegetation at several scales results in variable conclusions regarding the amount of potential source area of leaf detritus. The percentage of suspended whole leaves at sites in the Little Washita River, Oklahoma, U.S.A. was best explained by the percentage of riparian forest cover in 500 m and 1000 m reaches upstream of the sites, as viewed by remote sensing imagery. The amount of leaf fragments was best explained by distance downstream along the longitudinal gradient. Ash-free dry mass of suspended coarse particulate matter did not correlate with any measures of riparian cover.
4. Our results suggest that leaves originate over longer reach lengths than those generally considered as source areas. Scale is an important consideration in studies of riparian patterns and related instream processes because of the need to integrate point dynamics as well as upstream influences.     

Leaf litter decomposition and microbial activity in nutrient-enriched and unaltered reaches of a headwater stream

SUMMARY 1. Decomposition of red maple ( Acer rubrum ) and rhododendron ( Rhododendron maximum ) leaves and activity of associated microorganisms were compared in two reaches of a headwater stream in Coweeta Hydrologic Laboratory, NC, U.S.A. The downstream reach was enriched with ammonium, nitrate, and phosphate whereas the upstream reach was not altered.
2. Decomposition rate, microbial respiration, fungal and bacterial biomass, and the sporulation rate of aquatic hyphomycetes associated with decomposing leaf material were significantly higher for both leaf types in the nutrient-enriched reach. Species richness and community structure of aquatic hyphomycetes also exhibited considerable changes with an increase in the number of fungal codominants in the nutrient-enriched reach.
3. Fungal biomass was one to two orders of magnitude greater than bacterial biomass in both reaches. Changes in microbial respiration rate corresponded to those in fungal biomass and sporulation, suggesting a primary role of fungi in leaf decomposition.
4. Nutrient enrichment increased microbial activity, the proportion of leaf carbon channelled through the microbial compartment and the decomposition rate of leaf litter.     

Importance of Saprotrophic Freshwater Fungi for Pollen Degradation

Fungi and bacteria are the major organic matter (OM) decomposers in aquatic ecosystems. While bacteria are regarded as primary mineralizers in the pelagic zone of lakes and oceans, fungi dominate OM decomposition in streams and wetlands. Recent findings indicate that fungal communities are also active in lakes, but little is known about their diversity and interactions with bacteria. Therefore, the decomposer niche overlap of saprotrophic fungi and bacteria was studied on pollen (as a seasonally recurring source of fine particulate OM) by performing microcosm experiments with three different lake types. Special emphasis was placed on analysis of fungal community composition and diversity. We hypothesized that (I) pollen select for small saprotrophic fungi and at the same time for typical particle-associated bacteria; (II) fungal communities form specific free-living and attached sub-communities in each lake type; (III) the ratio between fungi or bacteria on pollen is controlled by the lake''s chemistry. Bacteria-to-fungi ratios were determined by quantitative PCR (qPCR), and bacterial and fungal diversity were studied by clone libraries and denaturing gradient gel electrophoresis (DGGE) fingerprints. A protease assay was used to identify functional differences between treatments. For generalization, systematic differences in bacteria-to-fungi ratios were analyzed with a dataset from the nearby Baltic Sea rivers. High abundances of Chytridiomycota as well as occurrences of Cryptomycota and yeast-like fungi confirm the decomposer niche overlap of saprotrophic fungi and bacteria on pollen. As hypothesized, microbial communities consistently differed between the lake types and exhibited functional differences. Bacteria-to-fungi ratios correlated well with parameters such as organic carbon and pH. The importance of dissolved organic carbon and nitrogen for bacteria-to-fungi ratios was supported by the Baltic Sea river dataset. Our findings highlight the fact that carbon-to-nitrogen ratios may also control fungal contributions to OM decomposition in aquatic ecosystems.     

LINKING BENTHIC ALGAL BIOMASS TO STREAM SUBSTRATUM TOPOGRAPHY1

The physical properties of substrata significantly influence benthic algal development. We explored the relationships among substratum surface texture and orientation with epilithic microphytobenthic biomass accumulation at the whole‐substratum and micrometer scales. Unglazed clay tiles set at three orientations (horizontal, vertical, and 45°), and six substrata of varying surface roughness were deployed in a prairie stream for 3 weeks. Substrata were analyzed for loosely attached, adnate, and total benthic algal biomass as chl a, and confocal laser scanning microscopy was used to measure substrata microtopography (i.e., roughness, microscale slope angles, and three‐dimensional surface area). At the whole‐substratum level, vertical substrata collected significantly (P < 0.05) less algal biomass, averaging 34% and 36% less than horizontal and 45° substrata, respectively. Benthic algal biomass was also significantly less on smoother surfaces; glass averaged 29% less biomass than stream rocks. At the microscale level, benthic algal biomass was the greatest at intermediate values, peaking at a mean roughness of approximately 17 μm, a mean microscale slope of 50°, and a projected/areal surface area ratio of 2:1. The proportion of adnate algae increased with surface roughness (26% and 67% for glass and brick, respectively), suggesting that substratum type changes the efficiency of algal removal by brushing. Individual substrata and microsubstrata characteristics can have a strong effect on benthic algae development and potentially affect reach scale algal variability as mediated by geomorphology.     

Storage and decomposition of particulate organic matter along the longitudinal gradient of an agriculturally-impacted stream

An agriculturally-impacted stream in northern Idaho was examined over a two-year period to determine seasonal and longitudinal patterns of the storage and decomposition of particulate organic matter. Biomass of benthic organic matter (BOM) was considerably less than values reported in the literature for comparable, undisturbed streams. Coarse, fine, and total benthic particulate organic matter were not correlated with parameters pertaining to stream size (e.g., stream order), but were correlated with sample site and amount of litterfall. The association of BOM with site and litterfall suggests that storage of particulate organic matter is a function of local characteristics rather than stream size. Low biomass of stored organic matter is a response to the low input of terrestrially-derived organic matter resulting from removal of climax vegetation.Leaf packs of alder, Alnus sp., were placed in the stream seasonally for 30 and 60 d. While there were significant differences for months, there was no significant difference among sites for leaf packs exposed for 30 d. Significant differences were observed among both sites and months for leaf packs exposed for 60 d; however, differences among sites accounted for only 5% of the variance. The absence of differences in decomposition of organic matter along the gradient of Lapwai Creek, despite heterogeneity of the drainage basin and availability of organic matter, may be in response to the overall low biomass of stored benthic organic matter. This study demonstrates that agricultural activity can substantially influence instream heterotrophic processes through reduced availability of organic matter and can shape community structure and ecosystem dynamics of streams flowing through agricultural drainage basins.     

Food resources of shredders and other benthic macroinvertebrates in relation to shading conditions in tropical Hong Kong streams

1. Gut content analyses (GCA) of benthic macroinvertebrates, supplemented by carbon and nitrogen stable isotope analyses (SIA), were used to determine the relative contribution of leaf litter and autochthonous food sources to consumer biomass in five shaded and five unshaded streams in tropical Hong Kong. 2. Only four obligate shredders and two facultative shredders were identified out of 58 morphospecies dissected. Non‐shredder taxa consumed little (<23% food eaten) coarse particulate organic matter (CPOM) in spite of its abundance in streams, and GCA revealed that fine particulate organic matter was the major food (25–99%) of most primary consumers. 3. Stable isotope analysis results were in general agreement with the findings of GCA, and confirmed that three of the four obligate shredders had a high dependence (55–78% of assimilated carbon) on CPOM. 4. Autochthonous energy sources were important in all streams: non‐shredding primary consumers examined, which accounted for 72% of total macroinvertebrate abundance in shaded streams, derived (on average) 61% of their biomass from autochthonous foods; the equivalent values for unshaded streams were 72% (abundance) and 71% (biomass).     

Nutrient limitation of epilithic and epixylic biofilms in ten North American streams

1. Nutrient diffusing substrata were used to determine the effect of added inorganic nitrogen (N) and phosphorus (P) on the development of epilithic and epixylic biofilms in 10 North American streams. Four treatments of diffusing substrata were used: Control (agar only), N addition (0.5 m NaNO₃), P addition (0.5 m KH₂PO₄), and N + P combined (0.5 m NaNO₃ + 0.5 m KH₂PO₄). Agar surfaces were covered with glass fibre filters (for epilithon) or discs of untreated white oak wood veneer (for epixylon). 2. We found that if algae showed significant response to nutrient addition, N limitation (either N alone or N with P) was the most frequent response both on GF/F filters and on wood. Despite the low dissolved nutrient concentrations in our study streams, more than a third of the streams did not show any response to N or P addition. In fact, P was never the sole limiting nutrient for algal biofilms in this study. 3. Nutrient addition influenced algal colonisation of inorganic versus organic substrata in different ways. The presence of other biofilm constituents (e.g. fungi or bacteria) may influence whether algal biomass on wood increased in response to nutrient addition. Algae on organic and inorganic substrata responded similarly to nutrient addition in only one stream. 4. Fungal biomass on wood was nutrient limited in six of 10 study streams. N limitation of fungal biomass (with or without secondary P limitation) was most frequent, but P limitation did occur in two streams. 5. Our results show that biomass responses to nutrient addition by the heterotrophic and autotrophic components of the epixylic biofilm were different, though both experienced the same stream nutrient conditions. For algae and fungi growing on wood, limiting nutrients were rarely similar. Only three of nine streams showed the same biomass response to nutrient addition, including two that showed no significant change in biomass despite added nutrients.     

Predictable changes in trophic community structure along a spatial disturbance gradient in streams

1. We assessed insect and resource standing stocks along a spatial gradient of flood disturbance in 19 sub-alpine Swedish streams to test the prediction that change in trophic structure arises from the joint action of disturbance, which affect basal resources, and resource-control, which ties the response of the consumers to the response of the resources.
2. Trophic structure, quantified as scores of non-metric multidimensional scaling based on the biomass of insect trophic groups, changed predictably along the disturbance gradient. In early summer, predators and algae feeders decreased relative to suspension feeders with increasing disturbance; in autumn, algae feeders decreased relative to leaf feeders with increasing disturbance.
3. Across the disturbance gradient, the biomass of algae-, deposit- and leaf-feeders was principally controlled by the availability of the respective resource (algae, fine detritus and coarse detritus), while disturbance only had subsidiary effects on algae feeders in early summer.
4. Overall, patterns in trophic-group biomass along the disturbance gradient were more likely to reflect indirect effects of disturbance via impact on the resources, which reverberated to the consumers because of resource-control, rather than direct effects. In contrast with the view that stream communities are the result of stochastic colonization following disturbance events, in the study streams the trophic structure of insect assemblages is predictable and partly organized by resource-control across a broad range of disturbance conditions.     

Evidence for the use of non-detrital dissolved organic matter by microheterotrophs on plant detritus in a woodland stream

SUMMARY 1. Recent studies provide evidence for the use of exudates from living plants by epilithic microheterotrophs in streams. This study investigated the possible use of such non-detrital sources of dissolved organic matter (DOM) by streatn microheterotrophs colonizing leaf litter. Biomass of bacteria and of fungi accumulating in situ on autumn-shed leaves in flow-through troughs from which light was excluded was compared to that accumulating on leaves in troughs open to natural illumination.
2. In experiments repeated at different times of year and in different stream sections, greater biomass of microheterotrophs consistently accumulated on the leaf detritus in troughs open to natural illumination. Differences in water temperature or in grazing of leaf surfaces by macroinvertebrates could not account for these consistent differences. Further, greater microheterotroph biomass accumulated on light- and dark-incubated leaves in a stream section relatively open to sunlight, compared to corresponding leaves in a section heavily shaded by canopy and understorey vegetation.
3. These and other results suggest that, to some yet undetermined extent, detritus-associated microheterotrophs use non-detrital DOM. This conclusion is consistent with a priori predictions based on consideration of microbial energetics involved in the use of detrital versus non-detrital DOM.
4. Studies of trophic pathways in streams and other aquatic habitats have failed to assess some potentially important sources of non-detrital DOM. The ability of available techniques to assess the relative roles of detrital and non-detrital sources of DOM is evaluated, and alternative approaches to this problem are suggested.     

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

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