Zooplankton carcasses stimulate microbial turnover of allochthonous particulate organic matter

Carbon turnover in aquatic environments is dependent on biochemical properties of organic matter (OM) and its degradability by the surrounding microbial community. Non-additive interactive effects represent a mechanism where the degradation of biochemically persistent OM is stimulated by the provision of bioavailable OM to the degrading microbial community. Whilst this is well established in terrestrial systems, whether it occurs in aquatic ecosystems remains subject to debate. We hypothesised that OM from zooplankton carcasses can stimulate the degradation of biochemically persistent leaf material, and that this effect is influenced by the daphnia:leaf OM ratio and the complexity of the degrading microbial community. Fresh Daphnia magna carcasses and ¹³C-labelled maize leaves (Zea mays) were incubated at different ratios (1:1, 1:3 and 1:5) alongside either a complex microbial community (<50 µm) or solely bacteria (<0.8 µm). ¹³C stable-isotope measurements of CO₂ analyses were combined with phospholipid fatty acids (PLFA) analysis and DNA sequencing to link metabolic activities, biomass and taxonomic composition of the microbial community. Our experiments indicated a significantly higher respiration of leaf-derived C when daphnia-derived OM was most abundant (i.e. daphnia:leaf OM ratio of 1:1). This process was stronger in a complex microbial community, including eukaryotic microorganisms, than a solely bacterial community. We concluded that non-additive interactive effects were a function of increased C–N chemodiversity and microbial complexity, with the highest net respiration to be expected when chemodiversity is high and the degrading community complex. This study indicates that identifying the interactions and processes of OM degradation is one important key for a deeper understanding of aquatic and thus global carbon cycle.Subject terms: Limnology, Biogeochemistry, Ecology, Microbiology     

Effects of atmospheric N deposition on coarse organic matter in a headwater stream

Input, storage, export potential, and system-level processing of coarse organic matter were investigated in the intermittent streams that drain the Bear Brook Watershed in Maine (BBWM). BBWM is a paired catchment study investigating ecosystem effects of atmospheric N and S deposition. We predicted that the increased N loading to the treatment catchment would elevate input of organic matter, result in higher levels of coarse organic matter biomass, and increase litter processing rates in the treatment stream relative to the reference stream. We found that the streams draining BBWM did not have statistically different coarse organic matter input, biomass, or processing rates and we found only modest differences in export potential. System-level processing rates for maple (Acer spp.) litter were similar to rates previously quantified using litterbag methods. However, system-level processing rates for American beech (Fagus grandifolia) litter were an order of magnitude faster than rates measured with litterbags. This difference was likely due to movements of these leaves from riffle/runs and pools into debris dams, rather than differences in measurements of leaf tissue processing rates between methods. Organic matter dynamics of the intermittent streams at BBWM were similar to other forested, headwater streams. Our results indicate that the long-term N manipulation experiment at BBWM has not altered input, storage or processing of coarse organic matter in the treatment stream. Physical characteristics of these stream ecosystems appear to regulate organic matter dynamics rather than differences in nutrient chemistry.     

Bacterial communities associated with benthic organic matter in headwater stream microhabitats

Bacterial communities associated with a variety of benthic detritus types were studied in three streams in the context of the chemical characteristics of the sediment material and the stream water. A cell purification assay was developed for a quantitative microscopic evaluation of bacterial community structure in detritus samples by fluorescence in situ hybridization (FISH). The efficiency of FISH with fluorescently monolabelled probes was compared with FISH with signal amplification by catalysed reporter deposition (CARD-FISH). In detritus types poor in organic carbon and nitrogen, the numbers of prokaryotes were related to the chemical characteristics of the stream water column, whereas no such relationship was found for detritus types rich in organic carbon and nitrogen. These results might help to provide criteria for the selection of detritus types for river ecosystem assessment and monitoring. The percentage of bacteria detected by FISH with monolabelled probes was correlated with the detritus total organic matter (OM). This is likely attributed to a higher ribosome content of microbial cells on substrates rich in OM. Cell detection by CARD-FISH did not show any correlation with OM content, indicating that this technique renders the results more independent from the activity state of cells. Fluorescence in situ hybridization with four group-specific probes suggested a relationship between substrate quality and the composition of the microbial assemblages on the various types of detritus. The improved protocol for cell purification and CARD-FISH may facilitate future investigations on the relationship between the riverine benthic detritus quality and microbial community composition.     

Flow extremes and benthic organic matter shape the metabolism of a headwater Mediterranean stream

1. Single‐station diel oxygen curves were used to monitor the oxygen metabolism of an intermittent, forested third‐order stream (Fuirosos) in the Mediterranean area, over a period of 22 months. Ecosystem respiration (ER) and gross primary production (GPP) were estimated and related to organic matter inputs and photosynthetically active radiation (PAR) in order to understand the effect of the riparian forest on stream metabolism. 2. Annual ER was 1690 g O₂ m^?2 year^?1 and annual GPP was 275 g O₂ m^?2 year^?1. Fuirosos was therefore a heterotrophic stream, with P : R ratios averaging 0.16. 3. GPP rates were relatively low, ranging from 0.05 to 1.9 g O₂ m^?2 day^?1. The maximum values of GPP occurred during a few weeks in spring, and ended when the riparian canopy was fully closed. The phenology of the riparian vegetation was an important determinant of light availability, and consequently, of GPP. 4. On a daily scale, light and temperature were the most important factors governing the shape of photosynthesis–irradiance (P–I) curves. Several patterns could be generalised in the P–I relationships. Hysteresis‐type curves were characteristic of late autumn and winter. Light saturation responses (that occurred at irradiances higher than 90 μE m^?2 s^?1) were characteristic of early spring. Linear responses occurred during late spring, summer and early autumn when there was no evidence of light saturation. 5. Rates of ER were high when compared with analogous streams, ranging from 0.4 to 32 g O₂ m^?2 day^?1. ER was highest in autumn 2001, when organic matter accumulations on the streambed were extremely high. By contrast, the higher discharge in autumn 2002 prevented these accumulations and caused lower ER. The Mediterranean climate, and in its effect the hydrological regime, were mainly responsible for the temporal variation in benthic organic matter, and consequently of ER.     

Colonization of fine particulate organic matter by invertebrates in a Central Florida stream

Abstract. Our study was conducted to determine whether small invertebrates of the shredder and collector feeding guilds would preferentially colonize fine particulate organic matter (FPOM) from fast-decaying species of leaves in central Florida streams. Bags of FPOM (0.2–1 mm) generated from maple, elm, sweetgum, and water oak leaves, as well as artificial Sty-rofoam, were suspended at two sites in Poley Creek, a 1st-order sand-bottomed stream. Three replicate bags of each type were collected from each site after 7, 14, and 28 d, and bacteria and invertebrates present were counted. Bacteria colonized maple and elm in significantly higher numbers than sweetgum, oak and Styrofoam. The amphipod Gammarus tigrinis accounted for 97% of all shredders in FPOM bags, and 63% of total invertebrates. Deposit-feeding chi-ronomids comprised 87% of collectors and 28% of total invertebrates. No other taxon comprised >2% of total invertebrates. Large numbers of invertebrates colonized in the first 7 d, but densities did not increase after Day 7. Both shredders and total invertebrates occurred in significantly greater densities on maple and elm than on sweetgum and oak. Amounts of FPOM remaining in bags after 28 d were significantly smaller for maple and elm (P<.05), suggesting that shredders were eating the FPOM. Collectors occurred in similar densities in all natural FPOM bags. Only a few individuals of either functional feeding group colonized Styrofoam bags, suggesting that FPOM was being colonized for its nutritional value rather than as a refuge from predation.     

Lateral input of particulate organic matter from bank slopes surpasses direct litter fall in the uppermost reaches of a headwater stream in Hokkaido, Japan

In forested streams, surrounding riparian forests provide essential supplies of organic matter to aquatic ecosystems. We focused on two pathways of particulate organic matter inputs: direct input from upper riparian forests and indirect lateral input from bank slopes, for which there are limited quantitative data. We investigated the inputs of coarse particulate organic matter (CPOM) and carbon and nitrogen in the CPOM into the uppermost reaches of a headwater stream with steep bank slopes in Hokkaido, Japan. CPOM collected by litter traps was divided into categories (e.g., leaves, twigs) and weighed. Monthly nitrogen and carbon inputs were also estimated. The annual direct input of CPOM (ash-free dry mass) was 472 g m⁻², a common value for temperate riparian forests. The annual lateral CPOM input was 353 g m⁻¹ and 941 g m⁻² when they were converted to area base. This value surpassed the direct input. Organic matter that we could not separate from inorganic sediments contributed to the total lateral input from the bank slopes (124 g m⁻¹); this organic matter contained relatively high amounts of nitrogen and carbon. At uppermost stream reaches, the bank slope would be a key factor to understanding the carbon and nitrogen pathways from the surrounding terrestrial ecosystem to the aquatic ecosystem.     

Kinetics and mechanisms of dissolved organic carbon retention in a headwater stream

Freeze-dried aqueous extracts of autumn-shed maple leaves, birch leaves, and spruce needles were added to a third-order reach of Bear Brook, New Hampshire at concentrations similar to those predicted to occur during peak leaf fall. Leachate from each species was rapidly removed from solution. With initial concentrations of added leachate of approximately 5 mgl^–1, dissolved organics (DOC) uptake ranged from 73 to 130 mg m^–2 h^–1 for the first five hours of travel downstream from the point of addition. There was no preferential removal of DOC of low molecular weight, or of monomeric carbohydrates relative to phenolics or unidentified DOC.Stream sediments and organic debris rapidly removed DOC from solution in laboratory experiments. No significant flocculation or microbial assimilation of sugar maple leachate occurred in stream water alone. Stream sediments showed small increases in respiration with addition of leaf leachate, but no increase in respiration occurred upon addition of leachate to organic debris. Abiotic adsorption due to the high concentrations of exchangeable iron and aluminium in stream sediments may be responsible for much of the rapid removal of leaf leachate observed in field experiments. Abiotic processes appear to retain DOC within the stream, thereby allowing subsequent metabolism of dissolved organic carbon by stream microflora.     

Dissolved organic matter of a low-coloured stream

SUMMARY. The dissolved organic matter (DOM) of White Clay Creek (Pennsylvania, U.S.A.), a low-coloured stream, was characterized by gel permeation chromatography, humic acid determination, compound classification analysis and gas chromatography.
Large polymers (approximate mol.wt above 5000) were virtually absent. The majority of the DOM consisted of fulvic acid-like material, of probable mol.wt less than 3000. Highly coloured material represented only a small portion of the total. Substances classifiable as phenols, carbohydrates, lipids, amino acids or proteins made up only a small fraction of the DOM. Sixteen low mol. wt compounds (carboxylic acids, amino acids, carbohydrates and an amide) were tentatively identified.
DOM concentration had a mean value of 6.4 mgC/l and showed annual fluctuations, with maxima in autumn and in late winter.     

Flood effects on invertebrates, sediments and particulate organic matter in the hyporheic zone of a gravel-bed stream

1. We investigated the effects of a flood on the fauna and physical habitat of the hyporheic zone of the Kye Burn, a fourth order gravel‐bed stream in New Zealand. 2. Freeze core hyporheic samples (to 50 cm depth) and benthic samples (to 10 cm) were taken, along with measurements of vertical hydrological gradient, before, 2 days after and 1 month after the flood (estimated return period: 1.5 years, estimated Q_max = 10.4 m³ s^?1). 3. The composition of the hyporheos differed over the three sampling occasions with fewer taxa collected immediately postflood than preflood. The equitability of the community was higher on both postflood occasions, consistent with the reduced densities of two abundant taxa (Leptophlebiidae and Copepoda). 4. Total invertebrate abundance was lower on the postflood occasions than preflood in both benthic (0–10 cm) and hyporheic (10–50 cm) sediments. Several taxa, including asellotan isopods and amphipods, recovered within 1 month of the event. Hyporheic densities of larval Hydora and nematodes did not differ among the three sampling occasions, but the water mite Pseudotryssaturus was more abundant 1 month after the flood than preflood. There was no evidence of vertical movements (to 50 cm) by any taxa in response to the flood. 5. The proportion of fine sediments (<1 mm) in the subsurface sediments (10–50 cm) increased over the three sampling occasions and median particle size declined, but sediment porosity did not change. More particulate organic matter was found in the sediments after the flood. 6. Our study provides little evidence that the hyporheic zone (to 50 cm) acted as a significant refuge during the flood event, although movements to or recolonisation from sediments deeper than 50 cm could explain the recovery of many crustacean and mite taxa within 1 month.     

Composition of riparian litter input regulates organic matter decomposition: Implications for headwater stream functioning in a managed forest landscape

Although the importance of stream condition for leaf litter decomposition has been extensively studied, little is known about how processing rates change in response to altered riparian vegetation community composition. We investigated patterns of plant litter input and decomposition across 20 boreal headwater streams that varied in proportions of riparian deciduous and coniferous trees. We measured a suite of in‐stream physical and chemical characteristics, as well as the amount and type of litter inputs from riparian vegetation, and related these to decomposition rates of native (alder, birch, and spruce) and introduced (lodgepole pine) litter species incubated in coarse‐ and fine‐mesh bags. Total litter inputs ranged more than fivefold among sites and increased with the proportion of deciduous vegetation in the riparian zone. In line with differences in initial litter quality, mean decomposition rate was highest for alder, followed by birch, spruce, and lodgepole pine (12, 55, and 68% lower rates, respectively). Further, these rates were greater in coarse‐mesh bags that allow colonization by macroinvertebrates. Variance in decomposition rate among sites for different species was best explained by different sets of environmental conditions, but litter‐input composition (i.e., quality) was overall highly important. On average, native litter decomposed faster in sites with higher‐quality litter input and (with the exception of spruce) higher concentrations of dissolved nutrients and open canopies. By contrast, lodgepole pine decomposed more rapidly in sites receiving lower‐quality litter inputs. Birch litter decomposition rate in coarse‐mesh bags was best predicted by the same environmental variables as in fine‐mesh bags, with additional positive influences of macroinvertebrate species richness. Hence, to facilitate energy turnover in boreal headwaters, forest management with focus on conifer production should aim at increasing the presence of native deciduous trees along streams, as they promote conditions that favor higher decomposition rates of terrestrial plant litter.     

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.     

The influence of dissolved nutrients and particulate organic matter quality on microbial respiration and biomass in a forest stream

1. Although dissolved nutrients and the quality of particulate organic matter (POM) influence microbial processes in aquatic systems, these factors have rarely been considered simultaneously. We manipulated dissolved nutrient concentrations and POM type in three contiguous reaches (reference, nitrogen, nitrogen + phosphorus) of a low nutrient, third‐order stream at Hubbard Brook Experimental Forest (U.S.A). In each reach we placed species of leaves (mean C : N of 68 and C : P of 2284) and wood (mean C : N of 721 and C : P of 60 654) that differed in elemental composition. We measured the respiration and biomass of microbes associated with this POM before and after nutrient addition. 2. Before nutrient addition, microbial respiration rates and biomass were higher for leaves than for wood. Respiration rates of microbes associated with wood showed a larger response to increased dissolved nutrient concentrations than respiration rates of microbes associated with leaves, suggesting that the response of microbes to increased dissolved nutrients was influenced by the quality of their substrate. 3. Overall, dissolved nutrients had strong positive effects on microbial respiration and fungal, but not bacterial, biomass, indicating that microbial respiration and fungi were nutrient limited. The concentration of nitrate in the enriched reaches was within the range of natural variation in forest streams, suggesting that natural variation in nitrate among forest streams influences carbon mineralisation and fungal biomass.     

Retention of particulate organic matter in a tropical headstream

Retention of particulate organic matter was investigated over a range of water discharges in pool and riffle zones of a tropical stream (Itaúna Stream, São Paulo, Brazil). A closed-system leaf release and capture method was used to quantify leaf retention in a 50-m reach of the stream. Instantaneous retention rates (k) were calculated by a negative exponential model and specific retention rates (k _e) computed for each 1m interval. The mean k was 0.0161±0.0101 (S.E.) and ranged from 0.005 to 0.036 during the study period. This suggests a low retention of particulate matter. The evidence provided by this study shows that there was a relationship between retention and discharge. At low discharges, greater percentages of leaves were retained in a 3-h period while high discharges resulted in low retention percentages within the studied reach. In the Itaúna Stream, no significant difference (p<0.05) was found between the k _e values of pool and riffle zones.     

Assessment of particulate organic matter in river water

Particulate organic matter in a downriver riffle of the Grand River, the largest Canadian Great Lakes tributary, was studied between June 1970 and April 1972. In winter and spring, concentrations of particulate organic matter (1.0–26.2 mg/l) varied with river flow. High summer levels (3.4–12.7 mg/l) were attributable to high autochthonous primary production. Mean chlorophyll a concentration in summer (29.8 mg/m³) was nearly 15 times higher than in winter, and 8 times the spring mean level. High algal cell counts (15,000–19,000 cells/l) also occurred in summer. Autochthonous and allochthonous contributions to the total particulate organic carbon input to the river in summer were estimated by daily organic input and river flow relationships, carbon to chlorophyll a and to pheopigments ratios. The allochthonous source accounted for 21.5% of the total organic carbon while the autochthonous contributed the remaining 78.5%. The latter included living algae (23.0%), senescent plant material (30.3%) and detritus (25.2% — including microbes). The study establishes a new approach whereby the various components of particulate organic matter in river water can be indirectly partitioned and their biomass estimated by using quantitative relationships among readily obtainable parameters of river flow, standing biomass, chlorophyll a and pheopigments.     

Daphnia performance on diets containing different combinations of high‐quality algae,heterotrophic bacteria,and allochthonous particulate organic matter

1. Filter‐feeding zooplankton in lakes feed on a mixture of phytoplankton, bacteria, and terrestrial particles and the proportions and nutritional value of these components can be highly variable. However, the extent to which food quality interacts with food quantity in affecting overall zooplankton performance is not yet fully resolved.
2. Here we performed laboratory feeding experiments to test how the performance of the unselective filter feeder Daphnia galeata was affected if various quantities of high‐quality food (the phytoplankton Rhodomonas) were diluted with low‐quality food such as heterotrophic bacteria (Pseudomonas) or terrestrial detritus particles (t‐POM) from the riparian zone of a boreal forest stream. We hypothesised: that increased proportions of bacteria and t‐POM in the diet will lead to decreased survival, somatic growth; and reproduction of Daphnia despite the presence of phytoplankton; that these effects are more pronounced for t‐POM than for heterotrophic bacteria; and that this response is stronger when phytoplankton availability is low.
3. Increasing the concentrations of Pseudomonas affected Daphnia survival, growth, and reproduction negatively when Rhodomonas was available at intermediate (0.37 mgC/L) and high (0.55 mgC/L) quantities. When Rhodomonas quantity was low (0.22 mgC/L), the addition of Pseudomonas generally resulted in better Daphnia performance except at very high concentrations of the bacterium relative to Rhodomonas. In contrast, the addition of t‐POM was detrimental for overall Daphnia performance at all Rhodomonas concentrations.
4. Daphnia performance was best described by a model including the interaction between food quality and quantity, with stronger negative effects on Daphnia when high‐quality food was supplemented with t‐POM than with Pseudomonas.
5. The results indicate that the ability of zooplankton to use low‐quality food is affected by the concurrent availability of high‐quality food. Furthermore, food sources that can be used but do not fulfil dietary requirements of grazers (e.g. bacteria), may still provide nutritional benefits as long as other complementary food components are available in sufficient quantities to compensate for biochemical deficiencies.
6. Therefore, we conclude that heterotrophic bacteria, but not peat layer t‐POM, can be an important component of zooplankton diets in boreal lakes, especially if the concentration of phytoplankton is low.

     

Sources of organic and inorganic carbon in a headwater stream: Evidence from carbon isotope studies

A combination of stable isotope studies and ¹⁴Cdating were used to identify the main sources andprocesses controlling streamwater DOC and TIC in atemperate non-forested watershed. ¹³Cvalues for terrestrial (–24.9 to –29.1) and aquatic(–30.5 to –33.5) plants were similar to valuesreported in the literature for similar ecosystems.¹³C values for DOC in soil solution andstreamwater were consistent with soil and terrestrialvegetation, indicating that the terrestrial ecosystemis the dominant source of aquatic DOC in thiswatershed. ¹³C values of soil atmosphereCO₂ (–17.2 to –25.2) were slightly lessnegative than would be expected for production viaaerobic soil microbial decomposition and rootrespiration. There was a close correspondence between¹³C values (–15.5 to –21.5) forstreamwater TIC and soil atmospheric CO₂ in thecentral part of the catchment where the stream drainsCO₂-rich peats. ¹⁴C dating showed thatalthough peat has been accumulating in the watershedfor at least 2700 years, DOC in soil pore water andstreamwater contains carbon of predominantly recentorigin (post-AD 1955).