Microbial heterotrophic utilization of dissolved organic matter in a piedmont stream

SUMMARY.

• 1 The microbial heterotrophic utilization of dissolved organic matter (DOM) was determined experimentally in microcosms using stream water and stream-bed sediments from a third order reach of White Clay Creek (Pennsylvania, U.S.A.).
• 2 Sources of DOM for the experiments included White Clay Creek water at baseflow and stormflow and cold water extracts of jewel weed (Impatiens capensis L.) and spicebush (Linderu henzoin (L.) Blume).
• 3 The heterotrophic activity of the sediments was measured as uptake of the following: dissolved organic carbon (DOC), molecular weight fractions within the DOC pool, carbohydrates, amino acids and peptides, phenolics, and dissolved oxygen (DO), all in the overlying water.
• 4 Concentrations of adenosine triphosphate (ATP), and direct microscopic counts of bacteria were used to estimate bacterial biomass in the surface sediments.
• 5 The microcosm experiments showed that specific DOC molecular size classes and DOM functional groups were selectively removed from solution, exposure to one DOM source affected responses to a different DOM source and certain DOM sources were more readily utilized than others.
• 6 Continued exposure to a DOM source increased microbial heterotrophic activity (a condition which persisted even after removal of the DOM source for several days).
• 7 Rates of biotic DOC uptake ranged from 3.6 to 242.8 mg Cm^-2h^-1.
• 8 Indirect estimates of biosynthesis calculated from DOC and DO data ranged from 1.6 at baseflow and 2.6–61.2 at stormflow to as high as 192.6 mg C m^-2 h^-1 when the community was repeatedly exposed to enriched DOM sources.
• 9 The mean generation times of bacteria in sediments, determined from direct microscopy data, ranged from 12.5 to 46.2 h at 15°C.

     

Mineralisation of dissolved organic matter by heterotrophic stream biofilm communities in a large boreal catchment

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Dissolved Organic Matter Concentration and Quality Influences upon Structure and Function of Freshwater Microbial Communities

Past studies have suggested that the concentration and quality of dissolved organic matter (DOM) may influence microbial community structure. In this study, we cross-inoculated the bacterial communities from two streams and a dystrophic lake that varied in DOM concentration and chemistry, to yield nine fully crossed treatments. We measured dissolved organic carbon (DOC) concentration and heterotrophic microbial community productivity throughout a 72-h incubation period, characterized DOM quality by molecular weight, and determined microbial community structure at the initial and final time points. Our results indicate that all bacterial inoculate sources had similar effects upon DOC concentration and DOM quality, regardless of the DOM source. These effects included an overall decrease in DOM M _W and an initial period of DOC concentration variability between 0-24h. In contrast, microbial communities and their metabolic rates converged to profiles that reflected the DOM source upon which they were growing, regardless of the initial bacterial inoculation. The one exception was that the bacterial community from the low-concentration and low-molecular-weight DOM source exhibited a greater denaturing gradient gel electrophoresis (DGGE) band richness when grown in its own DOM source than when grown in the highest concentration and molecular weight DOM source. This treatment also exhibited a higher rate of productivity. In general, our data suggest that microbial communities are selected by the DOM sources to which they are exposed. A microbial community will utilize the low-molecular-weight (or labile) DOM sources as well as parts of the high-molecular-weight (refractory) DOM, until a community develops that can efficiently metabolize the more abundant high-molecular-weight source. This experiment examines some of the complex interactions between microbial community selection and the combined factors of DOM quality and concentration. Our data suggest that the roles of aerobic aquatic heterotrophic bacteria in carbon cycling, as well as the importance of high-molecular-weight DOM as a carbon source, may be more complex than is conventionally recognized.     

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

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Urban infrastructure influences dissolved organic matter quality and bacterial metabolism in an urban stream network

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Low-level dissolved organic carbon subsidies drive a trophic upsurge in a boreal stream

1. Energy pathways in stream food webs are often driven by allochthonous basal resources. However, allochthonous dissolved organic carbon (DOC) is generally viewed as a minor if not insignificant basal resource because much of the DOC pool comprises high molecular weight, recalcitrant compounds and is inefficiently incorporated into biomass. Nevertheless, there is increasing evidence that the relatively small, labile fraction of DOC may indeed fuel microbial activity to a level that stimulates productivity across multiple trophic levels, resulting in a trophic upsurge. Here, we tested the trophic upsurge hypothesis by subsidising the labile DOC pool of an Alaskan boreal stream that had relatively high nutrient availability but low levels of naturally occurring DOC.
2. We continuously added ecologically relevant (0.250 mg C/L, c. 10% increase above ambient bulk DOC) concentrations of labile DOC (acetate-C) for 62 days to a treatment reach that was statistically indistinguishable in its channel form and chemistry from an upstream reference reach. We measured responses of periphyton production and biomass, whole reach metabolism and nutrient uptake, benthic invertebrate abundances, and juvenile salmonid (Dolly Varden, Salvelinus malma) abundance and growth.
3. Measurements of basal ecosystem responses collectively indicated increased energy mobilization at the base of the food web in response to labile DOC addition. Periphyton bacterial production in the treatment reach was generally >1.5× reference reach values, and periphyton ash-free dry mass, chlorophyll-a, and chlorophyll-a:ash-free dry mass were all greater in the treatment reach by the end of the study. Throughout dosing, ecosystem respiration was 1.3× greater in the treatment reach and dissolved inorganic nitrogen uptake was greater in the treatment reach on eight out of nine measurements.
4. Benthic invertebrate counts, dominated by Baetis spp. and Chironomidae, were c. 4× greater after 28 dosing days and c. 8× greater after 56 days in the upstream portion of the treatment reach. Abundance generally declined with increasing distance from the dosing station. Dolly Varden fry and parr age classes were nearly 2× more abundant in the upstream portion of the treatment reach than in any section of the reference reach and also declined with increasing distance from the dosing station. Further, Dolly Varden tagged with passive integrated transponders prior to the experiment had significantly higher instantaneous growth rates in the treatment reach than those recaptured in the reference reach.
5. The strong consumer responses to small quantities of labile DOC mirrored significant treatment reach increases in basal ecosystem function and therefore demonstrated a response consistent with a trophic upsurge. Terrestrial DOC has historically been viewed as contributing little to metazoan consumers, instead modulating the influence of nutrients and being respired out of a disconnected microbial loop. Because we dosed the treatment reach with a relevant concentration of labile DOC, based on measurements in nearby peatland-draining streams, we suggest that terrestrial DOC deserves more attention as a basal resource for whole food webs, akin to nutrients fuelling green (autochthonous) pathways.

     

The formation, structure and utilization of stone surface organic layers in two New Zealand streams

SUMMARY.

• 1 The development of stone surface organic layers was investigated in dark and light experimental channels at two field sites. Layer formation was monitored by measuring organic carbon, chlorophyll-a, ATP and rates of oxygen consumption, and using scanning electron microscopy.
• 2 In the darkened forest stream channel an organic layer consisting of slime, fine particles, bacteria and fungi developed and attained maximum biomass (=0.08 mg cm^-2) in about 2 months. At the second site, channels were fed by spring water low in dissolved and particulate organic matter (DOC < 0.5 g m^-3) and no organic layer developed on stones in the dark. Organic layers grown in channels subject to natural light intensities and photoperiods were dominated by diatoms and/or filamentous algae at both sites.
• 3 Laboratory experiments carried out in enclosed, recirculating stream channels demonstrated the importance of dissolved organic matter (DOM) as a prerequisite for layer formation. Also. DOM additions in the form of leaf leachates stimulated oxygen consumption by preformed layers. Uptake by microorganisms accounted for most of the reduction in water-column DOM.
• 4 Radiotracer experiments (¹⁴C and ¹⁴⁴Ce) showed that several common stream invertebrates could feed on ‘heterotrophic’ layers. Calculated assimilation efficiencies ranged from 18% to 74% and imply that nonautotrophic components of stone surface organic layers are likely to play a significant role in carbon transfer to the benthos, particularly in small, shaded streams.

     

Land‐use effects on resource net flux rates and oxygen demand in stream sediments

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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.     

Gradient-dominated ecosystems: sources and regulatory functions of dissolved organic matter in freshwater ecosystems

The emergent wetland and littoral components of the land-water zone are functionally coupled by the amounts and types of dissolved organic matter that are released, processed, transported to, and then further processed within the recipient waters. Operational couplings and integrations in freshwater ecosystems occur along physical and metabolic gradients of a number of scales from micrometer to kilometer dimensions. The operation and turnover of the microbial communities, largely associated with surfaces, generate the metabolic foundations for material fluxes along larger-scale gradients. Because of the predominance of small, shallow freshwater bodies, most dissolved organic carbon (DOC) of lacustrine and riverine ecosystems is derived from photosynthesis of higher plants and microflora associated with detritus, including sediments, and is only augmented by photosynthesis of phytoplankton. As the dissolved organic compounds generated in the wetland and littoral interface regions move toward the open-water regions of the ecosystems, partial utilization effects a selective increase in organic recalcitrance. Even though DOC from allochthonous and from interface sources is more recalcitrant than that produced by planktonic microflora, decomposition of the much larger interface quantities imported to the pelagic zone dominates ecosystem decomposition. The observed high sustained productivity of the land-water interface zone results from extensive recycling of essential resources (nutrients, inorganic carbon) and conservation mechanisms. On the average in lakes and streams, greater than 90 percent of the decomposition in the ecosystem is by bacteria utilizing DOM from non-pelagic sources of primary productivity. In addition to direct mineralization of DOC from non-pelagic sources, many of the organic compounds function indirectly to influence metabolism. New evidence is presented to demonstrate formation of complexes between humic and fulvic organic acids and extracellular enzymes. These complexes inhibit enzyme activity and can be transported within the ecosystem. The complex can be decoupled by mild ultraviolet photolysis with regeneration of enzyme activity in displaced locations.     

Browning reduces the availability—but not the transfer—of essential fatty acids in temperate lakes

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Structure and grazing of stone surface organic layers in some acid streams of southern England

SUMMARY.

• 1 Stone surface organic layers were investigated at five sites on small, acid streams in the Ashdown Forest, southern England. Sites differed in stream water pH (means 4.3–6.6) and some other physicochemical features.
• 2 Organic layers at the stream bed surface differed between sites in structure and in the amount of organic carbon present. Algae were abundant at the sites with higher conductivity and pH, iron bacteria (mainly Leptothrix sp.) predominated in the iron-rich Broadstone stream, whereas the surface layer at the most acid site was predominantly a rather structure less organic film with few living organisms.
• 3 Amounts of organic carbon on stones buried within the stream bed for 3 months were almost identical at all sites. When viewed with the scanning electron microscope (SEM), organic layers on buried stones were rather structure less with some amorphous, floceulent material. The paucity of microflora suggests that these layers may have been formed primarily by abiotic mechanisms.
• 4 X-ray microanalysis of organic layers indicated that complexes of organic matter with the metals iron, aluminium and manganese were present.
• 5 Laboratory experiments with seven species of invertebrates showed that all were able to remove and at least partially ingest organic layers and underlying sandstone. The weight of material removed was highest on microbially impoverished layers from the most acid site, and was substantially lower where algae were abundant. 6. The role of stone surface organic layers in stream systems is discussed. They may be important sites of dissolved organic matter (DOM) uptake and transfer to the benthos. A major impact of acidity on running water could be through its effect on the structure and function of organic layers.

     

Biogeochemical processes in the groundwater discharge zone of urban streams

The influence of biogeochemical processes on nitrogen and organic matter transformation and transport was investigated for two urban streams receiving groundwater discharge during the dry summer baseflow period. A multiple lines of evidence approach involving catchment-, and stream reach-scale investigations were undertaken to describe the factors that influence pore water biogeochemical processes. At the catchment-scale gaining stream reaches were identified from water table mapping and groundwater discharge estimated to be between 0.1 and 0.8 m³ m^?2 d^?1 from baseflow analysis. Sediment temperature profiles also suggested that the high groundwater discharge limited stream water infiltration into the sediments. At the stream reach-scale, dissolved organic carbon (DOC) and dissolved organic nitrogen (DON) concentrations were higher in stream water than in groundwater. However, DOC and DON concentrations were greatest in sediment pore water. This suggests that biodegradation of sediment organic matter contributes dissolved organic matter (DOM) to the streams along with that delivered with groundwater flow. Pore water ammonium (NH₄ ⁺) was closely associated with areas of high pore water DOM concentrations and evidence of sulfate (SO₄ ^2?) reduction (low concentration and SO₄:Cl ratio). This indicates that anoxic DOM mineralization was occurring associated with SO₄ ^2? reduction. However the distribution of anoxic mineralization was limited to the center of the streambed, and was not constrained by the distribution of sediment organic matter which was higher along the banks. Lower sediment temperatures measured along the banks compared to the center suggests, at least qualitatively, that groundwater discharge is higher along the banks. Based on this evidence anoxic mineralization is influenced by groundwater residence time, and is only measurable along the center of the stream where groundwater flux rates are lower. This study therefore shows that the distribution of biogeochemical processes in stream sediments, such as anoxic mineralization, is strongly influenced by both the biogeochemical conditions and pore water residence time.     

Sources and seasonal patterns of dissolved organic matter (DOM) in the hyporheic zone

The hyporheic zone is a region underneath streambeds that integrates surface and groundwater. Although its location is central to biogeochemical linkages between the riparian zone, dissolved nutrients, and benthic biota, the seasonal quality and likely sources of dissolved organic matter (DOM) in the hyporheic zone are not well understood. To investigate DOM characteristics in the hyporheic zone, water from the surface and subsurface (at depths 20, 60, and 100 cm below the streambed) was sampled every 4 weeks from 2007 to 2008 in a third-order stream in southern Ontario. Using UV spectroscopy, measures of spectral slopes, aromaticity, and A ₂₅₄/A ₃₆₅ ratios (molecular weight) were obtained. Temporal changes in these measures were consistent with watershed processes such as shedding of leaf litter in the fall, and photochemical and biofilm influence in the spring and summer. The fluorescence index (a measure of relative DOM source) suggested that at the surface and in the downwelling zone, DOM microbial sources increased with depth in the sediment, regardless of the season. Excitation–emission matrices (EEMs) showed seasonally distinct, protein-like DOM components of bacterial origin that were stronger in the fall. Leachates from specific allochthonous DOM sources—leaf litter from Betula papyrifera (white birch) and Thuja occidentalis (white cedar)—and an autochthonous source, biofilm, were isolated and incubated with unfiltered surface water. EEMs from these leachates indicated that these sources could indeed help explain observed patterns of DOM in surface and subsurface waters. These results suggest that although DOM sources were relatively constant, biogeochemical processing within the hyporheic zone resulted a DOM pool that was temporally dynamic and altered the nature of organic matter transported downstream into lakes and rivers.     

Fluorescence characteristics and biodegradability of dissolved organic matter in forest and wetland soils from coastal temperate watersheds in southeast Alaska

Understanding how the concentration and chemical quality of dissolved organic matter (DOM) varies in soils is critical because DOM influences an array of biological, chemical, and physical processes. We used PARAFAC modeling of excitation–emission fluorescence spectroscopy, specific UV absorbance (SUVA₂₅₄) and biodegradable dissolved organic carbon (BDOC) incubations to investigate the chemical quality of DOM in soil water collected from 25 cm piezometers in four different wetland and forest soils: bog, forested wetland, fen and upland forest. There were significant differences in soil solution concentrations of dissolved organic C, N, and P, DOC:DON ratios, SUVA₂₅₄ and BDOC among the four soil types. Throughout the sampling period, average DOC concentrations in the four soil types ranged from 9–32 mg C l⁻¹ and between 23–42% of the DOC was biodegradable. Seasonal patterns in dissolved nutrient concentrations and BDOC were observed in the three wetland types suggesting strong biotic controls over DOM concentrations in wetland soils. PARAFAC modeling of excitation–emission fluorescence spectroscopy showed that protein-like fluorescence was positively correlated (r ² = 0.82; P < 0.001) with BDOC for all soil types taken together. This finding indicates that PARAFAC modeling may substantially improve the ability to predict BDOC in natural environments. Coincident measurements of DOM concentrations, BDOC and PARAFAC modeling confirmed that the four soil types contain DOM with distinct chemical properties and have unique fluorescent fingerprints. DOM inputs to streams from the four soil types therefore have the potential to alter stream biogeochemical processes differently by influencing temporal patterns in stream heterotrophic productivity.     

Nutrients associated with terrestrial dissolved organic matter drive changes in zooplankton:phytoplankton biomass ratios in an alpine lake

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Seasonal relationships between planktonic microorganisms and dissolved organic material in an alpine stream

The relationships between the abundance and activity of planktonic, heterotrophic microorganisms and the quantity and characteristics of dissolved organic carbon (DOC) in a Rocky Mountain stream were evaluated. Peak values of glucose uptake, 2.1 nmol L⁻¹ hr⁻¹, and glucose concentration, 333 nM, occurred during spring snowmelt when the water temperature was 4.0°C and the DOC concentration was greatest. The turnover time of thein situ glucose pool ranged seasonally from 40–1110 hours, with a mean of 272 hr. Seasonal uptake of³H-glucose, particulate ATP concentrations, and direct counts of microbial biomass were independent of temperature, but were positively correlated with DOC concentrations and negatively correlated with stream discharge. Heterotrophic activity in melted snow was generally low, but patchy. In the summer, planktonic heterotrophic activity and microbial biomass exhibited small-scale diel cycles which did not appear to be related to fluctuations in discharge or DOC, but could be related to the activity of benthic invertebrates. Leaf-packs placed under the snow progressively lost weight and leachable organic material during the winter, indicating that the annual litterfall in the watershed may be one source of the spring flush of DOC. These results indicate that the availability of labile DOC to the stream ecosystem is the primary control on seasonal variation in heterotrophic activity of planktonic microbial populations.     

Response of dissolved organic matter in the forest floor to long-term manipulation of litter and throughfall inputs

Dissolved organic matter (DOM) contributes to organic carbon either stored in mineral soil horizons or exported to the hydrosphere. However, the main controls of DOM dynamics are still under debate. We studied fresh leaf litter and more decomposed organic material as the main sources of DOM exported from the forest floor of a mixed beech/oak forest in Germany. In the field we doubled and excluded aboveground litter input and doubled the input of throughfall. From 1999 to 2005 we measured concentrations and fluxes of dissolved organic C and N (DOC, DON) beneath the Oi and Oe/Oa horizon. DOM composition was traced by UV and fluorescence spectroscopy. In selected DOM samples we analyzed the concentrations of phenols, pentoses and hexoses, and lignin-derived phenols by CuO oxidation. DOC and DON concentrations and fluxes almost doubled instantaneously in both horizons of the forest floor by doubling the litter input and DOC concentrations averaged 82 mg C l⁻¹ in the Oe/Oa horizon. Properties of DOM did not suggest a change of the main DOM source towards fresh litter. In turn, increasing ratios of hexoses to pentoses and a larger content of lignin-derived phenols in the Oe/Oa horizon of the Double litter plots in comparison to the Control plots indicated a priming effect: Addition of fresh litter stimulated microbial activity resulting in increased microbial production of DOM from organic material already stored in Oe/Oa horizons. Exclusion of litter input resulted in an immediate decrease in DOC concentrations and fluxes in the thin Oi horizon. In the Oe/Oa horizon DOC concentrations started to decline in the third year and were significantly smaller than those in the Control after 5 years. Properties of DOM indicated an increased proportion of microbially and throughfall derived compounds after exclusion of litter inputs. Dissolved organic N did not decrease upon litter exclusion. We assume a microbial transformation of mineral N from throughfall and N mineralization to DON. Increased amounts of throughfall resulted in almost equivalently increased DOC fluxes in the Oe/Oa horizon. However, long-term additional throughfall inputs resulted in significantly declining DOC concentrations over time. We conclude that DOM leaving the forest floor derives mainly from decomposed organic material stored in Oe/Oa horizons. Leaching of organic matter from fresh litter is of less importance. Observed effects of litter manipulations strongly depend on time and the stocks of organic matter in forest floor horizons. Long-term experiments are particularly necessary in soils/horizons with large stocks of organic matter and in studies focusing on effects of declined substrate availability. The expected increased primary production upon climate change with subsequently enhanced litter input may result in an increased production of DOM from organic soil horizons.     

Uptake of Allochthonous Dissolved Organic Matter from Soil and Salmon in Coastal Temperate Rainforest Streams

Dissolved organic matter (DOM) is an important component of aquatic food webs. We compare the uptake kinetics for NH₄–N and different fractions of DOM during soil and salmon leachate additions by evaluating the uptake of organic forms of carbon (DOC) and nitrogen (DON), and proteinaceous DOM, as measured by parallel factor (PARAFAC) modeling of DOM fluorescence. Seasonal DOM slug additions were conducted in three headwater streams draining a bog, forested wetland, and upland forest using DOM collected by leaching watershed soils. We also used DOM collected from bog soil and salmon carcasses to perform additions in the upland forest stream. DOC uptake velocity ranged from 0.010 to 0.063 mm s⁻¹ and DON uptake velocity ranged from 0.015 to 0.086 mm s⁻¹, which provides evidence for the whole-stream uptake of allochthonous DOM. These findings imply that wetlands could potentially be an important source of DOM to support stream heterotrophic production. There was no significant difference in the uptake of DOC and DON across the soil leachate additions (P > 0.05), although differential uptake of DOM fractions was observed as protein-like fluorescence was removed from the water column more efficiently than bulk DOC and DON (P < 0.05). Moreover, PARAFAC analysis of DOM fluorescence showed that protein-like fluorescence decreased downstream during all DOM additions, whereas humic-like fluorescence did not change. This differential processing in added DOM suggests slow and fast turnover pools exist for aquatic DOM. Taken together, our findings argue that DON could potentially fill a larger role in satisfying biotic N demand in oligotrophic headwater streams than previously thought. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author contributions  J.B.F. conceived of or designed study, performed research, analyzed data, contributed new methods or models, and wrote the paper. E.H. conceived of or designed study and analyzed data. R.T.E. conceived of or designed study and analyzed data. J.B.J. contributed new methods or models and analyzed data.     

Effect of Native Vegetation Loss on Stream Ecosystem Processes: Dissolved Organic Matter Composition and Export in Agricultural Landscapes

Stream and river ecosystems are dependent on energetic inputs from their watersheds and thus shifts in land use from forest cover to agriculture will affect stream community composition and function. The disruption of forest-aquatic linkages alters the organic matter resources in agricultural streams. Dissolved organic matter (DOM) is the dominant form of organic matter in aquatic ecosystems, and a microbial energy source that is important for stream respiration. The concentrations and characteristics of DOM are regulated by both terrestrial (for example, terrestrial organic matter supply) and in-stream processes (for example, microbial respiration and periphyton production) that are influenced by land management. The effects of watershed land use and topographic, soil and climatic variables on DOM quantity (dissolved organic carbon concentration and load), source (terrestrial or in-stream) and quality (composition and lability) were measured in 14 streams across an agricultural land-use gradient. DOC concentration was positively correlated with watershed pasture cover and negatively correlated with watershed relief. No watershed variables were important correlates of DOC load. Stream DOM was primarily of terrestrial origin, but DOM in agricultural streams had a greater proportion of sources from in-stream sources. This may be due to reduced connection with riparian vegetation and increased in-stream primary production. We suggest that maintaining watershed tree cover greater than 52% and ensuring less than 10% of the length of riparian corridor is cleared for pasture could minimize changes to DOM composition. This is important to avoid flow-on effects for stream ecosystem processes that are mediated by DOM. Long-term DOM monitoring will be valuable for assessing the functional impacts of land-use change.