The stoichiometry of nitrogen and phosphorus spiralling in heterotrophic and autotrophic streams

1. Nutrient spiralling provides a conceptual framework and a whole‐system approach to investigate ecosystem responses to environmental changes. We use spiralling metrics to examine how the coupling of nitrogen and phosphorus uptake varies between streams dominated by either heterotrophic (i.e. bacteria‐dominated) or autotrophic (algal‐dominated) microbial communities. 2. Algae generally exhibit greater capacity to store nutrients than bacteria because of differences in cellular structures. These differences led us to hypothesise that the uptake of N and P in heterotrophic ecosystems should have reduced stoichiometric variation in response to changes in supply N : P compared to autotrophic ecosystems when assimilation dominates nutrient uptake. 3. To test this hypothesis, we used an array of serial nutrient additions in several streams in the South Fork Eel River watershed in Northern California. In one set of experiments, N and P were added alone and simultaneously in separate experiments to two small, heterotrophic streams to assess uptake rates and interactions between nutrient cycles. In a second set of experiments, N and P were added simultaneously at a range of N : P in one heterotrophic and one autotrophic stream to assess differences in uptake responses to changes in supply N : P. 4. Results of these experiments suggest two important conclusions. First, increased N supply significantly shortened P uptake lengths, while P addition had little impact on N uptake in both streams, indicating that uptake of non‐limiting nutrients is tightly coupled to the availability of the limiting element. Second, changes in P uptake and uptake ratios (U_N : U_P) with increased supply N : P supported our hypothesis that heterotrophic streams are more homeostatic in their responses to changes in nutrient supply than autotrophic streams, suggesting that physiological controls on nutrient use scale up to influence ecosystem‐scale patterns in nutrient cycling.     

The effect of grazing intensity on phosphorus spiralling in autotropic streams

The effect of grazing on primary productivity and phosphorus cycling in autotrophic streams was studied using the snail Goniobasis clavaeformes. Snails were added to each of three replicate laboratory stream channels, receiving once-through flow of groundwater, in densities of 2.1, 3.0, and 4.2 g ash free dry mass (AFDM)/m². A fourth channel received no snails and served as an ungrazed control. Presence of snail grazers resulted in a large reduction in aufwuchs biomass, primary productivity, and biotic phosphorus uptake; a modest reduction in fine particulate organic matter (FPOM); and an increase in the fraction of stream particulate organic matter (POM) exported as seston. Although primary production and aufwuchs biomass continued to decline with increasing snail density, phosphorus uptake increased. This increased phosphorus uptake is attributed to abiotic sorption to inorganic surfaces exposed as a result of efficient removal of aufwuchs at high snail densities. Although snail densities were chosen to bracket the density measured in a natural stream, the experimental densities may result in considerably higher grazing pressure on aufwuchs due to the absence of alternate food sources (e.g., coarse particulate organic matter) usually found in natural streams. Presence of snail grazers increased the spiralling length of phosphorus, primarily by reducing aufwuchs biomass and consequently reducing uptake of phosphorus from the water. Presence of snails also increased downstream transport velocity of phosphorus bound to organic particles. These results follow the patterns predicted in a previous theoretical analysis for mildly phosphorus-limited streams.     

Benthic organic matter dynamics in Texas prairie streams

Concentrations of benthic particulate organic matter (POM) in six Texas prairie streams (2nd–4th order, intermittent and perennial) were monitored over a 20 month period to determine temporal and spatial dynamics. Benthic POM mass was highly variable, having coefficients of variation (CV) in excess of 300%. Benthic POM mass in all streams was similar with the exception of the 4th order intermittent stream which had significantly higher concentrations. Benthic POM at all sites was dominated by coarse POM (CPOM), followed by fine POM (FPOM), ultrafine POM (UPOM), and medium POM (MPOM). The dominance by CPOM is especially noteworthy in the 4th order intermittent stream where it accounted for 83% of the annual POM mass. Seasonally, benthic POM was highest in summer and lowest in the fall.     

Storage and dynamics of organic matter in different springs of small floodplain streams

The flow of groundwater through the sediment layer (underflow or hyporheic zone) of streams and at the origin of streams can influence organic matter uptake dynamics of floodplain. The River Rhône floodplain has limestone foothills. Here we studied 2 karstic and 2 interstitial springs differing by aquifer geology. Organic matter, physico-chemical conditions were compared between these springs during two seasons (from March to September 1989) and at different depths (0, –20 cm, –40 cm).Temperatures indicated large differences in underflow between springs, in their relation to the surrounding environment, and between seasons. Springs are well oxygenated, with differences between layers. Cultivated fields supply interstitial springs with nitrates, and pools are nutrient traps. DOC was heterogeneous in space and time and correlates with VFPOC. Particulate nutrients were correlated with available surface area of sediment grains. Physical conditions of each spring were prominent in determining storage and turnover of organic matter. Each spring, by its own characteristics and dynamics regulating stability and turnover, had an effect or control on storage, transport and retention of organic matter (quality, quantity). These springs offer an example of the heterogeneity, and give a view of the diversity of patches within a floodplain. The data suggest that groundwater flow of springs may be a major factor in the functioning of floodplain tributaries.     

Allochthonous litter inputs, organic matter standing stocks, and organic seston dynamics in upland Panamanian streams: potential effects of larval amphibians on organic matter dynamics

Allochthonous inputs of detritus represent an important energy source for streams in forested regions, but dynamics of these materials are not well studied in neotropical headwater streams. As part of the tropical amphibian declines in streams (TADS) project, we quantified benthic organic matter standing stocks and organic seston dynamics in four Panamanian headwater streams, two with (pre-amphibian decline) and two without (post-decline) healthy amphibian assemblages. We also measured direct litterfall and lateral litter inputs in two of these streams. Continuous litterfall and monthly benthic samples were collected for 1 year, and seston was collected 1–3 times/month for 1 year at or near baseflow. Direct litterfall was similar between the two streams examined, ranging from 934–1,137 g DM m⁻² y⁻¹. Lateral inputs were lower, ranging from 140–187 g DM m⁻¹ y⁻¹. Dead leaves (57–60%), wood (24–29%), and green leaves (8–9%) contributed most to inputs, and total inputs were generally higher during the rainy season. Annual habitat-weighted benthic organic matter standing stocks ranged from 101–171 g AFDM m⁻² across the four study reaches, with ∼4 × higher values in pools compared to erosional habitats. Total benthic organic matter (BOM) values did not change appreciably with season, but coarse particulate organic matter (CPOM, >1 mm) generally decreased and very fine particulate organic matter (VFPOM, 1.6–250 μm) generally increased during the dry season. Average annual seston concentrations ranged from 0.2–0.6 mg AFDM l⁻¹ (fine seston, <754 μm >250 μm) and 2.0–4.7 mg AFDM l⁻¹ (very fine, <250 μm >1.6 μm), with very fine particles composing 85–92% of total seston. Quality of fine seston particles in the two reaches where tadpoles were present was significantly higher (lower C/N) than the two where tadpoles had been severely reduced (P = 0.0028), suggesting that ongoing amphibian declines in this region are negatively influencing the quality of particles exported from headwaters. Compared to forested streams in other regions, these systems receive relatively high amounts of allochthonous litter inputs but have low in-stream storage. Handling editor: J. Padisak     

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.

     

Dissolved organic matter in Cumbrian lakes and streams

SUMMARY. 1. A survey of dissolved organic carbon concentration [DOC] and optical absorbance at 340 nm (A340) was carried out for thirteen lakes and twenty-three streams in Cumbria over the period May 1986 to April 1987.
2. Values of [DOC] were in the range 0-6.2 mg 1-1-1, no water having a mean [DOC] greater than 3.6mgI-1. Streamwaters showed no gross seasonal dependence in [DOC], but there was a peak in [DOC] associated with the first major rain event of autumn 1986. Summer values of [DOC] in lakes were greater than winter values, probably because of the production of DOC by phytoplankton.
3. In streams, [DOC] was weakly positively correlated (r2 <0.3) with discharge (as estimated from rainfall); A340 was more strongly correlated (0.4 < r2 < 0.6). Linear regression of mean A340 against mean [DOC] for streams gave a slope consistent with the absorptivities of aquatic humic substances, and a positive intercept on the abscissa that suggests the presence of, on average 0.6 mg 1-1 of DOC that does not absorb at 340 nm. Lake samples taken in winter gave a similar A.340/[DOC] relationship to that for streams, but in summer A340/[DOC] ratios were lower, suggesting that phytoplankton-derived [DOC] absorbs weakly or not at all at 340nm.     

Uptake of ammonium and soluble reactive phosphorus in forested streams: influence of dissolved organic matter composition

Many microbes responsible for inorganic nutrient uptake and transformation utilize dissolved organic matter (DOM) as a nutrient or energy source, but little is known about whether DOM composition is an important driver of nutrient uptake in streams. Our goal was to determine whether incorporating DOM composition metrics with other more commonly considered biological, physical, and chemical variables improved our ability to explain patterns of ammonium (\({\text{NH}}_{4}^{ + }\)–N) and soluble reactive phosphorus (SRP) uptake across 11 Lake Superior tributaries. Nutrient uptake velocities (V_f) ranged from undetectable to 14.6 mm min^?1 for \({\text{NH}}_{4}^{ + }\)–N and undetectable to 7.2 mm min^?1 for SRP. Logistic regressions suggested that DOM composition was a useful predictor of where SRP uptake occurred (4/11 sites) and \({\text{NH}}_{4}^{ + }\)–N concentration was a useful predictor of where \({\text{NH}}_{4}^{ + }\)–N uptake occurred (9/11 sites). Multiple regression analysis revealed that the best models included temperature, specific discharge, and canopy cover, and DOM composition as significant predictors of \({\text{NH}}_{4}^{ + }\)–N V_f. Partial least squares revealed fluorescence index (describing the source of aquatic fulvic acids), specific ultraviolet absorbance at 254 nm (an indicator of DOM aromaticity), temperature, and conductivity were highly influential predictors of \({\text{NH}}_{4}^{ + }\)–N V_f. Therefore, streams with higher temperatures, lower solute concentrations, more terrestrial DOM signal and greater aromaticity had greater \({\text{NH}}_{4}^{ + }\)–N V_f. Our results suggest that DOM composition may be an important, yet often overlooked, predictor of \({\text{NH}}_{4}^{ + }\)–N and SRP uptake in deciduous forest streams that should be considered along with commonly measured predictors.     

Lagrangian tracking measurements revealed the temporal dynamics of nitrogen and phosphorus spiralling in a large Japanese river

Limnology - The in-stream processing of nutrients plays an important role in the fluvial nutrient transport from lands to the ocean, but few empirical studies have addressed the temporal dynamics...     

Transport and settlement of organic matter in small streams

1. After it enters streams, terrestrially derived organic matter (OM) rapidly absorbs water. Using field and laboratory experiments, we examined how this process affected the buoyancy, settling velocity, transport distance and retention locations of four types of organic matter typically found in Pacific coastal streams (‘flexible’ red alder leaves and three ‘stiff’ particle types – Douglas‐fir needles, red cedar fronds and Douglas‐fir branch pieces). 2. Immersion in water rapidly changed the physical characteristics of alder leaves, Douglas‐fir needles and red cedar fronds, which all reached constant still‐water settling velocities after only a few days of soaking. In contrast, the settling velocity of branch pieces continued to increase for 13 days, eventually reaching much higher values than any other OM type. Dried alder leaves became negatively buoyant after only two days of immersion, while other types took substantially longer (up to 24 days) before the specific gravity of all particles was >1. 3. We released saturated OM particles in an experimental channel and found that all particle types travelled further in a fast, shallow ‘riffle’ than a slow, deep ‘pool’. Comparisons with a passive settlement null model indicated that leaves were retained more rapidly than expected in the riffle (by large protruding stones), while the three stiff particle types travelled further than expected (probably due to turbulent suspension) and were retained when they settled in deeper water between larger stones. In pools, passive settlement appeared to dominate the retention of all OM types, with leaves travelling furthest. 4. These retention patterns corresponded well with those observed when saturated OM particles collected in the field were released in two pools and two riffles in a second‐order coastal stream. 5. When the experimental channel and in‐stream data were combined, the retention rates of the three stiff OM types were closely related to calculated Rouse numbers (Rouse number = particle settling velocity/shear velocity), whereas the retention rate of alder leaves was not. This suggests that different physical mechanisms are responsible for the retention of leaves and stiff OM types in shallow streams.     

Experimental evidence quantifying the role of benthic invertebrates in organic matter dynamics of headwater streams

SUMMARY. 1. The insecticide methoxychlor was applied seasonally to one of three small headwater streams in the southern Appalachian Mountains in North Carolina, U.S.A. The initial application caused massive invertebrate drift (>1,000,000 organisms/week) and resulted in a community with few shredders and reduced abundances of most insect taxa.
2. Bacterial densities and microbial respiration rates were not affected by treatment.
3. Disruption of the invertebrate community resulted in significant reductions in leaf litter processing rates (50–74% reduction depending on leaf species) and in the amount of leaf litter processed annually (reduction of 25–28%).
4. Reductions in leaf litter processing rates resulted in significant reductions in fine particulate organic matter (FPOM) export. Declines in both concentration and total export were detectable within 1 week of treatment. Annual FPOM export was reduced to 33% of pretreatment levels. Alteration to the invertebrate community had a much greater effect on FPOM export than a severe (50–200 year) drought.
6. Course particulate organic matter (CPOM) export was not significantly influenced by treatment but was influenced by hydrologic differences among years.     

The effect of deposition of organic matter on phosphorus dynamics in experimental marine sediment systems

The effect of deposition of organic matter on phosphorus dynamics in sandy marine sediments was evaluated using an experimental system (boxcosms) and three different strategies: (1) no supply (2) one single addition (3) weekly additions of a suspension of algal cells (Phaeocystis spec.). Macrofauna (3 species, 6 individuals of each) were added to half of the boxes. Both in the case of the single and weekly additions a clear effect of increased organic matter loading on phosphorus dynamics was found. Following the organic matter addition, porewater phosphate concentrations in the upper sediment layer increased, phosphate release rates from the sediment increased by a factor 3–5 and in the boxes to which a single addition was applied NaOH-extractable phosphorus increased substantially. The increase in phosphate release rates from the sediment was attributed to mineralization of the added material and to direct release from the algal cells. No clear effect of the presence of macrofauna on sediment-water exchange of phosphate could be discovered. The macrofauna were very effective at reworking the sediment, however, as illustrated by the organic carbon profiles. It is hypothesized that the sediment-water exchange rates of phosphate were regulated by the layer of algal material which was present on the sediment surface in the fed boxes. In the boxes to which the single addition was applied porewater phosphate concentrations were lower and NaOH-extractable phosphorus was higher in the presence of macrofauna, suggesting that macrofauna can stimulate phosphate binding in the sediment.Publication no. 40 of the project Applied Scientific Research Netherlands Institute for Sea Research (BEWON)     

Suspended organic matter in fast-flowing streams in Scotland

1. Samples of suspended microscopic particles were collected from a small stream (Shelligan Burn) over a distance of 3–6 km, in conditions of low and high flow, and from a longer stream (River Almond) over a distance of 27 km during average flow. 2. In the low-flow conditions in the Shelligan Burn the commonest material was non-cellular detritus, with living diatoms, all originating from the bed of the stream, as the only other relatively abundant component. The concentration of some species of diatoms varied considerably in the course of the stream, others were more evenly distributed. Zoospores of Chlorophyceae were In much higher concentration in one short stretch of the stream, whereas filaments of the same group were fairly evenly distributed. Backwaters contained lower concentrations of certain components, presumably due to local sedimentation in the decreasing rate of flow. No clear differ-ences were found between samples from pool stations and those from nearby riffles. A sample from the mouth of a small tributary contained more of certain materials originating from its denser bankside vegetation, and less moss because of its sparse-ness on the unstable bed. Its diatom flora was different from that of the main stream, having fewer of the epilithic and epiphytic species but more of the mud-surface ones. As there was no significant downstream increase in the volume of flow the absolute abundance of particles was relative to their concentration. 3. In one flood condition in the same stream there was a downstream increase in the concentration of suspended monocotyledenous tissue and non-cellular detritus. The absolute abundance of detritus at the most downstream station was at least forty-five times greater than it was in the low-flow conditions. 4. In the River Almond the commonest materials in suspension were again non-cellular detritus and diatom cells. There was a distinct downstream increase in the concentration of the diatoms and also of Chlorophyceae. The volume of flow increased downstream so that between the most upstream and downstream stations tbe absolute abundance of diatoms increased 100 times and Chlorophyceae increased 1000 times.     

Diet and growth of a leaf-shredding caddisfly in southern Appalachian streams of contrasting disturbance history

Diet and growth of leaf-shredding caddisfly larvae, Pycnopsyche spp.,were examined in streams draining a reference catchment and a 16-year-oldclear-cut (disturbed) catchment at Coweeta Hydrologic Laboratory insouthwestern North Carolina, USA. The objective was to explain why shredderproduction is higher in the disturbed streams despite the larvae having lessfood (i.e., leaves) available. We predicted larvae would grow faster onfast-decaying leaf material representative of the disturbed streams. Larvaeconsumed mostly leaf detritus in three streams draining each catchment overthree seasons (fall, winter, and spring), which showed larvae did notconsume higher quality foods (e.g., algae and animal material) in disturbedstreams. When fed 2-month-old conditioned black birch (Betula lenta L.) (afast-decaying leaf species) and white oak (Quercus alba L.) (a slow-decayingleaf species) leaves in the laboratory, larvae grew significantly faster onthe birch leaves. However, when larvae were fed the same leaf types after3-months conditioning, larvae grew significantly faster on oak leaves. Afield growth experiment conducted for 42 d using mixed-species leaf dietsrepresentative of each catchment and initially conditioned for 2 monthsfound that Pycnopsyche grew significantly better on the diet representativeof the reference catchment. The reference diet contained more oak leaveswhich apparently became a more acceptable food as the experiment proceeded.High shredder production in the disturbed streams could not be explained byhigh Pycnopsyche growth rates on fast-decaying leaves. Instead, larvae grewbetter on leaves that were apparently conditioned optimally regardless ofconditioning rate.     

Sequential bioavailability of sedimentary organic matter to heterotrophic bacteria

Aquatic sediments harbour diverse microbial communities that mediate organic matter degradation and influence biogeochemical cycles. The pool of bioavailable carbon continuously changes as a result of abiotic processes and microbial activity. It remains unclear how microbial communities respond to heterogeneous organic matrices and how this ultimately affects heterotrophic respiration. To explore the relationships between the degradation of mixed carbon substrates and microbial activity, we incubated batches of organic‐rich sediments in a novel bioreactor (IsoCaRB) that permitted continuous observations of CO₂ production rates, as well as sequential sampling of isotopic signatures (δ¹³C, Δ¹⁴C), microbial community structure and diversity, and extracellular enzyme activity. Our results indicated that lower molecular weight (MW), labile, phytoplankton‐derived compounds were degraded first, followed by petroleum‐derived exogenous pollutants, and finally by higher MW polymeric plant material. This shift in utilization coincided with a community succession and increased extracellular enzyme activities. Thus, sequential utilization of different carbon pools induced changes at both the community and cellular level, shifting community composition, enzyme activity, respiration rates, and residual organic matter reactivity. Our results provide novel insight into the accessibility of sedimentary organic matter and demonstrate how bioavailability of natural organic substrates may affect the function and composition of heterotrophic bacterial populations.     

Controls on periphyton biomass in heterotrophic streams

1. Headwater streams of the Hubbard Brook Experimental Forest (HBEF) are typically characterised by a periphyton assemblage of low biomass and diversity. However, periphyton blooms have been observed following catchment deforestation experiments and occasionally during the annual spring thaw before canopy leaf‐out. 2. There is pronounced seasonal variation in both nutrient and light availability in HBEF streams. Stream water nitrogen (N) concentrations and light levels are higher before canopy leaf‐out and after leaf senescence and are lower during the growing season. Periphyton accrual rates also change seasonally; they are highest in spring prior to leaf‐out and significantly lower during summer and in autumn. 3. Periphyton biomass rarely responded positively to in‐situ experimental enrichment with nitrogen or phosphorus. In the summer, nutrient enrichment overall had no effect on periphyton biomass, while outside the growing season N enrichment had inhibitory effects on periphyton. 4. Despite these experimental results, surveys of ambient chlorophyll a concentrations in streams across the HBEF demonstrated no relationship between streamwater dissolved inorganic N or P concentrations and benthic chlorophyll a. 5. Our results suggest that HBEF periphyton communities are not closely regulated by nutrient availability, even during periods of high light availability. The inhibitory effects of nutrient enrichment outside the growing season are interesting, but further research is necessary to elucidate the mechanisms driving these responses.     

Microbial biofilm structure and organic matter use in mediterranean streams

River and stream biofilms in mediterranean fluvial ecosystems face both extreme seasonality as well as arrhythmic fluctuations. The hydrological extremes (droughts and floods) impose direct changes in water availability but also in the quantity and quality of organic matter and nutrients that sustain the microbial growth. This review analyzes how these ecological pulses might determine unique properties of biofilms developing in mediterranean streams. The paper brings together data from heterotrophic and autotrophic community structure, and extracellular enzyme activities in biofilms in mediterranean streams. Mediterranean stream biofilms show higher use of peptides during the favorable period for epilithic algae development (spring), and preferential use of cellulose and hemicellulose in autumn as a response to allochthonous input. The drying process causes the reduction in bacterial production and chlorophyll biomass, but the rapid recovery of both autotrophs and heterotrophs with rewetting indicates their adaptability to fluctuations. Bacteria surviving the drought are mainly associated with sediment and leaf litter which serve as “humid refuges”. Some algae and cyanobacteria show resistant strategies to cope with the drought stress. The resistance to these fluctuations is strongly linked to the streambed characteristics (e.g., sediment grain size, organic matter accumulation, nutrient content).     

Hovenia dulcis Thunb. (Rhamnaceae) invasion in the riparian zone alters the dynamics and decomposition of organic matter in subtropical streams,but not of associated invertebrate assemblages

Limnology - Stream functioning is affected by allochthonous and autochthonous energy sources, organic matter decomposition and the structure and composition of the aquatic community. The presence...     

Sources of organic nitrogen,phosphorus and carbon in antarctic streams

Dissolved and particulate organic materials were analysed in 14 streamwaters of the McMurdo Sound region of Antarctica. These streams are fed by glacial meltwaters and pass through catchments largely devoid of terrestrial vegetation. Nonetheless they contained measurable amounts of organic material in both dissolved and particulate form. Most of the dissolved organic carbon (DOC) values lay in the range 1–3 g C m^–3. Higher values were recorded close to penguin rookeries on the coast. Dissolved organic nitrogen (DON) concentrations were generally two orders of magnitude less than DOC and in flowing waters with rich blue-green algal growth DON increased with distance downstream. Dissolved organic phosphorus levels were generally much lower than DON, but highly variable. Particulate organic carbon concentrations (both fine and coarse) were unexpectedly high. Five sources of organic matter were identified: birdlife (only near the coast), autochthonous algal production (especially important for DON), streambed soils (important at first flows), lacustrine and marine sediments, through which certain streams and glaciers cut, and the glacial ice, which received organic input from wind-blown particulates, snowfall and the underlying bedrock of sedimentary origin. Highest organic levels were recorded in the first melt down the glacier face, suggesting that winter deposition of organic materials may be especially important.