Probing whole‐stream metabolism: influence of spatial heterogeneity on rate estimates

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Periodical Cicada Detritus Impacts Stream Ecosystem Metabolism

The emergence of 17-year periodical cicadas in Maryland, USA, in 2004 provided a unique opportunity to study the effect of a large, but temporally limited, resource pulse of arthropod detritus on stream ecosystem function. Cicada emergence was quantified in the forests adjacent to two small streams with different histories of riparian disturbance (Intact and Disturbed sites). We estimated the input of cicada detritus to the streams, described its retention and breakdown dynamics, and measured whole-stream respiration over the cicada flight season (May–July). Average emergence density was significantly greater at the Intact site, but average cicada detritus input rates were greater at the Disturbed site. Cicada detritus was locally retained within both streams and rapidly broke down. Daily whole-stream respiration (CR₂₄) at both sites responded dramatically to the cicada pulse, with CR₂₄ doubling pre-cicada measurements following the period of greatest cicada input (Intact: 12.82 → 23.78 g O₂ m⁻² d⁻¹; Disturbed: 2.76 → 5.77 g O₂ m⁻² d⁻¹). CR₂₄ returned to baseline levels when cicada input decreased at the Intact site, but more than doubled again at the Disturbed site (13.14 g O₂ m⁻² d⁻¹), despite a decline in cicada input rate. Differences in respiration response may be a function of differences in cicada input rates as well as differences in microbial community activity. The strong effects on stream ecosystem function exerted by a short but intense input of periodical cicada detritus may provide insights regarding the response of streams to other irregular resource pulses. HM, MP, LC, and DR conceived and designed study; HM, LC, and DR performed research; HM, LC, and DR analyzed data; HM, MP, LC, and DR wrote the paper.     

Variations of vinblastine accumulation and redox state affected by exogenous H<Subscript>2</Subscript>O<Subscript>2</Subscript> in <Emphasis Type="Italic">Catharanthus roseus</Emphasis> (L.) G. Don

Effects of exogenous H₂O₂ application on vinblastine (VBL) and its precursors, vindoline (VIN), catharanthine (CAT) and α-3′,4′-anhydrovinblastine (AVBL), were measured in Catharanthus roseus seedlings in order to explore possible correlation of VBL formation with oxidative stress. VBL accumulation has previously been shown to be regulated by an in vitro H₂O₂-dependent peroxidase (POD)-like synthase. Experimental exposure of plants to different concentrations of H₂O₂ showed that endogenous H₂O₂ and alkaloid concentrations in leaves were positively elevated. The time-course variations of alkaloid concentrations and redox state, reflected by the concentrations of H₂O₂, ascorbic acid (AA), oxidative product of glutathione (GSSG) and POD activity, were significantly altered due to H₂O₂ application. The further correlation analysis between alkaloids and redox status indicated that VBL production was tightly correlated with redox status. These results provide a new link between VBL metabolisms and redox state in C. roseus.     

Beaver dams maintain fish biodiversity by increasing habitat heterogeneity throughout a low‐gradient stream network

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Effects of dilution stress on the functioning of a saline Mediterranean stream

The effects of seasonality and dilution stress on the functioning of Rambla Salada, a hypersaline Mediterranean stream in SE Spain, were evaluated. The stream is subject to diffuse freshwater inputs from the drainage of intensively irrigated agriculture in the catchment and periodic losses of water through an irrigation channel. Metabolic rates and the biomass of primary producers and consumers were estimated over a 2-year period. During the first year several dilution events occurred, while during the second year the salinity recovery reached predisturbance levels. Functional indicators were compared in the disturbance and recovery salinity periods. Primary production and respiration rates in the Rambla Salada ranged between 0.07–21.05 and 0.19–17.39 g O₂ m⁻² day⁻¹, respectively. The mean values for these variables were 7.35 and 5.48 g O₂ m⁻² day⁻¹, respectively. Mean net daily metabolism rate was 1.87 ± 0.52 g O₂ m⁻² day⁻¹ and mean production/respiration ratio was 2.48 ± 1.1, reflecting autotrophic metabolism. The metabolic rates showed the typical seasonal pattern of Mediterranean open canopy streams. Therefore, gross primary production (GPP) and ecosystem respiration (ER) registered maximum values in summer, intermediate values in spring and autumn and minimum values in winter. The metabolic rates and biomass of consumers were greater in the disturbance period than in the recovery period. However, they did not show significant differences between periods due to their important dependence on seasonal cycle. Seasonality accounted for much of the temporal variability in GPP and ER (76% and 83% in the multiregression models, respectively). Light availability seems to be the most important factor for GPP and ER in the Rambla Salada. Autotrophic biomass responded more to variations in discharge and conductivity than to seasonal variations. In fact, it was severely affected by freshwater inputs after which the epipelic biomass decreased significantly and Cladophora glomerata proliferated rapidly. Epipelic algal biomass was the most sensitive parameter to dilution disturbance. Handling editor: Luigi Naselli-Flores     

Macrophyte diversity and composition in relation to substratum characteristics in regulated and unregulated Danish streams

1. The objective of the present study was to examine how the physical stream environment in regulated and unregulated lowland streams affects the diversity and distribution of macrophyte communities. We analysed the abundance, distribution and composition of macrophytes, together with physical parameters, in seven regulated and seven unregulated unshaded Danish stream reaches. 2. Total macrophyte coverage was similar in the regulated and unregulated streams, but species richness and Shannon diversity were higher in the unregulated streams. Overall, we found fifty-two different species in the regulated stream reaches and sixty-two in the unregulated stream reaches. The spatial distribution of macrophytes on the stream bottom was more heterogeneous in the unregulated streams. 3. We found positive correlations between the coverage and diversity of macrophytes and the coverage of coarse-textured substratum types on the stream bottom, as well as between macrophyte coverage and diversity and substratum heterogeneity. We also found that the macrophytes were more heterogeneously distributed where substratum heterogeneity was greater. 4. The species growing both submerged and emergent were more abundant in the regulated streams, whereas species growing only submerged were more abundant in the unregulated streams. Species growing submerged, species growing both submerged and emergent, and species only growing emergent segregated differently in a canonical correspondence analysis ordination. The submerged species were primarily associated with coarser-textured substrata, whereas species growing both submerged and emergent, and species growing only emergent were associated with finer-textured substrata. 5. The most abundant species in the regulated streams, Sparganium emersum, accounting for almost one-third of the total macrophyte coverage, was primarily associated with clay and sandy bottom substrata, whereas the most abundant species in the unregulated streams, Batrachium peltatum, was primarily associated with gravel and stony substrata.     

Macrophyte beds contribute disproportionately to benthic invertebrate abundance and biomass in a sand plains stream

Macrophyte beds have been shown to influence organic matter retention and nutrient processing in streams. Less is known about the extent to which plant beds contribute to abundance, biomass, and diversity of macroinvertebrate assemblages in low-order streams. We measured aquatic invertebrate abundance, biomass, and diversity associated with plant beds and sand/gravel patches in a low-gradient second-order stream in the Central Sand Plains of Wisconsin, USA from March to October. Invertebrate abundance and biomass were higher on average in plant beds (2,552 m⁻² and 1,575 mg m⁻²) than in sand/gravel patches (893 m⁻² and 486 mg m⁻²). Although sand/gravel habitat was over three times more abundant than plant beds in the study reach, plant beds and sand/gravel patches contributed similarly to invertebrate abundance and biomass at the whole-reach scale. The abundance and biomass of invertebrates associated with plant beds decreased from spring to autumn. Non-insect invertebrates in the plant beds increased in relative abundance as the year progressed. Shannon–Weiner diversity and taxa richness of invertebrates were higher in the plant beds than in the sand/gravel habitat. Our results suggest that plant beds can represent hot spots for invertebrate abundance and production in low-gradient streams, and have implications for stream management and restoration in these types of ecosystems. Handling editor: S. I. Dodson     

Seasonal variation in nutrient limitation of microbial biofilms colonizing organic and inorganic substrata in streams

Humans have increased the availability of nutrients including nitrogen and phosphorus worldwide; therefore, understanding how microbes process nutrients is critical for environmental conservation. We examined nutrient limitation of biofilms colonizing inorganic (fritted glass) and organic (cellulose sponge) substrata in spring, summer, and autumn in three streams in Michigan, USA. Biofilms were enriched with nitrate (NO₃ ⁻), phosphate (PO₄ ³⁻), ammonium (NH₄ ⁺), NO₃ ⁻ + PO₄ ³⁻, NH₄ ⁺ + PO₄ ³⁻, or none (control). We quantified biofilm structure and function as chlorophyll a (i.e., primary producer biomass) and community respiration on all substrata. In one stream, we characterized bacterial and fungal communities on cellulose in autumn using clone library sequencing and denaturing gradient gel electrophoresis to determine if community structure was linked to nutrient limitation status. Despite oligotrophic conditions, primary producer biomass was infrequently nutrient limited. In contrast, respiration on organic substrata was frequently limited by N + P combinations. We found no difference between biofilm response to NH₄ ⁺ versus NO₃ ⁻ enrichment, although the response to both N-species was positively related to water column PO₄ ³⁻ concentrations and temperature. Molecular analysis for fungal community composition suggested no relationship to nutrient limitation, but the dominant members of the bacterial community on cellulose were different on NO₃ ⁻, PO₄³, and NO₃ ⁻ + PO₄ ³⁻ treatments relative to control, NH₄ ⁺, and NH₄ ⁺ + PO₄ ³⁻ treatments, which matched patterns for biofilm respiration rates from each treatment. Our results show discrete patterns of nutrient limitation dependent upon substratum type and season, and imply changes in bacterial community structure and function may be linked following nutrient enrichment in streams.     

Stream metabolism controls diel patterns and evasion of CO2 in Arctic streams

Streams play an important role in the global carbon (C) cycle, accounting for a large portion of CO₂ evaded from inland waters despite their small areal coverage. However, the relative importance of different terrestrial and aquatic processes driving CO₂ production and evasion from streams remains poorly understood. In this study, we measured O₂ and CO₂ continuously in streams draining tundra‐dominated catchments in northern Sweden, during the summers of 2015 and 2016. From this, we estimated daily metabolic rates and CO₂ evasion simultaneously and thus provide insight into the role of stream metabolism as a driver of C dynamics in Arctic streams. Our results show that aquatic biological processes regulate CO₂ concentrations and evasion at multiple timescales. Photosynthesis caused CO₂ concentrations to decrease by as much as 900 ppm during the day, with the magnitude of this diel variation being strongest at the low‐turbulence streams. Diel patterns in CO₂ concentrations in turn influenced evasion, with up to 45% higher rates at night. Throughout the summer, CO₂ evasion was sustained by aquatic ecosystem respiration, which was one order of magnitude higher than gross primary production. Furthermore, in most cases, the contribution of stream respiration exceeded CO₂ evasion, suggesting that some stream reaches serve as net sources of CO₂, thus creating longitudinal heterogeneity in C production and loss within this stream network. Overall, our results provide the first link between stream metabolism and CO₂ evasion in the Arctic and demonstrate that stream metabolic processes are key drivers of the transformation and fate of terrestrial organic matter exported from these landscapes.     

Consistent temperature dependence of respiration across ecosystems contrasting in thermal history

Ecosystem respiration is a primary component of the carbon cycle and understanding the mechanisms that determine its temperature dependence will be important for predicting how rates of carbon efflux might respond to global warming. We used a rare model system, comprising a network of geothermally heated streams ranging in temperature from 5 °C to 25 °C, to explore the nature of the relationship between respiration and temperature. Using this ‘natural experiment’, we tested whether the natal thermal regime of stream communities influenced the temperature dependence of respiration in the absence of other potentially confounding variables. An empirical survey of 13 streams across the thermal gradient revealed that the temperature dependence of whole‐stream respiration was equivalent to the average activation energy of the respiratory complex (0.6–0.7 eV). This observation was also consistent for in‐situ benthic respiration. Laboratory experiments, incubating biofilms from four streams across the thermal gradient at a range of temperatures, revealed that the activation energy and Q₁₀ of respiration were remarkably consistent across streams, despite marked differences in their thermal history and significant turnover in species composition. Furthermore, absolute rates of respiration at standardised temperature were also unrelated to ambient stream temperature, but strongly reflected differences in biofilm biomass. Together, our results suggest that the core biochemistry, which drives the kinetics of oxidative respiratory metabolism, may be well conserved among diverse taxa and environments, and that the intrinsic sensitivity of respiration to temperature is not influenced by ambient environmental temperature.     

Predictability of streamflow and particulate organic matter concentration as indicators of stability in prairie streams

Predictability of stream discharge and particulate organic matter (POM) in the water column was estimated, using Colwell's indices of constancy and contingency, for 6 Texas prairie streams (1 each of 2nd, 3rd, and 4th order with intermittent or perennial discharge). Stream discharge in these 6 prairie streams varied between 0 and 36000 1 s^–1, depending on the stream and season. Predictability (P) of discharge in these streams ranged from 0.45 to 0.62, within the range of values expected for North American streams. Predictability of stream discharge was not significantly different between streams. Particulate organic matter concentrations in these prairie streams are relatively low, ranging from 0.25 to 4.00 mg AFDM 1^–1. Predictability of POM concentration in these streams was high, ranging from 0.75 to 0.85, and was largely the result of constancy of POM concentrations. Within the different POM size classes, Fine POM (FPOM) had the highest predictability (P = 0.89–0.96). In spite of relatively unpredictable stream discharge, POM remained fairly constant providing a measure of habitat predictability and stability.     

Exchange between interstitial and surface water: Implications for stream metabolism and nutrient cycling

Metabolism of a Sonoran Desert stream was investigated by both enclosure and whole system oxygen techniques. We used recirculating chambers to estimate surface sediment metabolism and measured deep sediment respiration in isolated sediment cores. Metabolism of the stream ecosystem was determined for a 30-m reach as dark and light oxygen change with and without black plastic sheeting that darkened the stream and prevented diffusion. Average ecosystem respiration for two dates in August (440 mg O₂ m^-2 h^-1) exceeded respiration of either the surface sediment community (155 Mg O₂ m^-2 h^-1) or the hyporheic community (170 mg O₂ m^-2 h^-1) alone. Deep sediments show substantial oxygen and nitrate uptake when isolated. In the stream, this low nitrate interstitial water is exchanged with surface water. Metabolism of the isolated surface community suggests a highly productive and autotrophic system, yet gross production is balanced or exceeded by community respiration when ecosystem boundaries include the hyporheic zone. Thus, despite high rates of gross primary production (600–1200 mg O₂ m^-2 h^-1), desert streams may be heterotrophic (P_G < R) during summer.     

Spatial and temporal patterns of nitrogen concentrations in pristine and agriculturally-influenced prairie streams

Long-term data on nitrogen chemistry of streams draining Konza Prairie Biological Station (Konza), Kansas were analyzed to assess spatial and temporal patterns and examine the influence of agricultural activity on these patterns. Upland watersheds of Konza are predominantly tallgrass prairies, but agricultural fields and riparian forests border the lower reaches of the streams. We have up to 11 years of data in the relatively pristine upland reaches and 4 years of data on wells and downstream reaches influenced by fertilized croplands. Seasonal and spatial patterns in total nitrogen (TN) concentrations were driven largely by changes in the nitrate (NO₃ ^–) concentrations. A gradient of increasing NO₃ ^– concentrations occurred from pristine upland stream reaches to the more agriculturally-influenced lowland reaches. Nitrate concentrations varied seasonally and were negatively correlated with discharge in areas influenced by row-crop agriculture (p = 0.007). The NO₃ ^– concentrations of stream water in lowland reaches were lowest during times of high precipitation, when the relative influence of groundwater drainage is minimal and water in the channel is primarily derived from upland prairie reaches. The groundwater from cropland increased stream NO₃ ^– concentrations about four-fold during low-discharge periods, even though significant riparian forest corridors existed along most of the lower stream channel. The minimum NO₃ ^– concentrations in the agriculturally influenced reaches were greater than at any time in prairie reaches. Analysis of data before and after introduction of bison to four prairie watersheds revealed a 35% increase of TN concentrations (p < 0.05) in the stream water channels after the introduction of bison. These data suggest that natural processes such as bison grazing, variable discharge, and localized input of groundwater lead to variation in NO₃ ^– concentrations less than 100-fold in prairie streams. Row-crop agriculture can increase NO₃ ^– concentrations well over 100-fold relative to pristine systems, and the influence of this land use process over space and time overrides natural processes.     

Theoretical study of spin-spin coupling across the hydrogen (O-H⋯N) bond in adenosine derivatives

The study of spin-spin coupling constants across hydrogen bond provides useful information about configuration of complexes. The interesting case of such interactions was observed as a coupling across an intramolecular hydrogen bond in 8-bromo-2′,3′-O-isopropylideneadenosine between the -CH₂OH (at 5″ proton) group and the nitrogen atom of adenine. In this paper we report theoretical investigations on the ^4h J _NH coupling across the H″-C-O-H···N hydrogen bond in adenosine derivatives in various solvent models. Figure Coupling constants in 8-bromo-2′,3′-O-isopropylideneadenosine Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Respiration Rate of Stream Insects Measured <Emphasis Type="Italic">in situ</Emphasis> Along a Large Altitude Range

Field studies of respiration in stream insects are few in comparison with laboratory studies. To evaluate the influence of temperature and oxygen along altitudinal gradients we measured the respiration rate of fully acclimatized larval Trichoptera, Plecoptera and Ephemeroptera under similar field conditions in streams from 400 to 3800 m above sea level in tropical Ecuador. Mean active respiration rates of the animals at 3800 m were approximately half of those at 400 m. Trichoptera showed a slightly larger difference in respiration with altitude than Ephemeroptera. Comparative respiration measurements at 100 and 50% oxygen saturation indicated that highland animals reduced their oxygen uptake more than their counterparts in the lowland when oxygen availability decreased. The temperature response of respiration calculated between the insect assemblages at different altitudes showed a mean assemblage Q₁₀−value of 1.50. Trichopteran larvae had a slightly stronger temperature response (Q₁₀ of 1.68) than ephemeropterans (Q₁₀ of 1.30). These community Q₁₀-values are considerably lower than the mean value of 2.36 found in single species in the laboratory. The weak community-wide response of respiration to temperature in tropical streams is probably due to full acclimatization of the component species to stable and narrow temperature ranges. Adaptations to the low oxygen availability at high altitude probably consist of a suite of genetic physiological and behavioural features.     

Stream ecosystem properties and processes along a temperature gradient

We surveyed macrophyte community structure and measured community metabolism and nutrient uptake along a temperature gradient (9.7–17.4°C) in four Icelandic streams influenced by geothermal heating. The study streams are part of the geothermal area in Hengill that is uniquely characterised by streams with comparable water chemistry despite the geothermal influence. Stream metabolism was studied applying the diurnal upstream–downstream dissolved oxygen change technique. Nutrient uptake was studied by adding solutions of nitrogen and phosphorus together with a conservative tracer. Rates of primary production (GPP) and uptake of nitrate–N and phosphate-P increased with increasing stream temperature. GPP was 20 times higher (up to 12.99 g O₂ m⁻² day⁻¹) and rates of nutrient uptake were up to 30-times higher (up to 22.99, 13.31 and 7.94 mg m⁻² h⁻¹ for ammonium, nitrate and phosphate, respectively) in the warmest streams compared with the coldest. Furthermore, macrophytes, when present, were strongly controlling ecosystem processes. Our study implies that temperature may affect stream ecosystem processes both directly (i.e. physiologically) and indirectly (i.e. by changing other structural parameters).     

Multi‐scaled drivers of rural prairie stream metabolism along human activity gradients

1. Stream metabolism is increasingly used for monitoring and assessment of the biological condition of aquatic ecosystems. However, distal environmental drivers, such as land use, are typically not well connected to the proximate controls, such as stream chemistry, that are usually invoked as driving metabolism. This is particularly true for North American prairie streams and for grassland streams worldwide. 2. Stream metabolism was measured at the outflow of 19 subcatchments of the Red River in southern Manitoba, Canada. Subcatchments represented gradients of nutrient‐producing human activities present in the region, that is, wastewater treatment (WWT), livestock production and crop cultivation. Stream metabolism was estimated at all sites using diel changes in dissolved oxygen (DO) concentration over 1 week in the middle of summer. Environmental parameters hypothesised to control stream metabolism were sampled across three spatial scales (stream reach, stream segment and catchment). Model selection using Akaike’s information criterion (AIC) was used to determine linkages between environmental parameters and measures of stream metabolism. 3. Estimated rates of metabolism were within the range of past studies of metabolism in prairie streams, although most streams had negative values of net ecosystem metabolism. However, production‐to‐respiration ratios were >0.5, at all but three sites suggesting that autochthonous production was an important source of organic matter. 4. The a priori model that best predicted gross primary production (GPP) was the intensity of nutrient‐producing human activities (i.e. WWT, livestock and crop cultivation) measured at the catchment scale. Ecosystem respiration (ER) was best predicted by the a priori model comprised of GPP, total nitrogen (TN) and total phosphorus (TP). However, model averaging revealed that prediction of ER could be improved by including riparian cover and removing TP from the model. The positive association between GPP and ER suggested that heterotrophic compartments of the ecosystem were modest contributors to variation in respiration rates. 5. Overall, this study suggests that variation of metabolism in prairie streams of southern Manitoba is controlled by human activities occurring at the catchment scale, a finding consistent with current hierarchically structured riverine paradigms. Moreover, increased understanding of the hierarchical structure of stream metabolism drivers will help to ensure that assessment results can be used more effectively to inform management strategies for prairie ecosystems.     

Theoretical study of the hydroxylation of phenolates by the Cu2O2(N,N′-dimethylethylenediamine)2 2+ complex

Tyrosinase catalyzes the ortho hydroxylation of monophenols and the subsequent oxidation of the diphenolic products to the resulting quinones. In efforts to create biomimetic copper complexes that can oxidize C–H bonds, Stack and coworkers recently reported a synthetic μ-η²:η²-peroxodicopper(II)(DBED)₂ complex (DBED is N,N′-di-tert-butylethylenediamine), which rapidly hydroxylates phenolates. A reactive intermediate consistent with a bis-μ-oxo-dicopper(III)-phenolate complex, with the O–O bond fully cleaved, is observed experimentally. Overall, the evidence for sequential O–O bond cleavage and C–O bond formation in this synthetic complex suggests an alternative mechanism to the concerted or late-stage O–O bond scission generally accepted for the phenol hydroxylation reaction performed by tyrosinase. In this work, the reaction mechanism of this peroxodicopper(II) complex was studied with hybrid density functional methods by replacing DBED in the μ-η²:η²-peroxodicopper(II)(DBED)₂ complex by N,N′-dimethylethylenediamine ligands to reduce the computational costs. The reaction mechanism obtained is compared with the existing proposals for the catalytic ortho hydroxylation of monophenol and the subsequent oxidation of the diphenolic product to the resulting quinone with the aim of gaining some understanding about the copper-promoted oxidation processes mediated by 2:1 Cu(I)O₂-derived species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Dynamics of soil respiration in sparse <Emphasis Type="Italic">Ulmus pumila</Emphasis> woodland under semi-arid climate

Sparse Ulmus pumila woodlands play an important role in contributing to ecosystem function in semi-arid grassland of northern China. To understand the key attributes of soil carbon cycling in U. pumila woodland, we studied dynamics of soil respiration in the canopy field (i.e., the projected crown cover area) and the open field at locations differing in distance (i.e., at 1–1.5, 3–4, 10, and >15 m) to tree stems from July through September of 2005, and measured soil biotic factors (e.g., fine root mass, soil microbial biomass, and activity) and abiotic factors [e.g., soil water content (SWC) and organic carbon] in mid-August. Soil respiration was further separated into root component and microbial component at the end of the field measurement in September. Results showed that soil respiration had a significant exponent relationship with soil temperature at 10-cm depth. The temperature sensitivity index of soil respiration, Q ₁₀, was lower than the global average of 2.0, and declined significantly (P < 0.05) with distance. The rate of soil respiration was generally greater in the canopy field than in the open field; monthly mean of soil respiration was 305.5–730.8 mg CO₂ m⁻² h⁻¹ in the canopy field and 299.6–443.1 mg CO₂ m⁻² h⁻¹ in the open field from July through September; basal soil respiration at 10°C declined with distance, and varied from ~250 mg CO₂ m⁻² h⁻¹ near tree stems to <200 mg CO₂ m⁻² h⁻¹ in the open field. Variations in soil respiration with distance were consistent with patterns of SWC, fine root mass, microbial biomass and activities. Regression analysis indicated that soil respiration was tightly coupled with microbial respiration and only weakly related to root respiration. Overall, variations in SWC, soil nutrients, microbial biomass, and microbial activity are largely responsible for the spatial heterogeneity of soil respiration in this semi-arid U. pumila woodland.     

Oxygen and nutrient dynamics of the upside down jellyfish (Cassiopea sp.) and its influence on benthic nutrient exchanges and primary production

The oxygen and nutrient dynamics of the zooxanthellate, upside down jellyfish (Cassiopea sp.), were determined both in situ and during laboratory incubations under controlled light conditions. In the laboratory, Cassiopea exhibited a typical Photosynthesis–Irradiance (P–I) curve with photosynthesis increasing linearly with irradiance, until saturation was reached at an irradiance of ~400 μE m⁻² s⁻¹, with photosynthetic compensation (photosynthesis = respiration) being achieved at an irradiance of ~50 μE m⁻² s⁻¹. Under saturating irradiation, gross photosynthesis attained a rate of almost 3.5 mmol O₂ kg WW⁻¹ h⁻¹, whereas the dark respiration rate averaged 0.6 mmol O₂ kg WW⁻¹ h⁻¹. Based upon a period of saturating irradiance of 9 h, the ratio of daily gross photosynthesis to daily respiration was 2.04. Thus, photosynthetic carbon fixation was not only sufficient to meet the carbon demand of respiration, but also to potentially support a growth rate of ~3% per day. During dark incubations Cassiopea was a relatively minor source of inorganic N and P, with the high proportion of NO _X (nitrate + nitrite) produced indicating that the jellyfish were colonised by nitrifying bacteria. Whereas, under saturating irradiance the jellyfish assimilated ammonium, NO _X and phosphate from the bathing water. However, the quantities of inorganic nitrogen assimilated were small by comparison to carbon fixation rates and the jellyfish would need to exploit other sources of nitrogen, such as ingested zooplankton, in order to maintain balanced growth. During in situ incubations the presence of Cassiopea had major effects on benthic oxygen and nutrient dynamics, with jellyfish occupied patches of sediment having 3.6-fold higher oxygen consumption and 4.5-fold higher ammonium regeneration rates than adjacent patches of bare sediment under dark conditions. In contrast at saturating irradiance, jellyfish enhanced benthic photosynthetic oxygen production almost 100-fold compared to the sediment alone and created a small sink for inorganic nutrients, whereas unoccupied sediment patches were sources of inorganic nutrients to the water column. Overall, Cassiopea greatly enhanced the spatial and temporal heterogeneity of benthic fluxes and processes by creating “hotspots” of high activities which switched between being sources or sinks for oxygen and nutrients over diurnal irradiance cycles, as the metabolism of the jellyfish swapped between heterotrophy and net autotrophy.