Feeding,metabolism and body composition of the Antarctic copepod Calanus propinquus Brady with special reference to its life cycle

Summary In the Weddell Sea, several biochemical and physiological characteristics of copepodite stage III to adult females of Calanus propinquus from the surface layer (0–100 m) and the deep layer (500–1,000 m) were compared at a time of high phytoplankton abundance (February) and a month later, when chlorophyll concentration was much lower and older copepodite stages had migrated to their overwintering depth. Daily rations of surface animals, estimated by the gut fluorescence method, varied from 0.8% to 7.8% body carbon (increasing with phytoplankton concentration). Respiration rate (calculated from measurements of electron-transport-system activity) and lipid and protein body content of animals inhabiting the surface layer decreased from February to April. Deep water animals (stage V and adult females) were characterized by high lipid and protein content; their respiration and excretion rates were much lower than in surface copepods. Our calculation showed that they could survive without an additional source of energy for 8–9 months. Based on our own and published data on bio-chemistry and physiology the possibility of a two-year life cycle is suggested.     

Physiological characteristics of the antarctic copepod Calanoides acutus during late summer in the Weddell Sea

Ingestion, respiration and excretion rates as well as lipid and protein body content of the dominant Antarctic copepod Calanoides acutus (CIV to adult females) were studied during the period covering the end of phytoplankton bloom (February) to the beginning of transition to overwintering (March-April). Daily rations measured with gut fluorescence varied from 2.2 to 2.7% in surface C. acutus. Weight-specific respiration and excretion rates in deep C. acutus decreased by a factor of 11 and 3.5–3.8 compared to their surface counterparts. High lipid (up to 455 µg ind^–1) and protein (198 µg ind^–l) content was observed in surface C. acutus CV in February; a month later the animals with similar lipid and protein content were found in the depth (500–1000 m layer). Their lipid reserves were enough to overwinter and probably to ascend, molt and reproduce. At the same time some of the deep CVs had much lower protein and lipid content and could survive only for 4–5 months. Our own and literature data led to the conclusion that females of C. acutus reach adulthood at the age of more than one year while development of males could be completed in one year.     

Digestive gut structure and activity of protease, amylase, and alkaline phosphatase in Calanus sinicus during summer in the Yellow Sea and the East China Sea

Calanus sinicus aggregate at the depth of 40–60 m (ambient temperature is 16 °C) in the waters of the continental shelf of the Yellow Sea during summer. In animals found in near shore regions, there are changes in digestive gut cells structure, digestive enzyme activity (protease, amylase), and tissue enzyme (alkaline phosphatase (ALP)), which may represent adaptations by this cold-water animal to a sharp seasonal increase in temperature of 6–23 °C. The activities of the digestive enzymes (protease and amylase) are very low in animals at stations near the estuary of Yangtse River, whereas they are relatively high in animals at stations in the central Yellow Sea. During summer, B-cells of the intestine and the villi intestinalis disappear in animals that do not feed at stations near the estuary of the Yangtse River. Respiration rates were undetectable or quite low during summer in C. sinicus from stations near the estuary of the Yangtse River, whereas they were relatively high at stations in the central Yellow Sea. Based upon the morphological characteristics of the digestive gut structure, enzyme levels, respiration rates, and the distribution of C. sinicus, we concluded that C. sinicus might be dormant during summer in the near shore areas of the East China Sea while remaining active in the central Yellow Sea.     

Denitrification in the water column of an intermittently anoxic fjord

Denitrification was studied in the water column in the Bunnefjord, inner part of the Oslofjord in southern Norway, using a ¹⁵N-technique (the isotope pairing method). The fjord is 150 m deep and during our surveys in September–December 1998 hydrogen sulphide was present in the deep water below 80 m. No significant denitrification was found in water samples from the surface layer (4 m depth), but high rates were observed within a deep density gradient between 62 and 78 m depth. Oxygen concentration within this layer was low (<21 mmol m^–3), and the concentration of NO₃ decreased from ca. 15 mmolm^–3 at 62 m depth to not detectable below 78 m. Pronounced peaks of NO₂ up to 4.4 mmol m^–3 were observed at 70–78 m depth. The maximum denitrification rate of 1.5 mmol N m^–3 d^–1 was observed at 70 m depth. Integrated for the whole layer, the denitrification rate was 13 mmol N m^–2 d^–1. A significant linear correlation was found between the denitrification rate and the ambient nitrate concentration which indicated that the rate was primarily controlled by the availability of nitrate in the O₂-poor water. Compared to rates reported for coastal water, denitrification in the water column in the Bunnefjord was high and the process appears to be a major sink of bioavailable nitrogen in the fjord.     

Caesium-137 and lead-210 dating of recent sediments from Mondsee (Austria)

Mean annual sedimentation rates over the last 20–30 years were determined in the pre-alpine Mondsee (Upper Austria) using Cs-137 and Pb(Po)-210 profiles for sediment core dating and two natural sediment markers. Lower sedimentation rates of about 2–3 mm yr^–1 were observed in the central part of the lake near the shore at 18–20 m and in the southern part at 30 m depth. Higher sedimentation rates of 4–7 mm yr^–1 were found in the central part of the lake at 47 and 65 m and in the northern bay at 18 and 41 m depth. At both these sites the Pb-210 profiles were strongly disturbed in the upper zone of the sediment cores, whereas the Cs-137 pattern remained intact. The higher annual sediment accumulation rates can be explained only partly by deposition of allochthonous material discharged by the streams, enhanced eutrophication in these parts of the lake, erosion and sediment focusing by turbidity currents being also probable.     

Methane emissions from fen,bog and swamp peatlands in Quebec

A static chamber technique was used weekly from spring thaw to winter freezing to measure methane emissions from 10 sites representing subarctic fens and temperate swamps and bogs. Rates of < 200 mg CH₄ m^–2 d^–1 were recorded in subarctic fens: within-site emissions were primarily controlled by the evolution of the peat thermal regime, though significant releases during spring thaw were recorded at some sites. Between subarctic fens, topography and water table elevation were important controls on methane emissions, with the general sequence: pool = horizontal fen> string. Emission rates from the 2 swamp sites were lower (< 20 mg CH₄ m^–2 d^–1 ), except during the spring thaw and when the sites were saturated. The low water table ( < 80 cm depth) in abnormally dry years reduced emission rates; rates were also low from a swamp site which had been drained and cleared of vegetation for horticulture. Methane emission rates were also low (< 5 mg CH₄ m^–2 d^–1) from 2 ombrotrophic bog sites. Laboratory measurements of rates of methane production under anaerobic conditions and methane consumption under aerobic conditions revealed that production rates were generally highest in the surface layers (0 to 2.5 cm depth); production was high in the fens and very low in the bogs. The swamp samples were able to produce methane under anaerobic conditions, but were also able to consume methane under aerobic conditions. Annual methane emission rates are estimated to be 1 to 10 g CH₄ m^–2 from the fens, 1 to 4 g CH4 m^–2 from the swamps and <0.2 g CH₄ m^–2 from the bogs and drained swamp.     

Does uptake of Alexandrium fundyense cysts contribute to the levels of PSP toxin found in the sea scallop, Placopecten magellanicus?

Atlantic sea scallops, Placopecten magellanicus, in most areas of the Bay of Fundy, New Brunswick, Canada, have year-round concentrations of paralytic shellfish posioning (PSP) toxins greater than the regulatory concentration of 80 μg STX eq. 100 g⁻¹ wet weight. Scallops (mean shell height of 10.7 cm, age 3–5 years) were collected by SCUBA and individually tagged near Parker Island, Bay of Fundy. Half were hung 2 m below the low tide water level and the remainder were placed on the bottom (11 m depth at low tide) under the scallops held at 2 m. Scallop, water and sediment samples were collected monthly for determination of concentrations of PSP toxins and Alexandrium fundyense.In October, 1993, mean concentrations of PSP toxins in digestive gland, and mantle were 3205 and 1018 μg STX eq. 100 g⁻¹ wet weight, respectively. Eight months later (June 1994), PSP concentrations in digestive glands from the surface and bottom had declined to 504 and 682 μg STX eq. 100 g⁻¹ wet weight, respectively, whereas those in the mantle had declined to 802 and 681 μg STX eq. 100 g⁻¹ wet weight. During July 1994, A. fundyense concentrations observed at Parker Island and offshore were 320 cells l⁻¹ and 14,200 cells l⁻¹, respectively. Subsequently, toxin concentrations in surface and bottom scallop digestive glands increased to 12,720 and 11,408 μg STX eq. 100 g⁻¹ wet weight, whereas concentrations in mantles increased to 2126 and 1748 μg STX eq. 100 g⁻¹ wet weight, respectively. Concentrations of PSP toxins in these tissues in October 1994 were similar to those measured in October 1993. Concentrations of PSP toxin were less than the regulatory concentration in the gonads and non-detectable in adductor muscles of all scallops sampled.There were no statistically significant differences in profiles for uptake and depuration of PSP toxins in scallops held at the surface compared to those from bottom, suggesting that A. fundyense cysts at the concentrations found in the sediment (45 cysts cm⁻³) did not contribute significantly to the year-round presence of PSP toxins within scallop tissues. The year-round occurrence of PSP toxin is probably due to accumulation during summer blooms followed by a very slow rate of depuration.     

Vertical distribution in and isolation of bacteria from Lake Vanda: an Antarctic lake

Vertical distribution of bacteria in Lake Vanda, an Antarctic meromictic lake, was examined by the acridine orange epifluorescence direct count method. Total bacteria were 10⁴–10⁵ cells · ml^–1 in the water at 55 m depth and above, and increased drastically to 10⁷ cells · ml^–1 in the bottom water. Filamentous or long rodshaped bacteria occurred at a high frequency in the upper layers, but in the bottom layers most bacteria were coccoidal or short rods. Mean bacterial cell volume in water of between 10 m and 60 m deep was fairly large compared with common bacterial populations in seawater and lake water. Aerobic heterotrophic bacteria were recovered from the water of a depth of 30 m and above, and were assumed to belong to Caulobacter. Viable heterotrophic bacteria were not recovered from the high salinity deep water by media prepared with the same deep water. Phototrophic purple non-sulphur bacteria were isolated by enrichment cultures from water at 55 m depth.     

Variations saisonnières de la production primaire dans les eaux de surface de la rivière du Saguenay

Experiments of primary production were carried out at weekly intervals in the surface waters at one station (maximum depth of 20 m) in the Saguenay River, near Chicoutimi, during May–December 1978. The photic zone was very thin (maximum depth of 2 m). Phosphates are very low during the season sampling (maximum of 0.1 µat-g.^–1). Maximum of production rates and biomass are respectively 3.5 mg C.m^–3.h^–1 and 3.7 mg.m^–3. The river receives both industrial and urban runoff. Trace metals (Mercury, Copper, Lead, and Iron) seemed to be one of the important limiting factors for phytoplankton growth.

     

House-dust mite (Pyroglyphidae) populations in mattresses,and their control by electric blankets

A reduction in house-dust-mite populations of 19–84% in beds in regular use could be achieved by heating the mattresses with electric blankets, when the beds were not being slept on. The effect of the heating treatment on the temperature and relative humidity inside the mattress is discussed. Housedust mites in the heated portions of the mattress migrated deeper inside the mattress to a depth of 3.5–5.0 cm. Evidence was also found that house-dust mites in non-heated mattresses do not only live near the surface, but that they are also present deep inside the mattress.This investigation was supported by a research grant from the Nederlands Astma Fonds.     

Seasonal activity and energetics of two species of varanid lizards in tropical Australia

The field metabolic rates (FMR) and rates of water flux were measured in two species of varanid lizards over five periods of the year in tropical Australia. The energetics of these species were further investigated by directly measuring activity (locomotion) and body temperatures of free-ranging animals by radiotelemetry, and by measuring standard metabolic rate (over a range of body temperatures) and activity metabolism in the laboratory. Seasonal differences in the activity and energetics were found in these goannas despite similar, high daytime temperatures throughout the year in tropical Australia. Periods of inactivity were associated with the dry times of the year, but the onset of this period of inactivity differed with respect to habitat even within the same species. Varanus gouldii, which inhabit woodlands only, were inactive during the dry and late dry seasons. V. panoptes that live in the woodland had a similar seasonal pattern of activity, but V. panoptes living near the floodplain of the South Alligator River had their highest levels of activity during the dry season when they walked long distances to forage at the receding edge of the floodplain. However, during the late dry season, after the floodplain had dried completely, they too became inactive. For V. gouldii, the rates of energy expenditure were 196 kJ kg^–1 day^–1 for active animals and 66 kJ kg^–1 day^–1 for inactive animals. The rates of water influx for these groups were respectively 50.7 and 19.5 ml kg^–1 day^–1. For V. panoptes, the rates of energy expenditure were 143 kJ kg^–1 day^–1 for active animals and 56 kJ kg^–1 day^–1 for inactive animals. The rates of water influx for these two groups were respectively 41.4 and 21.0 ml kg^–1 day^–1. We divided the daily energy expenditure into the proportion of energy that lizards used when in burrows, out of burrows but inactive, and in locomotion for the two species during the different seasons. The time spent in locomotion by V. panoptes during the dry season is extremely high for a reptile (mean of 3.5 h/day spent walking), and these results provide an ecological correlate to the high aerobic capacity found in laboratory measurements of some species of varanids.     

Methane production in meromictic Ace Lake,Antarctica

Methane occurred in the monimolimnion, at depths greater than 11 m, of an antarctic meromictic lake, Ace Lake (depth 24.7 m). Although the water of the lake was of approximate marine salinity, bottom waters were depleted in sulfate (less than 1 mmol 1^–1). The temperature of the bottom waters of the lake were constantly between 1 °C and 2 °C. Rates of methanogenesis from ¹⁴C-labelled precursors (bicarbonate, formate and acetate) were determined in time course experiments with the detection of ¹⁴CH₄ produced by a gas chromatography-gas proportional counting system. Rates of ¹⁴CH₄ production were difficult to determine as the reactions were always near our limit of detection.Reliable determinations of rates of methanogenesis at some depths using some precursors were obtained, the fastest rate being 2.5 µmol kg^–1 day^–1 at depth 20 m. Assuming constant rates of methanogenesis with time, this would equate to a turnover of methane in the lake every two years.The slow rate of methanogenesis suggests that the methanogens in Ace Lake may be working at well below their optimum temperature although definitive statements regarding the presence of psychrophilic methanogens in this antarctic lake must await isolation attempts or longer field studies using alternative methodologies.     

Control of carbon mineralization to CH4 and CO2 in anaerobic,Sphagnum-derived peat from Big Run Bog,West Virginia

The mineralization of organic carbon to CH₄ and CO₂ inSphagnum-derived peat from Big Run Bog, West Virginia, was measured at 4 times in the year (February, May, September, and November) using anaerobic, peat-slurry incubations. Rates of both CH₄ production and CO₂ production changed seasonally in surface peat (0–25 cm depth), but were the same on each collection date in deep peat (30–45 cm depth). Methane production in surface peat ranged from 0.2 to 18.8 mol mol(C)^–1 hr^–1 (or 0.07 to 10.4 g(CH₄) g^–1 hr^–1) between the February and September collections, respectively, and was approximately 1 mol mol(C)^–1 hr^–1 in deep peat. Carbon dioxide production in surface peat ranged from 3.2 to 20 mol mol(C)^–1 hr^–1 (or 4.8 to 30.3 g(CO₂) g^–1 hr^–1) between the February and September collections, respectively, and was about 4 mol mol(C)^–1 hr^–1 in deep peat. In surface peat, temperature the master variable controlling the seasonal pattern in CO₂ production, but the rate of CH₄ production still had the lowest values in the February collection even when the peat was incubated at 19°C. The addition of glucose, acetate, and H₂ to the peat-slurry did not stimulate CH₄ production in surface peat, indicating that CH₄ production in the winter was limited by factors other than glucose degradation products. The low rate of carbon mineralization in deep peat was due, in part, to poor chemical quality of the peat, because adding glucose and hydrogen directly stimulated CH₄ production, and CO₂ production to a lesser extent. Acetate was utilized in the peat by methanogens, but became a toxin at low pH values. The addition of SO₄ ^2– to the peat-slurry inhibited CH4 production in surface peat, as expected, but surprisingly increased carbon mineralization through CH4 production in deep peat. Carbon mineralization under anaerobic conditions is of sufficient magnitude to have a major influence on peat accumulation and helps to explain the thin (< 2 m deep), old (> 13,000 yr) peat deposit found in Big Run Bog.     

Prostaglandin 15-hydroxy dehydrogenase and Δ reductase levels in the lungs of maternal, fetal and neonatal rabbits

The levels of prostaglandin 15-hydroxy dehydrogenase and reductase have been studied in the lungs of maternal, fetal and neonatal rabbits. Fetal lungs obtained at gestational age of 28–30 days (full term 31 days) had the same levels of prostaglandin dehydrogenase as the adults, while the reductase levels in the fetal lungs were only one fourth that in the adults. The lungs of maternal rabbits at near term possessed very high levels of prostaglandin dehydrogenase — approximately twenty-fold higher than in the adult non-pregnant female controls. The Δ¹³ reductase appeared slightly elevated during pregnancy. Neonatal animals at different ages showed the same levels of both enzymes as the near term fetus and/or the non-pregnant adults, which suggests that the development of the ability for prostaglandin metabolism is completed at least several days before birth. The high dehydrogenase levels in the near term maternal lungs indicated the requirement for extra protection against prostaglandin release during late pregnancy.     

Suspension feeding in Bithynia tentaculata (Prosobranchia,Bithyniidae), as affected by body size,food and temperature

The suspension feeding of Bithynia tentaculata was tested in laboratory experiments. The animals were fed in 1-1 aerated glass beakers, and filtration rates were calculated from changes in cell concentrations during the 6-h experiment. Temperature influenced the filtering rate, with minimum values of 5ml · ind^–1 · h^–1 at 5° C and maxima of 17.2 ml · ind^–1 · h^–1 at 18° C. Three food species of different size, motility and cell surface characteristics (Chlamydomonas reinhardii, Chlorella vulgaris and Chlorogonium elongatum) did not affect filtration rates. Suspension feeding increased with increasing food concentrations up to 12 nl · ml^–1, above which feeding rate was kept constant by lowering the filtering rates. Even the smallest animals tested (<4 mm body length) were found to be feeding on suspended food at a rate of 2.7 ml · ind^–1 · h^–1, and increasing rates up to 8.4 ml were found in the 6–7 mm size class. All size classes of Bithynia showed a circannual fluctuation of their filtration rates. The ecological consequences of Bithynia's ability to switch between two feeding modes, grazing and suspension feeding, are discussed.     

Scaling food chains in aquatic mesocosms: do the effects of depth override the effects of planktivory?

To assess the effects of physical dimension and planktivorous fish on phytoplankton standing crop, we repeated an experiment at different scales in plastic enclosures during summer 1995 in Lake Créteil, France. Enclosures were scaled for a constant surface (1.5 × 1.5 m) as depth was increased from 2.5 to 4.5 m. Even-link (zooplankton and phytoplankton) and odd-link (planktivorous fish, zooplankton and phytoplankton) food webs were established in both shallow and deep enclosures. Fish densities in the deep enclosures were scaled to allow comparisons with shallow ones for both in individuals m⁻² or individuals m⁻³. We explicitly designed this experiment to examine the scale-dependent behavior of the top-down mechanism of algal biomass control in lakes, and in particular to test the hypothesis of stronger cascading effects of fish on lower trophic levels at reduced depth. Both fish and enclosure size had highly significant effects on phytoplankton biomass over the duration of the experiment. No depth × fish interaction effects were observed. The presence of planktivorous fish enhanced phytoplankton biomass in both shallow and deep enclosures, although the reduction in depth generally produced a stronger effect. The mean concentration of chlorophyll a in the deep odd-link systems (ca 5 mg m⁻³) was lower than in the shallow even-link systems (ca 17 mg m⁻³). Statistical interpretation did not change when data were expressed as phytoplankton biomass per unit of surface area. Light limitation and zooplankton grazing are the most probable mechanisms explaining our results in these nutrient-enriched systems. Moreover, we found that the strength of the cascading effect of fish on plankton was not a function of depth. We believe that further studies on scaling effects should be conducted in order to improve our understanding of ecological patterns and to extrapolate results from micro/mesocosms to natural ecosystems. Received: 18 January 1999 / Accepted: 7 June 1999     

Bradyrhizobium Populations Occur in Deep Soil under the Leguminous Tree Acacia albida

Soil cores were drilled under the leguminous tree Acacia albida growing in two different ecoclimatic zones of West Africa: the Sahelian area (100 to 500 mm of annual rainfall) and the Sudano-Guinean area (1,000 to 1,500 mm of annual rainfall). Soil samples were collected at different depths from the surface down to the water table level and analyzed for the presence of rhizobia able to nodulate A. albida. In both areas, population densities of rhizobia were substantially greater near the water table than near the surface. In the Sahelian area, rhizobia were present as deep as 34 m at a concentration of 1.3 × 10³/g of soil. In the Sudano-Guinean area, population densities at 0.5 to 4.5 m depth were higher than in the Sahelian area and, at several depths, comparable to that of temperate soils supporting legume crops (10⁴ rhizobia per g of soil). Surface and deep soil isolates from all four sites were found to be slow-growing rhizobia (Bradyrhizobium sp.). The proportion of effective isolates was almost the same within surface and deep soils.     

Vertical Distribution of Denitrification Potential,Denitrifying Bacteria,and Benzoate Utilization in Intertidal Microbial Mat Communities

Aspects of denitrification and benzoate degradation were studied in two estuarine microbial mat communities on the California coast by measuring the depth distributions of potential denitrification rates, genetic potential for denitrification, nitrate concentration, benzoate mineralization rates, total bacterial abundance, and abundance of a denitrifying strain (TBD-8b) isolated from one of the sites. Potential denitrification was detected in microbial mat cores from both Elkhorn Slough and Tomales Bay. Maximum denitrification rates were more than two orders of magnitude higher at Elkhorn Slough (3.14 mmol N m⁻² d⁻¹) than at Tomales Bay (0.02 mmol N m⁻² d⁻¹), and at both sites, the maximum rates occurred in the 0–2 mm depth interval. Ambient pore [NO₃+NO₂] was substantially higher at Elkhorn Slough than at Tomales Bay. Incorporation and mineralization of benzoate was maximal near the mat surface at Elkhorn Slough. The areal rate of benzoate utilization was 1045 nmol C m⁻² d⁻¹, which represented utilization of 70% of the added substrate in 24 h. Total bacterial and TBD-8b abundances were greatest near the surface at both Tomales Bay and Elkhorn Slough, and TBD-8b represented less than 0.2% of the total. Genetic potential for denitrification, quantified by hybridization with a nitrite reductase gene fragment, was present below the mat surface at average levels representing presence of the gene in approximately 10% of the total cells.     

Characterisation of Yeasts Isolated from Deep Igneous Rock Aquifers of the Fennoscandian Shield

The diversity of prokaryotes in the groundwater deep below the surface of the Baltic Sea at the Äspö Hard Rock Laboratory (HRL) in southeast Sweden is well documented. In addition, there is some evidence that eukaryotes, too, are present in the deep groundwater at this site, although their origins are uncertain. To extend the knowledge of eukaryotic life in this environment, five yeast, three yeastlike, and 17 mold strains were isolated from Äspö HRL groundwater between 201 and 444 m below sea level. Phenotypic testing and phylogenetic analysis of 18S rDNA sequences of the five yeast isolates revealed their relationships to Rhodotorula minuta and Cryptococcus spp. Scanning and transmission electron microscopy demonstrated that the strains possessed morphological characteristics typical for yeast, although they were relatively small, with an average length of 3 µm. Enumeration through direct counting and most probable number methods showed low numbers of fungi, between 0.01 and 1 cells mL^–1, at some sites. Five of the strains were characterized physiologically to determine whether they were adapted to life in the deep biosphere. These studies revealed that the strains grew within a pH range of 4–10, between temperatures of 4°C and 25–30°C, and in NaCl concentrations from 0 to 70 g L^–1. These growth parameters suggest a degree of adaptation to the groundwater at Äspö HRL. Despite the fact that these eukaryotic microorganisms may be transient members of the deep biosphere microbial community, many of the observations of this study suggest that they are capable of growing in this extreme environment.     

Factors controlling phytoplankton primary productivity in Byram Lake,Mt. Kisco,N.Y., summer 1977

Phytoplankton productivity was measured in Byram Lake Reservoir during summer 1977. Depth integrated productivity (0–5 gC m^{– 2} d^–1) increased with station depth, which together with visibility measurements indicated that light did not limit deep station productivity (C1 and S2). Macrophytes at station C5 (shallow) reduced the euphotic zone to 0 in June.On a unit depth basis, C5 was the most productive station. Apparently changes in macrophyte growth, regulated by light and temperature, controlled phytoplankton production. At C1, productivity was related to levels of different nutrients at different depths, the thermocline influencing nutrient availability at mid-depth. At S2, NH₃-N controlled mid-depth productivity. Surface and mid-depth productivity appeared influenced by factors not measured in this study.