Seasonal Anoxia Research Articles

Geochemistry of a large impoundment, Part I: solute sources, mixing dynamics, and seasonal anoxia

Lake Texoma is a large impoundment on the border of Texas and Oklahoma, formed from the confluence of two river systems with different salinities, the Red River (total dissolved salt, TDS, of 2700–11 900 mg/l, average of 4862 mg/l) and the Washita River (TDS of 420–915 mg/l, average of 701.4 mg/l). Systematic analyses for major and trace elements were conducted of water samples collected spatially and with depth in the lake in different seasons. Overall, Lake Texoma waters are characterized by Na-Ca-Cl-SO 4 -type waters with spatial distribution shifting from Na-Cl type to Ca-SO 4 type from the Red River arm to the main lake and to the Washita River arm. In addition, vertical and seasonal variations in major and trace elements concentrations indicate major elements in the lake are mainly controlled by different bedrock weathering from the two river systems. Trace elements that exhibit different distribution patterns to the major species are associated with variable sources such as river inflow, summer stratification effects, biological effects and anthropogenic activities. In Lake Texoma, differential inflow volumes and summer stratification are principal factors controlling the variation and geochemistry of lake waters and mixing dynamics. Supplementary material: Summary of major and trace element chemistry in the five zones of Lake Texoma for summer, fall and spring seasons (2003–2004) is available at http://www.geolsoc.org.uk/SUP18771

Enhanced metabolic versatility of planktonic sulfur-oxidizing γ-proteobacteria in an oxygen-deficient coastal ecosystem

Sulfur-oxidizing Gamma-proteobacteria are abundant in marine oxygen-deficient waters, and appear to play a key role in a previously unrecognized cryptic sulfur cycle. Metagenomic analyses of members of the uncultured SUP05 lineage in the Canadian seasonally anoxic fjord Saanich Inlet (SI), hydrothermal plumes in the Guaymas Basin (GB) and single cell genomics analysis of two ARCTIC96BD-19 representatives from the South Atlantic Sub-Tropical Gyre (SASG) have shown them to be metabolically versatile. However, SI and GB SUP05 bacteria seem to be obligate chemolithoautotrophs, whereas ARCTIC96BD-19 has the genetic potential for aerobic respiration. Here, we present results of a metagenomic analysis of sulfur-oxidizing Gamma-proteobacteria (GSO), closely related to the SUP05/ARCTIC96BD-19 clade, from a coastal ecosystem in the eastern South Pacific (ESP). This ecosystem experiences seasonal anoxia and accumulation of nitrite and ammonium at depth, with a corresponding increase in the abundance of GSO representatives. The ESP-GSOs appear to have a significantly different gene complement than those from Saanich Inlet, Guaymas Basin and SASG. Genomic analyses of de novo assembled contigs indicate the presence of a complete aerobic respiratory complex based on the cytochrome bc1 oxidase. Furthermore, they appear to encode a complete TCA cycle and several transporters for dissolved organic carbon species, suggesting a mixotrophic lifestyle. Thus, the success of sulfur-oxidizing Gamma-proteobacteria in oxygen-deficient marine ecosystems appears due not only to their previously recognized anaerobic metabolic versatility, but also to their capacity to function under aerobic conditions using different carbon sources. Finally, members of ESP-GSO cluster also have the genetic potential for reducing nitrate to ammonium based on the nirBD genes, and may therefore facilitate a tighter coupling of the nitrogen and sulfur cycles in oxygen-deficient waters.

Geochemical records of decadal variations in terrestrial input and recent anthropogenic eutrophication in the Changjiang Estuary and its adjacent waters

Increasing eutrophication and seasonal anoxia in bottom water in the Changjiang Estuary and its adjacent waters has progressed in recent decades, caused by elevated anthropogenic N and P input. Sedimentary biogenic elements were investigated to determine whether the biogenic proxies could be used in paleo-environmental studies in an energetic estuary, as well as to reconstruct the histories of environmental changes in the East China Sea (ECS). Two Pb-210-dated cores from the coastal and offshore waters were analyzed for organic C (TOC) and its stable isotope (delta C-13), total N (TN), biogenic Si (BSi), total P (TP) and P species. In coastal sediment, the variations of P species, especially Fe-P, Al-P and detrital apatite P (Det-P), reflected the dry-wet oscillations in the Changjiang River for the past century, which has influenced the sediment grain size and terrestrial material input. Much lower BSi content (0.756%) at 16-22 cm likely recorded the pronounced decrease in silicate flux in the Changjiang River and its lower flow in the late 1980s. In offshore sediment, higher concentrations of TOC, TN, BSi, Ex-P, Fe-P and Lea-OP indicated higher primary productivity in response to the strong winter monsoons during the 1960s-1980s, and their 20-a fluctuations were in agreement with the decadal variations of the winter monsoon. Low contents and little variations of Al-P and Det-P indicated the slight influence of the terrestrial sediment input in offshore waters. The influence of human activities on the environment in recent decades has also been recorded in coastal sediment. Grain-size normalized concentrations of TOC, TN, TP, Ex-P, Fe-P and Lea-OP increased by 24%, 23%, 15%, 13% and 51% in the upper 16 cm of coastal sediment, indicating elevated P and N load and primary productivity since the 1990s. Elevated TN/TP ratios and decreased BSi/TOC recorded the changed nutrient structure and the decrease in the proportion of the diatom to phytoplankton community. However, the sediment record indicated that the eutrophication might actually have started from the end of the 20th century rather than the reported middle of 20th century. In contrast, biogenic elements in offshore sediment did not reflect disturbance by human activities. This study revealed that multi-nutrient proxies in sediment in the ECS could indicate natural environmental changes including runoff and the winter monsoon over the past century, as well as the influence of human activities in recent decades. Phosphorus species with distinct origins and biogeochemical behaviors could effectively reflect different aspects of past environmental conditions. (C) 2012 Elsevier Ltd. All rights reserved.

Production of dimethylsulphide during the seasonal anoxia off Goa

Subsurface waters over the western Indian continental shelf experience seasonal anoxia towards the end of the southwest monsoon season. During a 3-day study carried out at the Candolim time series site (off the coast of Goa), dimethylsulphide (DMS) concentrations showed a 40-fold increase to a maximum of 442 nM at 25 m depth compared to the oxygenated surface waters. This extremely high DMS was found to be associated with relatively low chlorophyll a, low phytoplankton cell counts and a high concentration of hydrogen sulphide. However, total dimethylsulphoniopropionate, total dimethylsulphoxide and methanethiol concentrations were quite low and unlikely to account for the DMS build-up through presently known pathways of DMS production. While there are several possible mechanisms for the observed accumulation of DMS, we were unable to pinpoint the exact pathway of DMS production. Future work will involve investigation of the source of DMS through sediment slurry experiments, to explore this interesting link between the carbon and sulphur cycles under anoxic conditions.

Cause of the chalcophile trace element enrichments marking the Holocene to Anthropocene transition in northern Chesapeake Bay sediments

In Chesapeake Bay sediments, concentrations of 15 chalcophile trace elements, half rarely determined in estuaries, display historical profiles having remarkably similar features. All element concentrations rose more or less simultaneously in the 1920–1940 interval, creating a chemostratigraphic marker of the Holocene to Anthropocene transition. Subsequently, concentration maxima occurred at ∼20-year intervals, suggesting a link to a documented climate cycle of similar period. These elements’ correlated profiles suggest that sediments approximate binary mixtures of one lithogenic and one multi-element anthropogenic component. The latter component is characterized by these mass ratios (±standard error):Co/Zn 0.071±0.003Cu/Zn 0.147±0.007Ag/Zn 0.0030±0.0002Cd/Zn 0.0050±0.0004In/Zn 0.00031±0.00004Sn/Zn 0.019±0.002Sb/Zn 0.0040±0.0002Te/Zn 0.00059±0.00003Tl/Zn 0.0016±0.0002Pb/Zn 0.242±0.013Bi/Zn 0.00087±0.00005Where comparisons are possible, these ratios differ from those of contaminants in the harbor of the region’s principal industrial city, Baltimore, but are surprisingly similar to those in sediment contaminants from the Susquehanna River, the Bay’s chief tributary. Thus both the anthropogenic and the lithogenic components in the Bay’s central channel appear to originate in the river basin. Many chalcophile element ratios in the anthropogenic component are similar to those in regional aerosols. If cumulative aerosol deposition on soils in the river basin is the source of the anthropogenic component, then the above ratios could be a regional anthropogenic signature that should be looked for more widely. Unlike Mo, the enrichment of these chalcophile elements in the Bay’s sediments is not controlled by seasonal anoxia; Mo apparently possesses a unique capacity to record past redox information about estuaries owing to its high seawater concentration.

Trends in benthic macroinvertebrate community biomass and energy budgets in Lake Sevan, 1928–2004

Water levels of Lake Sevan (Armenia) were artificially lowered by nearly 20 m between 1949 and 1997. Lowered water levels, combined with increased eutrophication, were associated with seasonally anoxic conditions (lasting 1-4 months) near the bottom of the profundal zone each year during 1976-2004. In addition, the extents of the macrophyte zone and of certain substrate types were severely reduced following drawdown. Maximal depth of occurrence decreased by 2-44 m for at least for 50 species of benthic macroinvertebrates between 1982 and 2004 compared to 1937-1961. Species richness of benthic macroinvertebrates declined from 25 to three species at depths where seasonal anoxia occurred. Total biomass increased by a factor of 10 from the period 1928-1948 to 1976-1979 then declined by a factor of 3 to 4 between 1987 and 2004. Energy flow through detritivores was more than tripled during 1976-2004 compared to 1928-1971, a result of increased plankton primary production. In contrast, energy flow through herbivorous benthic macroinvertebrates decreased by a factor of nearly 5, due to reduced areal coverage of macrophytes. Energy flow through filter feeders did not change over the time period examined, but energy flow through the entire zoobenthos community was nearly tripled. The biomasses of Oligochaeta, Chironomidae, and total zoobenthos showed a delayed response to changes in primary production of 7-9, 2, and 2-4 years, respectively. These patterns may provide a basis to predict results of restoration efforts based on the abundance of the zoobenthos in future years as the level of the lake is restored and water quality improves.

Preface: Lake restoration: an experimental ecosystem approach for eutrophication control

This special section of Hydrobiologia includes a series of papers that document the ecological effects of an intensive programme of management designed to combat eutrophication of a small, monomictic lake with a long history of cyanobacteria blooms (Walsby & McAllister, 1987) and bottom-water anoxia (McColl, 1972). In more than four decades of research on, and management of, lake eutrophication across the globe (e.g., Jeppesen et al., 2005; Davis & Kloop 2006; Smith & Schindler, 2009) and despite pressures from lake acidification, bio-invasions and climate change (Smol, 2008), the effect of human activities acting at a catchment scale remains arguably the most persistent threat to the sustainability and biodiversity of natural lake ecosystems. The potential for these stressors to act in a synergistic way, for example, through increased catchment nutrient loads and climate change exacerbating eutrophication (Trolle et al., 2010), necessitates continued emphasis and vigilance on catchment management measures to address lake trophic status and biodiversity via controls on point and diffuse sources of nutrients, sediments and other contaminants. Recent history has shown that with management practices that target removal of point sources of nutrients as well as changes in, or improvement of, land use practices, it is possible to effect re-oligotrophication of lake ecosystems on time scales of 1–2 decades (Anderson et al., 2005). In other cases, in-lake techniques have been used for eutrophication control, including applications of flocculants (Welch & Cooke, 1999; Cooke et al., 2005), artificial mixing (Schladow & Fisher, 1995) and biomanipulation (Robertson et al., 2000), both with and without active catchment management programmes. In the series of papers in this special issue, the main focus is on the effects of a novel material that was applied to attempt to bring about control of internal nutrient loads in a small eutrophic lake, which has also been subject to an intensive catchment management programme. Control of internal loads was deemed necessary to combat the disproportionately high internal phosphorus load occurring with prolonged seasonal anoxia in bottom waters of the lake. The novel material referred to in this issue was initially known as Z2G1 (subsequently renamed to Aqual P) and was sought to bind phosphorus (P) in sediment porewaters and prevent the release of P Guest editors: D.P. Hamilton, M.J. Landman / Lake Restoration: An Experimental Ecosystem Approach for Eutrophication Control

Natural and anthropogenic lead in sediments of the Rotorua lakes, New Zealand

Global atmospheric sources of lead have increased more than 100-fold over the past century as a result of deforestation, coal combustion, ore smelting and leaded petroleum. Lead compounds generally accumulate in depositional areas across the globe where, due to low solubility and relative freedom from microbial degradation, the history of their inputs is preserved. In lakes there is rapid deposition and often little bioturbation of lead, resulting in an excellent depositional history of changes in both natural and anthropogenic sources. The objective of this study was to use sediments from a regionally bounded set of lakes to provide an indication of the rates of environmental inputs of lead whilst taking into account differences of trophic state and lead exposure between lakes. Intact sediment gravity cores were collected from 13 Rotorua lakes in North Island of New Zealand between March 2006 and January 2007. Cores penetrated sediments to a depth of 16–30 cm and contained volcanic tephra from the 1886 AD Tarawera eruption. The upper depth of the Tarawera tephra enabled prescription of a date for the associated depth in the core (120 years). Each core showed a sub-surface peak in lead concentration above the Tarawera tephra which was contemporaneous with the peak use of lead alkyl as a petroleum additive in New Zealand. An 8 m piston core was taken in the largest of the lakes, Lake Rotorua, in March 2007. The lake is antipodal to the pre-industrial sources of atmospheric lead but still shows increasing lead concentrations from <2 up to 3.5 μg g − 1 between the Whakatane eruption (5530 ± 60 cal. yr BP) and the Tarawera eruption. Peaks in lead concentration in Lake Rotorua are associated with volcanic tephras, but are small compared with those arising from recent anthropogenic-derived lead deposition. Our results show that diagenetic processes associated with iron, manganese and sulfate oxidation-reduction, and sulfide precipitation, act to smooth distributions of lead from anthropogenic sources in the lake sediments. The extent of this smoothing can be related to changes in sulfate availability and reduction in sulfide driven by differences in trophic status amongst the lakes. Greatest lead mobilisation occurs in mesotrophic lakes during seasonal anoxia as iron and manganese are released to the porewater, allowing upward migration of lead towards the sediment–water interface. This lead mobilisation can only occur if sulfides are not present. The sub-surface peak in lead concentrations in lake sediments ascribed to lead alkyl in petroleum persists despite the diagenetic processes acting to disperse lead within the sediments and into the overlying water.

Nitrate suppresses internal phosphorus loading in an eutrophic lake

The presence of nitrate in the hypolimnion of the eutrophic, dimictic Upper Mystic Lake has been previously shown to suppress the release of arsenic from lake sediments during seasonal anoxia, in large part by oxidizing iron (II) and producing iron oxyhydroxides that sorb inorganic arsenic. Because of the importance of internal phosphorus loading in the phosphorus budget of many eutrophic lakes, the chemical similarities between phosphate and arsenate, and the need to account for internal phosphorus loading as part of many lake restoration strategies, we carried out measurements to determine if the presence of nitrate also suppressed the release of phosphorus from the sediments of this lake during anoxia. Observations showed that this was the case. Arsenic, phosphorus, and iron (II) concentrations were strongly correlated in the water column, as expected, and the depths below which phosphorus and iron concentrations increased relative to epilimnetic values was predicted by the depth at which nitrate concentration approached zero. The results suggest that knowledge of a lake’s nitrogen budget may be a useful tool in the design of lake remediation efforts, even though phosphorus is typically the limiting nutrient.

Voltammetry: An In Situ Tool to Monitor the Health of Ecosystems

AbstractA solid‐state Au/Hg voltammetric microelectrode was used to monitor the development of seasonal anoxia in Delaware coastal bays which have deep holes with stratification. Since fish kills and harmful algal blooms have occurred over the years here, water circulators were installed in October 2003 to improve water quality. We conducted in situ determination of temperature, salinity, pH, dissolved O2 and H2S to document the circulators' function which was evaluated by comparing in situ data obtained before and after their installation. Stratification at both sites was similar in 2004 to that in previous years. Furthermore, the circulators did not oxygenate the water column but did bring some H2S to the surface waters. We found that O2 concentrations in the surface waters and H2S concentrations in the bottom waters were similar in different years. Thus, the H2S conditions were not abated and water quality was still a problem.

Chemical and physical speciation of mercury in Offatts Bayou: A seasonally anoxic bayou in Galveston Bay

A chemical equilibrium model was used to predict the solution speciation of dissolved mercury (Hg) in the stratified water column of Offatts Bayou, a subestuary in Galveston Bay, Texas, which undergoes seasonal anoxia in bottom waters. Chemical equilibrium modeling was conducted using conditional stability constants and concentrations of Hg‐complexing organic ligands experimentally determined by competitive ligand equilibration methods. Dissolved Hg complexation was dominated by interactions with sulfide and dissolved organic matter (DOM) (HOHgHS0, HOHgHS(DOM), HgSHS−, and HgS22‐ at all depths. Sulfide and glutathione competed for methylmercury (MeHg) complexation in oxic layers; in anoxic waters, sulfide complexation dominated MeHg speciation. The particle‐water distribution coefficient (Kd) of Hg decreased in the anoxic layer of the water column, where the dissolved sulfide concentration increased, providing evidence that sulfide complexation influences the solubility of Hg. The solubility of MeHg was elevated in the anoxic as compared to the oxic layers, and this distributional feature was coincident with a change in the solution speciation of dissolved MeHg from glutathione/sulfide complexation in the oxic layers to a predominantly sulfide complexation in the anoxic layers. Maximum enrichment of Hg, MeHg, and iron (Fe) in suspended particulate matter was observed in the lower layer of the pycnocline, most likely resulting from formation of insoluble Fe oxide, which scavenged dissolved Hg‐sulfide and MeHg‐sulfide species. The concomitant decrease in dissolved inorganic Hg, Fe, and sulfide in the anoxic layers is suggested to result from scavenging of inorganic Hg by FeS, which is in accordance with the Hg speciation model. Overall, Hg cycling in the water column of Offatts Bayou was associated with sulfide and DOM complexation, Fe dissolution/precipitation, water column production of MeHg, and/or efflux of MeHg from anoxic sediment.

Factors contributing to the internal loading of phosphorus from anoxic sediments in six Maine, USA, lakes

Phosphorus (P) is the limiting macronutrient for primary production in most lakes. Seasonal anoxia in the hypolimnion of lakes has been strongly correlated with internal P loading to the water column. Gravimetric sediment cores were collected before and after the onset of anoxia in six Maine (USA) lakes during the summer of 2003. This study investigates the relative importance of P sequestration by aluminum hydroxide (Al(OH)3(s)), and ferric (oxy)hydroxide (Fe(OH)3(S)) dissolution with subsequent P release in lakes with varying trophic status. Two lakes, Pennesseewassee and Highland, are oligotrophic. The remaining lakes, China, Cobbosseecontee, Webber and Salmon, have varying levels of productivity. Sediment P, Al and Fe in the top 10 cm were extracted sequentially using ammonium chloride (NH4Cl), bicarbonate-dithionite (BD), and sodium hydroxide (NaOH) at 25 degrees C. The results suggest that a sediment [NH4Cl-Al+BD-Al+NaOH25-Al]:[NH4Cl-Fe+BD-Fe] molar ratio >3 and a sediment [NaOH25-Al]:[NH4Cl-P+BD-P] molar ratio >25 predict low P flux from sediments during the development of anoxia, as proposed by Kopácek et al. [Kopácek J, Borovec J, Hejzlar J, Ulrich K, Norton S, Amirbahman A. Aluminum control of Phosphorus Sorption in Lake Sediments. Environ Sci Technol 2005; 39: 8784-8789.], despite the development of anoxia in and the release of Fe(II) from the hypolimnia of the two study lakes. However, when these molar ratios are not exceeded the model does not adequately describe sedimentary P flux. The application of the model proposed by Kopácek et al. to Cobbosseecontee Lake suggests that its sediment may be a source of P to the water column. However, water column data indicate little to no sedimentary P flux. Therefore, the lack of P flux may be attributed to the absence of Fe(III) reduction in the Cobbosseecontee Lake sediment or perhaps to the slow diagenesis of organically-bound P.

Shift of algal community structure in dead end lagoons of the Delaware Inland Bays during seasonal anoxia

Development of seasonal anoxia and algal blooms was studied in Torquay Canal and Bald Eagle Creek, 2 dead end canals in the northern Rehoboth Bay, one of the 3 Delaware Inland Bays. Mean low water depth is ca. 2 m, but dredging has produced over a dozen holes with a water depth of 5.5 m. From May to September 2002, in situ temperature, salinity, pH, dissolved O 2 and H 2 S were measured in the water column. Nutrients (NO 3 - , NO 2 - , NH 4 + and PO 4 3- ) were analyzed and the dominant members of the phytoplankton community were identified and numerated from samples collected in conjunction with in situ depth profiles. In early May, a significant potential harmful Prorocentrum minimum bloom (275 μg 1 -1 chl a) was present in Torquay Canal. Dissolved O 2 was supersaturated in the surface water, but H 2 S developed below 2 m as the water column stratified. Diatom blooms were observed in late May and mid-July, the only times that O 2 penetrated deeper, but their biomass was not significant. In early September, a storm over 3 d partially mixed the water column, and a large Heterosigma akashiwo bloom (231 μg 1 -1 chl a) was observed. Our data indicate that nutrients accumulated in the water column from runoff, organic matter decomposition, and from Fe(III) (oxy)hydroxide reduction in sediments. High concentrations of H 2 S, NH 4 + and PO 4 3- were present in the bottom waters during summer. PO 4 3- and NH 4 + from the bottom water entered shallower waters as the oxic-anoxic interface moved up to the surface. The supply of nutrients from bottom to surface waters supported harmful algal blooms during seasonal anoxic conditions as dinoflagellates and flagellates dominated over diatoms in surface waters. Seasonal anoxia development is not only a potential threat to fish and shellfish but also causes shifts of algal species to potentially harmful taxa.

Removal of H 2S via an iron catalytic cycle and iron sulfide precipitation in the water column of dead end tributaries

The oxidation and precipitation of H 2S were investigated in Torquay Canal and Bald Eagle Creek, two tributaries of northern Rehoboth Bay, one of the Delaware Inland Bays. These man-made dead end canals develop seasonal anoxia and have been the site of past fish kills and harmful algal blooms. The canals have multiple holes over 5.5 m deep compared to an average low tide depth of 2 m. In situ determination for dissolved O 2, H 2S and other Fe and S redox species were conducted with a solid-state Au/Hg microelectrode in 2003 and 2004. Laboratory analyses of discrete samples were also performed to measure dissolved and particulate Fe, Mn, and S 8 to follow the seasonal dynamics of O, S, Fe and Mn redox species. Our results indicate that the water in the holes becomes stratified with O 2 decreasing with depth and H 2S increasing with depth. Dissolved Fe was as high as 30 μM whereas dissolved Mn was only 0.2 μM in the water column, indicating that Fe is the dominant metal involved in S redox cycling and precipitation. In surface oxic waters, the dominant form of Fe was particulate Fe(III) (oxy)hydroxides. When seasonal anoxia developed, Fe(III) (oxy)hydroxides were reduced by H 2S to Fe(II) at the oxic–anoxic interface. The Fe(II) reduced from particulate Fe can be re-oxidized to Fe(III) by O 2 above and at the interface to form a catalytic cycle to oxidize H 2S. Elemental S is the predominant oxidation product and was as high as 30 μM level (as S 0) at the interface. When the system was stable, the Fe catalytic cycle prevented H 2S from being released into surface waters during seasonal anoxia. However, when storms came, the water column was overturned and H 2S was released to the surface water. The reaction rates for the Fe catalytic cycle are not fast enough and the concentration of Fe was not high enough to regulate the high concentration of H 2S in surface waters during storm and mixing events.

Seasonal changes in glycogen content and Na+-K+-ATPase activity in the brain of crucian carp

Changes in the number of Na+-K+-ATPase alpha-subunits, Na+-K+-ATPase activity and glycogen content of the crucian carp (Carassius carassius) brain were examined to elucidate relative roles of energy demand and supply in adaptation to seasonal anoxia. Fish were collected monthly around the year from the wild for immediate laboratory assays. Equilibrium dissociation constant and Hill coefficient of [3H]ouabain binding to brain homogenates were 12.87+/-2.86 nM and -1.18+/-0.07 in June and 11.93+/-2.81 nM and -1.17+/-0.06 in February (P>0.05), respectively, suggesting little changes in Na+-K+-ATPase alpha-subunit composition of the brain between summer and winter. The number of [3H]ouabain binding sites and Na-K-ATPase activity varied seasonally (P<0.001) but did not show clear connection to seasonal changes in oxygen content of the fish habitat. Six weeks' exposure of fish to anoxia in the laboratory did not affect Na+-K+-ATPase activity (P>0.05) confirming the anoxia resistance of the carp brain Na pump. Although anoxia did not suppress the Na pump, direct Q10 effect on Na+-K+-ATPase at low temperatures resulted in 10 times lower catalytic activity in winter than in summer. Brain glycogen content showed clear seasonal cycling with the peak value of 203.7+/-16.1 microM/g in February and a 15 times lower minimum (12.9+/-1.2) in July. In winter glycogen stores are 15 times larger and ATP requirements of Na+-K+-ATPase at least 10 times less than in summer. Accordingly, brain glycogen stores are sufficient to fuel brain function for about 8 min in summer and 16 h in winter, meaning about 150-fold extension of brain anoxia tolerance by seasonal changes in energy supply-demand ratio.

Multi-proxy Reconstruction of Trophic State, Hypolimnetic Anoxia and Phototrophic Sulphur Bacteria Abundance in a Dimictic Lake in Northern Germany over the past 80 Years

During monthly investigations from 1996 to 2000, a hypolimnetic layer of phototrophic sulphur bacteria (Chromatium spp.) were observed in Lake Dudinghausen, a small dimictic lake in northern Germany. This paleolimnological study was initiated to detect if the occurrence of sulphur bacteria was related to cultural eutrophication or reflected natural conditions. Therefore, diatoms, algal pigments, okenone, geochemical proxies, and 210Pb and 137Cs were used in four sediment cores to investigate historical changes in trophic development, hypolimnetic redox conditions, anoxia and phototrophic sulphur bacteria abundances. Fossil diatoms, pigments, the ratio of chlorophyll derivatives to total carotenoids and the ratio of chlorophyll a to its derivatives suggest two phases of eutrophication coupled with hypolimnetic anoxia over the last ~80 years: a first phase from about 1923–1932 and a second from 1952 to 1982. In the first phase the ratios of Fe–Mn as well as Fe–Ca increased, suggesting seasonal anoxia. However, hypolimnetic anoxia was only weak because low levels of okenone suggest no mass development of sulphur bacteria. In contrast, sulphur bacteria increased during the early stages of the second eutrophication phase, suggesting increased temporal and spatial hypolimnetic anoxia. Surprisingly, the ratios of Fe–Mn as well as Fe–Ca decreased during this time. Possibly Fe, Mn and Ca were equally reduced through the intense anoxia. In the final stage, sulphur bacteria decreased again. As these bacteria need both anoxic conditions and a certain amount of light, the increased nutrient load probably led to low Secchi depth and therefore insufficient light conditions. In more recent years, diatoms and pigments suggest a decrease in nutrient levels. A second mass development of sulphur bacteria occurred, probably due to improved light conditions and continued anoxia in the upper hypolimnion. We conclude that the recent development of phototrophic sulphur bacteria do not represent natural conditions in Lake Dudinghausen. Furthermore, the upper sediments contain a completely new diatom flora that never occurred in older sediments of Lake Dudinghausen. Therefore, nutrient levels may eventually reach natural conditions, however they may not represent biological background reference conditions.

NO ARREST NEEDED TO PREVENT CARDIAC ARREST

Most vertebrates die within a few minutes when deprived of molecular oxygen(anoxia) because the heart and brain depend on a continuous supply of oxygen. However, some cold-blooded vertebrates are remarkably anoxia-tolerant and can survive for months without oxygen. One of the keys to surviving prolonged anoxia is being able to balance energy supply and demand. One way to achieve this is to reduce energy-consuming processes of cells to a level that can be supported by the decreased amount of energy available from anaerobic respiration. Since ion pumping is energetically costly, energy can potentially be conserved by reducing the density of cellular membrane ion transport channels in excitable tissues such as the heart. Matti Vornanen and Vesa Paajanen of the University of Joensuu, Finland, set out to test this `channel arrest' hypothesis in the crucian carp (Carassius carassius), an extremely anoxia-tolerant fish that spends a third of its life in anoxic water under the ice of shallow Finno-Scandinavian ponds.The oxygen content of the water in the ponds where crucian carp live is too low for the fish to produce energy aerobically for five and a half months throughout the winter, so carp need to conserve energy during winter. Vornanen and Paajenen reasoned that since L-type Ca2+ channels regulate the force of heart muscle contraction through changes in the amount of free intracellular Ca2+, these channels are good targets for channel arrest. They hypothesized that, if crucian carp use channel arrest to conserve energy during anoxia in winter, the number of dihydropyridine receptors(subunit of the L-type Ca2+ cardiac channel that triggers channel opening) and density of L-type Ca2+ current should both be decreased in winter-captured, anoxic carp compared with summer-captured,normoxic animals.Vornanen and Paajanen captured wild carp monthly throughout an entire year and measured the number of dihydropyridine receptors and the density of L-type Ca2+ current in the ventricles of the carp's hearts to see how these changed with the seasons. Unexpectedly, they found that the number of dihydropyridine receptors in winter fish was the same as in summer fish. Thus,cardiac L-type Ca2+ channels were not downregulated by seasonal anoxia in the natural environment, suggesting that carp do not use differential expression of the Ca2+ channel protein to save energy in ion pumping or in reduced cardiac contractility during anoxia. This finding, in conjunction with the team's earlier finding that inward rectifier K+ current density of crucian carps' heart cells is also unaffected by prolonged anoxia, led Vornanen and Paajanen to conclude that crucian carp do not use channel arrest as an energy-conserving mechanism in the heart during prolonged anoxia.However, Vornanen and Paajanen did observe a 6.1-fold reduction in Ca2+ current in winter compared with summer fish, indicating that low temperatures depress Ca2+ current. They suggest that when temperatures drop, the resulting decreased Ca2+ current may allow for sufficient savings in ATP-dependent ion pumping and reduced cardiac contractility to attain a balance between energy supply and demand, providing protection against anoxia.

Decapod crustaceans from the Middle Jurassic Opalinus Clay of northern Switzerland, with comments on crustacean taphonomy

Four species of decapod crustaceans from the Middle Jurassic Opalinus Clay (Aalenian) of Northern Switzerland are described. Of these, Mecochirus cf. eckerti is the most common one, while Eryma cf. bedelta, Glyphea sp. and Aeger sp. were present as individuals, or only a few specimens. The preservation of these crustaceans ranges from moderate to excellent, reflecting the favourable taphonomic conditions of the depositional environment. An interesting aspect of the taphocoenosis in the Opalinus Clay is that the decapod crustaceans are by far outnumbered by small peracarid crustaceans (isopods and tanaids). This is interpreted as reflecting the original differences in abundance. Yet this distribution is not frequently encountered in sedimentary sequences where decapods (although rare) are far more common than isopods and tanaids. In rare instances, this reflects the original predominance of decapods, more often it is a consequence of the differential taphonomic behaviour of these two groups. A new model relating the ecology to the taphonomic behaviour of decapod and peracarid crustaceans is proposed. According to this model, decapods dominate in settings that were deposited under extremely dysoxic (peracarids wiped out by seasonal anoxia) as well as under fully oxic conditions (peracarids destroyed by taphonomic processes). Only in muddy dysoxic depositional environments are peracarid crustaceans frequently preserved. In these settings with equal preservation potential of decapods and peracarids, the original composition of the crustacean fauna would show a predominance of peracarid crustaceans. Examples from some well known fossiliferous settings are provided to illustrate the use of the new model.

Seasonality of dihydropyridine receptor binding in the heart of an anoxia-tolerant vertebrate, the crucian carp (Carassius carassius L.).

Prolonged anoxia tolerance of facultative anaerobes is based on metabolic depression and thus on controlled reduction of energy-utilizing processes. One proposed survival mechanism is the closing of ion channels to decrease energetic cost of ion pumping (Hochachka PW. Science 231: 234-241, 1986). To test this hypothesis, the involvement of L-type Ca2+ channels in seasonal anoxia tolerance of the vertebrate heart was examined by determining the number of [methyl-3H]PN200-110 (a ligand of L-type Ca2+ channel alpha-subunit) binding sites of the cardiac tissue and the density of Ca2+ current in ventricular myocytes of an anoxia-resistant fish species, the crucian carp. In their natural environment, the fish were exposed for > 3 mo of hypoxia (O2 < 2.5 mg/l) followed by almost 8 wk of anoxia that resulted in abrupt depletion of cardiac glycogen stores in late spring. Unexpectedly, however, the number of [methyl-3H]PN200-110 binding sites did not decline in hypoxia/anoxia as predicted by the channel arrest hypothesis but remained constant for most of the year. However, in early summer, the number of [methyl-3H]PN200-110 binding sites doubled for a period of approximately 2 mo, which functionally appeared as a 74% larger Ca2+ current density. Thus the anoxia tolerance of the carp heart cannot be based on downregulation of Ca2+ channel units in myocytes but is likely to depend on suppressed heart rate, i.e., regulation of the heart at the systemic level, and direct depressive effects of low temperature on Ca2+ current to achieve savings in cardiac work load and ion pumping. The summer peak in the number of functional Ca2+ channels indicates a short period of high cardiac activity possibly associated with reproduction and active perfusion of tissues after the winter stresses.

The roles of anoxia, H2S, and storm events in fish kills of dead-end canals of Delaware inland bays

In 2001, the development of seasonal anoxia was studied in two waterways located at the head of Delaware’s northern inland bay, Rehoboth Bay. Bald Eagle Creek is a northern tributary of the bay, which has tidal exchange with Torquay Canal (a dead-end canal) via a short channel with a 1.4 m sill. Mean low water depth in Torquay Canal is about 2 m, but dredging produced over a dozen depressions with a total water depth of 5.5 m. During the summer of 2000, four major fish kills were reported in Torquay Canal and Bald Eagle Creek with more than 2.5 million juvenile menhaden (Brevoortia tyrannus) killed. Low O2 concentration was assumed to be the problem but production of toxic H2S is more likely. From late spring 2001, we conducted in situ determination of temperature, salinity, pH, dissolved O2, and H2S in Torquay Canal and Bald Eagle Creek. During spring, water column stratification began in the depressions with warmer and less salty water observed in the upper layer, and cooler, saltier water below 2 m. O2 was at saturation levels in the surface waters but was not detectable below 2 m by the end of May. The depressions were anoxic with H2S accumulating to mM concentrations in June. A storm event prior to July 12 mixed these two layers with a subsequent loss of H2S. The H2S levels again increased in the deep water due to stratification and reached another maximum in late August. Another storm event occurred at this time resulting in no detectable O2 and up to 400 μM H2S in surface waters. H2S appears to be the primary reason for fish kills in these tributaries. Aerators installed in Torquay Canal on June 21 had no significant effect on abating stratification and anoxic conditions beyond their immediate area.