CLODRONATE LIPOSOMES: PERSPECTIVES IN RESEARCH AND THERAPEUTICS

ABSTRACT

Liposomes encapsulating the bisphosphonate clodronate can be used for the transient suppression of macrophage functions. Given the important role of macrophages in various disorders, the application of clodronate liposomes has been studied in several models of rheumatoid arthritis, neurological disorders such as experimental allergic encephalomyelitis and spinal cord injury, autoantibody mediated disorders such as immune thrombocytopenic purpura (ITP) and for the improvement of the efficacy of gene transfer and drug targeting.     

BIODIVERSITY AND CONSERVATION IN CHINA: A VIEW FROM ENTOMOLOGSTS

Abstract China is among the twelve megadiversity countries in the world and one of the two countries with the highest biodiversity and endemism in Asia and South-Pacific region. China has the highest mountain system of the world and different types of environments inhabited by living organisms, ranging from the Palaearctic to Oriental Realms. These characteristics account for the magnificent species diversity and high level of endemism. Insects of 45 000 species have been recorded in China by now. The total endemic elements are obviously much more than the neighbouring countries. Especially in the south-west mountainous region and tropical areas the endemic elements occupy high proportions. The human disturbances, original habitat losses due to reclamation, deforestation, pollution, etc., are driving undetermined number of species to be endangered or extinct. Recommendations of China's insect conservation are hereon presented: identifying conservation priorities, investigating the biology and ecology of endangered species, monitoring habitat and diversity losses, compiling the "Red List" and the database of rare and endangered species, establishing reserves for insect conservation, and developing techniques of insect farming and ranching.     

THERMOREGULATORY ADAPTATIONS IN MARINE MAMMALS: INTERACTING EFFECTS OF EXERCISE AND BODY MASS. A REVIEW1

A review of thermoregulation in marine mammals led to the following conclusions: very little is known about thermoregulation in large cetaceans. The only measured value for the metabolic rate of a whale, albeit a young one, was substantially higher than the predicted value for a terrestrial mammal of similar size. Very small and newborn marine mammals rely on a high metabolic heat production to sustain their body temperature during exposure to cold or in the water. The considerable insulation of some adult marine mammals may absolve them from the need for a high level of heat production. One marine mammal in tropical or subtropical waters is hypometabolic. There is evidence for a powerful control of thermoregulatory mechanisms by the anterior hypothalamic/preoptic region of the brain in two species. Thermoregulation in marine mammals during exercise remains paradoxical.     

CONSERVATION IN THE MARINE ENVIRONMENT: SOME PROBLEMS WITH THE MANAGEMENT OF SHORE-ANGLING IN THE SOUTHWESTERN CAPE

Summary

Recreational shore angling is a very important sector of South Africa's diverse marine fisheries. The number of anglers participating in this sport is increasing at 6% per annum and currently stands at approximately 380 000. This number is expected to exceed one million within 20 years. The catch rates of many species caught from the shore have declined markedly for at least 50 years and the species composition of the catch has changed. This indicates that stocks of many species are overexploited and that effort increases cannot be sustained. Measures introduced to conserve these stocks appear to have been largely unsuccessful because they do not have the support of many anglers and they are ineffectively policed. It is concluded that, in order to sustain this fishery, current legislation needs to be improved and that a licensing system for anglers should be introduced through which an education campaign can be channelled.     

嵌套性:研究方法、形成机制及其对生物保护的意义

岛屿或者“生境岛”中的生物区系常常显示出一种嵌套结构 ,即物种较贫乏的岛屿中的物种是物种较丰富的岛屿中的物种的一个适当的子集 ,如果将各个岛屿中的生物区系排列起来就形成一个嵌套的序列。与种 面积关系一样 ,嵌套结构在很多生境类型和生物类群中也都存在。嵌套性对生物保护也有一定的意义 ,特别是与SLOSS争论 (是单个大的还是几个小的保护区能保护更多的物种 )有一定关系。在过去的十几年中 ,已经提出了一些方法 ,可以对嵌套性进行定量刻画和统计检验。同时 ,对嵌套性的形成机制也进行了大量的研究 ,其中选择性的迁移和选择性的灭绝是两个主要的原因。由于嵌套性分析只需要物种的存在 /不存在数据 ,使得很多调查数据都能够利用起来 ,因此 ,这是一个值得深入研究的领域     

FISH CONSERVATION IN SOUTH AFRICAN ESTUARIES: PRESSURES,PROBLEMS AND PROSPECTS

Summary

A review of the conservation status of fish in the estuarine environment around the South African coastline reveals that some species face serious problems associated either with habitat destruction, and its associated biological, physical and chemical components or exploitation. The 65 species considered fall into three categories; truly estuarine species, which are dependent on estuaries for their entire life; marine species dependent on estuaries during the juvenile phase of their life cycle; and marine species whose juveniles occur mainly in estuaries but are also found at sea. Included in the first two categories are 14 species of fish which are on the South African Red Data list. These comprise one species which is endangered, five which are vulnerable and eight which are rare. All groups are considered in relation to factors in estuaries which are affecting their populations. A conservation strategy is suggested for certain estuarine types or for specific estuaries which could ensure the survival of the entire range of estuarine faunas.     

DIEL OSCILLATIONS IN THE PHOTOSYNTHESIS-IRRADIANCE RELATIONSHIP OF A PLANKTONIC MARINE DIATOM1

The amplitude of diel oscillations in photosynthesis as a function of irradiance varied with the growth phase in a marine phytoplankton species. The common centric diatom (Bacillariophyta), Ditylum brightwellii (West) Grun., showed strong periodicity in the photosynthesis-irradiance (P-I) relationship, which damped progressively from early to late exponential and stationary phase. These findings suggest that short-term temporal characteristics of phytoplankton production depend on factors which affect growth, and that the amplitude is most enhanced at maximal growth rates likely to be encountered in the natural environment.     

D-VITAMINS AND THEIR PRECURSORS AS GROWTH REGULATORS IN AXENICALLY CULTIVATED MARINE MACROALGAE1

Light petroleum extracts from thalli of Fucus vesiculosus (L.) and Laminaria digitata (Huds.) Lamour. stimulate growth of axenically cultured Fucus spiralis L. and Enteromorpha compressa (L.) Grev. in amounts per litre culture medium corresponding to one to 10 mg wet weight of the brown algae. This indicates the presence of a lipophilic, very active growth regulator in the blades. Extracts from the meristematic zone of Laminaria are much less active. Vitamins D₃, D₂ and 1,25-(OH)₂-D₃ at 1·10^?8– 1·10^?7 M increase the fresh weight of Enteromorpha compressa and Fucus spiralis up to 100% or more. The dry weight of Nemalion helminthoides (Veil, in With.) Batt. is also increased under the same conditions. Ergosterol and cholesterol are also growth-stimulating but higher concentrations (e.g. 1·10^?6 M) are necessary. The effects of the D vitamins manifest themselves without any auxins being added to the medium.     

PATTERNS OF ALGAL SUCCESSION IN A PERTURBATED MARINE INTERTIDAL COMMUNITY1

Patterns of algal succession for a sewage-polluted and an unpolluted habitat near Wilson Cove, San Clemente Island, California, were studied from December 1974 to June 1977. Resident populations were analyzed for 56 fully denuded and 34 undisturbed control quadrats during 11 assessment periods. The denuded quadrats in the perturbated (polluted) habitat showed recovery within 1.0 mo as determined by cover, percent similarity and species diversity comparisons with control plots. The short recovery times of the algal populations dominating the perturbated habitat indicate that these species maintain relatively constant overall abundances due to their potential for rapid recruitment and growth. Denuded quadrats in the unpolluted habitat did not show recovery even after 30.0 mo. These quadrats were dominated during the first 1.3 mo by algae characteristic of the perturbated area, including filamentous Ectocarpaceae, colonial diatoms and bluegreen algae. Petalonia fascia (Müll.) Kuntze (1.3–3.0 mo), and Scytosiphon lomentaria (Lyngb.)J. Ag. and Colpomenia sinuosa (Roth) Derb. & Sol. (1.3–5.0 mo) were abundant shortly following plot denudation and provided thick growths that may have excluded other algae. The similarity between the species occupying the sterilized plots during the first few months and those that provide the majority of cover in the perturbated area supports the hypothesis that the dominant algae of the upper and midintertidal regions of this habitat consist largely of early successional or opportunistic species with high capacities for growth and reproduction. Additionally, these experiments suggest that algal populations described for other perturbated epilithic systems also represent resilient subclimax associations.     

LIGHT ABSORPTION AND QUANTUM YIELD FOR GROWTH IN FIVE SPECIES OF MARINE MACROALGA1

Light utilization efficiency in five species of marine macroalgae was measured in laboratory growth experiments (13–41 days duration) at different irradiances at 7°C. All species acclimated to irradiance by changing their light absorption, resulting in a peak in light absorption between 2 and 15 μmol·m^?2.s^?1. Light absorption increased with thallus-specific chlorophyll and carbon content according to linear inverse relationships between chlorophyll content (chlarea^?1) and log[transmission] and between log[carbon content, Carea^?1] and log[transmission]. Quantum yields for light-limited growth and estimated gross photosynthesis were calculated based on incident and absorbed light. Quantum yield for photosynthesis based on light absorbed by pigments was high (mean = 114 mmol C·mol^?1 photons) and similar among the species. Quantum yield for net growth based on incident light was also high but more variable, between 22 and 75 mmol C·mol^?1 photons. Differences among species were mainly due to differences in light absorption. In conclusion, all species acclimated to low light by increasing light absorption to the maximum attainable, and growth efficiencies based on absorbed light were close to the maximum theoretically possible.     

A MODEL APPROACH FOR SIZE-SELECTIVE COMPETITION OF MARINE PHYTOPLANKTON FOR FLUCTUATING NITRATE AND AMMONIUM1

Phytoplankton size-selective competition for fluctuating nutrients was studied with the use of a numerical model, which describes nitrate and ammonium uptake, nitrate reduction to ammonium, and growth as a function of cell she under fluctuating nitrogen limitation. The only size-dependent parameter in the model was the cell nutrient quota. Related to this, the cell surface area per biomass was negatively correlated to cell volume, and the vacuole volume per biomass ratio was positively correlated to cell volume. Simulations showed an inverse correlation between the maximum specific growth rate and cell size under steady-state conditions. With nitrate as the limiting nitrogen source, nitrogen quotas were always higher than with ammonium at the same specific growth rate. Net passive transport of ammonium due to unspecific diffusion of ammonia across the plasma membrane decreased the affinity for ammonium, whereas the affinity for nitrate was not influenced. Transient state-specific ammonium uptake was not dependent on cell size. However, small algae always have the highest specific growth rate in ammonium-controlled systems according to our model. Transient state nitrate uptake rate was positively correlated to cell size because larger algae have a higher vacuole volume per biomass, in which nitrate can be stored. Despite their lower maximum growth rate, larger algae became dominant during simulations under fluctuating nitrate supply when amplitude of and the period between nitrate pulses were high enough. Results from model simulations were qualitatively validated by earlier observations that large diatoms become dominant under fluctuating conditions when nitrate is the main nitrogen source.     

USING ORGANOCHLORINE POLLUTANTS TO DISCRIMINATE MARINE MAMMAL POPULATIONS: A REVIEW AND CRITIQUE OF THE METHODS1

Organochlorine pollutants are potentially useful for identifying discrete populations of marine mammals that overlap in geographic distribution. However, many factors unrelated to geographical distribution may affect the chemical burden of individual animals or of entire population components even within a homogeneously distributed population. These factors include. among others, nutritional state, sex, age, trophic level, distance of habitat from mainland and pollution source, excretion. metabolism, and tissue composition. Sample storage and analytical methodology may also be an important source of variation. These, and any other factors, must be identified and their effect ascertained before attempting any comparison between populations. This paper critically examines the nature and magnitude of the effects of these factors on organochlorine tissue loads in marine mammals. Pollutant concentrations can be strongly biased if carefully designed sampling regimes are not followed, but they are affected only moderately by sample treatment after collection. Conversely, ratios between concentrations of compounds, such as the DDE/tDDT or the tDDT/PCB ratios, seem less dependent on sampling regime but more affected by storage. analytical procedures and ecological variations such as distance from pollutant source or trophic level. Taking these effects into account, advice is provided about sampling and strategies for selection of variables that will improve the reliability of the comparisons between populations.     

SEASONAL PERIODICITY OF CHRYSOPHYCEAE AND SYNUROPHYCEAE IN A SMALL NEW ENGLAND LAKE: IMPLICATIONS FOR PALEOLIMNOLOGICAL RESEARCH1

The seasonal periodicity of taxa of Chrysophyceae and Synurophyceae from a small New England lake is described for the period September 1983 through June 1988. We found 51 taxa, including 29 that accounted for over 10% of the total in at least one collection. The taxa were fitted into one of five seasonal patterns. Patterns I and II represented taxa restricted to warm (pattern I) or cold (pattern II) months, respectively. Pattern HI represented organisms that began growth in the summer, persisted through autumn and disappeared with the onset of an ice cover. Pattern IV was an extension of pattern III, in which the taxon remained in the plankton throughout the winter and disappeared soon after ice out. Species without a clear seasonal pattern were grouped as pattern V. The seasonal periodicity of the flora, as examined with ordination analyses, was found to remain remarkedly similar during the 58–month study. Except for episodes of low pH during spring snow melt and unseasonally warm or cold weather, sample scores followed a fairly consistent pattern along the first and second primary axes. Water temperature, specific conductance, and pH were important variables that controlled changes in the species composition during the course of a given year. The flora was used to develop an inference model for water temperature. According to the analyses, the remains of a surface sediment sample represented a flora that grew primarily during the late autumn period at 7.6 ° C. Ways in which seasonal data could be utilized to improve paleolimnological inference work are discussed.     

AN IN VITRO NITRATE REDUCTASE ASSAY FOR MARINE MACROALGAE: OPTIMIZATION AND CHARACTERIZATION OF THE ENZYME FOR FUCUS GARDNERI (PHAEOPHYTA)1

Measurement of the activity of the enzyme nitrate reductase (NR) may provide a useful index of nitrogen metabolism in marine macroalgae. In several species, including Fucus gardneri P. C. Silva, in vitro assays previously failed to detect NR activity, necessitating the use of in situ (or so-called“in vivo”) assays, which are more loosely controlled and lead to dafficulties in assessing enzyme characteristics such as the half-saturation constant (K_m). In this paper, we describe an in vitro NR assay developed for F. gardneri, in which tissue was homogenized using liquid nitrogen prior to the assay. In contrast to previous studies, enzyme activity was always detectable in F. gardneri collected directly from the field at levels up to 30 nmol nitrate converted to nitrite·min^?1·g^?1 wet weight. The effect of a variety of compounds, commonly added to NR extraction buffers, were tested. Additions of protease inhibitors, bovine serum albumin, and ethylenediamine tetraacetic acid had no consistent effects on NR activity, while polyvinyl pyrrolidone, potassium ferricyanide, and flavin adenine dinucleotide significantly decreased activity. The half-saturation constant (K_m) for NADH was 0.18 (± 0.05) mM and for nitrate, K_m=0.99 (±0.41) mM. Significant NR activity was detected without the addition of nitrate, suggesting that internal pools of nitrate averaging approximately 20 μmol NO₃^?·g^?1 wet weight were present in F. gardneri in February. The distribution of NR activity within the plant was highly variable between individuals, but activities were approximately 5-fold lower in the stipe than in midregions. In plants freshly sampled from the field, NR activity increased 7-fold from February to March, then fell to near-February levels by April. These changes in activity may correspond to seasonal changes in growth rate. The assay, optimized for F. gardneri, was used in several different macroalgal species from different taxa: Porphyra sp., Coralina vancouveriensis Yendo, Ulva sp., Enteromorpha intestinalis (Linnaeus) Nees, Macrocystis integrifolia Bory; and Costaria costatum (C. Agardh) Saunders. For all species tested, NR activity was detectable and, except for one species (Porphya sp.), was equal to or greater than activities measured by other workers using in vivo or in vitro assays for plants under similar conditions.     

DISTRIBUTION OF INTRACELLULAR NITROGEN IN MARINE MICROALGAE: BASIS FOR THE CALCULATION OF SPECIFIC NITROGEN-TO-PROTEIN CONVERSION FACTORS

The utilization of nitrogen-to-protein conversion factors (N-Prot factors) is a widely accepted and practical way to determine total protein content. The accuracy of protein determination depends on the establishment of specific N-Prot factors, since the conventional factor of 6.25 may be unsuitable for all species. This study was designed to determine the concentrations of the main nitrogenous compounds and to establish N-Prot factors specific for the following marine microalgae: Chlorella minutissima, Dunaliella tertiolecta, Hillea sp., Isochrysis galbana, Nannochloropsis oculata, Phaeodactylum tricornutum, Prorocentrum minimum, Skeletonema costatum, Synechococcus subsalsus, and Tetraselmis gracilis. Cultures were maintained under a 12-h photoperiod (300 μmol photons·m^?2·s^?1) at temperatures of 20.0°± 1.0° C (dark) to 23.0°± 2.0° C (light) in Walne’s culture medium without additional external carbon sources. The distribution of intracellular nitrogen was studied by determining total nitrogen (TN, by CHN [carbon, hydrogen, and nitrogen] analysis), protein N (PN, by analysis of total amino acids), and nonprotein N (NPN, determined by analysis of DNA, RNA, chlorophylls (chl) a,b, and c, and intracellular inorganic nitrogen—NO₃^?, NO₂^?, and NH₃+ NH₄⁺) in logarithmic and stationary growth phases of cultures. Variations occurred in both accumulation and distribution of PN and NPN among the species, as well as in each species during the different growth phases. Inorganic nitrogen compounds were observed to be the most important NPN source (from 6.4 ± 0.1% to 41.8 ± 4.2% of total N) in all species (except D. tertiolecta), followed by nucleic acids (from 0.8 ± 0.1% to 26.1 ± 2.4% of TN) and chlorophylls (from 0.2 ± 0.0% to 3.1 ± 0.3% of TN). Total amino acid residues ranged from 63.1 ± 4.6% up to 88.1 ± 11.2% of TN, which is in agreement with the presence of high NPN concentrations. N-Prot factors are proposed for each growth phase in the studied species, based on the ratio of amino acid residues to TN, establishing specific N-prot factors ranging from 3.60 ± 0.27 to 4.99 ± 0.64. The mean N-Prot factor for all species/growth phases was 4.58 ± 0.11. The present study shows that the use of the traditional factor 6.25 is not suitable for these marine microalgae, and possibly for other species, because it overestimates their actual protein content.     

DNA:ATP RATIOS IN MARINE MICROALGAE AND BACTERIA: IMPLICATIONS FOR GROWTH RATE ESTIMATES BASED ON RATES OF DNA SYNTHESIS1

DNA: ATP and carbon: DNA (C:DNA) ratios were measured in a total of 14 species of marine microalgae and bacteria. Comparison of several DNA assay methods with results obtained with cultures uniformly labeled with ³³P indicated that by far the most accurate results were obtained using diaminobenzoic acid (DABA) or diphen-ylamine, with DABA having the highest precision. Both the Hoechst and DAPI methods seriously underestimated DNA concentrations in algal cultures. Average DNA: ATP ratios in the algal and bacterial cultures were I7 and 34 by weight, respectively, with almost all values lying in the range of 10–40. DNA: ATP ratios in the microalgae showed no correlation with growth conditions but varied by about a factor of 3 among species. C:DNA ratios for individual species of microalgae and bacteria ranged from 21 to 155 by weight and averaged 50 for the microalgae and bacteria taken together. Growth rates of microalgal species grown in cyclostats were estimated to within 8% of dilution rates when calculated from the uptake of ³H-adenine and the DNA: ATP ratio of the species. Use of the ³H-adenine method for estimating microalgal growth rates in the field may thus be a useful tool for investigating the physiology of microalgae in nature.     

LECTIN-LIKE COMPOUNDS AND LECTIN RECEPTORS IN MARINE MICROALGAE: HEMAGGLUTINATION AND REACTIVITY WITH PURIFIED LECTINS1

We detected lectin-like compounds and lectin receptors in microalgae by hemagglutination, competitive inhibition with sugars, and reactivity with lectins isolated from other sources. Cell extracts from eight species of Dinophyceae and from one species each of Raphidophyceae and Bacillariophyceae exhibited hemagglutination toward trypsinized rabbit erythrocytes. In addition, the culture media of the dinoflagellate Alexandrium cohorticula and the raphidophyte Chattonella antiqua displayed similar hemagglutination. These activities were not inhibited by any monosaccharides or oligosaccharides tested but were inhibited by some specific glycoproteins. This suggests that the active factors were lectin-like compounds. Upon exposing intact, healthy cells of 12 species of Dinophyceae and one species each of Raphidophyceae, Cryptophyceae, Bacillariophyceae, and Chlorophyceae to lectins isolated from either macroalgae or terrestrial plants, most species were adversely affected. The negative effects included one or more of the following: impaired motility, disappearance of motility, agglutination, abnormal morphology, and cell rupture or lysis. Some species, even after freezing, thawing, and washing with saline solution, still agglutinated with macroalgal or terrestrial plant lectins. This study suggests that lectins and carbohydrate-containing lectin receptors may commonly occur on the cell surfaces of various species of microalgae.     

INTERACTIVE EFFECTS OF COPPER AND SILICIC ACID ON RESTING SPORE FORMATION AND VIABILITY IN A MARINE DIATOM1

The toxic effects of copper on resting spore formation and viability in the marine diatom Chaetoceros protuberans Lauder were determined both with and without silicic acid added to the medium. With silicic acid available, partial inhibition of resting spore formation occurred only at the highest cupric ion activity (pCu 8.6), while the percentage of cells forming spores at pCu's 10.2 and 11.3 was nearly the same as in the controls. Without silicic acid added to the medium, sporulation was completely inhibited at pCu 8.6 and greatly inhibited at pCu 10.2. At pCu 11.3 and in the controls, the rate of spore formation was less than 50%. The results indicate that the inhibition of resting spore formation by copper is related to the concentration of silicic acid available to cells of C. protuberans. This is consistent with previous studies which show that copper toxicity during vegetative growth involves interference with silicification in diatoms and is a function of the silicic acid concentration of the medium. Viable resting spores of C. protuberans were still present in cultures following exposure to elevated copper concentrations during a 100-day incubation period. This indicates that resting spores can serve to enhance diatom survival in areas polluted by heavy metals.     

INTERACTIVE EFFECTS OF COPPER AND SILICIC ACID ON RESTING SPORE FORMATION AND VIABILITY IN A MARINE DIATOM1

The toxic effects of copper on resting spore formation and viability in the marine diatom Chaetoceros protuberans Lauder were determined both with and without silicic acid added to the medium. With silicic acid available, partial inhibition of resting spore formation occurred only at the highest cupric ion activity (pCu 8.6), while the percentage of cells forming spores at pCu's 10.2 and 11.3 was nearly the same as in the controls. Without silicic acid added to the medium, sporulation was completely inhibited at pCu 8.6 and greatly inhibited, at pCu 10.2. At pCu 11.3 and in the controls, the rate of spore formation was less than 50%. The results indicate that the inhibition of resting spore formation by copper is related to the concentration of silicic acid available to cells of C protuberans. This is consistent with previous studies which show that copper toxicity during vegetative growth involves interference with silicification in diatoms and is a Junction of the silicic acid concentration of the medium. Viable resting spores of C. protuberans were still present in cultures following exposure to elevated copper concentrations during a 100-day incubation period. This indicates that resting spores can serve to enhance diatom survival in areas polluted by heavy metals.     

INTRACELLULAR DIMETHYLSULFOXIDE (DMSO) IN UNICELLULAR MARINE ALGAE: SPECULATIONS ON ITS ORIGIN AND POSSIBLE BIOLOGICAL ROLE

Recent studies have established that aqueous phase concentrations of dimethylsulfoxide (DMSO) often exceed those of dimethylsulfoniopropionate (DMSP) and dimethylsulfide (DMS). Yet, in comparison to DMSP and DMS, DMSO remains a poorly understood component of the marine sulfur cycle. Much of what is known about the mechanisms for the formation and loss for DMSO is inferred from laboratory experiments, and no explanation exists to rationalize how a large pool of DMSO is maintained. One formation pathway that, until very recently, has been ignored involves the direct synthesis of DMSO by marine phytoplankton. This review examines some of the circumstantial evidence for DMSO in marine particulate material and recent reports containing preliminary data for particulate DMSO (DMSO_p) in the marine environment. Drawing on literature from a range of scientific disciplines, speculations on the possible origins and biological functions of intracellular DMSO are also made. On the basis of its physicochemical properties, intracellular DMSO could have a potential role as a cryoprotectant, a specialist cryo-osmoregulator in extreme environments, an intracellular electrolyte modifier, and a free-radical scavenger. The review also assesses the impact of DMSO_p at both the organism and the global level. Consideration is given to the marine biogeochemical cycling of sulfur and potential links to climate control.