Trichodesmium provides new nitrogen (N) to marine surface waters via N 2 fixation, a process that... more Trichodesmium provides new nitrogen (N) to marine surface waters via N 2 fixation, a process that requires a substantial amount of iron (Fe). Organic ligands in seawater that bind Fe could either increase or reduce the bioavailability of Fe. Electrochemical techniques indicate that these naturally occurring ligands have Fe-binding constants similar to those of siderophores and porphyrins, suggesting that these chelators play an important role in determining the bioavailability of Fe to cyanobacteria. We conducted Fe uptake experiments using model ligands labeled with 55 Fe to compare the bioavailability of inorganic Fe(III), porphyrin-bound Fe(III), and siderophorebound Fe(III) to field-collected Trichodesmium colonies. Inorganic Fe(III) and siderophore-bound Fe(III) were more bioavailable to Trichodesmium colonies than was porphyrin-bound Fe(III). Furthermore, the bioavailability of the siderophore-bound Fe(III) can be characterized by the functional groups of the siderophore. The dihydroxamate siderophore and an uncharacterized ligand from a cultured Synechococcus sp. increased the bioavailability of Fe compared to the trihydroxamate siderophores. Except for experiments with desferrioxamine B, dark incubations resulted in lower Fe uptake rates for all treatments, relative to parallel lighted incubations. This suggests that light enhances the photochemical dissociation of most of the ligand complexes or that light energy is required for the active transport of Fe complexed to the model ligands. The Fe uptake rate of Trichodesmium colonies also differed slightly on the basis of colony morphology, with higher uptake rates with ''puffs'' than ''tufts.'' These experiments show that Trichodesmium colonies are capable of discriminating between Fe bound to different organic complexes.
Viral lysis of phytoplankton constrains marine primary production, food web dynamics and biogeoch... more Viral lysis of phytoplankton constrains marine primary production, food web dynamics and biogeochemical cycles in the ocean. Yet, little is known about the biogeographical distribution of viral lysis rates across the global ocean. To address this, we investigated phytoplankton group-specific viral lysis rates along a latitudinal gradient within the North Atlantic Ocean. The data show large-scale distribution patterns of different virus groups across the North Atlantic that are associated with the biogeographical distributions of their potential microbial hosts. Average virus-mediated lysis rates of the picocyanobacteria Prochlorococcus and Synechococcus were lower than those of the picoeukaryotic and nanoeukaryotic phytoplankton (that is, 0.14 per day compared with 0.19 and 0.23 per day, respectively). Total phytoplankton mortality (virus plus grazer-mediated) was comparable to the gross growth rate, demonstrating high turnover rates of phytoplankton populations. Virus-induced morta...
Background/Question/Methods Bacteria typically live in environments which are depleted in resourc... more Background/Question/Methods Bacteria typically live in environments which are depleted in resources such as nitrogen (N) and phosphorus (P). These organisms have advanced nutrient uptake and retention mechanisms to cope with nutrient limiting situations, but these same structural features may be sites of attack by viral predators (i.e., phage). Phage play an important role in the ‘top-down’ regulation and structuring of bacterial communities, but also influence the evolutionary trajectory of host populations on ecologically relevant time scales. The interaction between virus predation and nutrient limitation, however, has not been extensively examined within an evolutionary context. Using the marine cyanobacterium, Synechococcus sp. WH 7803 and a T4-like phage, we explored the fitness tradeoffs and eco-evolutionary dynamics associated with phage attack and nutrient limitation. Through an eight month chemostat experiment, we manipulated virus exposure and the N:P supply ratio to crea...
Here we examined the impact of a commonly employed method used to measure nitrogen fixation, the ... more Here we examined the impact of a commonly employed method used to measure nitrogen fixation, the acetylene reduction assay (ARA), on a marine sediment community. Historically, the ARA technique has been broadly employed for its ease of use, in spite of numerous known artifacts. To gauge the severity of these effects in a natural environment, we employed high-throughput 16S rRNA gene sequencing to detect differences in acetylene-treated sediments vs. non-treated control sediments after a 7 h incubation. Within this short time period, significant differences were seen across all activity of microbes identified in the sediment, implying that the changes induced by acetylene occur quickly. The results have important implications for our understanding of marine nitrogen budgets. Moreover, because the ARA technique has been widely used in terrestrial and freshwater habitats, these results may be applicable to other ecosystems.
We announce the complete genome sequences of two temperate Podoviridae, Sulfitobacter phages ΦCB2... more We announce the complete genome sequences of two temperate Podoviridae, Sulfitobacter phages ΦCB2047-A and ΦCB2047-C, which infect Sulfitobacter sp. strain 2047, a member of the Roseobacter clade. This is the first report of temperate podophage infecting members of the Sulfitobacter genus of the Roseobacter clade.
Little is known about the molecular and physiological function of co-occurring microbes within fr... more Little is known about the molecular and physiological function of co-occurring microbes within freshwater cyanobacterial harmful algal blooms (cHABs). To address this, community metatranscriptomes collected from the western basin of Lake Erie during August 2012 were examined. Using sequence data, we tested the hypothesis that the activity of the microbial community members is independent of community structure. Predicted metabolic and physiological functional profiles from spatially distinct metatranscriptomes were determined to be >90% similar between sites. Targeted analysis of Microcystis aeruginosa, the historical causative agent of cyanobacterial harmful algal blooms over the past ϳ20 years, as well as analysis of Planktothrix agardhii and Anabaena cylindrica, revealed ongoing transcription of genes involved in microcystin toxin synthesis as well as the acquisition of both nitrogen and phosphorus, nutrients often implicated as independent bottom-up drivers of eutrophication in aquatic systems. Transcription of genes involved in carbon dioxide (CO 2 ) concentration and metabolism also provided support for the alternate hypothesis that high-pH conditions and dense algal biomass result in CO 2 -limiting conditions that further favor cyanobacterial dominance. Additionally, the presence of Microcystis-specific cyanophage sequences provided preliminary evidence of possible top-down virus-mediated control of cHAB populations. Overall, these data provide insight into the complex series of constraints associated with Microcystis blooms that dominate the western basin of Lake Erie during summer months, demonstrating that multiple environmental factors work to shape the microbial community. Citation Steffen MM, Belisle BS, Watson SB, Boyer GL, Bourbonniere RA, Wilhelm SW. 2015. Metatranscriptomic evidence for co-occurring top-down and bottomup controls on toxic cyanobacterial communities. Appl Environ Microbiol 81:3268 -3276.
We announce the complete genome sequence of a lytic podovirus, ⌽CB2047-B, which infects the bacte... more We announce the complete genome sequence of a lytic podovirus, ⌽CB2047-B, which infects the bacterium Sulfitobacter sp. strain 2047, a member of the Roseobacter clade. Genome analysis revealed ⌽CB2047-B to be an N4-like phage, with its genome having high nucleotide similarity to other N4-like roseophage genomes.
We present the genome of a cyanosiphovirus (KBS2A) that infects a marine Synechococcus sp. (strai... more We present the genome of a cyanosiphovirus (KBS2A) that infects a marine Synechococcus sp. (strain WH7803). Unique to this genome, relative to other sequenced cyanosiphoviruses, is the absence of elements associated with integration into the host chromosome, suggesting this virus may not be able to establish a lysogenic relationship.
Proceedings of the National Academy of Sciences, 2014
The supply and bioavailability of dissolved iron sets the magnitude of surface productivity for ∼... more The supply and bioavailability of dissolved iron sets the magnitude of surface productivity for ∼40% of the global ocean. The redox state, organic complexation, and phase (dissolved versus particulate) of iron are key determinants of iron bioavailability in the marine realm, although the mechanisms facilitating exchange between iron species (inorganic and organic) and phases are poorly constrained. Here we use the isotope fingerprint of dissolved and particulate iron to reveal distinct isotopic signatures for biological uptake of iron during a GEOTRACES process study focused on a temperate spring phytoplankton bloom in subtropical waters. At the onset of the bloom, dissolved iron within the mixed layer was isotopically light relative to particulate iron. The isotopically light dissolved iron pool likely results from the reduction of particulate iron via photochemical and (to a lesser extent) biologically mediated reduction processes. As the bloom develops, dissolved iron within the surface mixed layer becomes isotopically heavy, reflecting the dominance of biological processing of iron as it is removed from solution, while scavenging appears to play a minor role. As stable isotopes have shown for major elements like nitrogen, iron isotopes offer a new window into our understanding of the biogeochemical cycling of iron, thereby allowing us to disentangle a suite of concurrent biotic and abiotic transformations of this key biolimiting element.
Macronutrients in sinking phytoplankton are typically remineralized at different rates, but less ... more Macronutrients in sinking phytoplankton are typically remineralized at different rates, but less is known about the fate of micronutrient metals associated with sinking cells. Scavenging, the presence of co-occurring abiotic particles, and inadvertent contamination limit the utility of bulk analytical approaches to study remineralization of trace metals in sinking phytoplankton. We used synchrotron x-ray fluorescence mapping to measure macronutrients (P, S, and Si) and trace metals (Fe, Ni, and Zn) in individual cells of the diatom Asterionellopsis glacialis during a spring bloom in subtropical waters off New Zealand. P, S, Zn, and Ni were released significantly faster than Fe and Si from sinking cells in the upper 200 m. Bulk particulate element fluxes to sediment traps indicated similar trends, but biogenic silica flux was attenuated much faster than Si was lost from intact sinking cells collected in the traps. The metals were spatially co-located with P and S in upper ocean cells, but this association with P and S (based on a spatial resolution of 450 nm) was largely absent in sinking cells. In contrast, Fe retained a weak spatial association with Si, suggesting that remineralized Fe may be re-scavenged onto cell surfaces. As a result, dissolved Fe : macronutrient stoichiometries in the water column likely underestimate stoichiometries in sinking cells. We propose linkages between the selective loss of diatom cellular components (e.g., ribosomes or phospholipid membranes, Zn-finger proteins, and urease) and the observed recycling of specific elements (P, Zn, and Ni, respectively), which set the stoichiometry of macro-and micronutrient supply to surface waters.
| In marine environments, virus-mediated lysis of host cells leads to the release of cellular car... more | In marine environments, virus-mediated lysis of host cells leads to the release of cellular carbon and nutrients and is hypothesized to be a major driver of carbon recycling on a global scale. However, efforts to characterize the effects of viruses on nutrient cycles have overlooked the geochemical potential of the virus particles themselves, particularly with respect to their phosphorus content. In this Analysis article, we use a biophysical scaling model of intact virus particles that has been validated using sequence and structural information to quantify differences in the elemental stoichiometry of marine viruses compared with their microbial hosts. By extrapolating particle-scale estimates to the ecosystem scale, we propose that, under certain circumstances, marine virus populations could make an important contribution to the reservoir and cycling of oceanic phosphorus.
Marker genes for viruses are typically amplified from aquatic samples to determine whether specif... more Marker genes for viruses are typically amplified from aquatic samples to determine whether specific viruses are present in the sample, or to examine the diversity of a group of related viruses. In this chapter, we will provide an overview of common methods used to amplify, clone, sequence, and analyze virus marker genes, and will focus our discussion on viruses infecting algae, bacteria, and heterotrophic flagellates. Within this chapter, we endeavor to highlight critical aspects and components of these methods. To this end, instead of providing a detailed experimental protocol for each of the steps involved in examining virus marker gene libraries, we have provided a few key considerations, recommendations, and options for each step. We conclude this chapter with a brief discussion of research on a major capsid protein (g20) of cyanomyoviruses using this work as a case study for polymerase chain reaction primer design and development. By building on the experience of numerous labs, this chapter should not only be useful to the new virus ecologist, but also serve as a valuable resource to established research groups.
Microcystins are secondary metabolites produced by cyanobacteria that act as hepatotoxins in high... more Microcystins are secondary metabolites produced by cyanobacteria that act as hepatotoxins in higher organisms. These toxins can be altered through abiotic processes, such as photodegradation and adsorption, as well as through biological processes via metabolism and bacterial degradation. Some species of bacteria can degrade microcystins, and many other organisms metabolize microcystins into a series of conjugated products. There are toxicokinetic models used to examine microcystin uptake and elimination, which can be difficult to compare due to differences in compartmentalization and speciation. Metabolites of microcystins are formed as a detoxification mechanism, and little is known about how quickly these metabolites are formed. In summary, microcystins can undergo abiotic and biotic processes that alter the toxicity and structure of the microcystin molecule. The environmental impact and toxicity of these alterations and the metabolism of microcystins remains uncertain, making it difficult to establish guidelines for human health. Here, we present the current state of knowledge regarding the alterations microcystins can undergo in the environment.
Trichodesmium provides new nitrogen (N) to marine surface waters via N 2 fixation, a process that... more Trichodesmium provides new nitrogen (N) to marine surface waters via N 2 fixation, a process that requires a substantial amount of iron (Fe). Organic ligands in seawater that bind Fe could either increase or reduce the bioavailability of Fe. Electrochemical techniques indicate that these naturally occurring ligands have Fe-binding constants similar to those of siderophores and porphyrins, suggesting that these chelators play an important role in determining the bioavailability of Fe to cyanobacteria. We conducted Fe uptake experiments using model ligands labeled with 55 Fe to compare the bioavailability of inorganic Fe(III), porphyrin-bound Fe(III), and siderophorebound Fe(III) to field-collected Trichodesmium colonies. Inorganic Fe(III) and siderophore-bound Fe(III) were more bioavailable to Trichodesmium colonies than was porphyrin-bound Fe(III). Furthermore, the bioavailability of the siderophore-bound Fe(III) can be characterized by the functional groups of the siderophore. The dihydroxamate siderophore and an uncharacterized ligand from a cultured Synechococcus sp. increased the bioavailability of Fe compared to the trihydroxamate siderophores. Except for experiments with desferrioxamine B, dark incubations resulted in lower Fe uptake rates for all treatments, relative to parallel lighted incubations. This suggests that light enhances the photochemical dissociation of most of the ligand complexes or that light energy is required for the active transport of Fe complexed to the model ligands. The Fe uptake rate of Trichodesmium colonies also differed slightly on the basis of colony morphology, with higher uptake rates with ''puffs'' than ''tufts.'' These experiments show that Trichodesmium colonies are capable of discriminating between Fe bound to different organic complexes.
Viral lysis of phytoplankton constrains marine primary production, food web dynamics and biogeoch... more Viral lysis of phytoplankton constrains marine primary production, food web dynamics and biogeochemical cycles in the ocean. Yet, little is known about the biogeographical distribution of viral lysis rates across the global ocean. To address this, we investigated phytoplankton group-specific viral lysis rates along a latitudinal gradient within the North Atlantic Ocean. The data show large-scale distribution patterns of different virus groups across the North Atlantic that are associated with the biogeographical distributions of their potential microbial hosts. Average virus-mediated lysis rates of the picocyanobacteria Prochlorococcus and Synechococcus were lower than those of the picoeukaryotic and nanoeukaryotic phytoplankton (that is, 0.14 per day compared with 0.19 and 0.23 per day, respectively). Total phytoplankton mortality (virus plus grazer-mediated) was comparable to the gross growth rate, demonstrating high turnover rates of phytoplankton populations. Virus-induced morta...
Background/Question/Methods Bacteria typically live in environments which are depleted in resourc... more Background/Question/Methods Bacteria typically live in environments which are depleted in resources such as nitrogen (N) and phosphorus (P). These organisms have advanced nutrient uptake and retention mechanisms to cope with nutrient limiting situations, but these same structural features may be sites of attack by viral predators (i.e., phage). Phage play an important role in the ‘top-down’ regulation and structuring of bacterial communities, but also influence the evolutionary trajectory of host populations on ecologically relevant time scales. The interaction between virus predation and nutrient limitation, however, has not been extensively examined within an evolutionary context. Using the marine cyanobacterium, Synechococcus sp. WH 7803 and a T4-like phage, we explored the fitness tradeoffs and eco-evolutionary dynamics associated with phage attack and nutrient limitation. Through an eight month chemostat experiment, we manipulated virus exposure and the N:P supply ratio to crea...
Here we examined the impact of a commonly employed method used to measure nitrogen fixation, the ... more Here we examined the impact of a commonly employed method used to measure nitrogen fixation, the acetylene reduction assay (ARA), on a marine sediment community. Historically, the ARA technique has been broadly employed for its ease of use, in spite of numerous known artifacts. To gauge the severity of these effects in a natural environment, we employed high-throughput 16S rRNA gene sequencing to detect differences in acetylene-treated sediments vs. non-treated control sediments after a 7 h incubation. Within this short time period, significant differences were seen across all activity of microbes identified in the sediment, implying that the changes induced by acetylene occur quickly. The results have important implications for our understanding of marine nitrogen budgets. Moreover, because the ARA technique has been widely used in terrestrial and freshwater habitats, these results may be applicable to other ecosystems.
We announce the complete genome sequences of two temperate Podoviridae, Sulfitobacter phages ΦCB2... more We announce the complete genome sequences of two temperate Podoviridae, Sulfitobacter phages ΦCB2047-A and ΦCB2047-C, which infect Sulfitobacter sp. strain 2047, a member of the Roseobacter clade. This is the first report of temperate podophage infecting members of the Sulfitobacter genus of the Roseobacter clade.
Little is known about the molecular and physiological function of co-occurring microbes within fr... more Little is known about the molecular and physiological function of co-occurring microbes within freshwater cyanobacterial harmful algal blooms (cHABs). To address this, community metatranscriptomes collected from the western basin of Lake Erie during August 2012 were examined. Using sequence data, we tested the hypothesis that the activity of the microbial community members is independent of community structure. Predicted metabolic and physiological functional profiles from spatially distinct metatranscriptomes were determined to be >90% similar between sites. Targeted analysis of Microcystis aeruginosa, the historical causative agent of cyanobacterial harmful algal blooms over the past ϳ20 years, as well as analysis of Planktothrix agardhii and Anabaena cylindrica, revealed ongoing transcription of genes involved in microcystin toxin synthesis as well as the acquisition of both nitrogen and phosphorus, nutrients often implicated as independent bottom-up drivers of eutrophication in aquatic systems. Transcription of genes involved in carbon dioxide (CO 2 ) concentration and metabolism also provided support for the alternate hypothesis that high-pH conditions and dense algal biomass result in CO 2 -limiting conditions that further favor cyanobacterial dominance. Additionally, the presence of Microcystis-specific cyanophage sequences provided preliminary evidence of possible top-down virus-mediated control of cHAB populations. Overall, these data provide insight into the complex series of constraints associated with Microcystis blooms that dominate the western basin of Lake Erie during summer months, demonstrating that multiple environmental factors work to shape the microbial community. Citation Steffen MM, Belisle BS, Watson SB, Boyer GL, Bourbonniere RA, Wilhelm SW. 2015. Metatranscriptomic evidence for co-occurring top-down and bottomup controls on toxic cyanobacterial communities. Appl Environ Microbiol 81:3268 -3276.
We announce the complete genome sequence of a lytic podovirus, ⌽CB2047-B, which infects the bacte... more We announce the complete genome sequence of a lytic podovirus, ⌽CB2047-B, which infects the bacterium Sulfitobacter sp. strain 2047, a member of the Roseobacter clade. Genome analysis revealed ⌽CB2047-B to be an N4-like phage, with its genome having high nucleotide similarity to other N4-like roseophage genomes.
We present the genome of a cyanosiphovirus (KBS2A) that infects a marine Synechococcus sp. (strai... more We present the genome of a cyanosiphovirus (KBS2A) that infects a marine Synechococcus sp. (strain WH7803). Unique to this genome, relative to other sequenced cyanosiphoviruses, is the absence of elements associated with integration into the host chromosome, suggesting this virus may not be able to establish a lysogenic relationship.
Proceedings of the National Academy of Sciences, 2014
The supply and bioavailability of dissolved iron sets the magnitude of surface productivity for ∼... more The supply and bioavailability of dissolved iron sets the magnitude of surface productivity for ∼40% of the global ocean. The redox state, organic complexation, and phase (dissolved versus particulate) of iron are key determinants of iron bioavailability in the marine realm, although the mechanisms facilitating exchange between iron species (inorganic and organic) and phases are poorly constrained. Here we use the isotope fingerprint of dissolved and particulate iron to reveal distinct isotopic signatures for biological uptake of iron during a GEOTRACES process study focused on a temperate spring phytoplankton bloom in subtropical waters. At the onset of the bloom, dissolved iron within the mixed layer was isotopically light relative to particulate iron. The isotopically light dissolved iron pool likely results from the reduction of particulate iron via photochemical and (to a lesser extent) biologically mediated reduction processes. As the bloom develops, dissolved iron within the surface mixed layer becomes isotopically heavy, reflecting the dominance of biological processing of iron as it is removed from solution, while scavenging appears to play a minor role. As stable isotopes have shown for major elements like nitrogen, iron isotopes offer a new window into our understanding of the biogeochemical cycling of iron, thereby allowing us to disentangle a suite of concurrent biotic and abiotic transformations of this key biolimiting element.
Macronutrients in sinking phytoplankton are typically remineralized at different rates, but less ... more Macronutrients in sinking phytoplankton are typically remineralized at different rates, but less is known about the fate of micronutrient metals associated with sinking cells. Scavenging, the presence of co-occurring abiotic particles, and inadvertent contamination limit the utility of bulk analytical approaches to study remineralization of trace metals in sinking phytoplankton. We used synchrotron x-ray fluorescence mapping to measure macronutrients (P, S, and Si) and trace metals (Fe, Ni, and Zn) in individual cells of the diatom Asterionellopsis glacialis during a spring bloom in subtropical waters off New Zealand. P, S, Zn, and Ni were released significantly faster than Fe and Si from sinking cells in the upper 200 m. Bulk particulate element fluxes to sediment traps indicated similar trends, but biogenic silica flux was attenuated much faster than Si was lost from intact sinking cells collected in the traps. The metals were spatially co-located with P and S in upper ocean cells, but this association with P and S (based on a spatial resolution of 450 nm) was largely absent in sinking cells. In contrast, Fe retained a weak spatial association with Si, suggesting that remineralized Fe may be re-scavenged onto cell surfaces. As a result, dissolved Fe : macronutrient stoichiometries in the water column likely underestimate stoichiometries in sinking cells. We propose linkages between the selective loss of diatom cellular components (e.g., ribosomes or phospholipid membranes, Zn-finger proteins, and urease) and the observed recycling of specific elements (P, Zn, and Ni, respectively), which set the stoichiometry of macro-and micronutrient supply to surface waters.
| In marine environments, virus-mediated lysis of host cells leads to the release of cellular car... more | In marine environments, virus-mediated lysis of host cells leads to the release of cellular carbon and nutrients and is hypothesized to be a major driver of carbon recycling on a global scale. However, efforts to characterize the effects of viruses on nutrient cycles have overlooked the geochemical potential of the virus particles themselves, particularly with respect to their phosphorus content. In this Analysis article, we use a biophysical scaling model of intact virus particles that has been validated using sequence and structural information to quantify differences in the elemental stoichiometry of marine viruses compared with their microbial hosts. By extrapolating particle-scale estimates to the ecosystem scale, we propose that, under certain circumstances, marine virus populations could make an important contribution to the reservoir and cycling of oceanic phosphorus.
Marker genes for viruses are typically amplified from aquatic samples to determine whether specif... more Marker genes for viruses are typically amplified from aquatic samples to determine whether specific viruses are present in the sample, or to examine the diversity of a group of related viruses. In this chapter, we will provide an overview of common methods used to amplify, clone, sequence, and analyze virus marker genes, and will focus our discussion on viruses infecting algae, bacteria, and heterotrophic flagellates. Within this chapter, we endeavor to highlight critical aspects and components of these methods. To this end, instead of providing a detailed experimental protocol for each of the steps involved in examining virus marker gene libraries, we have provided a few key considerations, recommendations, and options for each step. We conclude this chapter with a brief discussion of research on a major capsid protein (g20) of cyanomyoviruses using this work as a case study for polymerase chain reaction primer design and development. By building on the experience of numerous labs, this chapter should not only be useful to the new virus ecologist, but also serve as a valuable resource to established research groups.
Microcystins are secondary metabolites produced by cyanobacteria that act as hepatotoxins in high... more Microcystins are secondary metabolites produced by cyanobacteria that act as hepatotoxins in higher organisms. These toxins can be altered through abiotic processes, such as photodegradation and adsorption, as well as through biological processes via metabolism and bacterial degradation. Some species of bacteria can degrade microcystins, and many other organisms metabolize microcystins into a series of conjugated products. There are toxicokinetic models used to examine microcystin uptake and elimination, which can be difficult to compare due to differences in compartmentalization and speciation. Metabolites of microcystins are formed as a detoxification mechanism, and little is known about how quickly these metabolites are formed. In summary, microcystins can undergo abiotic and biotic processes that alter the toxicity and structure of the microcystin molecule. The environmental impact and toxicity of these alterations and the metabolism of microcystins remains uncertain, making it difficult to establish guidelines for human health. Here, we present the current state of knowledge regarding the alterations microcystins can undergo in the environment.
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Papers by Steven Wilhelm