Arsenobetaine in Atlantic salmon (Salmo salar L.): influence of seawater adaptation

Glycine betaine has been suggested to improve the maintenance of ionic and osmotic homeostasis during seawater adaptation in teleost fish. Arsenobetaine may also behave as an osmolyte, due to its structural similarity to glycine betaine. The influence of seawater adaptation on intestinal uptake and muscle accumulation of arsenobetaine in the teleost Atlantic salmon (Salmo salar L.) was investigated. Atlantic salmon (freshwater and seawater adapted) were given a single oral dose of arsenobetaine, which was absorbed over the intestine within 6 h after exposure. Seawater adapted Atlantic salmon had significantly higher levels of accumulated arsenobetaine in blood compared to the freshwater adapted salmon. However, seawater adaptation had no effect on the levels of accumulated arsenobetaine in muscle tissue. Similar retention of the administered dose was found in muscle tissue in both freshwater and seawater adapted salmon, with 49+/-6% and 50+/-10% retention after 144 h, respectively. Results indicate that muscle retention was not influenced by salinity in seawater adapting teleosts.     

Arsenic accumulation in livers of pinnipeds, seabirds and sea turtles: subcellular distribution and interaction between arsenobetaine and glycine betaine

Concentrations of total arsenic and individual arsenic compounds were determined in liver samples of pinnipeds (northern fur seal Callorhinus ursinus and ringed seal Pusa hispida), seabirds (black-footed albatross Diomedea nigripes and black-tailed gull Larus crassirostris) and sea turtles (hawksbill turtle Eretmochelys imbricata and green turtle Chelonia mydas). Among these species, the black-footed albatross contained the highest hepatic arsenic concentration (5.8+/-3.7 microg/g wet mass). Arsenobetaine was the major arsenic species found in the liver of all these higher tropic marine animals. To investigate the cause of high accumulation of arsenobetaine, subcellular distribution of arsenic and relationship between arsenobetaine and glycine betaine concentrations were examined in the livers of these animals. There was no relationship between total arsenic concentration and its subcellular distribution in liver tissues. However, a significant negative correlation was found between arsenobetaine and glycine betaine concentrations in the liver of six species examined. This result may indicate that arsenobetaine is accumulated in these marine animals as an osmolyte along with glycine betaine, which is a predominant osmolyte in marine animals because the chemical structure and properties of arsenobetaine are similar to those of glycine betaine.     

Marine intrusion and freshwater discharge as opposite forces driving fish guilds distribution along coastal plain streams

This work investigated variations in fish guilds along marine surf zone to freshwater reaches of coastal plain streams and their relationships with environmental factors. Fish and abiotic data were collected monthly during 1 year and an automatic image-recording system was used to monitor marine intrusion events. Aside ephemeral salinity gradients produced by storm surges, freshwater conditions prevailed inside streams. Despite of that, fish habitat use guilds were spatially distributed according to their salinity tolerance, with marine species occurring mainly in the stream site near the adjacent surf zone and non-salinity tolerant freshwater fish in the upstream site. Marine intrusion was the main factor correlated with the entrance of marine-related fish into streams during summer. In contrast, higher rainfall during colder months prevented the dominance of marine species. This work highlighted that spatial segregation in fish habitat use guild could occur even in the absence of long-lasting salinity gradients, since active colonization by euryhaline fish and sporadic marine intrusions can lead to the occurrence of marine-related fish in coastal plain streams. Future studies should evaluate if this marine intrusion role on fish of coastal plain streams would be affected by changes in sea level and rainfall in a global warming scenario.     

An origin for arsenobetaine involving bacterial formation of an arsenic-carbon bond

Lysed-cell extract of a Pseudomonas sp. was shown to catalyse bioconversion of dimethylarsinoylacetate to arsenobetaine and dimethylarsinate. Provision of the universal methyl donor S-adenosylmethionine to bioconversion mixtures promoted both the rate and extent of arsenobetaine formation. These findings suggest that in the proposed biosynthesis of arsenobetaine from dimethylarsinoylethanol, oxidation (i.e. the formation of the carboxymethyl group of dimethylarsinoylacetate) would precede the reduction and methylation at the arsenic atom. The presence of enzyme(s) capable of methylating dimethylarsinoylacetate in a bacterial isolate from marine mussel (Mylitus edulis), highlights a possible direct involvement of prokaryotic organisms in the biosynthesis of organoarsenic compounds within marine animals.     

Why are there so few freshwater fish species in most estuaries?

The freshwater fish assemblage in most estuaries is not as species rich as the marine assemblage in the same systems. Coupled with this differential richness is an apparent inability by most freshwater fish species to penetrate estuarine zones that are mesohaline (salinity: 5·0–17·9), polyhaline (salinity: 18·0–29·9) or euhaline (salinity: 30·0–39·9). The reason why mesohaline waters are avoided by most freshwater fishes is difficult to explain from a physiological perspective as many of these species would be isosmotic within this salinity range. Perhaps, a key to the poor penetration of estuarine waters by freshwater taxa is an inability to develop chloride cells in gill filament epithelia, as well as a lack of other osmoregulatory adaptations present in euryhaline fishes. Only a few freshwater fish species, especially some of those belonging to the family Cichlidae, have become fully euryhaline and have successfully occupied a wide range of estuaries, sometimes even dominating in hyperhaline systems (salinity 40+). Indeed, this review found that there are few fish species that can be termed holohaline (i.e. capable of occupying waters with a salinity range of 0–100+) and, of these taxa, there is a disproportionally high number of freshwater species (e.g. Cyprinodon variegatus, Oreochromis mossambicus and Sarotherodon melanotheron). Factors such as increased competition for food and higher predation rates by piscivorous fishes and birds may also play an important role in the low species richness and abundance of freshwater taxa in estuaries. Added to this is the relatively low species richness of freshwater fishes in river catchments when compared with the normally higher diversity of marine fish species for potential estuarine colonization from the adjacent coastal waters. The almost complete absence of freshwater fish larvae from the estuarine ichthyoplankton further reinforces the poor representation of this guild within these systems. An explanation as to why more freshwater fish species have not become euryhaline and occupied a wide range of estuaries similar to their marine counterparts is probably due to a combination of the above described factors, with physiological restrictions pertaining to limited salinity tolerances probably playing the most important role.     

Comparative approaches to understand metal bioaccumulation in aquatic animals

Over the past decades, comparative physiology and biochemistry approaches have played a significant role in understanding the complexity of metal bioaccumulation in aquatic animals. Such a comparative approach is now further aided by the biokinetic modeling approach which can be used to predict the rates and routes of metal bioaccumulation and assist in the interpretation of accumulated body metal concentrations in aquatic animals. In this review, we illustrate a few examples of using the combined comparative and biokinetic modeling approaches to further our understanding of metal accumulation in aquatic animals. We highlight recent studies on the different accumulation patterns of metals in different species of invertebrates and fish, and between various aquatic systems (freshwater and marine). Comparative metal biokinetics can explain the differences in metal bioaccumulation among bivalves, although it is still difficult to explain the evolutionary basis for the different accumulated metal body concentrations (e.g., why some species have high metal concentrations). Both physiological/biochemical responses and metal geochemistry are responsible for the differences in metal concentrations observed in different populations of aquatic species, or between freshwater and marine species. A comparative approach is especially important for metal biology research, due to the very complicated and potentially variable physiological handling of metals during their accumulation, sequestration, distribution and elimination in different aquatic species or between different aquatic systems.     

Small Changes in Gene Expression of Targeted Osmoregulatory Genes When Exposing Marine and Freshwater Threespine Stickleback (Gasterosteus aculeatus) to Abrupt Salinity Transfers

Salinity is one of the key factors that affects metabolism, survival and distribution of fish species, as all fish osmoregulate and euryhaline fish maintain osmotic differences between their extracellular fluid and either freshwater or seawater. The threespine stickleback (Gasterosteus aculeatus) is a euryhaline species with populations in both marine and freshwater environments, where the physiological and genomic basis for salinity tolerance adaptation is not fully understood. Therefore, our main objective in this study was to investigate gene expression of three targeted osmoregulatory genes (Na⁺/K⁺-ATPase (ATPA13), cystic fibrosis transmembrane regulator (CFTR) and a voltage gated potassium channel gene (KCNH4) and one stress related heat shock protein gene (HSP70)) in gill tissue from marine and freshwater populations when exposed to non-native salinity for periods ranging from five minutes to three weeks. Overall, the targeted genes showed highly plastic expression profiles, in addition the expression of ATP1A3 was slightly higher in saltwater adapted fish and KCNH4 and HSP70 had slightly higher expression in freshwater. As no pronounced changes were observed in the expression profiles of the targeted genes, this indicates that the osmoregulatory apparatuses of both the marine and landlocked freshwater stickleback population have not been environmentally canalized, but are able to respond plastically to abrupt salinity challenges.     

A Novel Kauranoid Isolated from Gibberella fujikuroi

A novel arseno-sugar was isolated from the brown alga Sargassum thunbergii. Instead of the dimethylarsinoyl group reported for algal arseno-sugars, this has a tri-methylarsonium group, which is borne by arsenobetaine, a ubiquitous organoarsenic compound in marine animals. This may be an intermediate between arseno-sugars and arsenobetaine.     

On the conspecificity of marine and freshwater Bangia in Britain

The salinity responses of marine and freshwater Bangia have been studied in laboratory culture. Although both isolates possessed salinity tolerances which were dependent upon the salinity regimes of their original habitats, neither was restricted in its salinity tolerances to that salinity range alone. The marine isolate was able to proliferate after a single step transfer into a freshwater-based medium. Certain cells within the thallus appeared to be resistant to the sudden decrease in salinity and these developed into dwarf plantlets, following a period of dormancy. An hypothesis is proposed that Bangia occurring in freshwater is a well-adapted ecotype of the marine form, thus supporting the conspecific theory.     

Life History Traits of Two Chaetonotids (Gastrotricha) Under Different Experimental Conditions

Summary

A marine and a freshwater species of Chaetonotida (Gastrotricha) were reared under laboratory conditions. Their life tables and principal demographic parameters were determined at 2 different temperatures (20°and 25°C). At 25°C the data relative to the marine species were collected from 5 cohorts kept at 5 different salinity levels (15, 25, 35, 45, 55°/oo).

A higher temperature increases reproductive activity while shortening its duration in both species, whereas the length of the lifespan remains unaffected. Extreme salinity values (15 and 55°/oo) reduce the maximum longevity of the marine species and have opposite effects on reproductive activity, which is higher at low salinity and becomes lower at high salinity. The postparthenogenetic phase is remarkably long relative to the life cycle: this was observed in all experimental conditions and may be related to the existence of a second reproductive phase, which is hermaphroditic and follows the parthenogenetic one, as recently postulated from morphological data.     

The salinity preference of members of the genus Macroplea (Coleoptera, Chrysomelidae, Donaciinae), fully aquatic leaf beetles that occur in brackish water

Reed beetles (Donaciinae) of the genus Macroplea Samouelle, 1819 live permanently submerged. Literature indicates that Macroplea mutica occurs in brackish water, whereas Macroplea appendiculata is restricted to freshwater. The salinity preference of these two species was tested in a linear and a circular device that offered a continuous salinity gradient. The distribution of animals in the devices was monitored over at least 3 h in each of the 21 experiments. Both species preferred freshwater (salinity 0) over brackish water (salinity 10). In particular, this holds true for specimens collected in brackish water. Likewise, immediate reactions could be observed when during such experiments the direction of the gradient was reversed. While M. mutica can be regarded as a truly marine insect, this marine environment does not strictly reflect its fundamental niche with respect to salinity preference. This is in line with accumulating evidence that M. mutica can be found in freshwater habitats (and M. appendiculata in brackish water). This indicates that the species’ distribution might be influenced by other factors like host plant preference or dispersal mechanisms. It is discussed if—in spite of similar fundamental niches—differences in salinity tolerance (and hence performance in brackish water) may have contributed to speciation in the genus Macroplea.     

Phylogeography and environmental diversification of a highly adaptable marine amphipod, Gammarus duebeni

Genetic diversity and phylogeographic population structure in the gammarid amphipod, Gammarus duebeni, were investigated across its broad latitudinal distribution in the NE and NW Atlantic by analysis of mitochondrial DNA sequence. Gammarus duebeni has exceptional tolerance of salinity change and inhabits environments ranging from marine to freshwater. The longstanding debate on whether there are distinct marine and freshwater subspecies was assessed by sampling populations from sites characterized by different salinities. Our sequence data demonstrates that there are two major lineages, with little internal geographic structuring. Evidence is provided to suggest a pre-glacial divergence of these two clades, involving segregation between a region historically associated with the freshwater form and the majority of the marine localities on both sides of the Atlantic. A modern contact zone between the marine and freshwater forms is proposed in western Britain.     

Sulfur and phytoplankton: acquisition, metabolism and impact on the environment

Sulfur emission from marine phytoplankton has been recognized as an important factor for global climate and as an entry into the biogeochemical S cycle. Despite this significance, little is known about the cellular S metabolism in algae that forms the basis of this emission. Some biochemical and genetic evidence for regulation of S uptake and assimilation is available for the freshwater model alga Chlamydomonas. However, the marine environment is substantially different from most fresh waters, containing up to 50 times higher free sulfate concentrations and challenging the adaptive mechanisms of primary and secondary S metabolism in marine algae. This review intends to integrate ecological and physiological data to provide a comprehensive view of the role of S in the oceans.     

The osmorespiratory compromise in sculpins: impaired gas exchange is associated with freshwater tolerance

We acclimated two species of sculpin, the freshwater prickly sculpin (Cottus asper) and the closely related marine Pacific staghorn sculpin (Leptocottus armatus) to freshwater ( approximately 0 g/L), brackish water (15 g/L), and seawater (30 g/L) for at least 4 wk and examined the relationships between respiration, ion regulation, gill morphology, and freshwater tolerance. The prickly sculpin successfully acclimated to all three salinities and did not experience appreciable changes in plasma osmolality, [Cl-], or mortality. Gill Na+/K+-ATPase activity was lowest in prickly sculpins acclimated to freshwater, their native salinity, and increased during acclimation to seawater. Furthermore, prickly sculpins acclimated to freshwater had a 30% higher P(crit) than fish acclimated to brackish water or seawater; P(crit) is the environmental P(O2) below which an animal can no longer maintain a routine (.-)M(O2), and an increase in P(crit) represents a compromise of respiratory gas exchange. The higher P(crit) observed in prickly sculpins acclimated to freshwater is likely a consequence of their having small, relatively thick gills that increase in thickness (by approximately 1 microm) during freshwater exposure. In contrast, the marine Pacific staghorn sculpin successfully acclimated to brackish water and seawater, but high mortality (25%) was observed after 3 wk of exposure to freshwater. Pacific staghorn sculpins exposed to freshwater suffered significant, 15%-20%, reductions in plasma osmolality and [Cl-], and these losses in plasma ions resulted in a 1.4-fold increase in gill Na+/K+-ATPase activity. Pacific staghorn sculpins have large, thin gills that are not modified in response to salinity acclimation, and as a result, these animals show no respiratory compromise during freshwater acclimation, as evidenced by the lack of change in P(crit), but show significant ion regulatory disturbance. Overall, this study suggests that gill thickening and the resulting respiratory compromise are necessary for freshwater tolerance in sculpins.     

Evolution of growth by genetic accommodation in Icelandic freshwater stickleback

Classical Darwinian adaptation to a change in environment can ensue when selection favours beneficial genetic variation. How plastic trait responses to new conditions affect this process depends on how plasticity reveals to selection the influence of genotype on phenotype. Genetic accommodation theory predicts that evolutionary rate may sharply increase when a new environment induces plastic responses and selects on sufficient genetic variation in those responses to produce an immediate evolutionary response, but natural examples are rare. In Iceland, marine threespine stickleback that have colonized freshwater habitats have evolved more rapid individual growth. Heritable variation in growth is greater for marine full-siblings reared at low versus high salinity, and genetic variation exists in plastic growth responses to low salinity. In fish from recently founded freshwater populations reared at low salinity, the plastic response was strongly correlated with growth. Plasticity and growth were not correlated in full-siblings reared at high salinity nor in marine fish at either salinity. In well-adapted lake populations, rapid growth evolved jointly with stronger plastic responses to low salinity and the persistence of strong plastic responses indicates that growth is not genetically assimilated. Thus, beneficial plastic growth responses to low salinity have both guided and evolved along with rapid growth as stickleback adapted to freshwater.     

Attached and Free-Floating Bacteria in a Diverse Selection of Water Bodies

The contribution of attached and free-floating bacteria to the bacterial numbers and heterotrophic uptake in 44 diverse aquatic environments was studied. A factor analysis reduced the variability of the raw data base to three major factors explaining 53.6% of total variance. These factors were (i) salinity, (ii) heterotrophic uptake, and (iii) particulate load. A cluster analysis categorized the 44 habitats into five distinct environmental types based on these three factors. There was no significant pattern in the distribution of attached versus free-floating bacteria when assessed by epifluorescent microscopy. However the contribution of attached bacteria to the uptake of an amino acids mix was reduced in marine waters. Heavy particulate loads resulted in an increased percentage uptake of amino acids and glucose from the attached bacteria. Uptake response was found to be substrate specific especially in oliogotrophic freshwater. Amino acid uptake was more associated with the attached fraction, whereas glucose uptake was mediated more by the free-floating fraction.     

Arsenic in Seafood: Speciation Issues for Human Health Risk Assessment

Major sources of arsenic exposure for humans are foods, particularly aquatic organisms, which are called seafood in this report. Although seafood contains a variety of arsenicals, including inorganic arsenic, which is toxic and carcinogenic, and arsenobetaine, which is considered nontoxic, the arsenic content of seafood commonly is reported only as total arsenic. A goal of this literature survey is to determine if generalizable values can be derived for the percentage of total arsenic in seafood that is inorganic arsenic. Generalizable values for percent inorganic arsenic are needed for use as default values in U.S. human health risk assessments of seafood from arsenic-contaminated sites. Data from the worldwide literature indicate the percent of inorganic arsenic in marine/estuarine finfish does not exceed 7.3% and in shellfish can reach 25% in organisms from presumably uncontaminated areas, with few data available for freshwater organisms. However, percentages can be much higher in organisms from contaminated areas and in seaweed. U.S. site-specific data for marine/estuarine finfish and shellfish are similar to the worldwide data, and for freshwater finfish indicate that the average percent inorganic arsenic is generally < 10%, but ranges up to nearly 30%. Derivation of nationwide defaults for percent inorganic arsenic in fish, shellfish, and seaweed collected from arsenic-contaminated areas in the United States is not supported by the surveyed literature.     

Cryptic freshwater amoeba species in the bottom sediments of Nivå Bay (Øresund, Baltic Sea)

An aquarium containing a portion of freshly collected brackish-water sediment from Nivå Bay (Øresund, Baltic Sea) was sampled to determine the species of amoebae present and refilled with inorganic salt medium to achieve freshwater salinity. After 2 months incubation, the diversity of amoebae was again studied, and the salinity was restored to the original level. The aquarium was incubated for a further 2 months and the amoeba fauna was studied for the last time. A number of freshwater species appeared in the sample after the first salinity shift, while some marine species disappeared. Most marine species did not re-appear after the salinity in the aquarium was restored, but one marine species not noted previously was recorded. The experiment illustrates the presence of ‘cryptic diversity’ of amoebae in natural habitats and demonstrates that laboratory manipulation of the salinity of a sample prior to inoculation may achieve an increased recovery of species from a brackish-water habitat.