Mytilid mussels: global habitat engineers in coastal sediments

Dense beds of mussels of the family Mytilidae occur worldwide on soft-bottoms in cold and warm temperate coastal waters and have usually been considered hot spots of biodiversity. We examined intertidal mussel beds at four distant locations around the globe with the same sampling method, to find out whether this “hot spot” designation holds universally. We studied species assemblages within the matrices of byssally interconnected mussels engineered by Mytilus edulis in the North Sea, by mixed Perumytilus purpuratus and Mytilus chilensis at the southern Chilean coast, by Musculista senhousia in the Yellow Sea and by Xenostrobus inconstans at the coast of southern Australia. In all cases, species assemblages inside mussel beds were significantly different from those outside with many species being restricted to one habitat type. However, species richness and diversity were not generally higher in mussel beds than in ambient sediments without mussels. In the North Sea (M. edulis) and at the Chilean coast (P. purpuratus, M. chilensis), mussel beds have markedly higher species numbers and diversities than surrounding sediments, but this was not the case for mussel beds in Australia (X. inconstans) and the Yellow Sea (M. senhousia) where numbers of associated species were only slightly higher and somewhat lower than in adjacent sediments, respectively. In conclusion, although soft bottom mytilid mussels generally enhance habitat heterogeneity and species diversity at the ecosystem level, mussel beds themselves are not universal centres of biodiversity, but the effects on associated species are site specific.     

Habitat structure is more important than nutrient supply in modifying mussel bed assemblage in an upwelling area of the Peruvian coast

Upwelling intensity modifies coastal primary production and influences individual traits of habitat-forming species. Along the Peruvian coast, beds of the mytilid Perumytilus purpuratus provide structurally complex habitats that harbour many organisms. We predict that in the nutrient-rich system of Central Peru, the modification of structural complexity would have stronger effects on the Perumytilus community than nutrient addition. We experimentally examined the effects of nutrient addition on the Perumytilus-dominated assemblage and we evaluated the potential effect of varying shell size on the Perumytilus-dominated assemblage. Nutrient addition to the mussel bed with slow-release fertilizers caused no changes in the total macro- and microalgal biomass and did not affect abundances or composition of the assemblage. To explore the effect of structural complexity on the Perumytilus assemblage, we manipulated mussel size with experimental bags containing small and large individuals. Predators, grazers and mobile organisms were more abundant among smaller mussels, with smaller gap volume, whereas the barnacle Jehlius cirratus was more abundant on larger mussels. In conclusion, point-source nutrient addition to the mussel bed did not enhance primary production. However, the modification of structural characteristics related to mussel size induced changes in the faunal assemblage. Thus, it seems that in this nutrient-rich system, nutrient enhancement would not significantly affect Perumytilus and its assemblage, whereas structural habitat seems to play an important role in shaping this community.     

Phosphoenolpyruvate metabolism in Teladorsagia circumcincta: A critical junction between aerobic and anaerobic metabolism

Nematodes which have adapted to an anaerobic lifestyle in their adult stages oxidise phosphoenolpyruvate (PEP) to oxaloacetate rather than pyruvate as the final product of glycolysis. This adaptation involves selective expression of the enzyme phosphoenolpyruvate carboxykinase (PEPCK), instead of pyruvate kinase (PK). However, such adaptation is not absolute in aerobic nematode species. We have examined the activity and kinetics of PEPCK and PK in larvae (L₃) and adults of Teladorsagia circumcincta, a parasite known to exhibit oxygen uptake. Results revealed that PK and PEPCK activity existed in both L₃s and adults. The enzymes had differing affinity for nucleotide diphosphates: while both can utilise GDP, only PK utilised ADP and only PEPCK utilised IDP. In both life cycle stages, enzymes showed similar affinity for PEP. PK activity was predominant in both stages, although activity of this enzyme was lower in adults. When combined, both the activity levels and the enzyme kinetics showed that pyruvate production is probably favoured in both L₃ and adult stages of T. circumcincta and suggest that metabolism of PEP to oxaloacetate is a minor metabolic pathway in this species.     

Studies on metabolic properties in the Northern Krill, Meganyctiphanes norvegica (Crustacea, Euphausiacea): influence of nutrition and season on pyruvate kinase

The specific activity and the kinetic properties of partly purified pyruvate kinase (PK) (EC 2.7.1.40) from the Northern Krill, Meganyctiphanes norvegica, were investigated in relation to varying food resources. In order to evaluate the effect of starvation on the total energy metabolism, the respiration rates of fed and unfed krill were determined. The FPLC–elution profile of PK displayed two distinct peaks — PK I and II. The first isoform represented 80% of the total PK activity in the organism, and 20% was contributed by the second isoform. PK I was inhibited by ATP but was not influenced by fructose–1,6–bisphosphate (FBP). In contrast, PK II showed ATP inhibition and up to 2.5-fold increased activity by addition of 17 μmol·l⁻¹ FBP. The Michaelis–Menten constants of both isoforms were 2–10-fold higher for ADP than for phosphoenolpyruvate (PEP). Alanine showed no regulatory effect on PK I and II. In specimens starved for 7 days oxygen consumption decreased by 20%. Neither the feeding experiments nor the animals captured in the field during low and high productive seasons indicate that PK properties of M. norvegica are modified in relation to food supply. Accordingly, alternative mechanisms are involved in the depression of the metabolic rate in terms of oxygen consumption.     

Highly divergent mitogenomes of Geukensia demissa (Bivalvia,Mytilidae) with extreme AT content

Complete sequences of two lineage-specific mitogenomes from mytilid bivalve Geukensia demissa are reported, confirming the existence of doubly uniparental inheritance system in this species. The reported mitogenomes show extreme sequence divergence; at protein level, it is in the range of 12%–55%, exceeding the highest values known from this family to date. Moreover, these mitogenomes are also extraordinarily AT-rich (~72%) making them the most compositionally biased mitogenomes from this family. The compositional bias is even more extreme at neutral sites, reaching 80% AT there. Despite high-sequence divergence, the mitogenomes are both compositionally and structurally similar, with only four trn genes relocated and overall gene order very similar to the phylogenetically close mitogenomes of Perumytilus purpuratus. Lineage-specific differences are limited to the non-coding regions and a short cox2 extension present in the paternally inherited M mitogenome. Phylogenetic analysis shows deeper separation of M and F lineages in Geukensia, than in Perumytilus consistent with higher protein divergence. It can be speculated that stronger mutational pressure in Geukensia is driving faster evolution of its mitogenomes.     

Rocky Intertidal Zonation Pattern in Antofagasta,Chile: Invasive Species and Shellfish Gathering

Background

Biological invasions affecting rocky intertidal zonation patterns, yield information on species interactions. In the Bay of Antofagasta, northern Chile, the non-indigenous tunicate Pyura praeputialis, originally from Australia, has invaded (in the past century or so) and monopolized a major portion of the mid-intertidal rocky shore, displacing upshore the native mussel Perumytilus purpuratus. In Antofagasta the tunicate is subjected to intensive exploitation. Monitoring protocols show that in the past 10 years Antofagasta''s tunicate population has experienced a drastic decline, affecting the intertidal zonation pattern.

Methodology/Principal Findings

A 12.5 km of coastline, on the southern eastern shore of the Bay of Antofagasta, was studied. Eight sites were systematically (1993–1994) or sporadically (2003–2014) monitored for the seaward-shoreward expansion or reduction of the tunicate Pyura praeputialis, and native mussel and barnacle bands. A notable reduction in the mid-intertidal band of P. praeputialis and a seaward expansion of the mussel, Perumytilus purpuratus, and barnacle bands was observed. We suggest that the major cause for the decline in the tunicate is due to its intensive exploitation by rocky shore Pyura-gathers. The rate of extraction of tunicates by professional Pyura-gathers ranged between 256–740 tunicates hour⁻¹. Between 2009–2014 the density of professional Pyura-gather ranged between 0.5–4.5 km⁻¹ per low tide. Hence, 10 professional Pyura-gathers working 1 h for 10 low tides per month, during 6 months, will remove between 307–888 m² of tunicates. A drastic decline in tunicate recruitment was observed and several P. praeputialis ecosystems services have been lost.

Conclusion and Significance

In Antofagasta, the continuous and intensive intertidal gathering of the invasive tunicate Pyura praeputialis, has caused a drastic reduction of its population modifying the zonation pattern. Thereby, native mussel Perumytilus purpuratus has regained its ecological center in the intertidal zone. We recorded a Pyura recruitment failure and loss of ecosystem services.     

Enzymes of the normothermic and hibernating bat,Myotis lucifugus: Metabolites as modulators of pyruvate kinase

Summary The previous paper (Borgmann, A., and Moon, 1976) suggested that temperature differentially affected the binding of the substrates phosphoenol pyruvate (PEP) and ADP to bat tissue pyruvate kinases (PK) under hibernating and normothermic conditions. Since the regulatory properties of most mammalian type-L PKs are temperature dependent, a study of these properties for the bat enzymes was initiated.M. pectoralis PK ofM. lucifugus is modulator insensitive, although ATP does increase theK _m(PEP) slightly (Table 1). This effect is most pronounced at low temperatures for HM-PK (Fig. 1) and may be of some regulatory significance.Modulators affect bat liver PK in a manner analogous to other mammalian type-L enzymes, and marked quantitative differences exist between the NL-and HL-enzymes. TheK _m(PEP) is increased only slightly by alanine and ATP (Table 3), althoughV _max decreases markedly for NL-PK; fructose-1,6-diphosphate (FDP) decreases theK _m(PEP) and overrides the inhibitory action of ATP and alanine (Table 3). As temperature decreases, the proportional change inK _m(PEP) with or without effectors is unchanged (Table 3).HL-PK is markedly affected by these modulators. The extent of this interaction, as indicated byn _H-values (Table 3) andK _i orK _a values (Table 2; Figs. 1, 2, 3), between the effectors and the binding of PEP to HL-PK is relatively greater than for the NL-enzyme. However, the temperature sensitivity of these interactions is reduced (Table 3).Therefore, the strategies associated with enzymes of hibernating and normothermic bats are similar to those previously reported by Hochachka and Somero (1973) for certain enzymes of poikilotherms, and argues for the existence of distinct enzyme forms in the two physiological states.Abbreviations ala alanine - FDP fructose-1,6-diphosphate,HL, HM, hibernator liver, and muscle, respectively - LDH lactate dehydrogenase - NL, NM normothermic liver, and muscle, respectively - PEP phosphoenol pyruvate - PK pyruvate kinase     

Bioinvasion threatens the genetic integrity of native diversity and a natural hybrid zone: smooth‐shelled blue mussels (Mytilus spp.) in the Strait of Magellan

Smooth‐shelled blue mussels of the Mytilus edulis species complex are widely distributed bivalve molluscs whose introductions threaten native marine biodiversity (non‐indigenous species – NIS). The aim of the present study was to identify the species and hybrids of Mytilus present in the Magellan Region (southern Chile). Results indicate that three mussel species of the Mytilus edulis complex are found in the region – M. edulis, M. chilensis (or the Southern Hemisphere lineage of Mytilus galloprovincialis), and M. galloprovincialis of Northern Hemisphere origin. For the first time, alleles of the introduced M. trossulus are reported from the Southern Hemisphere. In the Strait of Magellan the native Pacific blue mussel, Mytilus chilensis and the native Atlantic blue mussel, Mytilus edulis, meet and mix at a natural hybrid zone (about 125 km in length). This is the first record of a natural Mytilus hybrid zone in the Southern Hemisphere and is also the first record of the co‐occurrence of genes from all four Mytilus species in any one region. These results contribute to the knowledge of the biodiversity and delimitation of mussel species in southern South America, and highlight how introduced species may threaten the genetic integrity of native species through hybridization and introgression.     

Woodward''s reagent K reacts with histidine and cysteine residues inEscherichia coli andSaccharomyces cerevisiae phosphoenolpyruvate carboxykinases

The reaction of Woordward's reagent K (WRK) with model amino acids and proteins has been analyzed. Our results indicate that WRK forms 340-nm-absorbing adducts with sulfhydryl- and imidazol-containing compounds, but not with carboxylic acid derivatives, in agreement with Llamaset al. [(1986),J. Am. Chem. Soc. 108, 5543–5548], but not with Sinha and Brewer [(1985),Anal. Biochem. 151, 327–333]. The chemical modification ofEscherichia coli andSaccharomyces cerevisiae phosphoenolpyruvate carboxykinases with WRK leads to an increase in the absorption at 340 nm, and we have demonstrated its reaction with His and Cys residues in these proteins. These results caution against claims of glutamic or aspartic acid modification by WRK based on the absorption at 340 nm of protein-WRK adducts.Abbreviations HPLC high-performance liquid chromatography - MES 2-(N-morpholino)ethanesulfonic acid - PEP phosphoenolpyruvate - PEPCK phosphoenolpyruvate carboxykinase - PTH phenylthiohydantoin - WRK Woordward's reagent K (2-ethyl-5-phenylisoxazolium-3-sulfonate)     

不同生境盐地碱蓬对氮饥饿的响应

为了探讨不同生境盐地碱蓬对低氮生境的适应机制,测定了盐渍环境下(200 mmol/L Na Cl)不同浓度硝态氮(0.3、5 mmol/L NO~-_3-N)预处理两种生境盐地碱蓬经氮饥饿后的NO~-_3含量、硝酸还原酶(NR)活性、光合特性及生长状况。结果表明,0.3和5 mmol/L NO~-_3-N处理以及进行氮饥饿时,潮间带生境盐地碱蓬叶片NO~-_3含量均高于内陆生境盐地碱蓬。与内陆生境盐地碱蓬相比,氮饥饿后,潮间带生境盐地碱蓬叶绿素含量、NR活性和光合放氧速率下降幅度均小于内陆生境盐地碱蓬,在0.3mmol/L NO~-_3-N预处理进行氮饥饿时趋势更加明显。0.3 mmol/L NO~-_3-N预处理后氮饥饿对潮间带生境盐地碱蓬根冠比没有影响,却降低内陆生境盐地碱蓬根冠比。上述结果表明,低氮条件下潮间带生境盐地碱蓬具有较高的NO~-_3储存能力,在环境持续氮素缺乏时具有较高的NO~-_3-N再利用能力,能更好地维持氮代谢以及光合性能。说明潮间带生境盐地碱蓬能更好地适应低氮生境。     

Effects of copper on the fitness of the Chilean scallop Argopecten purpuratus (Mollusca: Bivalvia)

Exposure to high concentrations of metals can affect populations and individuals at morphological, physiological, biochemical or genetic levels. Metal pollution is a source of environmental stress that can have deleterious effects on organisms and generate selective pressure upon populations. This work attempts to establish whether concentrations of copper, above physiological requirements, can affect the genetic structure of a cohort of Chilean scallop Argopecten purpuratus. Following the determination of LC₅₀ for 96 h experiments, other acute toxicity tests were carried out, exposing the juveniles to solutions of 150 ppb copper for 120 h. Dead, surviving and control individuals from these bioassays were genetically characterized for five polymorphic loci, Isocitrate dehydrogenase (Idh), Octopine dehydrogenase (Ocdh), Phosphogluconate dehydrogenase (Pgd), Leucine aminopeptidase (Lap) and Phosphoglucose isomerase (Pgi). Results showed no significant differences in allele and genotypic frequencies between surviving and dead individuals, although for the same groups significant differences were found in estimated mean heterozygosity. Pgi and Ocdh showed significantly different heterozygosity values for dead and surviving individuals. A positive relationship between multilocus heterozygosity and survival was found for young A. purpuratus exposed to high copper concentrations; thus, we found evidence of a differential response to exposure to high copper concentrations, related to degree of heterozygosity.     

Mytilus galloprovincialis (Mollusca: Bivalvia) in a warm-temperate South African estuarine embayment

The distribution, population structure, condition and gonad indices, and the attachment strengths of the invasive Mediterranean mussel Mytilus galloprovincialis are described from the Knysna Estuary, South Africa. Mussels were found in the rocky intertidal at the mouth of the estuary and on all man-made hard substrata as far as 12?km from the mouth in 2013. However, mussel beds were found only in the lower reach of the estuary, where mussel densities were as high as 120 per 0.01?m². When compared to mussels from the embayment regions of the estuary, those at the estuary mouth, where wave action was high, were generally smaller, had significantly lower condition and gonad indices, and greater attachment strength. This suggests that mussels within the sheltered regions of the estuary invest more energy in growth and reproduction, and that this is one reason for their success as an invasive species in the Knysna estuarine embayment.     

The role of the predator Trophon geversianus in an intertidal population of Mytilus chilensis in a rocky shore of the Beagle Channel,Tierra del Fuego,Argentina

Predation influences changes in the abundance and the limits of spatial distribution of organisms on rocky shores. We quantified the effect of the predation pressure of the gastropod Trophon geversianus on the mytilid Mytilus chilensis in a rocky intertidal community of the Beagle Channel, Tierra del Fuego. We quantified the density of T. geversianus along with the density of T. geversianus preying on M. chilensis and examined the potential relationship between sizes of both T. geversianus and M. c hilensis. Additionally, we carried out laboratory experiments to quantify the drilling and ingestion rates of T. geversianus on M. c hilensis at different aerial exposure times. At mid intertidal levels, unlike at low intertidal levels, T. geversianus consumed more frequently the available size range of mussels. On average, sizes of predator and prey consumed were positively correlated. The drilling rate decreased with increasing time of aerial exposure. Additionally, permanently submerged predators invested less time consuming the same amount of food than individuals exposed for 4 out of 12?h (4/12?h). Predators exposed for 9/12?h invested less time and consumed less food than those exposed 4/12?h. Our results suggest that T. geversianus is an efficient predator of M. chilensis, due to both its high density and feeding rate. Our results suggest that the strong predation pressure of T. geversianus on M. chilensis prevents this key competitor from monopolizing space at the low level through controlling their abundance, and further limits their vertical distribution.     

Ecophysiological responses of the euhalophyte Suaeda salsa to the interactive effects of salinity and nitrate availability

Effects of salinity and nitrate nitrogen (NO₃⁻-N) on ion accumulation and chlorophyll fluorescence were monitored for two populations of Suaeda salsa grown from seeds in a greenhouse experiment. One population inhabits the intertidal zone and the other occurs on inland saline soils. Ion contents in soils and in leaves of the two populations were also investigated in field. In the greenhouse, seedlings were exposed to a NaCl concentration of 0.6 and 35.1 ppt, with 0.1 or 5 mM NO₃⁻-N treatments for 20 days. The contents of Na⁺ and Cl⁻ were higher, but NO₃⁻ was lower in soils of the intertidal zone than at the inland site. In the field, ion concentrations and the estimated contribution of these ions to osmotic potential in leaves showed no difference between the two populations, except that the estimated contribution of Na⁺ to osmotic potential in leaves of the intertidal population was lower than that in the inland population. In the greenhouse, in contrast, the concentration of Cl⁻ was lower, but NO₃⁻ concentration and the estimated contribution of NO₃⁻ to osmotic potential were higher, in the leaves of plants from the intertidal zone. Salinity had no effect on the maximal efficiency of PSII photochemistry (Fv/Fm) and the actual PSII efficiency (ΦPSII). The results indicated that S. salsa from the intertidal zone was better able to regulate Cl⁻ to a lower level, and accumulate NO₃⁻ even with low soil NO₃⁻ concentrations. Tolerance of the PSII machinery to high salinity stress may be an important characteristic for the studied species supporting growth in highly saline environments.     

Feeding behavior and differential absorption of biochemical components by the infaunal bivalve Mulinia edulis and the epibenthic Mytilus chilensis in response to changes in food regimes

We measured changes in the feeding rate and food absorption efficiency of two suspension feeding bivalves, cross-trasplanted between habitats with special emphasis on their capacity for differential absorption of biochemical components from their food supply. Mulinia edulis were moved from the intertidal zone to the subtidal zone, and Mytilus chilensis from the subtidal to the intertidal zone for a period of 7 days, and then compared with animal that had not been transplanted. Experimentally prepared diets similar to those available in the two different environments were offered to the bivalves, and their rates of feeding and differential uptake of biochemical components were determined and statistically compared. The two species did not achieve complete acclimation of their feeding behaviour during the transplant period since the highest ingestion rates for biochemical components occurred under dietary conditions that reflected their habitats of origin. Absorption efficiency showed greater acclimation than the other physiological parameters measured, indicating the capacity of these species to modulate their enzymatic-digestive activity depending on food composition. We conclude that both Mytilus and Mulinia have a certain degree of physiological plasticity in their feeding behaviour and assimilatory balance of biochemical components, being greater in Mytilus. When both species encounter ambient food conditions characteristic of their normal habitats, they show maximum values of food absorption, while under conditions where their typical diets are exchanged (Mytilus in intertidal and Mulinia in subtidal), the energy absorbed declines in each, but in ways very different between the two species. Thus, Mytilus exposed to high concentrations of low quality seston reduced the energy absorbed by 31.7% compared to its normal habitat, while Mulinia exposed to low concentrations of high-quality food reduced their energy absorption by 64%.     

Spatial and temporal variation of photosynthesis in intertidal Mazzaella laminarioides (Bory)Fredericq (Rhodophyta, Gigartinales)

The red alga Mazzaella laminarioides is an economically important species with an extended latitudinal distribution along the Chilean coast. Its populations form mid-intertidal stands, several meters wide, and therefore are differentially exposed to environmental variables that result in temporal and spatial variability in productivity. We evaluated the effect of latitude and intertidal height on productivity by in situ measurement of photosynthetic performance. Daily and seasonal variations of O₂-evolution rate and maximal quantum yield (F _v/F _m) were determined in plants from the upper and lower intertidal zone at two localities 1500,km apart. Results suggest that plant responses were mainly affected by irradiation, temperature and desiccation. At local level, upper intertidal plants showed a reduced photosynthetic rate and quantum efficiency as compared to those displayed by plants from the lower intertidal, indicating their higher level of excitation energy acclimation. Stronger acclimation differences between upper and lower intertidal plants were observed in spring and summer. Differences in photosynthetic parameters between reproductive phases were recorded in autumn and winter, regardless of the position of the individuals in the intertidal zone. The effects of tidal elevation on seasonal patterns of photosynthesis were also influenced by latitude. Seasonal variation in photosynthetic efficiency was observed in plants from the northern population at both intertidal elevations, but only at the upper intertidal level in the southern population. This study shows that production variability in M. laminarioides results from differences in the intensity of environmental factors observed seasonally at local (intertidal) and latitudinal scales.     

Erythrocruorins (extracellular haemoglobins) from the cirratulid polychaetes cirriformia tentaculata (Montagu) and Cirratulus cirratus (Müller) with special reference to saturation dependent characteristics of the oxygen equilibria

Erythrocruorins from the intertidal cirratulid polychaetes Cirriformia tentaculata (Montagu) and Cirratulus cirratus (Müller) have a high affinity for oxygen and large Bohr shifts. At low pH, the O₂-equilibrium curves from both species are hyperbolic. In alkaline solution both erythrocruorins show marked increases in O₂-affmity with the retention of the hyperbolic form in Cirriformia. The presence of a pronounced Bohr shift in a non-cooperative erythrocruorin is unusual and its possible physiological rôle is discussed.In Cirratulus, the shape of the curve becomes sigmoidal at high pH. Moreover, the cooperativity coefficient, n, increases at high O₂-saturation. The Bohr effect thus increases in magnitude with rising pH and it is saturation dependent. The significance of this complex oxygen transport system is discussed. The findings illustrate the remarkable elasticity of functions possible in erythrocruorins from closely related species in response to different environmental or metabolic dictates.