Food sources, behaviour, and distribution of hydrothermal vent shrimps at the Mid-Atlantic Ridge

Five species of bresilioid shrimp were investigated at seven hydrothermal sites on the Mid-Atlantic Ridge: Menez Gwen, Lucky Strike, Rainbow, Broken Spur, TAG, Snake Pit and Logatchev. Samples were prepared for analysis of stable isotopes, elemental composition and lipids. Shrimp behaviour was observed from the submersible ‘Alvin’ and in the laboratory aboard RV ‘Atlantis’. The distribution and zonation of the shrimp species was recorded. Juvenile shrimp of all species arrive at the vents carrying reserves of photosynthetic origin, built-up in the pelagic larval stages. These reserves are used while the shrimp metamorphose to the adult form and, in Rimicaris exoculata and Chorocaris chacei, while they develop epibiotic bacteria supporting structures, the modified mouthparts and the inside of the carapace. The main food of adult R. exoculata is filamentous bacteria that grow on these structures. The intermediate sizes of C. chacei also feed on such bacteria, but the final stage gets some food by scavenging or predation. Mirocaris species scavenge diverse sources; they are not trophically dependent on either R. exoculata or mussels. Adults of Alvinocaris markensis are predators of other vent animals, including R. exoculata. The dense swarms of R. exoculata, with their exosymbionts, can be compared to endosymbiont-containing animals such as Bathymodiolus and the vestimentiferan tube-worms of the Pacific vents. Such associations, whether endo- or ectosymbiotic, may be necessary for the development of flourishing communities at hydrothermal vents.

Different energy sources for three symbiont-dependent bivalve molluscs at the Lagatchve hydrothermal site (Mid Atlantic Ridge)

The vent mussel Bathymodiolus puteoserpentis, a large vesicomyid clam and a smaller thyasirid were collected from an area of sediment subject to diffuse hydrothermal flow. The mussels live on the surface, the vesicomyids are partly buried and the thyasirids burrow in the sediment. The fine structure of the gills differs in the three bivalves. Bathymodiolus puteoserpentis hosts two types of bacterial symbiont, one methanotrophic, and another probably thiotrophic. The other two bivalves have single types of symbiont of different shapes. Stable isotope ratios of carbon and nitrogen indicate thiotrophy in the vesicomyid and thyasirid, but a predominance of methanotrophy in the mussel. This is the first time that such an assemblage has been found at a hydrothermal site on the Mid-Atlantic Ridge (MAR), with the different faunistic elements exploiting different energy resources

Coordinated Rhythms Of Digestion Absorption And Excretion In Mytilus-Edulis (Bivalvia Mollusca)

Sequential alternation of extracellular digestion in the stomach and intracellular digestion in the diverticula appears widespread among bivalves. The present study documents some physiological consequences of such processes in Mytilus edulis L. collected during 1981 from Whitsand Bay, Cornwall, England. Pronounced temporal fluctuations in faecal deposition are described that relate, in terms of amplitude and period, to both sinusoidal rhythmicity established for ammonia excretion and changes in the morphology of digestive tubules. Although at least partially synchronised among replicate groups of mussels, these cycles bore no consistent relationship with exogenous influences. Hourly fluctuation in the net absorption efficiency for nitrogen, as evidenced by the mean percentage ±2 SE, measured over 24 h sampling periods, was considerable (16.0±53.7, 49.3±10.9 and 52.8±6.6 for mussels acclimated in March, June and October, respectively). This variation in absorption derived from an inverse relationship between the percentage nitrogen within faeces and the rate of faecal egestion. Accordingly, peaks of faecal deposition presumably represented the pulsed remnants of intracellular digestion. Co-ordinated rhythms of digestion, absorption and excretion were thus evident in M. edulis. These processes displayed seasonally dependent periodicities of approximately 8, 3 and 4 h in March, June and October, respectively. It was concluded that, at least for M. edulis, this previously unquantified rhythmicity of physiological processes warrants careful consideration during assays commonly undertaken in the complication of nutrient and energy budgets.

Reproductive Effort And Value In Different Populations Of The Marine Mussel Mytilus-Edulis-L

Fecundity, reproductive effort (estimated both from production measurements and from physiological data), the energetic costs of reproduction and the reproductive value of different size classes were measured for mussels at different sites and related to age and to tissue weight. Variability between sites was considerable and differences as great as 10 x were recorded between minimum and maximum values for egg production, reproductive effort and reproductive value. However, similarities between mussels from different sites were also apparent, as regards egg size, the estimated metabolic costs of egg production (based on measurements of oxygen consumption), the relationship (isometric) between egg production and body size, the fact of an increase in reproductive effort with increase in size, and the age at which maximum residual reproductive values was expressed. These relationships are discussed in terms of the fundamental reproductive strategy of the species and the degree of environmental stress imposed on the mussels at the different sites.

Effects Of Aromatic-Hydrocarbons On The Metabolism Of The Digestive Gland Of The Mussel Mytilus-Edulis-L

1. The results presented in this paper show that the exposure of mussels to a sublethal concentration of oil-derived aromatic hydrocarbons (30 μg 1−1) for a period of 4 months significantly decreases the protein level in the digestive gland of the animals (−17%). 2. The activity of the nuclear RNA polymerase I and II is also significantly decreased in the digestive gland of hydrocarbon-exposed mussels (−64% and −18%, respectively). 3. The RNAase(s) activity present in the nuclei from the digestive gland cells increases following the exposure of the mussels to aromatic hydrocarbons. This effect is particularly evident at high ionic strength [200 mM (NH4)2SO4]. 4. The analysis of some characteristics of the nuclear RNAase(s) (most of which is soluble and shows a maximum of activity at pH 4−5) could indicate that part of this hydrolytic enzyme may have a lysosomal origin. 5. This fact appears to be in agreement with the finding that in the mussels exposed for 4 months to aromatic hydrocarbons the lysosomal stability decreases drastically and the total content of lysosomal enzymes is significantly increased (+42.4%).

Lysosomes And The Response By Mytilus-Edulis-L To An Increase In Salinity

Cytochemical observations and measurements on cell-free suspensions of lysosomes from the digestive gland of Mytilus edulis showed a reduced latency of the lysosomal enzyme beta -N-acetyl-hexosaminidase 12h after mussels were transferred from 21 to 35%o salinity, but showed no change up to 6 h after transfer. There was a transient alteration in the form of the latency curve after 6 h at high salinity, signifying a gradual change in membrane integrity. Free hexosaminidase activity increased, 12 h after the salinity rise. The lysosomes were permeable to amino acids when ATP was present; permeability increased following the rise in salinity. The concentration of ninhydrin-positive substances in the lysosomes increased 6 h after transfer and then, between 6 and 12 h, the concentration declined. The results are consistent with the hypothesis that lysosomal hydrolysis is a source of free amino acids during the adaptation of mussels to increased salinity.

Mixed-Function Oxygenases And Xenobiotic Detoxication-Toxication Systems In Bivalve Mollusks

Components of a xenobiotic detoxication/toxication system involving mixed function oxygenases are present inMytilus edulis. Our paper critically reviews the recent literature on this topic which reported the apparent absence of such a system in bivalve molluscs and attempts to reconcile this viewpoint with our own findings on NADPH neotetrazolium reductase, glucose-6-phosphate dehydrogenase, aldrin epoxidation and other reports of the presence of mixed function oxygenases. New experimental data are presented which indicate that some elements of the detoxication/toxication system inM. edulis can be induced by aromatic hydrocarbons derived from crude oil. This includes a brief review of the results of long-term experiments in which mussels were exposed to low concentrations of the water accommodated fraction of North Sea crude oil (7.7–68 µg 1−1) in which general stress responses such as reduced physiological scope for growth, cytotoxic damage to lysosomal integrity and cellular damage are considered as characteristics of the general stress syndrome induced by the toxic action of the xenobiotics. In addition, induction in the blood cells of microsomal NADPH neotetrazolium reductase (associated with mixed function oxygenases) and the NADPH generating enzyme glucose-6-phosphate dehydrogenase are considered to be specific biological responses to the presence of aromatic hydrocarbons. The consequences of this detoxication/toxication system forMytilus edulis are discussed in terms of the formation of toxic electrophilic intermediate metabolites which are highly reactive and can combine with DNA, RNA and proteins with subsequent damage to these cellular constituents. Implications for neoplasms associated with the blood cells are also discussed. Finally, in view of the increased use of mussel species in pollutant monitoring programmes, the induction phenomenon which is associated with microsomal enzymes in the blood cells is considered as a possible tool for the detection of the biological effects of environmental contamination by low concentrations of certain groups of organic xenobiotics.

A Possible Role Of Alpha-Amylase Isoenzymes From The Style Of The Mussel Choromytilus-Meridionalis (Krauss) Following Thermal-Acclimation

Separation of the proteins comprising the crystalline style of the mussel Choromytilus meridionalis (Krauss) by anion exchange chromatography shows that there are three fractions displaying α-amylase activity in both warm- and cold-acclimated mussels. These fractions correspond with one or more proteins which remain unbound to the resin (Peak I), a bound fraction which is eluted at 100–150 mM NaCl (Peak II) and a further fraction which is eluted at 200–250 mM NaCl (Peak III) but which may represent contamination carried over from Peak II. Cold-acclimation to 8°C results in the appearance of a fourth α-amylase fraction (Peak IV) which is eluted from the column between 300–400 mM NaCl. Thermal acclimation also results in changes in the activities of Fractions I–IV such that a specific activity of 0.47 mg glucose liberated per A280 unit of protein per 8 min incubation at 8°C in Fraction IV is increased nearly 10-fold to a specific rate of 4.10 in protein Fraction I following acclimation to 22°C. It is suggested that an increased of digestive activity may be of equal importance to a suppression of metabolic costs in the maintenance of energy flow into growth and reproduction in ectothermic organisms which experience an increase of environmental temperature, especially in bivalves such as C. meridionalis which do not show a compensatory increase in filtration rate.

Measurement Of The Responses Of Individuals To Environmental-Stress And Pollution - Studies With Bivalve Mollusks

Certain physiological differences between individuals in different populations of the mussel, Mytilus edulis, are described. In particular, the scope for growth differs in space and time and may be used to assess the animals' physiological condition. When the required measurements are made in the field, the rates of growth predicted from the physiological data agree well with observed rates of growth. An alternative approach utilizes mussels transplanted to various waters, with indices of condition then measured in the laboratory under standard conditions; an example of this approach is illustrated. Laboratory experiments are used to equate various levels of physiological condition with fecundity, in an attempt to equate physiological effects on the individual with likely population damage. A cytochemical index of stress is described, based on the latency of lysosomal enzymes; spatial variability in this index, and its relation with the scope for growth, are discussed. Finally, the results of some experiments on the effects of petroleum hydrocarbons on mussels are described and the presence of inducible activity of NADPH-dependent tetrazolium reductase in the blood cells is demonstrated. Certain considerations that apply in adopting similar measurements of biological effects of pollution in environmental monitoring programmes are discussed.

Population-Dynamics Of Mytilicola-Intestinalis In Mytilus-Edulis In South West England

The population dynamics of Mytilicola intestinalis Steuer in mussels (Mytilus edulis L.) from the River Lynher, Cornwall, England, have been studied over 3 years. By transplanting uninfested mussels from the River Erme, South Devon, into the Lynher mussel bed, the study was extended to the growth and development of new infestations under natural conditions. Female Mytilicola intestinalis are shown to breed twice, and two generations of parasites coexist for most of the year, with recruitment taking place in summer and autumn. One generation contributes its first brood to the autumn recruits before overwintering and contributing its second brood to the following summer's recruits. The other generation overwinters as juvenile and immature stages to contribute its two broods successively to the summer and autumn recruits. Environmental temperatures are believed to control the rates of development at all stages rather than acting as triggers in the onset or cessation of breeding at specific times. There is no evidence to support the contention that heavily infested mussels are killed, and parasite mortality is shown to be density-independent.

Relationship Between Infestation By Mytilicola-Intestinalis Steuer (Copepoda Cyclopoidea) And Condition Index Of Mytilus-Edulis In Southwest England

Seasonal cycles in the condition index of Mytilus edulis from three sites in southwest England are described. These are analysed in relation to host length, stage of gonad development and parasite burden by linear regression analysis. An effect on the condition index due to the presence of Mytilicola intestinalis can be detected only in the sublittoral mussels in those few winter months when the mean intensity of infestation is over about 25 parasites per host. In all cases studied, the magnitude of the effect due to variation in host length, stage of gonad development, seasonal cycles and environmental factors is greater than that due to parasitism.

Responses Of Mytilus-Edulis L To Low Oxygen-Tension - Acclimation Of Rate Of Oxygen-Consumption

1. Mytilus edulis acclimated its rates of oxygen consumption when maintained at reduced oxygen tensions for periods in excess of five days. 2. Acclimation was complete down to approximately 55 mm Hg PO2 at slightly lower oxygen tensions (51, 49 and 43 mm Hg) acclimation was complete in one experiment and partial in two others. 3. The capacity to acclimate oxygen consumption was not affected by a reduction in ration nor by an increase in temperature (10 to 22 °C). 4. Mussels that were acclimated to reduced oxygen tension (40–80 mm Hg), and then exposed to P O 2 of less than 20 mm Hg for two or five hours, had depressed rates of oxygen uptake when subsequently “recovered” to 40–80 mm Hg. 5. These results are discussed in the context of biochemical studies of anaerobic metabolism in mussels from the same experiments.

Rates Of Nitrogen-Excretion By Species Of Mytilus (Bivalvia - Mollusca)

The results of experiments recorded by Bayne & Scullard (1977) confirmed earlier studies (Bayne, 1973) in describing a decline in the rate of oxygen uptake (Vo2) by Mytilus edulis during starvation, eventually reaching a steady-state value, called the standard rate of oxygen consumption. Earlier experiments had also shown that if such starved mussels were fed, oxygen uptake increased rapidly to a high level called the active rate of oxygen consumption (Thompson & Bayne, 1972; Bayne, Thompson & Widdows, 1973). Some of this increase in metabolic rate is undoubtedly due to an increased filtration rate that is stimulated by the presence of food (the ‘mechanical cost of feeding’ discussed by Bayne et al. 1976), and part is due to the ‘physiological costs of feeding’, which includes energy utilized in digestion and assimilation of the food, and energy that is lost during deamination and other catabolic processes that accompany digestion (Warren & Davis, 1967). Increases in metabolic rate associated with feeding have been called the specific dynamic action (SDA) of the ration (see Harper, 1971, for a discussion) or the apparent SDA (Beamish, 1974)5 and they have been related to aspects of protein metabolism (Krebs, 1964). This paper describes the results of some experiments designed to examine the relationships between SDA and ammonia excretion in Mytilus edulis L.

Apparent Specific Dynamic Action In Mytilus-Edulis L

The results of experiments recorded by Bayne & Scullard (1977) confirmed earlier studies (Bayne, 1973) in describing a decline in the rate of oxygen uptake (Vo2) by Mytilus edulis during starvation, eventually reaching a steady-state value, called the standard rate of oxygen consumption. Earlier experiments had also shown that if such starved mussels were fed, oxygen uptake increased rapidly to a high level called the active rate of oxygen consumption (Thompson & Bayne, 1972; Bayne, Thompson & Widdows, 1973). Some of this increase in metabolic rate is undoubtedly due to an increased filtration rate that is stimulated by the presence of food (the ‘mechanical cost of feeding’ discussed by Bayne et al. 1976), and part is due to the ‘physiological costs of feeding’, which includes energy utilized in digestion and assimilation of the food, and energy that is lost during deamination and other catabolic processes that accompany digestion (Warren & Davis, 1967). Increases in metabolic rate associated with feeding have been called the specific dynamic action (SDA) of the ration (see Harper, 1971, for a discussion) or the apparent SDA (Beamish, 1974)5 and they have been related to aspects of protein metabolism (Krebs, 1964). This paper describes the results of some experiments designed to examine the relationships between SDA and ammonia excretion in Mytilus edulis L.

Occurrence Of Mytilicola-Intestinalis Steuer An Intestinal Copepod Parasite Of Mytilus In Southwest Of England

The occurrence of Mytilicola intestinalis in populations of mussels in south-west England is recorded and compared with previous data. Since 1955 there have been two main changes in the distribution of Mytilicola: (a) it has invaded all the major estuarine mussel populations on the Bristol Channel coast, and (b) many previously uninfested open-coast populations all round the peninsula are now lightly infested. It is suggested that differences in infestation levels between estuarine and open-coast populations of mussels are due primarily to differences in the degree of exposure to wave action although factors such as size, population density and location of the hosts also influence infestation. The chance of the establishment of breeding pairs of Mytilicola depends on the parasite population size and its distribution through the host population.