Feeding and digestion in Corophium volutator (Crustacea: Amphipoda)

New observations on the feeding behavior of Corophium volutator, collected from the Menai Strait, Anglesey, UK in 1981, show that detritus drawn into one entrance of the U-shaped burrow is pushed out at the opposing entrance either uningested or as faeces. Periodically, the amphipod turns around in the burrow and recommences feeding, so both the uningested material and the faeces may be reworked by the mouthparts. A feeding individual produces, on average, 16 faecal pellets every 10 min, at a seawater temperature of 18°C. When fed on detritus dyed with azo-carmine, approximately 12 pellets are produced before the foregut, intestine and hindgut are completely clear of previously ingested material and the dye appears in the faeces. Dye which enters the ventral caeca takes between 12 and 48 h to clear this tissue and, furthermore, it is only cleared when the amphipod is feeding. It is concluded that ingested coarse material is subjected only to a primary phase of digestion in the stomach, and that it takes 4 to 24 min to clear the gut. Fine material which enters the ventral caeca is subjected to a secondary phase of digestion and absorption which takes 24 to 48 h. Particles of thorium dioxide and ferritin incorporated in the diet are absorbed by the B- and ageing RF-cells in the ventral caeca. Non-specific esterases occur in the apical regions of the same cells, the mature cells of the anterior dorsal caeca and the cells at the anterior of the intestine. Protease activity is greatest in the lumen of the ventral caeca and the large vacuole of the B-cells. Apart from the lumen of the posterior caeca, there is some protease activity throughout the lumen of the gut. Carbohydrates were localised primarily in the R/F-cells towards the proximal region of the ventral caeca. Lipid was confined to the R/F-cells in the same region.     

Accumulation of copper within the “Hepatopancreatic” caeca of Corophium volutator (Crustacea: Amphipoda)

The concentrations of various metals in specimens of Corophium volutator (Pallas) which have been taken from a site that receives freshwater drainage with a high concentration of copper, zinc and manganese, are compared with those in individuals from a site with a normal concentration of these metals for coastal waters. The amphipods at the contaminated site contain 259 g g⁺¹ dry weight of copper, which is three times that of individuals from the uncontaminated site. The concentrations of the other metals do not differ substantially between the two populations. The hepatopancreatic caeca of the midgut of amphipods with a high concentration of copper have numerous granules of homogeneous electron-dense material which can aggregate within the multivesicular bodies. Some of the amphipods with a lower concentration of copper have similar granules, but these are fewer in number. X-ray microporbe analysis shows that these granules contain copper and sulphur, and occasionally calcium. Various tests on the granules in frozen section indicate that they are relatively insoluble in a wide range of reagents and that the copper is probably complexed with organic matter.     

Studies on differential fitness of PGI genotypes with regard to temperature in Gammarus insensibilis (Crustacea: Amphipoda)

This paper deals with an experimental study of survival, combined with estimates of biochemical activity, of different genotypes at the PGI (phosphoglucose isomerase) locus in Gammarus insensibilis, in relation to temperature. Samples were collected in the lagoon of Venice during 1987. No mortality occurred at 10°C whereas at 27°C, where mortality reached the value of 50%, heterozygotes exhibited significantly higher survival than homozygotes. Experiments conducted in order to evaluate the PGI biochemical activity of homo- and heterozygote genotypes at three different temperatures (4°, 20° and 37°C) showed in all genotypes an increased activity from 4° to 20°C and a fall of activity from 20° to 37°C. Heterozygotes exhibited higher activity at all temperatures tested. The difference between homo- and heterozygotes became more obvious at 37°C. Our results suggest that in G. insensibilis the PGI locus, as already shown in other organisms, may be subject to selection and that the heterozygous genotypes possess superior fitness. The biochemical bases of the observed differences between genotypes are briefly discussed.     

Effects of cadmium on Pontoporeia affinis (Crustacea: Amphipoda) in laboratory soft-bottom microcosms

Laboratory soft bottom microcosms, containing sediment with natural populations of meio-and microfauna and the amphipod Pontoporeia affinis Lindström, were used to investigate chronic effects of cadmium on a typical Baltic Sea benthic community. Temperature, salinity, organic content of the sediment and the life cycles of the amphipods in laboratory control microcosms were in good agreement with conditions in the Baltic Sea. In Experiment 1, P. affinis was exposed to four assayed cadmium concentrations, 0 (control), 6.3, 41 and 127 g Cd·1^-1, for 265 d. At the lowest cadmium concentration (6.3 g Cd·1^-1), the sediment contained about 700 times, and P. affinis about 1 700 times, the water concentration at the end of the experiments. In microcosms without P. affinis, the sediments accumulated less cadmium. Effects on P. affinis embryogenesis were detected after 105 d of exposure in all tested cadmium concentrations. Mortality of juveniles was significantly increased after 265 d of exposure in mean assayed concentrations of 41 and 127 g Cd·1^-1. In Experiment 2, P. affinis was exposed to cadmium in assayed concentrations, 0, 6.5 and 46 g Cd·1^-1 for 460 d. At the lowest water concentration (6.5 g Cd·1^-1), the prolonged exposure did not increase the sediment concentration, which indicates a steady state, while concentrations in P. affinis reached about 3 500 times the lowest water concentration after 460 d. Juvenile P. affinis matured sexually and reproduced normally in all microcosms, but their offspring died in all microcosms with added Cd. This occurred at water concentrations not uncommon in polluted marine areas. The effects are serious in comparison to those found in conventional short-term laboratory experiments with the same concentrations and demonstrate the importance of long-term experiments in ecotoxicology.     

The ecophysiological complex of Bathyporeia pilosa and B. pelagica (Crustacea: Amphipoda). II. Effects of exposure

The resistance to high and low temperatures, starvation and desiccation effects in the sand dwelling amphipods Bathyporeia pilosa Lindström and B. pelagica (Bate) have been tested. B. pilosa was the more tolerant species in all experiments. Gravid females showed a greater resistance to high temperatures than adult males in both species, and starvation tolerance was in the order gravid females>juvenile males> adult males. Time-temperature relationships, however, would seem to be of greater ecological significance than upper or lower lethal temperatures, and desiccation effects become obvious well within any limits set by starvation. Exposure would appear to be an important concept limiting the intertidal distribution of sand dwelling animals, particularly those confined to the surface layers of sediment. The amplitude, rate and degree of environmental change and its effect on feeding and reproduction offer severe limitations to the distribution of the two Bathyporeia species studied.     

Combined effects of temperature and salinity on the complete development ofEurytemora velox (Crustacea: Calanoidea)

The calanoid copepodEurytemora velox was collected from rock pools at Castletown, Isle of Man, UK. Its optimum environmental requirements, particularly temperature and salinity, were determined, with a view to its possible future use as living food in intensive fish and shellfish farming. The species was cultured in 21 different temperature and salinity combinations. Investigations covered a period of two years from December 1983 to December 1985. Complete development from hatching to adult stage was followed in 21 temperature and salinity combinations. Nauplii suffered relatively high mortalities, indicating the sensitivity of this development stage to variations in temperature and salinity. Highest nauplii survival was observed in the combinations 15°C with 25 and 20 S and 20°C with 20 S, the highest copepodite survival at 10°C and 20 S. Lower salinities were tolerated better at higher temperatures and higher salinities at lower temperatures. Development time varied with the temperature and salinity combinations. Lower salinities at the lower temperatures of 10° and 15°C and both lower and higher salinities at 20°C prolonged development, particularly of the naupliar stage. Highest Q₅ values (i.e., rate of change of development with a 5 C° increase in temperature) were recorded for the naupliar stage. Statistical analysis indicated that salinity influences the survival of both nauplii and copepodites; however, this effect is not linear.     

Physiological responses to fluctuation in temperature of salinity in invertebrates. Adaptations ofAlpheus viridari (Decapoda,Crustacea),Terebellides parva (Polychaeta) andGolfinigia cylindrata (Sipunculida) to the mangrove habitat

The snapping shrimpAlpheus viridari (Armstrong, 1949), the polychaeteTerebellides parva Solis-Weiss, Fauchald and Blankensteyn 1990, and the sipunculanGolfingia cylindrata (Keferstein, 1865) are commonly found in the same mangrove habitat, where they experience frequent, acute fluctuations in temperature and salinity. Ecological studies indicate a temporal variation, including occasional absence, in the distribution of bothG. cylindrata andT. parva; this fed us to examine the physiological adaptations of the three species (collected at Western Bay, Twin Cays, Belize in 1985, 1986 and 1988). Each was subjected to acute, repeated exposure to either control (35 S) and decreased (25 S) salinity or to control and increased (45 S) salinity. Ability to regulate water and ion content (g H₂O or mol g^-1 solute free dry wt) was examinedA. viridari behaved as a hyperosmotic conformer at decreased salinity but as an osmoconformer at increased salinity. Regardless of direction of salinity change,A. viridari regulated water content through change in Na⁺, K⁺, and Cl^– contents. In contrast,G. cylindrata behaved as an osmoconformer and did not demonstrate ability to regulate water content.T. parva behaved as an osmoconformer, showed incomplete regulation of water content via change in Na⁺, K⁺, and Cl^– contents but had limited survival following exposure to 45 S. Each species was also exposed to change in temperature. Species were subjected to acute, repeated exposure either to control (28°C) and decreased (21°C) temperature or to control and inereased (35°C) temperatureA. viridari regulated water and ion content under both experimental conditions. In contrast,T. parva did not regulate water and ion content under either experimental temperature.G. cylindrata did not regulate water and ion content during exposure to decreased temperature and did not survive exposure to increased temperature. ForA. viridari, weight specific oxygen uptake rates (mg O₂ g^-1 ash-free dry wt) were determined. Exposure to decreased salinity or to increased temperature resulted in a small sustained elevation in O₂ uptake. It is concluded that, unlikeA. viridari, T. parva andG. cylindrata are only marginally adapted to withstand the salinity and temperature stresses, respectively, of the mangrove habitat. The inability ofT. parva andG. cylindrata to fully adapt to extremes in the mangrove habitat could well explain the temporal variation seen in the distribution of these two species.Contribution number 380. Caribbean Coral Reef Ecosystems, National Museum of Natural History, Smithsonian Institution     

Environmental influence on the locomotor activity rhythm of Talitrus saltator (Crustacea: Amphipoda)

The persistence and precision of the endogenous, nocturnal, circadian locomotor rhythm of the sand-beach amphipod Talitrus saltator is characterised, and the influence of substrate availability, photoperiod, temperature and relative humidity as potential environmental synchronizers of the rhythm is assessed. Individual, cyclic light-dark and temperature regimes synchronize and modify the laboratory activity pattern, but substrate availability and relative humidity have no significant long-term effect. Under simulateneous experimental regimes simulating field conditions of photoperiod, temperature and relative humidity the rhythm is entrained solely by photoperiod. The implications are discussed in relation to field behaviour.     

The ecophysiological complex of Bathyporeia pilosa and B. pelagica (Crustacea: Amphipoda). I. Respiration rates

The metabolic rates of high and low shore estuarine populations of Bathyporeia pilosa Lindström and an open coast population of B. pelagica (Bate) have been determined over a range of temperatures during January and February, and June and July, 1968. Changes in oxygen uptake have also been measured monthly at 15°C. During the winter, oxygen uptake was in the order: high shore B. pilosa>low shore B. pilosa>B. pelagica. During the summer, high shore B. pilosa and B. pelagica had similar metabolic rates, but both were significantly higher than low shore B. pilosa. Both populations of B. pilosa had lower metabolic rates in summer than in winter, whereas the metabolic rate of B. pelagica remained much the same. Seasonal changes in metabolic rate are closely correlated with reproductive cycles. The possible influences of environmental parameters are discussed.     

Genetic adaptation to local habitats and speciation processes within the genus Gammarus (Amphipoda: Crustacea)

A recent event of allopatric speciation (a time span of about 4 000 years) is reported from a comparison between two guilds of amphipod gammarids from the central Baltic Sea and from the Limfjord (inner Danish waters), respectively. For at least two of the five species populations involved, a genetically fixed habitat specialization has led to the establishment of reproductive barriers. It has, however, not been possible to correlate this speciation event with electrophoretically determined isozyme changes. It is argued that the lack of correlation is due to the presence of an initial time lag before a speciation event can be detected as isozyme differences between newly formed species populations as predicted by the neutrality hypothesis for evolution at the molecular level.     

Lipid biochemistry and reproductive biology in two species of Gammaridae (Crustacea: Amphipoda)

Gammarus oceanicus Segerstråle, 1947 and Echinogammarus marinus (Leach, 1815) were sampled during the breeding season from Oslofjord in 1984, and their lipid composition examined in relation to reproductive condition. In G. oceanicus, female lipid content increased as the ovary matured. Both the amount of lipid stored and the rate of accumulation were greater in spring than in winter. Spring eggs contained 12.4 g lipid, of which 63% was triacylglycerol and 27% phospholipid. Both fractions decreased steadily during embryonic development. Winter eggs contained 19.2g lipid, of which 52% was triacylglycerol and 43% phospholipid. During the early stages of embryonic development the amount of phospholipid decreased sharply, whereas that of triacylglycerol increased, suggesting that some of the fatty acid released from phospholipid was sequestered temporarily as triacylglycerol. When newly spawned, both winter and spring eggs were richer in monoenoic fatty acids than adult amphipods and these acids were the major fuel consumed during development. 6 fatty acids were utilised more slowly than 3 acids, and egg carotenoid pigment content remained constant. Female E. marinus increased in lipid content as the ovary matured. Spring eggs contained 14.7 g lipid when newly spawned and this increased to 16.6 g during the early stages of development. This increase was entirely triacylglycerol, which declined in later stages; the source of the extra lipid was unclear. Eggs contained very little phospholipid or sterol, and both of these components remained at a steady low level during development. E. marinus eggs were not significantly rich in thonoenoic acids compared with adults, and saturated, monoenoic and polyenoic acids were utilised about equally during development. Both adults and eggs were rich in 20.46, which was utilised at a slower rate than the 3 polyunsaturated acids during embryonic development; again, egg carotenoid pigment content remained constant. In both species there was a decrease in the size of the egg (and as a result, of the newly hatched juvenile), but an increase in total reproductive output (i.e., the total weight of the egg clutch) per female as the breeding season proceeded. The reproductive output of an individual female is probably related to food availability during the period of ovarian maturation, whereas the size of an individual egg is dictated largely by feeding conditions for the juveniles once they are independent of the female. The different patterns of lipid utilisation during development found in this study emphasize the flexibility of response in the reproductive biology of gammarid amphipods. It is not yet possible, however, to relate the differing patterns in a simple way either to egg size or total female reproductive output. Two outstanding problems are the source of extra triacylglycerol during the early stages of development of E. marinus and the metabolic cost of brooding eggs.     

Trace element distribution and mobilization in Scottish soils with particular reference to cobalt,copper and molybdenum

Total and extractable trace element contents have been determined in about 1000 soil profiles representing the main soil series occurring throughout Scotland. The frequency distributions of a number of trace elements in these soils are described and some relationships between total and extractable contents discussed. The geological nature of the soil parent material, soil texture, organic matter content and environmental contamination are the principal factors controlling soil total contents. Soil drainage class, because of its effect on mineral weathering and soil oxidation-reduction conditions, has a major influence on soil extractable contents, availability to plants and crop uptake. Particular attention is paid to cobalt, copper and molybdenum because of their importance for animal health in Scotland.     

Effects of temperature and salinity on larval development ofNecora puber (Brachyura: Portunidae)

Temperature and salinity affected both length of larval development and mortality inNecora puber collected in the Ría de A Coruña during December 1984 and January 1985. Development time decreased considerably with increased temperature. This decrease was sharper when temperature increased from 15° to 20°C than when it increased from 20° to 25°C. At 35S, average development took 48, 32 and 28 d at 15°, 20° and 25°C, respectively. At the three salinities tested (25, 30 and 35), larval development was completed only at 15°C, at 20°C/30 and 35S, and at 25°C/35S. Development times at 15° and 20°C were highly significantly different at both 35 and 30S (P 0.01). However, there were no significant differences between development times at 20° and 25°C (P > 0.05). Within any one specific temperature series, no significant difference was observed between the salinity values tested (P > 0.05). The duration of each of the five zoeal stages was similar within each and the same temperature/salinity combination, whereas the duration of the megalop was twice as long as any of the zoeal stages. The combination of the lowest temperature (15°C) and the highest salinity (35) tested resulted in the greatest larval survival of 28%. Highest mortality occurred at 25°C, at which temperature development was completed only at 35S. A sharp drop in larval survival was observed in the transition period Zoea V — megalop in all combinations of temperature and salinity tested. Within the limits of tolerance to temperature and salinity, the former effected more pronounced differences in the duration of larval development, while salinity appeared to constitute a limiting factor for survival.     

Life history and production of <Emphasis Type="Italic">Corophium urdaibaiense</Emphasis> (Crustacea: Amphipoda) in the Urdaibai estuary (NE Spain)

The population of species Corophium urdaibaiense has been studied for a year in the Urdaibai estuary (NE Spain). The new species is characterised by its preference for muddy sand sediments (15% of mud) and the long body size (with a maximum length of 10.24 mm). The maximum abundance occurs in November (11,600 ind m⁻²); it almost disappears at the end of spring. The juveniles represent 79% of the population and the recruitment is concentrated within the October–December period. The breeding period begins in May, when the temperature is above 15°C, and it ceases in November, when the temperature is below 15°C. Bivoltinism is probable, taking into account the latitude (43°), temperature range (12–25°C) and breeding period (May and August–October), but the data are not sufficiently consistent to confirm the hypothesis. The production ranges from 2.93 to 5.85 g m⁻² year⁻¹, considering one and two generations per year, respectively. Some aspects concerning the food availability, predation and intraspecific competition could have an influence on the dynamics of the population and are discussed.     

Anatomy of structures associated with air-breathing in Orchestia gammarellus (Crustacea: Amphipoda: Talitridae): coxal plates and gills

The structure of the coxal gills and coxal plates of the semi-terrestrial beachflea Orchestia gammarellus (Pallas) (Crustacea: Amphipoda: Talitridae) is described in relation to their possible use for aerial gas exchange and ion exchange. Anatomical evidence is presented to support the hypothesis that the medial surface of the coxal plates functions as an extrabranchial aerial gas-exchange site in O. gammarellus. Thus, the effective diffusion distance across the medial (or inside-facing) surface of O. gammarellus coxal plates (mean±SD=5.4±0.3 m; n=9, cuticle thickness 4.4±0.5 m, n=21) is only a third of the equivalent distance across both the coxal gills (18.4±6.0 m, n=10; cuticle thickness 1.7±0.6 m, n=7) and the lateral (or external) surface of the coxal plates (19.4±0.7 m, n=5; cuticle thickness 8.7±0.8 m, n=7). Chloride-ion-permeable areas were located using a silver-staining technique. All ten coxal gills appeared to be equally permeable to chloride ions after examination with a light microscope. However, the coxal plates and the rest of the integument do not appear to be chloride-permeable.     

Advantages and a disadvantage of large size for male gammarus pulex (Crustacea: Amphipoda)

Summary The breeding system of the freshwater amphipod Gammarus pulex includes a precopulatory guarding phase by a male. The length of this guarding phase is investigated with respect to a male's size and the number and size of his competitors. As the absolute number of competitors increases, so does the guarding time but as the absolute number of available females increases, the guarding time decreases. Takeovers of the females by unpaired males are more frequent in longer precopulas (Table 2). In contests for females, larger males have two advantages over smaller males; they are better able to make a takeover (Table 2) and better able to resist takeover attempts while paired (Table 3). Males increase the length of the guarding phase as the mean size of their competitors increases (Table 4). When not paired males are usually searching for available females, perhaps in the stream current. Females are unaffected by current speed but increasing current causes decreased male survivorship (Table 5) and increased precopula duration (Table 5). Searching in currents is more dangerous for larger males than smaller ones. It is proposed that the male size distribution observed is the result of selection pressure to increase size from male-male competition balanced by selection against large size while searching for females in the current.     

Synergistic effects of salinity,temperature and heavy metals on mortality and osmoregulation in marine and estuarine isopods (Crustacea)

The effects of cadmium (3CdSO₄·8H₂O), zinc (ZnSO₄·7H₂O) and lead [Pb (NO₃)₂] on mortality, and cadmium, zinc and mercury (HgCl₂) on osmoregulation, have been recorded for marine and estuarine species of isopods (Crustacea). The marine species studied were Idotea baltica, I. neglecta, I. emarginata and Eurydice pulchra, which were adapted to 100, 80, 60 and 40% sea water (SW) (100% SW э 34‰ S). The estuarine species used were Jaera albifrons sensu stricto and J. nordmanni, which were adapted to 100, 50, 10 and 1% SW. Both groups of isopods have low mortalities in 100% SW with 10 and 20 ppm of cadmium, zinc and lead, but a decrease in salinity caused an increase in the toxicities of these metals and reduced the LT₅₀ values (time, in hours, to 50% mortality). Mortalities at 10°C were generally higher than those recorded at 5°C. Cadmium had no significant effect on the osmoregulation of I. baltica and I. emarginata in 100 and 80% SW at 5°C, but this metal significantly lowered the blood osmotic concentration of I. neglecta in 80% SW. Zinc did not alter the haemolymph osmotic concentration of I. neglecta in 100 and 80% SW, but significantly lowered the blood osmotic concentration of I. baltica in 100% SW. Cadmium, zinc and mercury also significantly altered the osmoregulatory ability of J. albifrons in dilute saline.     

An opinion on the distribution and behavior of chemicals in response to climate change,with particular reference to the Asia-Pacific region

There is a general lack of knowledge as regards the effects of climate change on pollutant behavior. This is particularly true of the Asia-Pacific Region (APR). This region has major significance in terms of global pollutant emission and also displays a wide variety of environments. This review presents the authors’ opinions on possible implications of climate change for pollutant behavior in the APR. Although differing responses can be expected across the region, there are clear implications as regards the short- and long-term behavior of pollutants. Effects can be predicted through modeling, but further data are required for model calibration. Nevertheless, it can be predicted that climate change will affect processes including global distillation of persistent organic pollutants, airborne transport of heavy metals, half-life of readily degradable pollutants, and eutrophication in water bodies. Particulates are expected to play a central role in mediating the effects of climate change, and successful predictive models will need to be based on particulate-mediated transport and behavior. Climate change also has the potential to cause an increase in the intensity and frequency of harmful algal blooms in aquatic environments throughout the region, with significant implications for supply of both food and drinking water.