Production of Gammarus pulex L. (Amphipoda) in a small Danish stream

Quantitative samples of Gammarus pulex L. taken from a small Danish stream during 1975 showed mean annual population densities varying from 500 m^–2 in early May to 5 500 m^–2 in late September. The mean annual biomass was 1.5 g dry weight m^–2. No discrete cohorts could be distinguished from the size frequency distributions. Annual production, estimated by the size-frequency method, was 3.9 g dry weight M^-2 and P/B ratio was 2.6. The contribution to trout energetics may have been as much as 17%.     

Calcium balance in premoult and post-moult Gammarus pulex (Amphipoda)

SUMMARY. Adult Gammarus pulex lose about 42% body calcium into solution over a 2–3-day period preceding the moult. A further 54% body calcium is lost with the exuviae, leaving c . 4% in the newly moulted animal.
Nearly all of the body calcium in a stage A (post moult) animal is contained in the hepatopancreas and a transient increase in the calcium content of this tissue is seen in some individuals. The haemolymph calcium compartment is heavily depleted at this stage. In stage B animals the hepatopancreas calcium level has returned to, or even fallen below, the intermoult level while the haemolymph calcium concentration remains lower than in intermoult animals.
Animals switch from a strongly negative calcium balance to a state of rapid calcium uptake immediately following moult. Uptake at 10°C proceeds at a rate of between 3 and 10 μmol g⁻¹ h⁻¹ depending on the calcium concentration of the external medium. The lower rate is found in starved animals in 0.1 mM calcium and this concentration is probably close to a minimum for the satisfactory restoration of body calcium. Recalcification is completed in 10–14 days in 0.1 mM calcium and is apparently enhanced by the presence of food in the form of a portion of oak leaf together with the cast exuviae. In 1.0 mM calcium the recalcification period is shortened to 3–4 days. This is considerably longer than the time recorded for French populations of this species (Vincent, 1969), although reasons for this are offered.     

Life-cycle, drift and production of Gammarus pulex L. (Amphipoda) in a Danish spring

SUMMARY. Gammarus pulex L. was studied at three sites in a Danish spring. Breeding took place during the summer and therefore cohorts could be followed. Sexual maturity was not reached until the next summer and maximum age appeared to be two years.
Changes in population size differed between stations and was tentatively related to differences in drift and numbers of predators. Generally the drift was non-selective between cohorts and within cohorts.
Yearly production estimated from production curves was 2.6 g dry weight m⁻² (the young cohort) and 1.2 g dry weight (the old cohort) at Station 1. The respective P / ratios were 3.35 and 1.22. Production of the youngest cohort at the three stations in the period September-May was 0.9–1.3 g m⁻² and P / ratios were 2.14–2.31. Total yearly production at the three stations was estimated at 4–7 g dry weight m⁻².     

Experiments on feeding and growth of the amphipod Gammarus pulex (L.) related to its distribution in the River Duddon

Individuals of Gammarus pulex were kept at 15°C for periods of 23–70 days on diets comprised of elm or oak leaves, Tricladium and Clavariopsis (Hyphomycete fungi), Molinia (grass), Zygogonium (alga), Nardia (liverwort). Both the mean interval between moults (m_i), and daily increments in body wet weight, were related to diet. The lowest value for m_i was 14·6 days on a diet of naturally decaying elm and oak leaves. The largest weight gains also occurred on this diet; the average daily gain in weight (D_w) = 130·8 μg/day, the mean specific gain in weight (G_w) = 1·69 μg %/day. On green Molinia and fungi diets D_w ranged from nil to 60·7 μg/day, and m_i= 17·4–18·4 days was significantly (P<0·01) longer than m_i on leaf diets. Survival and growth were poor on brown Molinia. Nardia did not support growth or survival. A mixed diet of Tricladium, Zygogonium, Nardia and decaying grasses was not sufficient to promote an increase in body weight, and m_i= 21·8 days. This diet represents the commonly available food materials in Mosedale Beck, an acid stream in the upper Duddon catchment. Survival and growth of G. pulex on fungi and leaf diets in media containing low concentrations of potassium ions are also described and discussed briefly in relation to the distribution of G. pulex in the upper catchment of the R. Duddon.     

Parasite altered micro-distribution of Gammarus pulex (Crustacea: Amphipoda)

In a river survey, Gammarus pulex amphipods both unparasitised and parasitised with the acanthocephalan Echinorhynchus truttae were distributed similarly with respect to flow regimen, tending to be more abundant in faster, shallower, riffle patches. However, there was a higher prevalence of parasitism in faster, shallower areas than in slower, deeper areas and abundance correlated with macrophyte coverage for unparasitised but not parasitised amphipods, indicating subtle differences in habitat usage. A laboratory 'patch' simulation indicated that parasitism influenced micro-distribution. There were higher proportions of unparasitised amphipods in/under stone substrates and within weed. In contrast, there were higher proportions of parasitised amphipods in the water column and at the water surface. As the experiment progressed, unparasitised but not parasitised amphipod habitat usage shifted from those micro-habitats above the substrate and in the water column to those in/under the substrates. Experiments also demonstrated that parasitised amphipods were more active and had a greater preference for illumination. Previous studies of the effects of acanthocephalan parasitism of amphipod hosts have focussed on how drift behaviour is altered, now we show that subtle differences in micro-habitat usage could translate to greatly increased vulnerability to fish predation. We discuss how aggregation of parasitised individuals within specific habitats could promote parasite transmission.     

Effects of population density and of presence of Gammarus pulex L. (Amphipoda) on the growth in larvae of Potamophylax cingulatus steph. (Trichoptera)

Growth rates and mortality of Potamophylax cingulatus were studied at different densities, temperatures, and food items, as well as in the presence and absence of Gammarus pulex. When alder leaves were supplied as food as well as beech leaves at 15° C, an interspecific competition was observed. The potential influence of competition in a free living population as a component in the heavy ‘natural mortality’ was discussed.     

Effects of pH,humic substances and animal interactions on survival and physiological status of Asellus aquaticus L. and Gammarus pulex (L.)

Summary The mortality and physiological status (body water content) of Asellus aquaticus (Isopoda) and Gammarus pulex (Amphipoda) were measured after 25 days exposure in 20 natural streams with a pH range of 4.3–7.5 and a colour range of 8–280 mg Pt L^–1. In addition, the effects of keeping the animals as single species or together were studied. The response of Gammarus to low pH was an increased mortality and lower physiological status of surviving individuals in streams with a pH lower than 6.0. In Asellus the physiological status was correlated with pH, while the mortality was not pH dependent. The effects of humus on the physiological status of Asellus was significant when fitted to a second order polynomial function. The influence of humus can, however, be regarded as small relative to pH. The interactions between the species could be described as asymmetric under optimal conditions of high pH and low humus concentrations, where the presence of Gammarus decreased the survival and physiological status of Asellus. Acid stress did not seem to reverse the direction of this asymmetry, but the presence of Gammarus improved the physiological status of Asellus at pH lower than 6.0. Since the presence of Asellus did not increase the mortality or decrease the physiological status of Gammarus, this could be explained by Asellus feeding on Gammarus that died from physiological stress solely. This mechanism suggests that food quality, and thus effects of diffuse competition, can be important for the ability to withstand acid stress. The results, though, give no support for the hypothesis that competition from Asellus is important for the disappearance of Gammarus during the acidification of streams.     

Gut passage times in Gammarus pulex (Crustacea,Amphipoda) and aspects of summer feeding in a stony stream

Gut passage times in Gammarus pulex (Crustacea, Amphipoda) were calculated by counting and measuring voided faecal pellets. Following feeding on elm leaves conditioned in a stream, or summer fine detritus from the same source, calculated gut passage times were usually short, about 2 h or less at 15 °C. Using harder oak and beech leaves, not previously conditioned in the stream, apparent gut passage times were much longer, 8–48 h, but since intermittent feeding occurred these figures are not comparable to the others.In connection with summer feeding of the animal in the stream, the microbiological status of the fine detritus there was studied. It contained fungal hyphae, which were mostly empty, viable fungal spores of terrestrial derivation, and bacteria, all largely carried on fragments of vascular plant tissues. In experiments, such fungal spores (of Aureobasidium pullulans and of Mucor sp.), incorporated into fine detritus, were not digested by the animal. The bacterium Bacillus cereus was present in the fine detritus and when the latter was consumed it survived passage through the gut of the animal. Using B. cereus as a proportion marker it was concluded that other unicellular bacteria in the fine detritus were not digested either. In other experiments, not immediately related to the summer season, nutrient extraction by the animal from hyphae of the fungus Nectria lugdunensis was examined; it probably occurs through sub-microscopic pores connecting the cells. This mode of extraction may apply when the animal consumes vascular plant tissues.     

Factors affecting the microdistribution of Gammarus pulex (Amphipoda): an experimental study

SUMMARY. 1. The tnierodistribution of Gammarus species is size assortative: larger animals are associated with larger substrate particles. Using an artificial stream with scrubbed substrate we investigate the hypothesis that the assortative mechanism is stimulated by current avoidance rather than food searching.
2. In uniformly mixed substrate, G. pulex tend to move upstream, but in patchy substrate they are found in a predictable substrate grade. There is no discrimination between patches by different size classes of G. pulex in static water. The size assortative field pattern can only be reproduced under flow conditions in the laboratory. It occurs in the absence of food and irrespective of the sequence or location of substrate patches.
3. As density increases, G. pulex are displaced from the optimal patches but move into the open stream rather than into the cover of alternative patch types.
4. We discuss the significance of these results and suggest that some earlier models explaining mechanisms for microdistribution patterns tnay have been unnecessarily complex. Contrasts between natural and simulated stream situations and their experimental advantages are noted.     

State-dependent pairing behaviour in male Gammarus pulex (L.) (Crustacea, Amphipoda): effects of time left to moult and prior pairing status

Because mating can be costly in terms of time and energy, an individual's propensity to engage in courtship and mating activities might be modulated by its physiological state. However, so far, state-dependent mate choice has received little attention The present study examined the effect of both prior pairing status and time left to the moult on the ability of male Gammarus pulex (Crustacea, Amphipoda) to enter in precopula with receptive females. In the lab, males that were freshly collected in precopula pairs in the field had a higher probability of re-pairing and were quicker to enter in precopula with receptive females compared to males of similar size that were freshly collected unpaired. In addition, unpaired males found in the field were closer to their moult than paired males. Considered together, our results strongly suggest that time left to the moult and prior mating status directly influence male propensity to pair in G. pulex.     

Isolation and characterization of the first eight microsatellite loci in Gammarus fossarum (Crustacea, Amphipoda) and cross-amplification in Gammarus pulex and Gammarus orinos

We characterized the first microsatellite markers for Gammarus fossarum. Eight loci gave satisfactory amplification patterns in two stream populations (Southern France) with number of alleles ranging from 2 to 10 and expected heterozygosity from 0.076 to 0.857. We performed cross-amplification in two closely related gammarid species, Gammarus pulex and Gammarus orinos. Among the eight tested microsatellite loci, four correctly amplified in G. pulex and three in G. orinos.     

Effects of two acanthocephalan parasites on the fecundity and pairing status of female Gammarus pulex (Crustacea: Amphipoda)

Acanthocephalan parasites are known to alter the reproductive biology and physiology of their hosts in various ways. In this study we investigated the influence of two acanthocephalan parasites, Pomphorhynchus laevis and Polymorphus minutus, on the fecundity and pairing success of female Gammarus pulex. The results show that P. laevis and P. minutus affect female intermediate host reproduction in different ways. Females infected with P. minutus were totally castrated, whereas those infected with P. laevis only showed reduced fecundity. The oocytes of P. laevis-infected females showed a similar structure to those of uninfected females, although infected females had a higher proportion of oocytes that had failed to reach complete maturity. In comparison, the oocytes of P. minutus-infected females demonstrate a clearly altered structure that suggests a major disruption to the process of vitellogenesis. In the field, males paired more frequently with uninfected females than with infected ones, and is a stronger effect for P. minutus-infected females than P. laevis-infected females. We suggest that the difference in pairing success of P. minutus-infected and P. laevis-infected females is a direct result of the different effects that the two parasites have on female fecundity.     

The digestion of cellulose and other dietary components, and pH of the gut in the amphipod Gammarus pulex (L.)

SUMMARY. Gut extracts from Gammarus pulex hydrolysed native and other cellulose substrates in vitro. Digestive fluid cellulase is probably endogenous as cell-free fluid mediated cellulose hydrolysis, but no bacteria were isolated from the fluid which produced a detectable extra-cellular cellulase. There was no apparent digestion of plant cell walls during their passage along the digestive tract, which took about 5–7 h at 10°C. The pH sensitivities of the digestive enzymes and the pH of the various regions of the gut suggest that carbohydrate digestion occurs in the proventriculus, midgut glands and anterior midgut, but protein digestion may be largely limited to the posterior midgut. The pH of the digestive fluid was altered slightly, but significantly, by the consumption of different natural and artificial test diets and by starvation. The most probable reason for the non-digestion of plant cell-walls is the lack of necessary enzymes other than cellulase. The role of cellulase may be confined to digesting the many small, non-cellular particles which are present in the gut.     

Presence of sculpins (Cottus gobio) reduces drift and activity of Gammarus pulex (Amphipoda)

Introduction of sculpins into a stream previously devoid of these predators significantly reduced drift rate of Gammarus pulex. The drift of insect larvae was not affected. High amounts of exudates after implantation of sculpins were probably responsible for the low number of drifting G. pulex specimens. Laboratory experiments confirmed reduced locomotory activity of G. pulex when exposed to caged sculpins, an observation that excludes reduced drift activity as a result only of predation.The average size of drifting G. pulex specimens was larger during the night than during the day. This result is in accordance with the hypothesis that large individuals should, in relation to small ones, turn nocturnal because of greater predation risk during daytime. Presence of sculpins did not alter the size composition of drifting G. pulex.     

Population characteristics of Gammarus pulex (L.) from five English streams

Gammarus pulex were sampled from five English streams during April 1992. The population density, number of precopula pairs and incidence of parasitic infection were recorded, and the biomass was estimated from subsamples by relating body area to dry weight. Physical and chemical measurements were taken from each stream. The abundance and standing crop biomass differed significantly between streams, probably due to the influence of pollutants or the physical structure of the stream bed. The size of individual G. pulex also differed significantly between streams, although there was no obvious causal explanation for this. Few individuals were visibly parasitised in any of the populations. Males were significantly larger than females, both in precopula pairs and in the general populations. The sex ratio differed between populations and may explain inter-stream differences in the relationship between precopula male and female size.     

Comparative ecology of Gammarus pulex (L.) and Asellus aquaticus (L.) II: fungal preferences

This paper continues to explore niche differentiation in Gammarus pulex and Asellus aquaticus by analysis of their food preferences. Individuals from both species discriminated between leaf discs colonized by different fungal species and exhibited strong preferences for Anguillospora longissima and Heliscus lugdunensis. Fungal preferences were not correlated with the relative abundance of fungi in the field and there was considerable intra-population variability in food preferences — both between individuals and for the same individual through time. Niche overlap between animals from all four study populations was high and there was no evidence of differences in the potential trophic niches of animals from sympatric and allopatric populations.     

Anaerobic capacity of a crustacean sensitive to low environmental oxygen tensions,the freshwater amphipod Gammarus pulex (L.)

We investigated the accumulation of haemolymph L-lactate in response to anoxia, progressive hypoxia and activity, in a species that rarely encounters low PO₂ in its natural environment, the freshwater amphipod Gammarus pulex (L.). Individuals survived <1.5 h anoxia and accumulated L-lactate in the haemolymph at a rate of 10.26 mmol l^–1 h^–1. No lactate was excreted into the medium during this time. Despite the fact that most individuals of G. pulex were oxyconformers, exposure to acutely declining PO₂s did not result in a switch to anaerobic pathways until PO₂ < 4.4 kPa. Even then the concentrations accumulated remained very low until PO₂ < 0.46 kPa. There was no accumulation of L-lactate in the haemolymph as a result of exhaustive activity. Consequently, it was suggested that (a) it cannot be automatically assumed that anaerobic metabolism is switched on only at the point at which oxyregulation breaks down, (b) that the poor anaerobic capacity observed plays little, if any, role in the metabolic response to hypoxia by Gammarus pulex.     

Effect of zinc on four populations and two generations of Gammarus pulex (L.)

1. The amphipod Gammarus pulex (L.) is increasingly used in toxicity assessments and is usually obtained from wild populations. Interpopulation variability in response to toxicants may be due to genetic or phenotypic differences and could be large in wild-caught organisms exposed to different environmental conditions. This paper describes an investigation designed to assess the extent of lethal and sublethal variability between four different G. pulex populations when exposed to zinc, and to determine whether differences observed in the parental generation were also apparent in offspring bred in the laboratory. 2. The mortality of G. pulex from one of the four field-caught populations (Crags Stream) was significantly lower than for animals from the other three populations for the first 2 days of exposure. After 6 days’ exposure the LC₅₀ values for all four populations did not differ significantly, and were approximately 1.0 mg Zn 1-^?1. Sublethal effects on feeding rate after 6 days’ exposure were also similar for all four populations (feeding rate EC₅₀ approximately 0.5 mg Zn 1-^?1). 3. Lethal effects on the F₁ generation were similar among the four populations, in contrast to the short-term differences observed in the parental generation. These results suggest that interpopulation differences in zinc tolerance in G. pulex are short-lived, and phenotypically rather than genetically based.