A. F. Pasternak

Feeding, metabolism and body composition of the Antarctic copepod Calanus propinquus Brady with special reference to its life cycle

SummaryIn the Weddell Sea, several biochemical and physiological characteristics of copepodite stage III to adult females of Calanus propinquus from the surface layer (0–100 m) and the deep layer (500–1,000 m) were compared at a time of high phytoplankton abundance (February) and a month later, when chlorophyll concentration was much lower and older copepodite stages had migrated to their overwintering depth. Daily rations of surface animals, estimated by the gut fluorescence method, varied from 0.8% to 7.8% body carbon (increasing with phytoplankton concentration). Respiration rate (calculated from measurements of electron-transport-system activity) and lipid and protein body content of animals inhabiting the surface layer decreased from February to April. Deep water animals (stage V and adult females) were characterized by high lipid and protein content; their respiration and excretion rates were much lower than in surface copepods. Our calculation showed that they could survive without an additional source of energy for 8–9 months. Based on our own and published data on bio-chemistry and physiology the possibility of a two-year life cycle is suggested.

Gut fluorescence in herbivorous copepods : an attempt to justify the method

Recently the gut fluorescence technique has been critisized because of the possible degradation of chlorophyll into nonfluorescent derivatives during passage through copepod guts and changes of the gut passage time with food concentration. Here pigment budgets have been calculated in 6 experiments with Calanus finmarchicus CIV caught 2 km offshore of the Murmansk Marine Biological Institute (the Barents Sea, Dalnije Zelentsi) in September 1992. Copepods were fed with culture of Platymonas viridis at different concentrations. Gut pigment and ingestion rate increased with food concentration in a similar way. On average between 78% and 89% of ingested chlorophyll was recovered in the guts and faecal pellets. No trend for a greater loss of fluorescence at low food concentration than at high was observed. Pigment content of faecal pellets incubated in filtered seawater decreased by 20–30% in the first 7–12 h and by up to 60% in 48 h. The decline of pigment content was accompanied by a rapid bacterial growth (by a factor of 3 in 48 h). Gut passage time increased with decreasing food concentration (from 40 min at 9 µg pigm l−1 to 64 min at 0.9 µg pigm l−1). These results together with some data by other authors suggest that the gut fluorescence method can be used to estimate in situ grazing rate providing gut passage time is measured properly and there are no losses of faecal material. However, careful consideration should be given to the previous feeding history of copepods.

EFFECT OF TRIAENOPHORUS CRASSUS (CESTODA) INFECTION ON BEHAVIOR AND SUSCEPTIBILITY TO PREDATION OF THE FIRST INTERMEDIATE HOST CYCLOPS STRENUUS (COPEPODA)

Some parasites have been shown to manipulate host behavior so that parasite transmission to the next host is enhanced. Infection with Triaenophorus crassus Forel (Cestoda) caused alterations in the activity and microhabitat selection of the first intermediate host Cyclops strenuus Fischer (Copepoda) in the laboratory. Infected copepods made more starts to swim but spent less time swimming than uninfected copepods. These changes were independent of the intensity of infection. In a water column illuminated from above, infected copepods approached the surface, whereas uninfected ones remained close to the bottom. In the dark both infected and uninfected copepods stayed near the bottom. Finally, infection with T. crassus increased the probability of C. strenuus being eaten by the second intermediate host, whitefish (Coregonus lavaretus L. s.l.), in the laboratory. In experimental infections, 10-day-old procercoids had significantly lower infectivity for whitefish than older (12-, 14-, and 21-day-old) procercoids. Behavioral changes were detected in infected copepods containing procercoids 12 days old or older but not in experiments with 10-day-old procercoids. These results may indicate that T. crassus changes the behavior of the copepod host only after it has become infective to the next host, which is consistent with the active manipulation hypothesis.

Feeding patterns of dominant Antarctic copepods: an interplay of diapause, selectivity, and availability of food

Gut contents and feeding activity of five dominant Antarctic copepods (Calanus propinquus, Calanoides acutus, Rhincalanus gigas, Metridia gerlachei and Microcalanus pygmaeus) were studied from samples collected during several cruises of the RV Polarstern to the eastern Weddell Sea. In summer, feeding activity, estimated as percentage of copepods with food in the guts, was high in all the species, and diatoms dominated all gut contents. In winter, C. acutus was trophically inactive, and C. propinquus and R. gigas considerably decreased their feeding activity, while a decrease in feeding of M. gerlachei and M. pygmaeus was less pronounced. Unidentified mass dominated gut contents in winter, supplemented by phytoplankton and protozoans. Prior to the spring bloom, feeding activity of C. acutus was low, with unidentified food predominating, while carnivory was important in actively feeding C. propinquus. Rhincalanus gigas tended to be more carnivorous than C. acutus, however with less feeding activity than C. propinquus. Seasonal changes in feeding patterns are discussed.

Tuning host specificity during the ontogeny of a fish ectoparasite: behavioural responses to host-induced cues

The choice between two alternative hosts, brown trout (Salmo trutta) and roach (Rutilus rutilus), and the response to visual and olfactory cues were studied in the ontogeny of Argulus coregoni. The initial preference of the smallest parasites for brighter roach changed at the age of 2xa0weeks, at the size of about 2xa0mm, for trout, a typical salmonid host. Younger argulids were attracted by a non-specific visual stimulus (white disc over dark background), and they did not respond to olfactory stimulation (fish-conditioned water). Later, the response to visual stimuli was modulated by trout-conditioned water, but not by that conditioned by roach. The primary role of vision, particularly in early ontogeny, is emphasized as an adaptation of A. coregoni to habitats in boreal latitudes, clear and running water with a sparse fish population. In sub-adult and adult parasites, vision is complimented by olfaction that enables them to choose hosts more precisely. The nature and adaptive significance of the ontogenetic shift in host choice by A. coregoni is discussed.

Light-mediated host searching strategies in a fish ectoparasite, Argulus foliaceus L. (Crustacea: Branchiura)

Argulus foliaceus, an obligate fish ectoparasite, can search for its hosts in both light and dark conditions and uses vision in the light. We have examined what searching mode is used at night, when the infection rate was at its highest, and which stimuli produced by the fish are most important. A change of illumination produced a clear difference in the searching behaviour of adult Argulus females. The mean swimming speed and the area explored were 3-4 times higher in the dark, when the parasite employed a cruising search strategy. This changed to an ambush (hover-and-wait) strategy in the light. The swimming activity is accompanied by changes in metabolic costs; the activity of the electron transport system being approximately 25 % lower in the light. The most pronounced light-induced differences in host-searching behaviour took place in moderately hungry parasites (starved for 24-96 h). Less motivated (just having left a fish) or exhausted animals did not exhibit any clear differences in swimming speed. Among the external signals tested, fish smell, from both perch (Perca fluviatilis) and roach (Rutilus rutilus), induced an elevated swimming speed of the parasite. Periodic water movements caused similar but weaker effects. The effects of these stimuli were observed under both light and dark conditions. We conclude that host-searching behaviour of A. foliaceus is under internal (state of hunger) and external (illumination and host-induced signals) control and involves all its sensory equipment (vision, olfaction and mechano-reception). Perch (but not roach) reduced their swimming speed in the dark, which make them more susceptible to cruising Argulus. Thus the behavioural interplay between hosts and parasites can also influence the infection rate of A. foliaceus found on perch and roach in Finnish lakes.

Parasite-induced aggression and impaired contest ability in a fish host

BackgroundSuccess of trophically transmitted parasites depends to a great extent on their ability to manipulate their intermediate hosts in a way that makes them easier prey for target hosts. Parasite-induced behavioural changes are the most spectacular and diverse examples of manipulation. Most of the studies have been focused on individual behaviour of hosts including fish. We suggest that agonistic interactions and territoriality in fish hosts may affect their vulnerability to predators and thus the transmission efficiency of trophically transmitted parasites. The parasite Diplostomum spathaceum (Trematoda) and juvenile rainbow trout, Oncorhynchus mykiss, were used to study whether infection can alter aggression rates and territorial behaviour of intermediate fish hosts.ResultsThe changes in behaviour of rainbow trout, Oncorhynchus mykiss, infected with an eye fluke Diplostomum spathaceum (Trematoda), was monitored over the course of an experimental infection for 1.5 months. At the beginning of their development, not yet infective D. spathaceum metacercariae decreased the aggressiveness of rainbow trout. By the time that metacercariae were fully infective to their definitive hosts, the aggressiveness increased and exceeded that of control fish. Despite the increased aggressiveness, the experimentally infected fish lost contests for a territory (dark parts of the bottom) against the control fish.ConclusionsThe results obtained indicate that the parasitized fish pay the cost of aggressiveness without the benefit of acquiring a territory that would provide them with better protection against predators. This behaviour should increase transmission of the parasite as expected by the parasite manipulation hypothesis.

Seasonal dynamics of egg laying and egg-laying strategy of the ectoparasite Argulus coregoni (Crustacea: Branchiura)

Substrate preferences, spatial aggregation patterns and seasonal dynamics in the egg laying of ectoparasitic Argulus coregoni were studied at a commercial fish farm in Finland. Pilot experiments showed that A. coregoni females selected specific types of substrates for egg laying. Significantly more A. coregoni eggs were laid on dark substrates than on light ones suggesting the use of visual cues. Therefore, egg-laying plates of dark colour were constructed for further experiments. Most A. coregoni eggs were deposited in locations in shadow and in the deepest water in a 2 m deep farming canal. Relatively more eggs were laid on bottom stones situated near each egg-laying trap than on artificial egg-laying plates indicating a preference for irregular stones in the deeper locations in the canal. The plates were located 20 cm above the bottom. However, a total of 5,863 A. coregoni egg clutches, corresponding approximately to 1.5 million unhatched metanauplii, were successfully destroyed with the plates indicating that egg-laying traps can be used as an ecological control method against argulids in certain situations. For traps to be effective, ponds should not contain stones or any other hard substrata attracting female lice. The egg laying of A. coregoni in this study started on 5 July in 2001 and extended over 3.5 months up to mid-October. The egg-laying pattern of A. coregoni population was unimodal, supporting the view that only a single A. coregoni generation occurred annually in Central Finland.

Gut contents and diel feeding rhythm in dominant copepods in the ice-covered Weddell Sea, March 1992

Zooplankton was sampled at a diurnal station during the drift of the Ice Station Weddell-1 in the western Weddell Sea in March 1992. Gut contents of Metridia gerlachei, Calanoides acutus, Calanus propinquus and Rhincalanus gigas were studied. Calanoides acutus was trophically inactive, but 18%, 33% and 45% of Calanus propinquus, R. gigas and M. gerlachei, respectively, contained food in their guts. Diel changes in gut levels and percentage of individuals with full guts with maximum during the dark period were observed in M. gerlachei copepodite stage V and females. The possible reasons for observed differences in foraging patterns of the four copepod species are discussed.

Grouping facilitates avoidance of parasites by fish

BackgroundParasite distribution is often highly heterogeneous, and intensity of infection depends, among other things, on how well hosts can avoid areas with a high concentration of parasites. We studied the role of fish behaviour in avoiding microhabitats with a high infection risk using Oncorhynchus mykiss and cercariae of Diplostomum pseudospathaceum as a model. Spatial distribution of parasites in experimental tanks was highly heterogeneous. We hypothesized that fish in groups are better at recognizing a parasitized area and avoiding it than solitary fish.MethodsNumber of fish, either solitary or in groups of 5, was recorded in different compartments of a shuttle tank where fish could make a choice between areas with different risk of being infected. Intensity of infection was assessed and compared with the number of fish recorded in the compartment with parasites and level of fish motility.ResultsBoth solitary fish and fish in groups avoided parasitized areas, but fish in groups avoided it more strongly and thus acquired significantly fewer parasites than solitary fish. Prevalence of infection among grouped and solitary fish was 66 and 92 %, respectively, with the mean abundance two times higher in the solitary fish. Between-individual variation in the number of parasites per fish was higher in the “groups” treatment (across all individuals) than in the “solitary” treatment. Avoidance behaviour was more efficient when fish were allowed to explore the experimental arena prior to parasite exposure. High motility of fish was shown to increase the acquisition of D. pseudospathaceum.ConclusionFish in groups better avoided parasitized habitat, and acquired significantly fewer parasites than solitary fish. We suggest that fish in groups benefit from information about parasites gained from other members of a group. Grouping behaviour may be an efficient mechanism of parasite avoidance, together with individual behaviour and immune responses of fishes. Avoidance of habitats with a high parasite risk can be an important factor contributing to the evolution and maintenance of grouping behaviour in fish.