The histopathological changes in swimbladders of European eels naturally and experimentally infected with Anguillicola crassus were studied using transmission and scanning electron rnicroscopy. During the course of probably several infections swimbladders undergo characteristic changes. In addition to the thickening of the entire swimbladder wall, and to the folded internal surface of this organ, inflammation, migration of white blood cells, fibrosis and changes in the epithelial cells are frequently seen. Epithelia1 cells tend to proliferate heavily and form hyperplastic tissues; these processes are accompanied by changes in the internal structure of the cells. The normally cubic cells become spherical or columnar and form folds facing the lumen of the swimbladder. As a consequence, most of these cells lose contact with the blood vessels and show no strict polarity. In heavily affected swimbladders the basal labyrinth of the epithelial cells is reduced, i.e. becomes shorter and less densely packed. The lamina propria shows severe fibrosis with infiltration of white blood cells. Larvae of A. crassus, inhabiting the wall of the swimbladder, were found to be surrounded by cell debris, but this local necrosis does not affect the entire swimbladder in its overall structure. These histological findings can partly explain changes in the gas composition in eels infected with A. crassus.

The effect of Anguillicola crassus (Nematoda) on gas composition in the swimbladder of eels (Anguilla anguilla) was studied using mass spectrometry in feral eels from two German rivers and in experimentally infected eels. In both naturally and experimentally infected eels significant correlations were observed between the proportion of oxygen in the swimbladder and level of infection with Anguillicola crassus. In swimbladders of naturally infected feral eels the contribution of oxygen to swimbladder gas was reduced by 36-62.9% and in experimentally infected eels it was reduced by 11.4-57% compared to uninfected controls. The proportion of CO2 appeared to be lower in infected swimbladders compared to uninfected ones. However, this change was not significant. The findings are discussed in relation to an altered structure of the swimbladder wall due to the parasite.

ABSTRACT. The effect of low water temperatures on the development and viability of larval and adult Ang~lillicola crassus (Nematoda) in the final host Anguilla ang~lilla was studied. European eels were experimentally infected with A. crassirs and then maintained for 4 mo at 4,9, 10 and 19°C. Larval development showed a temperature-dependent pattern and was significantly retarded at low temperatures. Third-stage larvae survived a 4 mo period at 4'C without belng affected, although they were not able to invade the swimbladder wall at this temperature. In contrast, adult worms were severely harmed during a 4 mo period at 4'C, as reflected by increased mortality and decreased growth and reproductivity compared to the worms maintained for Lhe same period at 18°C. Starvation of the eels for 4 mo at 19°C dld not affect the development and growth of the nematode The experimentally obtained results support the hypothesis that the spread of A . crasslrs in boreal regions, e.g. Northern Europe, I S restricted by the natural ambient temperature regimes.

A total of 121 European eels (Anguilla anguilla) from 2 sampling sites on the River Rhine were investigated in respect of their parasite communities. Special attention was given to the swim bladders, intestines, gills and fins of the fish. Twelve different parasite species were found to live in and on the eels. Data from each sampling site were kept separate. Parasites found in descending order of prevalence were: Anguillicola crassus, Trypanosoma granulosum, Myxobolus sp., Paratenuisentis ambiguus, Pseudodactylogyrus sp., Bothriocephalus claviceps, Myxidium giardi, Pomphorhynchus laevis, Trichodina sp., Raphidascaris acus, Acanthocephalus lucii and Acanthocephalus anguillae. Significantly different prevalences were reported for L3 larvae of A. crassus, adult P. ambiguus, B. claviceps and Myxobolus sp. at the 2 sampling sites. The highest number of parasite species was recorded from the intestine, which contained up to 6 different helminths. The coexistence of the acanthocephalans P. laevis and P. ambiguus, which showed clear patterns of distribution within the intestine of the respective hosts, was reported for the first time. Up to 3 different helminth species were found in the intestine of individual fish. Among those, acanthocephalans were the most prevalent worms with the eel-specific parasite P. ambiguus as the dominant species not only of the intestinal but also of the total component communities. Both infra and component communities exhibited low diversity and were dominated by this single species. The evenness reached only approximately 50% or less and it remained unclear why the helminth communities of the eels from the River Rhine with its huge catchment area exhibit such a low parasite diversity and high dominance.

The ability of the nematode Anguillicola crassus to infect eel larvae (glass-eel stage) was tested. The results show that glass-eels fed on infected copepods, the natural intermediate host of the nematode, can be infected. Light microscopical examination of the infected developing swimbladder tissue revealed that the infection results in a significant thickening of the connective tissue. The basolateral labyrinth of gas gland cells is very much reduced in infected swimbladders, and the distance of gas gland cells to blood capillaries is enlarged. Critical swimming speed, defined as the speed where the larvae were no longer able to swim against the current, was similar in infected and uninfected animals. At intermediate speeds (about 60-80% of critical swimming speed) infected eels showed a slightly higher swimming activity than control animals. Resting oxygen consumption, measured as an index of metabolic activity, within the first 2 months of infection was higher in control animals, which may be due to a reduced rate of activity in infected glass-eels. By 4-5 months after the infection, however, it was significantly higher in infected animals. This may indicate that at this stage a higher activity of the animals is required to compensate for the increase in body density, but swimming performance of infected and non-infected glass-eels was not significantly different. Oxygen consumption during swimming activity, measured in a swim tunnel at 50% of maximal swimming speed, also was not affected. The results thus show that even glass-eels can be infected with A. crassus, and this probably contributes to the rapid spread of the nematode in Europe. While aerobic metabolism during swimming activity is not affected at this stage of infection, the swimbladder tissue shows severe histological changes, which most likely will impair swimbladder function.