A proximate cue for oviposition site choice in the bitterling (Rhodeus sericeus)

1. We investigated two possible proximate cues used for oviposition site choice by females of the bitterling ( Rhodeus sericeus ), a freshwater fish that spawns on the gills of live unionid mussels. The two cues were the flow velocity and/or oxygen content of water emerging from the exhalant siphon of a mussel.
2. Field observations showed that female bitterling always inspected the exhalant siphons of mussels before they spawned in them. Siphon inspection was not always a prelude to spawning and it may serve as a means of assessing mussel quality. Female skimming behaviour, swimming over a mussel without spawning, may also be used to assess mussel quality, although the mechanism for this is unclear.
3. Measurements of the flow velocity of water emerging from the exhalant siphons of four mussel species ( Anodonta anatina, A. cygnea, Unio pictorum and U. tumidus ) showed a significant difference among species, with U. tumidus having the highest mean flow velocity and U. pictorum the lowest.
4. Measurements of the change in oxygen concentration of water entering a mussel inhalant siphon and leaving its exhalant siphon in field and laboratory studies showed a significant difference among the four mussel species, with A. cygnea exhibiting a significantly higher change in oxygen concentration than the other species.
5. The presence of bitterling embryos in the gills of a mussel significantly increased its oxygen consumption whereas larval glochidia had no significant effect. We discuss oxygen availability as a possible proximate cue for oviposition site choice in bitterling.     

Acquired resistance of bluegill sunfish Lepomis macrochirus to glochidia larvae of the freshwater mussel Utterbackia imbecillis (Bivalvia: Unionidae) after multiple infections

The effects of multiple infections on the host-parasite relationship between bluegill sunfish (Lepomis macrochirus) and parasitic glochidial larvae of the freshwater mussel Utterbackia imbecillis were examined. Na?ve, young-of-the-year bluegills were infected with glochidia and placed in individual observation chambers. Each day, water was drained from each chamber and the numbers of dead glochidia, live glochidia, partially metamorphosed glochidia, and fully metamorphosed juvenile mussels were counted. The same fishes were infected a total of 4 times. After 2 infections, the fish began to exhibit evidence of acquired resistance to glochidia. During the third and fourth infections, this resistance was clearly evidenced by the marked increase in the percentage of dead and live glochidia shed during the first 5 days of the infection and by the significant decrease in the success of metamorphosis. The total number of glochidia that successfully attached to the fish decreased significantly during the fourth infection relative to the first. The number of larvae attached to the host fish was positively correlated with the size of the fish during the first infection but was negatively correlated during all subsequent infections. Variance to mean ratios indicated that larvae were aggregated among host fishes during the infections. This study has important implications in propagation and conservation efforts of this endangered group of organisms.     

Survival of glochidial larvae of the freshwater pearl mussel, Margaritifera laevis (Bivalvia: Unionoida), at different temperatures: a comparison between two populations with and without recruitment

The viability of free-living glochidia of the freshwater pearl mussel (Margaritifera laevis) was studied in the laboratory at water temperatures of 10 degrees C, 15 degrees C and 20 degrees C. To obtain glochidia, gravid female mussels were collected from the Chitose River, inhabited by adult and juvenile mussels, and from the Abira River, where only adult mussels were found. Daily survival rates of glochidia from each population at various water temperatures were significantly different, and survival time was longest at the lowest temperature in each population. Maintenance of some field mussel populations might become difficult at higher water temperatures due to the short survival time of glochidia and expected low density of host fish. Daily survival rates of glochidia were compared between the Abira population at 15 degrees C and the Chitose population at 20 degrees C, since these temperatures were close to the mean water temperature during the period of glochidial release in the respective rivers. Daily mean survival rates were significantly different between the Abira population at 15 degrees C and the Chitose population at 20 degrees C. Mean glochidial survival rate for the Chitose population changed from 85.3% to 66.2% from 9 to 13 h, whereas that for the Abira population dropped suddenly from 80.4% to 34.2% from 10 to 14 h after the initiation of experiment. Absence of juveniles in the Abira River might have been caused by the low glochidial viability. Survival times of free-living glochidia in Margaritiferidae tend to be shorter than in other families in Unionoida. A trade-off is suggested between high fertility and low glochidial survival rate in Margaritiferidae.     

Influence of cortisol on the attachment and metamorphosis of larval Utterbackia imbecillis on bluegill sunfish (Lepomis macrochirus)

The larvae of unionid freshwater mussels (i.e., glochidia) undergo a parasitic stage requiring their attachment to the external epithelia of fish hosts, where they metamorphose into free-living juveniles. We describe the physiological effects in bluegill sunfish (Lepomis macrochirus) of infection with glochidia from the paper pondshell (Utterbackia imbecillis). Glochidia accumulation on bluegill increased dramatically at concentrations of 2000 glochidia liter(-1) and above, reaching a maximum attachment density of about 30 glochidia g(-1) fish at 4000 glochidia liter(-1). Plasma cortisol was the most sensitive indicator of biological effect to glochidial exposure, increasing significantly in hosts exposed to 2000 glochidia liter(-1) or greater. Glochidia were 31% more likely to undergo successful juvenile metamorphosis when attached to bluegill with elevated plasma cortisol, largely due to the enhanced survivorship of these larvae during the first 48 h after infection. We tested the hypothesis that glochidial attachment and juvenile metamorphosis were stimulated directly by plasma cortisol in fish hosts. Bluegill were given an intraperitoneal injection of cortisol, then infected with 1000 glochidia liter(-1) at 48 h after hormone supplementation. Cortisol-injected fish had a 42% increase in the number of attached glochidia g(-1) fish and a 28% increase in larval metamorphosis compared to sham-injected and control fish. We provide evidence that cortisol enhances glochidial metamorphosis on hosts by improving the retention of attached glochidia. This study gives insights into the influence of host physiology on glochidial attachment and juvenile mussel transformation.     

Do individual Activity Patterns of Brown Trout (Salmo trutta) alter the Exposure to Parasitic Freshwater Pearl Mussel (Margaritifera margaritifera) Larvae?

The hypothesis that interindividual differences in the activity of brown trout alter the exposure to parasitic freshwater pearl mussel glochidia was tested in a Swedish stream. Wild yearling brown trout (N = 103) were caught, individually tagged for identification and scored for open‐field activity during standardized laboratory tests in June. Fifty gravid freshwater pearl mussels were relocated to the stream, where after the trout were released back into the stream. The fish were recaptured in October (N = 35), checked for glochidia encystment (infested individuals: n = 6) and re‐scored for open‐field activity traits. Swimming velocity during the test was higher in fish infected with glochidia, suggesting that high activity could increase their exposure to glochidia. Potentially, as metabolism rate and ventilation rate typically increase with activity, elevated activity may lead to an increased likelihood of glochidia passing over the gills. This novel finding suggests that glochidia infestation is non‐random and that the behaviour of the host fish can influence the likelihood of glochidia infestation.     

Parasite‐induced alterations of host behaviour in a riverine fish: the effects of glochidia on host dispersal

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Resident spawning of Japanese smelt,Hypomesus nipponensis,along gravel shorelines in Lake Nojiri,Central Japan

In Lake Nojiri, the Japanese smelt, Hypomesus nipponensis McAllister, 1963, population has been thought to be maintained by artificial spawning, and stocking efforts as natural spawning in in-flowing streams is unlikely due to the lack of inflowing stream habitat. In this study, novel resident Japanese smelt spawning was observed along the lakeshore in shallow areas with clean gravel and flow. Spawning occurred at night during early March to mid-April and eggs, confirmed on gravel substrate, progressed to the eyed state in 5–6 weeks. Although natural spawning was observed, the ratio of eggs that progressed to the eyed stage was minimal, possibly due to thick algae cover in areas with low flow velocity. Therefore, resident Japanese smelt spawning along the lake shoreline is physically possible and potentially contributes to the overall population in Lake Nojiri, but in-lake spawning alone presumably is not a biologically viable method for population sustainability due to low survival rates and egg density at present. Environmental improvements such as construction of fish ladders to suitable in-flowing spawning habitat, introduction of clean gravel to shorelines, and water level management adjusted to the smelt spawning run would contribute to higher recruitment by natural smelt reproduction, and consequently enhance the smelt production in Lake Nojiri.

     

Virus-associated spawning papillomatosis in smelt, Osmerus eperlanus L., in the River Thames

Smelt were collected from the R. Thames, at Lots Road Power Station, Fulham between October 1989 to June 1990. Smooth, white, ovoid tumours were observed on the fins of smelt during their anadromous spring migration and recognized as a form of spawning papillomatosis. Papillomatous fish constituted about 21% of the population in March and about 19% in April but were not present in any other samples. Up to 13 tumours were found on a single fish. Papilloma prevalence and intensity were both raised in larger fish. Ultrastructural examination revealed that a high proportion of tumour cells contained virus particles with herpesvirus group characteristics. Epithelial cells of fins without tumours revealed no such virus particles. Evidence of the herpesvirus as a likely aetiological agent of spawning papillomatosis is discussed.     

Toxicity of mosquitocides on freshwater mussel larvae

The effect of four insecticides also used for mosquito-killing (Fyfanon ULV, K-Othrin ULV, Unitox 7 ULV and Unitox 20 ULV) was studied on the larvae (glochidia) of two freshwater mussel species, Anodonta cygnea and Anodonta anatina. In addition to the determination of the 24-96 h LC50 values of the insecticides, studies were also performed on their effect exerted on a physiological process, the tryptamine-(TA)-induced activation of the larvae in normal and 24 h test. On the basis of the LC50 values, Fyfanon and Unitox 20 were found to be the most toxic to the studied mussel larvae, however, the TA-induced adductor muscle activity, as physiological process, was also inhibited by Unitox 7 in relatively low concentrations. This effect may reduce the chance of further development of the larvae, by exerting harmful effect on the number of freshwater mussels.     

Persistence of host response against glochidia larvae in Micropterus salmoides

Host fish acquire resistance to the parasitic larvae (glochidia) of freshwater mussels (Unionidae). Glochidia metamorphose into juvenile mussels while encysted on host fish. We investigated the duration of acquired resistance of largemouth bass, Micropterus salmoides (Lacepède, 1802) to glochidia of the broken rays mussel, Lampsilis reeveiana (Call, 1887). Fish received three successive priming infections with glochidia to induce an immune response. Primed fish were held at 22-23 degrees C and were challenged (re-infected) at intervals after priming. Metamorphosis success was quantified as the percent of attached glochidia that metamorphosed to the juvenile stage and were recovered alive. Metamorphosis success at 3, 7, and 12 months after priming was significantly lower on primed fish (26%, 40%, and 68% respectively) than on control fish (85%, 93%, and 92% respectively). A second group of largemouth bass was similarly primed and blood was extracted. Immunoblotting was used to detect host serum antibodies to L. reeveiana glochidia proteins. Serum antibodies were evident in primed fish, but not in naive control fish. Acquired resistance of host fish potentially affects natural reproduction and artificial propagation of unionids, many of which are of conservation concern.     

The pathology and seawater performance of farmed Atlantic salmon infected with glochidia of Margaritifera margaritifera

The pathology of glochidial infection of the freshwater mussel Margaritifera margaritifera was examined in farmed Atlantic salmon Salmo salar in fresh water and for 150 days after transfer of salmon to sea water. Prevalence of infection in fresh water was 95%, mean abundance 134 per fish and mean infection intensity 140. Prevalence in sea water was 80–94% in the first 7 weeks after transfer but glochidia were absent, apart from remains, after 50 days in sea water. Glochidia on salmon in fresh water were associated with localized hyperplasia and fusion of secondary gill lamellae. Focally extensive epithelial hyperplasia and fusion of secondary lamellae were present 4–10 days after transfer to sea water. Twenty-three days after transfer, small nodules with a more discrete appearance were present suggesting partial resolution of tissue response; hyperplastic responses associated with glochidia were much reduced after 50 days. Plasma chloride in infected fish 10 days after transfer was 153 mmol. 1⁻¹, significantly higher than fish without infection, suggesting poorer adaptation to sea water. No mortalities due to glochidial infection in sea water were recorded and there was no significant difference in mean weight between infected and control fish.     

THE BITTERLING–MUSSEL COEVOLUTIONARY RELATIONSHIP IN AREAS OF RECENT AND ANCIENT SYMPATRY

Host–parasite relationships are often characterized by the rapid evolution of parasite adaptations to exploit their host, and counteradaptations in the host to avoid the costs imposed by parasitism. Hence, the current coevolutionary state between a parasite and its hosts is predicted to vary according to the history of sympatry and local abundance of interacting species. We compared a unique reciprocal coevolutionary relationship of a fish, the European bitterling (Rhodeus amarus) and freshwater mussels (Unionidae) between areas of recent (Central Europe) and ancient (Turkey) sympatry. Bitterling parasitize freshwater mussels by laying their eggs in the gills of mussel and, in turn, mussel larvae (glochidia) parasitize the fish. We found that all bitterling from both regions avoided one mussel species. Preferences among other mussel species tended to be related to local mussel abundance rather than duration of sympatry. Individual fish were not consistent in their oviposition choices, precluding the evolution of host‐specific lineages. Mussels were demonstrated to have evolved strong defenses to bitterling parasitism in the area of ancient sympatry, but have no such defenses in the large areas of Europe where bitterling are currently invasive. Bitterling avoided glochidia infection irrespective of the duration of sympatry.     

Analyses of growth processes of pond smelt, Hypomesus nipponensis, in Lake Ogawara, Japan, through the use of daily otolith increments

It has been reported that pond smelt, Hypomesus nipponensis, in Lake Ogawara, Japan, appear in small and large size groups during spawning despite being an annual fish. It is hypothesized that pond smelt have a bimodal life history, anadromous for large size groups and resident in the lake for small size groups. We calculated the body length and growth rate of the small and large size groups through the use of daily otolith increments and compared growth parameters between groups. In addition, the growth processes of resident fish in Lake Ogawara and anadromous fish in the adjacent sea were examined and compared with small and large size groups, respectively. We found that the two size groups diverged after 40–50 days from hatching with significant size groups present after 50 days. Through otolith increment analysis the growth processes of the small and large size groups were correlated with resident and anadromous forms respectively. These results revealed the utility of otolith increment analyses to clarify population structure of this species.     

Cross-resistance of largemouth bass to glochidia of unionid mussels

We tested whether host fish that acquired resistance to glochidia of one mussel species were cross-resistant to glochidia of other species. Largemouth bass (Micropterus salmoides) were primed with 4-5 successive infections of glochidia of Lampsilis reeveiana. The percentage of attached glochidia that survived and transformed to the juvenile stage (transformation success) was compared between primed fish and na?ve controls. Transformation success of L. reeveiana, Lampsilis abrupta, Villosa iris, and Utterbackia imbecillis was significantly lower on primed fish (37.8%, 43.5%, 67.0%, and 13.2%, respectively) than on control fish (89.0%, 89.7%, 90.0%, and 22.2% respectively). Immunoblotting was used to analyze the binding of serum antibodies from primed fish with glochidia proteins. Antibodies bound to glochidia proteins of similar molecular weight from L. reeveiana and L. abrupta. Bound proteins of V. iris differed in molecular weight from those of the Lampsilis species. There was no binding to specific glochidia proteins of U. imbecillis or Strophitus undulatus. Our results indicate that host-acquired resistance can extend across mussel genera and subfamilies and might involve both specific and nonspecific mechanisms. Understanding the specificity of acquired resistance of hosts to glochidia could enhance understanding of the evolutionary and ecological relationships between mussels and their host fishes.     

Physiological and ecological hosts of Popenaias popeii (Bivalvia: Unionidae): laboratory studies identify more hosts than field studies

1. Freshwater mussels are critically endangered in North America, making it important to understand their environmental requirements at all life stages. As glochidia (larvae), they attach to fish hosts where they undergo substantial mortality, making this transition important in their life cycle. Larval host fish requirements have typically been described using data from laboratory infestations to determine suitable hosts. 2. Laboratory infestations circumvent many natural barriers that prevent infestation of physiologically compatible fishes by mussel larvae. While such methods are invaluable for identifying ‘physiological hosts,’ they cannot fully describe realised ‘ecological hosts’ in the field. 3. We studied Popenaias popeii in the Black River in New Mexico, because it is of conservation concern and it is the only mussel species present, facilitating identification of glochidial infestation. To explore the difference between physiological hosts and ecological hosts, we conducted a 3‐year field study of fishes infested by P. popeii glochidia. 4. Substantially fewer fish species were infested by P. popeii in the wild (10 of 20 observed) than had been identified as physiological hosts in laboratory trials (24 of 31). We combined data on fish abundance, proportion of fish hosts infested (prevalence) and the number of glochidia per fish (intensity) and identified three fish species that probably contributed substantially more to mussel recruitment by carrying more glochidia than other host species. 5. Similarities in behaviour among these fishes allowed us to hypothesise routes of infestation, such as benthos‐feeding by catostomids, that allow glochidia to infest these hosts at higher rates than other suitable hosts. Overall, this approach provides a method of quantifying the relative importance of different species of host fish in the mussel lifecycle.     

Habitat traits, population structure and host specificity of the freshwater pearl mussel Margaritifera margaritifera in the Waldaist River (Upper Austria)

In European streams and rivers, the freshwater pearl mussel (Margaritifera margaritifera L., 1758) faces extinction. This is also true for the Waldaist River, with 20,000 specimens recorded in the early 1990s then Austria’s most important pearl mussel river. Nowadays, there is only a single 320 m stretch with noteworthy mussel densities. During an in-depth survey of this river stretch in 2010, we detected a total of 2,774 specimens. Mussel microhabitats were confined to patches of sand and fine gravel (psammal and akal) at run sections of the river, stabilized by large boulders. Pearl mussels avoided large accumulations of fine sediments. Typically situated at undercut slopes, preferred microhabitats were 0.25–0.50 m deep at baseflow with current velocities (at 40% depth) of 0.2–0.6 m s^?1. A comparison of the present stock with data from 1997 revealed a rapid decline in mussel density down to 27%. We also noticed strongly reduced growth and a high mortality of medium age classes. Juvenile mussels were completely lacking. With respect to host specificity in terms of glochidia survival, the brook char Salvelinus fontinalis (Mitchill, 1814), a suitable host in North America, shed glochidia within eight days. In the brown trout Salmo trutta L., 1758, two strains were investigated. Glochidia survival, growth and prevalence were significantly higher in the Danish than in the Austrian hatchery strain.     

Encapsulation of attached ectoparasitic glochidia larvae of freshwater mussels by epithelial tissue on fins of naive and resistant host fish

To metamorphose into juveniles and subsequently mature into adults, the glochidia larvae of freshwater mussels in the order Unionoida must temporarily parasitize the gills, fins, or other external structures of fish. Once attached to the fish, the glochidium is encapsulated by host fish epithelial tissue. The migration of epithelial cells of the bluegill sunfish Lepomis macrochirus over glochidia of Utterbackia imbecillis was examined by time-lapse video microscopy, and the morphology was examined by scanning electron microscopy. Initially, the leading edge epithelial cells migrating over the larvae became rounded and the cells moved as a sheet until the attached glochidium was completely covered. Cyst formation on host fish that had been repeatedly exposed to mussel larvae was significantly delayed and morphologically irregular compared to that on na?ve fish. Cyst formation on other species of fish that are less successful as hosts was examined. In general, it took longer for glochidia to become encapsulated on these less suitable potential hosts. The delay and irregularities in cyst formation on resistant fish and nonhost fish species may result in increased mortality and reduced success of metamorphosis of glochidia.     

Timing of spawning and glochidial release in Scottish freshwater pearl mussel (Margaritifera margaritifera) populations

1. The timing of reproduction was investigated in six Scottish freshwater pearl mussel populations from 1993 to 2002. Gravid females were examined and the release of mussel larvae (glochidia) was monitored. 2. Annual spawning (oviposition) and spat (glochidial release) events occurred during June to July and June to September, respectively. 3. Between‐river differences in timing seem to be related to water temperature. Mussels in the warmest rivers tend to spawn and spat first, and vice‐versa. 4. Thermal variations also seem to influence the timing of reproduction within rivers, which can be delayed by several weeks during cold years. At least 3000°‐days occur between annual episodes of glochidial release. 5. The timing of spawning is determined gradually, probably by a thermal summation effect. 6. The release stage occurs as a sudden, synchronised event, with most of the glochidia spat over 1–2 days, indicating that it is triggered by an environmental cue. Sudden changes in water temperature and/or river level often result in spats, and the underlying mechanism may be respiratory.     

Morphological and morphometrical description of the glochidia of Westralunio carteri Iredale, 1934 (Bivalvia: Unionoida: Hyriidae)

Most freshwater mussel (Bivalvia: Unionoida) larvae (glochidia in Margaritiferidae, Hyriidae and Unionidae) are fish parasites. Knowledge of the larval morphology and the mechanism of release in freshwater mussels is useful in species systematics and ecology. Westralunio carteri is the only unionoid from south-western Australia. Little information is available on its biology and its glochidia have never been described. The aim of this study was to describe the glochidia of W. carteri and method of their release. Glochidia within vitelline membranes were embedded in mucus which extruded from exhalent siphons of females during spring/summer; they then hatched from vitelline membranes but remained tethered by a larval thread and began characteristically “winking”. Shells (n=120) were subtriangular, 308 μm long (±0.83 SE), 251 μm high (±0.73 SE) and had a hinge length of 212 μm (±0.78 SE). Larval teeth were singular with interlocking cusps and convex or concave basal protuberances on opposing valves.     

三角帆蚌钩介幼虫在两种寄主鱼上的寄生效果及其对寄主鱼血浆生化指标的影响

利用黄颡鱼(Pelteobagrus fulvldraco)、尼罗罗非鱼(Tilapia nilotica)对三角帆蚌(Hyriopsis cumingii)钩介幼虫进行了寄生实验,分析了幼虫在寄主鱼上的寄生周期和脱落数量与寄主种类、寄主鱼单尾体重的相关关系.以未寄生幼虫的罗非鱼为空白对照,测定分析了罗非鱼在幼虫寄生前、脱落后主要血浆生化指标的变化.实验结果显示:黄颡鱼、罗非鱼平均每千克体重脱落稚贝的数量分别为(26 435±6 430)只、(26 327±3 091)只,两种寄主鱼间不存在显著差异;脱落的稚贝数量随着寄主鱼体重的增加而增加,并呈显著线性相关.在水温(24±1)℃条件下,黄颡鱼组、罗非鱼组的稚贝脱落高峰分别出现在寄生后的第8 d、第7 d,罗非鱼组表现为明显的"早脱落"现象.罗非鱼在寄生前后血浆总蛋白(TP)、白蛋白(Alb)、总糖(Glu)、总胆固醇(Tc)和高密度脂蛋白(HDL)含量均未出现显著变化,而血浆甘油三酯(TG)、低密度脂蛋白(LDL)含量显著下降(P=0.001,0.020).未寄生组罗非鱼的7项指标均未出现明显变化.实验结果将为三角帆蚌钩介幼虫变态发育相关营养因子的探讨提供理论参考.