The Hydrobia ulvae-Maritrema subdolum association: influence of temperature,salinity, light,water-pressure and secondary host exudates on cercarial emergence and longevity

The effects of environmental factors and exudates from the amphipod Corophium volutator on the emergence of Maritrema subdolum cercariae (Digenea: Microphallidae) from the snail Hydrobia ulvae were investigated in the laboratory. Increasing the temperature (15 to 25 degrees C) caused an overall 11-fold increase in emergence rate under varying salinities (24 to 36 per thousand). The effect of salinity depended on the experimental temperature. Emergence increased with increasing salinity at higher temperatures, but decreased with increasing salinity at 15 degrees C.Whereas the different levels of salinity had no effect, increasing the temperature significantly reduced the life span of cercariae. In comparison with complete darkness, light caused a two-fold increase in emergence, whereas an increment of the water pressure from 1.0 to 1.3 ATM (corresponding to 0 and 3 m of depth) left the shedding rate unaffected. Unidentified exudates from the second intermediate host, C. volutator, significantly depressed the cercarial emergence rate. The main transmission window of M. subdolum seems to occur during low water in tidal pools where light levels are high and solar radiation rapidly elevates the water temperature, as well as salinity through evaporation. The consequence of such a transmission strategy is discussed in relation to the impact of M. subdolum on the population dynamics of the second intermediate host.     

Effects of sediment type and intra- and interspecific competition on growth rate of the marine snails Hydrobia ulvae and Hydrobia ventrosa

The combined effects of sediment grain size and intra- and interspecific competition on growth rate of Hydrobia ulvae and H. ventrosa were investigated in a 2-month laboratory experiment. Individuals of H. ulvae grew faster than H. ventrosa at all competition intensities. Growth rate did not differ between fine- and coarse-grained sediments in any of the species. This suggests that other factors than the sediment quality cause the often observed size differences between Hydrobia snails from these sediment types in nature. Competition from 1.6 to 1.9 times larger competitors affected growth of the smaller target snails. Competition between the species was asymmetrical and, in both species, per capita effect of competition was stronger from H. ulvae than from H. ventrosa, regardless of sediment type. From growth, Lotka-Volterra competition coefficients were estimated to 0.2 from H. ventrosa on H. ulvae and 1.8 from H. ulvae on H. ventrosa. Since competition between the species seems to be independent of sediment grain size, the shift in numerical dominance in nature, from H. ulvae on coarse-grained sediment to H. ventrosa on fine-grained, could not be explained with different competitive ability in different environments.     

Measurement of ingestion rate of Hydrobia ulvae (Pennant) on intertidal epipelic microalgae: the effect of mud snail density

The individual mean ingestion rate of Hydrobia ulvae was measured experimentally in controlled microcosms, in the dark to avoid primary production during measurement and at constant temperature. The experimental design was based on the addition of prelabelled epipelic microalgae to microcosms in a constant proportion with unlabelled diatoms, and in such a way that algal food availability was not a limiting factor within the range of tested densities (0.3 to 4.1 snails cm(-2)). Results show that the individual mean ingestion rate decreased significantly from 26.6+/-1.1 ng Chl a snail(-1) h(-1) to 22.4+/-1.0 ng Chl a snail(-1) h(-1) between 0.7 and 3 snails cm(-2). We hypothesize that this sharp decrease (the threshold density was between 1.4 and 2.5 snails cm(-2)) may account for a density-dependent effect. We have tested this hypothesis by using a simple random walk model including basic behavioural processes such as a break in feeding activity when two individuals contact each other. The model represents quantitatively well the threshold effect, suggesting that behavioural processes have to be taken into account for estimating a global feeding activity of H. ulvae populations.     

Physiological and behavioral responses of the mud snails Hydrobia glyca and Hydrobia ulvae to extreme water temperatures and salinities: implications for their spatial distribution within a system of temperate lagoons

Physiological responses (oxygen consumption) and behavioral responses (feeding and activity) of the mud snails Hydrobia ulvae and Hydrobia glyca at different salinities (20 per thousand-80 per thousand) and temperatures (20 degrees and 30 degrees C) were studied. After 24 h under experimental conditions, both Hydrobia species already showed maximal activities (>90%) for a wide salinity range (30 per thousand-70 per thousand), with significant differences in activity between species only outside the usual salinity range of the studied lagoon. In contrast, egestion rates of H. glyca were significantly higher at the lowest salinities tested (30 per thousand and 40 per thousand) irrespective of water temperature, whereas egestion rates of H. ulvae were always significantly higher (57% on average) at 20 degrees C than at 30 degrees C and at the usual salinities found in the field (40 per thousand and 50 per thousand). Both species showed an oxyregulatory response to dissolved oxygen concentrations ranging from saturation to 1.5 mg O(2) L(-1), although specific oxygen consumption rates were significantly higher at 30 degrees C than at 20 degrees C (Q(10)=1.47+/-0.08 for H. ulvae and Q(10)=12.1+/-0.06 for H. glyca) and at the lowest salinities (30 per thousand-50 per thousand for H. ulvae and 30 per thousand-40 per thousand for H. glyca). On average, specific rates were higher for the smaller-sized H. glyca (1.64+/-0.03 microg O(2) mg(-1) ash-free dry weight [AFDW]) than for H. ulvae (1.35+/-0.03 microg O(2) mg(-1) AFDW). Despite the overlapping of their tolerances to high temperatures and salinities, the observed interspecies differences could play a certain role in the distribution of H. ulvae and H. glyca in the studied habitat. In particular, the decreasing feeding activity but increasing respiration of H. ulvae at 30 degrees C for salinities that usually occur in the studied lagoon could represent disadvantages to H. glyca during the warm period.     

The influence of infections with trematodes Bunocotyle progenetica (Hemiuridae) and Cryptocotyle cancavum (Heterophyidae) onto mortality of littoral mollusks Hydrobia ulvae (Gastropoda: Prosobranchia) after freezing

The mortality of prosobranch snails Hydrobia ulvae uninfected and infected with the parthenites of two species of Trematodes (Bunocotyle progenetica and Cryptocotyle cancavum) after freezing in seawater of different salinity with following thawing has been studied. Reduced salinity exceeds mortality of snails under freezing significant (p < 0.01). The mortality of mollusks increases significant (p < 0.01) with the increase of the exposure time. The mortality of females exceeds the mortality of male significant (p < 0.01) within the group of 2-years individuals. The negative effect of trematode invasion to the vitality of mollusks after freezing was not showed. Trematode infestation registered in March 2004 and in September 2003 is not significant differ (Levakin, 2004). Thus any important differential mortality of individuals H. ulvae infected with parthenites of this Trematodes is not present during a winter time. The conclusion is conformed by the results of our experiments and well accords to features of realization of life cycles of these Trematodes under conditions of the White Sea intertidial zone, where infection is possible only within the worm season.     

Component community of larval trematodes in the mudsnail Hydrobia ventrosa: temporal variations in prevalence in relation to host life history

Temporal variations in the prevalence of larval trematodes in the short-lived prosobranch mudsnail Hydrobia ventrosa (Montagu) were investigated in relation to host life history and season for 4 successive years in temperate windflats of the southern Baltic Sea. The component community of trematode larvae in H. ventrosa comprises at least 10 species; families (and species) represented include Notocotylidae (1), Echinostomatidae (1 or 2), Heterophyidae (2), Monorchidae (1), Microphallidae (3 or 4), Psilostomatidae (1), and Hemiuridae (1). The notocotylid Paramonostomum alveatum was the most prevalent species, followed by the microphallids Maritrema subdolum and Microphallus sp. Trematode prevalence in H. ventrosa fluctuated seasonally. Prevalence usually peaked in summer between July and September-October and decreased in late winter-early spring. This seasonal change is chiefly explained by the life history patterns of the semelparous snail host. Hydrobia ventrosa has a maximum life span of about 2 yr and reproduces between June and November of its second calendar year. The first trematode infections appeared annually in May when the most abundant cohort of H. ventrosa, the second-calendar-year snails, mature. The prevalence continued to increase until August-September, throughout the reproductive period of the second-calendar-year snails, Prevalence decreased during winter, when most of the second-calendar-year snails died after reproduction. On the basis of longterm laboratory experiments, it has been shown that the late autumn-winter mortality was not the result of trematode infections. Seasonal patterns of prevalence were similar among the trematode species except for the monorchid Asymphylodora demeli, the only one using fish definitive hosts. Species-specific differences in the seasonal occurrence of prepatent infections and the predominance of certain larval stages in winter are interpreted as different strategies of the trematode species to survive the harsh winter conditions, or to survive the death of the first intermediate host in autumn-winter, or both.     

Vertical distribution of trematode cercariae Cryptocotyle concavum (Heterophyidae) and Maritrema subdolum (Microphallidae) in the water column

The effect of gravity and light on vertical distribution of Cryptocotyle concavum (Heterophyidae) and Maritrema subdolum (Microphallidae) cercariae in the water column was experimentally studied. Maximal larvae concentration was found in zones where the highest probability of contact with potential host is considered.     

Patterns and processes of larval emergence in an estuarine parasite system

Trematode parasites in intertidal estuaries experience constantly varying conditions, with the presence or absence of water potentially limiting larval transport between hosts. Given the short life spans (< or =24 h) of cercariae, emergence timing should be optimized to enhance the probability of successful transmission. In the present study, field measurements and laboratory experiments identified processes that regulate the emergence of cercariae from their first intermediate snail hosts in an intertidal marsh. Larvae emerged over species-specific temperature ranges, exclusively during daylight hours, and only when snails were submerged. The three factors operate over different temporal scales: temperature monthly, light diurnally (24-h period), and water depth tidally (12-h period). Each stimulus creates a necessary condition for the next, forming a hierarchy of environmental cues. Emergence as the tide floods would favor transport within the estuary, and light may trigger direct (downward or upward) swimming toward host habitats. Abbreviated dispersal would retain asexually reproduced cercariae within the marsh, and local mixing would diversify the gene pool of larvae encysting on subsequent hosts. In contrast to the timing of cercarial release, emergence duration was under endogenous control. Duration of emergence decreased from sunrise to sunset, perhaps in response to the diminishing lighted interval as the day progresses. Circadian rhythms that control cercarial emergence of freshwater species (including schistosomes) are often set by the activity patterns of subsequent hosts. In this estuary, however, the synchronizing agent is the tides. Together, exogenous and endogenous factors control emergence of trematode cercariae, mitigating the vagaries of an intertidal environment.     

The influence of infections with trematodes Bunocotyle progenetica (Hemiuridae) and Cryptocotyle cancavum (Heterophiidae) onto mortality of littoral molluscs Hydrobia ulvae (Gastropoda: Prosobranchia) in condition of extremely high temperature

The effect of invasion of two trematode species, Bunocotyle progenetica (Hemiuridae: Bunocotylinae) and Cryptocotyle cancavum (Heterophiidae) on the host vitality is estimated. The mortality of infected and uninfected individuals of Hydrobia ulvae after different exposure time (3, 6, 9 and 12 hours) under extremely high temperature (+42 degrees C) was compared. The total death of the host was observed after 12 hours of experiment. The death rate of molluscs infected with B. progenetica ranged from 60 to 100% and was significantly higher (p < 0.01) than mortality of individuals infected with C. cancavum and uninfected snails in each case of exposure (3, 6 and 9 hours). At the same time the mortality of individuals infected with C. cancavum was significantly higher (p < 0.05) than the mortality of uninfected snails only after 9 hours of exposure (13.3 and 3.1%, respectively). Difference in pathogen effects B. progenetica and C. cancavum is expressed in different temperature resistance of the host that may be explained by different localisation of redia, their motile activity and manner of feeding. Besides, the invasion by C. cancavum causes the sterilization and accelerated growth of hosts. Low decreasing of the resistance to overheating in snails infected with this trematode could be explained by a compensation effect of this infection. This compensation realizes due to the redistribution of some part of the reproductive energy after the sterilization. An absence of parasitic castration in the case of infection with B. progenetica is a key reason of such considerable declining of their vitality. High lethality of the host appears to be adaptive for the parasite with the monoxenous life cycle.     

Laboratory Studies of Parasite Transmission Aspects between Hydrobia spp. (Gastropoda) and Corophium volutator (Amphipoda)

Aspects of parasite transmission between Hydrobia spp. and Corophium volutator, first and second intermediate host of digenetic trematodes, were investigated under laboratory conditions. H. ventrosa is used as an intermediate host by several trematode species. Under laboratory conditions the most frequently observed emergence from H. ventrosa was of cercariae of Maritrema subdolum. The number of cercariae shed per day varied considerably. It was observed that 30 cercariae on average and up to 450 cercariae at maximum can emerge from a single H. ventrosa per day. Cercarial production continued until the death of the snails. The life-span of cercariae of the species M. subdolum decreased as the water temperature increased. It can be concluded that under natural conditions the cercariae, after emerging, have a maximum period of 1 day in which to seek out their second intermediate host C. volutator. Almost all specimens of C. volutator exposed to cercariae of the species M. subdolum died within the test period of 6 days. High average cercarial densities caused short life-spans (<50 h), while at lower densities longer survival times were possible. For cercariae of other Trematoda species, we were unable to find any equally clear evidence of a reduction in the survival rate of C. volutator within the test period. Mortality of C volutator, and other effects of infestation, as observed in our experiment, can be assumed to be a result of the penetration process of the cercariae but they also can be attributed to the mesocercariae.     

Resource partitioning on basis of hydrolytic enzymes in deposit-feeding mud snails (Hydrobiidae)

Summary Crude extracts of hydrolytic enzymes from the related mud snailsHydrobia ulvae, H. ventrosa andH. neglecta are compared by use of six different methods and 18 natural carbohydrates. Synthetic substrates for linkage specific carbohydrases and trypsin-like activity were studied in addition to lysozyme-like activity.A significant hydrolysis was only observed with the reserve carbohydrates amylose, glycogen and laminaran. Many algal carbohydrates were, however, digested to some extent.The qualitative spectra are almost identical for the three species but significant quantitative differences were found. The findings are discussed in relation to information on the chemical composition of potential food items and it is concluded that the commonly observed coexistence of the snails can in part be explained by selection for microalgae and detritus particularly meeting their enzymatic potentials.     

Effect of light intensity on Opisthorchis viverrini cercarial shedding levels from Bithynia snails--a preliminary study

Opisthorchis viverrini requires Bithynia snails as the first intermediate host and cyprinid fish as the second intermediate host. Very low natural infection rates have been reported in Bithynia snails, but very high rates have been found in cyprinid fish in the same endemic region. This study investigated the effect of light intensity, the most important stimulus, on the quantity of O. viverrini cercariae shed from naturally infected Bithynia (Digoniostoma) siamensis goniomphalos snails. Snails were evaluated for cercariae output every hour after exposure to various light intensities for a total period of 7 h. The same infected snail was tested under different intensities of light: in the dark, and at 1000, 3000 and 5000 lx. The data showed that under exposure to 1000 and 3000 lx of light, the average percentage and number of cercariae released were higher than that exposed to 5000 lx during the first 2 h of the experiment. In contrast, under higher illumination (5000 lx) a longer time (6 h) was required to stimulate the peak emergence of cercariae. Darkness was not able to induce O. viverrini cercariae emergence. Among the three intensities of light, exposure at 1000 lx induced the highest average number of released cercariae per snail and the highest percentage of cercarial emergence within the first 2 h (125, 54.86%), followed by exposure at 3000 lx (69, 25.58%) and 5000 lx (12, 7.78%). The results suggest that the light intensity of 1000 lx for 2 h would be optimal for O. viverrini cercarial shedding from naturally infected B. (D.) siamensis goniomphalos snails.     

Differential effects of microorganism-invertebrate interactions on benthic nitrogen cycling

Infaunal invertebrate activity can fundamentally alter physicochemical conditions in sediments and influence nutrient cycling. However, despite clear links between invertebrate activity and microbially mediated processes such as nitrification, the mechanisms by which bioturbating macrofauna affect microbial communities have received little attention. This study provides strong evidence for differential stimulation of microbial nitrogen transformations by three functionally contrasting species of macrofauna (Hediste diversicolor, Corophium volutator, Hydrobia ulvae). Despite increased nitrification, abundance of ammonia-oxidising bacteria (AOB) and ammonia-oxidising archaea (AOA) at the sediment-water interface did not significantly change in the presence of macrofauna. However, species-specific differences in macrofaunal activity did influence ammonia oxidiser community structure, increasing AOB abundance relative to AOA in the presence of C.?volutator or H.?ulvae, but with no change in H.?diversicolor and no-macrofauna treatments. Denaturing gradient gel electrophoresis profiles were similar between macrofaunal treatments, although one AOB band increased in relative intensity in the presence of C.?volutator, decreased in the H.?diversicolor treatment and was unchanged in the H.?ulvae treatment. These data suggest that links between bioturbating macrofauna and nutrient cycling are not expressed through changes in the abundance of ammonia oxidisers in surface sediments, but are associated with changes in the AOA?:?AOB ratio depending on the invertebrate species.     

Circadian Pattern of Cercarial Emergence in Schistosoma Mansoni(Platyhelminthes: Digenea) from Isolated Biomphalaria Glabrata

The present work aimed to compare the acrophases (peak hours) of emergence of Schistosoma mansoni cercariae among isolated individuals of the snail Biomphalaria glabrata. Laboratory stocks of melanic B. glabrata from the same biotope as the S. mansoni strain (Belo Horizonte, Minas Ger-ais) were used. Twenty-two snails individually exposed to five miracidia were tested. Chronobiological trials were performed outdoors after an acclimation period of at least a week. Three groups of snails were tested between November 1989 and April 1991. Cercarial emergence from individual isolated snails was quantified every 3 h for 3 consecutive days. In all trials, most cercariae were found to emerge during daytime (94.9%). Time series and chronograms showed recurrent peaks during the daytime. The periodogram suggested that 24 h was the period that best fitted cercarial emergence data in 90.9% of the snails. The single cosinor analysis confirmed 24-h rhythms in 95.5% of the snails. Acrophases of cercarial emergence among individual snails occurred between 14:15 and 17:02. They did not differ significantly. The population cosinor analysis indicated greater homogeneity in the 24-h rhythms of cercarial emergence than in the snail groups of each chronobiological trial. Acrophases of cercarial emergence occurred between 14:53 and 15:27 and did not differ significantly among all trials. Data from the three trials were pooled and analyzed using the population cosinor. This statistical method indicated a homogeneity in the 24-h rhythms of cercarial emergence from all snails, with acrophase occurring around 15:00. Results showed that the acrophases of cercarial emergence of S. mansoni are similar among isolated B. glubrura specimens. Data support the hypothesis of a “gate” rhythm in the dynamics of cercarial production and emergence. It is suggested that the adaptive importance of the “gate” mechanism is associated with the concentration of cercariae in the water at times when the vertebrate is present, optimizing the contact between the parasite and the host. The emergence of some cercariae at night (5.1% of the total number of emerged cercariae) suggests a flexible “gate” that could be associated with a residual light effect or with experimental procedures in the laboratory.     

Parasites on an intertidal Corophium-bed: factors determining the phenology of microphallid trematodes in the intermediate host populations of the mud-snail Hydrobia ulvae and the amphipod Corophium volutator

The phenology of microphallid trematodes within their intermediate hostpopulations has been studied on an intertidal mud flat. The parasites usethe mud snail Hydrobia ulvae and the infaunal amphipod Corophium volutatoras first and secondary intermediate host, respectively. Migratory shorebirdsact as final hosts. Our results show a general trend of decline in thedensity of infected intermediate hosts during both spring and autumn, whichcould mainly be ascribed to shorebird predation. During summer the densityof both infected snails and infected amphipods increased considerably, witha culmination in June within the snail population (1000 infectedm-² and in August within the amphipod population (40 000infected m-². This time lag in parasite occurrence could berelated to (1) the development time of larval trematodes within the snails,(2) higher ambient temperatures in late summer increasing parasitetransmission between snails and amphipods during this period, and (3) ageneral increase in the Corophium population during late summer. Fromsamples collected between 1990 and 1995 it is shown that microphallidtrematodes occasionally may give rise to mass mortality in the amphipodpopulation. The prerequisites for such an event are a high parasiteprevalence within the first intermediate host population and unusually highambient temperatures, facilitating parasite transmission to the secondaryintermediate host, C. volutator.     

Rediae of echinostomatid and heterophyid trematodes suppress phagocytosis of haemocytes in Littorina littorea (Gastropoda: Prosobranchia)

A modulation of the phagocytic activity of hemocytes from the common periwinkle Littorina littorea by secretory-excretory products (SEP) released by trematode rediae during axenic in vitro cultivation was studied. The SEP released by the parasites Himasthla elongata (Echinostomatidae) and Cryptocotyle lingua (Heterophyidae) were found to inhibit the phagocytosis of zymozan particles by periwinkle hemocytes. The specificity of SEP effects was assessed: SEP of Himasthla militaris and Cryptocotyle concavum, two trematodes belonging to the same genera but infecting another closely related prosobranch snail Hydrobia ulvae, were also shown to be able to suppress L. littorea hemocytes phagocytic activity. However, no decrease in phagocytosis rate was observed when SEP of H. elongata and C. lingua were applied to monolayers of hemocytes from the bivalve mollusc Mytilus edulis. SEP from H. elongata was fractionated; only those fractions containing proteins of molecular weight more than 50 kDa were shown to possess inhibitory activity. Different H. elongata SEP concentrations were tested in for their ability to suppress phagocytosis by L. littorea hemocytes. Even very low SEP concentrations were shown to retain their ability to decrease phagocytosis rate, the inhibitory effect being dose-dependent. Hemocytes derived from snails naturally infected with H. elongata were also found to have lower phagocytic ability as compared to healthy individuals.     

Infraspecific mitochondrial sequence diversity in Hydrobia ulvae and Hydrobia ventrosa (Hydrobiidae: Rissooidea: Gastropoda): Do their different life histories affect biogeographic patterns and gene flow?

Infraspecific relationships and population parameters are largely unknown in the ecologically significant mud snail genus Hydrobia s.I. We therefore studied infraspecific variation, population structure and gene flow in two Hydrobia species with different life history strategies: the marine, planktonic H. ulvae and the brackish-water, directly developing H. ventrosa. Based on sequencing data of a 638 bp fragment of the mtDNA gene for cytochrome oxidase 1, we found considerable differences between the two species. H. ulvae shows high average pairwisc nucleotide diversity, low population level differentiation (F_st). AND high average gene flow (Nm) between populations. Dispersal appears to accord with Wright's island model. In contrast, many populations of H. ventrosa have high population level differentiation and low gene flow. The average pairwise nucleotide diversity is relatively low; this species disperses according to Wright's isolation by distance model. Differences in dispersal modes and gene flow could be partly due to differences in type of early ontogeny and quantitative differences in passive dispersal. However, the fact that H. ulvae is a marine species with high tolerance to environmental stress and therefore less sensitive to migration barriers than H. ventrosa may better explain these differences. The extant lineages of H. ulvae and H. ventrosa most likely evolved in the northeastern Atlantic during the Pleistocene.     

滨鹬马蹄吸虫(复殖目:微茎科)的尾蚴和囊蚴期的季节动态

对科威特湾微茎科滨鹬马蹄吸虫幼虫期的中间宿主双带盾桑椹螺(Clypeomorus bifasciata)及小相手蟹(Nanosesarma minutum)的季节动态进行了研究。调查期超过一年,在检查的1 600只螺和415只蟹中, 11.8 %的螺感染了8种马蹄属线虫中的一种,且以滨鹬马蹄吸虫的感染占优势(9.9 %螺感染) ; 80 %的蟹感染滨鹬马蹄吸虫囊蚴。虽然一年四季两种宿主都会感染,但吸虫的流行和尾蚴(指成熟期感染)在夏季呈现高蜂。从螺体排出的尾蚴具有明显季节性,在此海湾必须要超过最低温度20℃。总的感染率在较大(较老)的螺里有所下降,显示吸虫影响宿主生存并随之影响宿主群体结构。囊蚴的感染丰度与蟹的个体大小有明显相关性;较大的蟹感染较多的囊蚴,显示宿主能耐受更多的吸虫。调查显示,囊蚴的感染率与蟹的大小或性别无相关性。囊蚴体外脱囊以及产卵吸虫的释放证明,成熟虫体终年存在于所有大小和性别不同的蟹里,显示从蟹到鸟的持续感染是可能的。总的来说,滨鹬马蹄吸虫在海湾的传播动态是由这两种无脊椎动物宿主来协调,并似乎是被一系列依赖于温度的活动控制,这些活动影响易感宿主种群及感染性幼虫期尾蚴和囊蚴的存在。     

Water-seeking behavior in worm-infected crickets and reversibility of parasitic manipulation

One of the most fascinating examples of parasite-induced host manipulation is that of hairworms, first, because they induce a spectacular "suicide" water-seeking behavior in their terrestrial insect hosts and, second, because the emergence of the parasite is not lethal per se for the host that can live several months following parasite release. The mechanisms hairworms use to increase the encounter rate between their host and water remain, however, poorly understood. Considering the selective landscape in which nematomorph manipulation has evolved as well as previously obtained proteomics data, we predicted that crickets harboring mature hairworms would display a modified behavioral response to light. Since following parasite emergence in water, the cricket host and parasitic worm do not interact physiologically anymore, we also predicted that the host would recover from the modified behaviors. We examined the effect of hairworm infection on different behavioral responses of the host when stimulated by light to record responses from uninfected, infected, and ex-infected crickets. We showed that hairworm infection fundamentally modifies cricket behavior by inducing directed responses to light, a condition from which they mostly recover once the parasite is released. This study supports the idea that host manipulation by parasites is subtle, complex, and multidimensional.     

Patterns of swimming of cercariae in some trematode species

Swimming activity of cercariae in the trematode species Cryptocotyle lingua, Himasthla elongata, Maritrema subdolum, and Cercaria parvicaudata (Renicola sp.) is studied using video microfilming. The larvae of C. lingua demonstrate a typical intermittent swimming with regularly alternating periods of active movement and passive floating. The duration of both periods during the first 30 min of cercaria's life retains equal, and later the passive period becomes longer. In 15 h the duration of the active period is 0.5-1 s and the passive one--20-30 s. The cercariae of H. elongata during the first 10-15 min of their free life make short stops (1-2 s) in every 10-15 s of swimming. Later these stops become rare and their duration reduces to 0.5-1 s. The cercariae of M. subdolum and C. parvicaudata are continuous swimmers, having left the molluscan host they move persistently. If stops take place they are very rare (one case in 5-15 min) and very short (0.5-1 s). The movements of cercariae body and their tail during swimming are similar in all species studied. The proximal part of the tail undulates in lateral plane, and the middle and distal parts of this organ in the top view circumscribe a figure of eight. In the side view the middle and distal parts of the tail being rotate form a figure like a screw. The cercariae move along the screw axis. The linear swimming rate in the beginning of cercariae's life is 4.8 mm/s for C. lingua, 2.4 mm/s for H. elongata, and 0.35 mm/s for M. subdolum and C. parvicaudata. By the aging the cercariae swimming rate decreases down to the full cease of swimming activity, and the larvae sediment onto the bottom. The age dynamics of swimming rate is different in cercariae of different species and is determined by the morphological functional features of the larvae.