Factors affecting abundance of Triaenophorus infection in Cyclops strenuus, and parasite-induced changes in host fitness

Factors affecting the abundance of Triaenophorus crassus and Triaenophorus nodulosus procercoids in their copepod first intermediate host, Cyclops strenuus, and effects of infection on feeding behaviour, reproduction and survival of the host were studied experimentally. When exposed to the same number of coracidia, copepods harboured considerably less procercoids in the trials where ciliates or Artemia salina nauplii were given as alternative food items. The prevalence of infection was higher in adult copepods as compared with copepodite stage IV and stage V, and higher in stage V than in stage IV. The prevalences in adult females and males did not differ significantly from each other. The frequency of females carrying egg sacs was lower among infected than among exposed uninfected and unexposed copepods. The rate of feeding on Artemia nauplii remained at the same level in uninfected copepods, but decreased strongly in infected copepods during 7 days p.i. The survival of unexposed, exposed uninfected and infected copepods did not differ significantly from each other for the first 11 days post-exposure, but the mortality of infected copepods increased significantly after 3 weeks post-exposure. However, the rate of development and mortality of copepods might have been affected by the apparently arrested development of stage IV copepodites found in the experiment. Some of the contradictions between these results and earlier observations are suggested to be caused by the differences in the duration of exposure, intensity of infection and duration of observation post-exposure in the present study as compared with other experiments.     

Changes in body-length, and in shape of the furcal ramus, of a freshwater cyclopoid copepod

Some morphological relationships during larval development of a freshwater cyclopoid copepod (Crustacea), Cyclops strenuus abyssorum Sars (Gurney, 1933) have been studied. The proportion which the cephalothorax forms of total length decreases at each copepodid moult whilst that which the furcal ramus forms increases. However, within each instar, each of these ratios is a constant at different seasons and in different lakes. The amount by which total length in one instar is greater than that in the preceding instar tends to decrease at each moult from a maximum of 1.31 in the first copepodid moult to lower values, especially in males. The furcal ramus is twice as long as wide in the first copepodid instar but seven times as long as wide in the adult female. This change is enhanced in the last two moults not only by a proportionately greater increase in length over width but by an absolute reduction in width itself.     

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.     

The helminth parasites of the bullhead Cottus gobio (L.) and the stone loach Noemacheilus barbatulus (L.) from the River Avon, Hampshire

Specimens of Cottus gobio and Noemacheilus barbatulus from the River Avon, Hampshire, were examined between July 1970 and July 1971. Both fish were found to act as hosts for Nicolla gallica (adult), Pomphorhynchus laevis (adult) and Triaenophorus nodulosus (plerocercoid). The incidence and intensity of infection were lower in N. barbatulus than in C. gobio , and this was related to differences in host diet. A small number of N. barbatulus were also found to harbour Proteocephalus torulosus . Seasonal cycles of incidence and intensity were evident in N. gallica , with a peak occurring in the winter. P. laevis and T. nodulosus showed no such cycles. Increasing host size was associated with a higher infection of both N. gallica and P. laevis , but not T. nodulosus . The diet of the host was considered to be important in influencing this increase. Differences in host reproductive condition affected the population levels of N. gallica only, where an increased parasite burden was evident in gravid female fish. This was considered to influence the observed increase in incidence and intensity of this parasite during the winter. None of the parasites showed any seasonal maturation cycles. There was no evidence of any interactions between the parasite populations. Host diet and the availability of infective larvae were considered to be the most important features influencing the number of parasites which became established, and the rate at which this occurred. Temperature probably acted indirectly on the establishment of N. gallica , by controlling the availability of the larval stages, and also, by influencing the host reproductive condition, on the length of time for which the parasite was retained by the host.     

The life-cycle of Ergasilus briani Markewitsch, 1993 (Copepoda: Poecilostomatoida)

The life cycle of Ergasilus briani is shown to consist of six nauplius stages, five copepodid stages and the adults. Only the adult female is parasitic. The free-living stages are described in detail. The presence of three aesthetascs on the antennules of all copepodid stages and the adult is noted for the first time. Unusual aspects of E. briani development include: the antennule is sexually dimorphic from copepodid IV onwards, the dimorphism being expressed in the numbers of setae present on the more proximal segments, and the first appearance of the male maxilliped is at the copepodid II stage. The endopodal setation of the male maxilliped is the maximum known within the Poecilostomatoida. The striking pigmentation of the developmental stages of ergasilids is noted and discussed.     

Dormancy, dispersal and the survival of cyclopoid copepods (Cyclopoida, Copepoda) in a lowland floodplain

1. The survival of cyclopoid copepods was investigated in a floodplain for 2 years where flooding occurred during the cold season. The cyclopoid community was studied in three waterbodies with distinct hydroperiods: a permanent pond connected to the flooded area during inundation, a temporary pool that is part of the flooded area and an isolated temporary pool.
2. Field studies, including data obtained from samples of water, sediment and soil, showed the overall predominance of species with a summer diapause over those with a winter diapause or without diapause. Emergence of cyclopoid copepods at the onset of flooding, examined using emergence traps and data from recently filled or still isolated temporary pools, showed that some species can survive several months of drying.
3. The ability of the diapausing fourth copepodid stages of Cyclops strenuus and C. insignis , the two cyclopoids most abundant during winter and spring flooding, to survive terrestrial conditions was tested in laboratory experiments. Both species survived for several months, but rates differed among the species. A higher percentage of C. strenuus survived for a longer period, possibly explaining why this species was relatively more abundant in more temporary habitats.
4. Both dormancy and dispersal facilitated survival of cyclopoid copepods in transient habitats connected to each other during flood periods. Dormancy was the most important survival strategy, whereas dispersal could be more important following prolonged periods without flooding.     

EVOLUTION OF INCOMPATIBILITY-INDUCING MICROBES IN SUBDIVIDED HOST POPULATIONS

Many insects, other arthropods, and nematodes harbor maternally inherited bacteria inducing "cytoplasmic incompatibility" (CI), reduced egg hatch when infected males mate with uninfected females. Although CI drives the spread of these microbes, selection on alternative, mutually compatible strains in panmictic host populations does not act directly on CI intensity but favors higher "effective fecundity," the number of infected progeny an infected female produces. We analyze the consequences of host population subdivision using deterministic and stochastic models. In subdivided populations, effective fecundity remains the primary target of selection. For strains of equal effective fecundity, if population density is regulated locally (i.e., "soft selection"), variation among patches in infection frequencies may induce change in the relative frequencies of the strains. However, whether this change favors stronger incompatibility depends on initial frequencies. Demographic fluctuations maintain frequency variation that tends to favor stronger incompatibility. However, this effect is weak; even with small patches, minute increases in effective fecundity can offset substantial decreases in CI intensity. These results are insensitive to many details of host life cycle and migration and to systematic outbreeding or inbreeding within patches. Selection acting through transfer between host species may be required to explain the prevalence of CI.     

Physiological specificity of the interaction between Traenophorus nodulosus (Cestoda) and its host--fishes

The influence of the T. nodulosus plerocercoid invasion on the liver of the intermediate fish hosts (perch) was studied. The water, protein, glycogen and glucose contents was examined in adult worms inhabiting the intestine of definitive hosts (pike), the rate of glucose accumulation along the strobile of parasite and an ability of adult worms to hydrolyze the main nutrients was also investigated. As a result of the plerocercoid infection, the glucose and glycogen contents in the liver of perch juveniles decreased, while the wet weight of liver increased. It was found, that the water and protein contents from the anterior to posterior section of strobile in T. nodulosus adults was not significantly different, while the glucose contents and the rate of its accumulation decreased. The data confirming the possibility to realize the final stages of protein and carbohydrate hydrolisis with the participation of membrane digestion on the surface of helminths and proving the ability of cestodes. The problems of relations of T. nodulosus with intermediate and definitive hosts at the level of the whole organisms are discussed.     

An empirical model of the optimal timing of reproduction for female amphipods infected by trematodes

Life-history theory predicts that hosts should reproduce when first infected by parasites if hosts are capable and if parasites have a lower cost on current than on future reproduction of hosts. We constructed an empirical model to explore fitness of females of the intertidal amphipod Corophium volutator that reproduced soon versus long after infection by the trematode Gynaecotyla adunca. For uninfected females, the optimal time to reproduce was at their maximum body length. However, for females infected by low or high intensities of trematode metacercariae, reproductive potential (realized fecundity) was highest for females that mated immediately after becoming infected. Even after removing a high cost of delaying reproduction for infected amphipods (high likelihood of depredation by sandpipers, which are final hosts of G. adunca), realized fecundity remained highest if reproduction occurred immediately following infection by trematodes. Results from our model support the view that early reproduction of female amphipods following infection by G. adunca is an adaptive life-history response to parasitism.     

Postembryonic development of Drescheriella glacialis Dahms & Dieckmann (Copepoda,Harpacticoida) reared in the laboratory

Summary All 6 naupliar and 5 copepodite stages of Drescheriella glacialis, a tisbid from Antarctic fast ice, are described. A key for the identification of the naupliar stages is given. Sexual dimorphism makes its first appearance in the copepodid IV. The setae configuration of the exopod of the second antenna in copepodid I is considered to be an ancestral state, and subsequent ontogenetic reductions are proposed to be apomorphic processes.     

Chalimus stages of Caligus latigenitalis (Copepoda:Caligidae) parasitic on blackhead seabream from Japanese waters, with discussion of terminology used for developmental stages of caligids

The first and third chalimus stages and chalimus adult (previously known as young adult) of Caligus latigenitalis are described based on new material collected from the body surface of a heavily infected wild blackhead seabream Acanthopagrus schlegelii schlegelii from Hiroshima Bay, Japan. The second and fourth chalimus stages of the same species are redescribed. Adults of C. latigenitalis are characterized by possessing 2 stout marginally indented processes at the base of 2 terminal spines at distal exopodal segment of leg 4. The chalimi were identified to stage based on the structure of the frontal filament and the discrete ranges in body length. Sexual dimorphism is first observed at the third chalimus stage in the shape of the distal segment of the antenna. The total number of postnaupliar stages of C. latigenitalis is 6, including 4 semaphoronts, i.e., 1 copepodid stage consisting of 1 infective copepodid and the chalimus copepodid, 4 chalimus stages, and 1 adult stage with 1 chalimus adult and 1 mobile adult. New terminology for the developmental stages of caligid copepods is proposed herein by amending the definition of chalimus as the postnaupliar stages, as well as semaphoronts having a frontal filament for host attachment.     

Postembryonic development ofParalabidocera antarctica (I. C. Thompson) (Copepoda,Calanoida) from the fast ice near Syowa Station,Antarctica

Six nauplius and five copepodid stages as well as adults ofParalabidocera antarctica (I. C. Thompson, 1898) (Copepoda:Calanoida) are described based on specimens obtained from fast ice and collected by a plankton net near Syowa Station (69°00 S, 39° 35 E), Antarctica. The adult male and female are redescribed in detail. Nauplius stages ofP. antarctica are very similar to the previously describedAcartia species. Sexual dimorphism becomes apparent from copepodid IV onwards in the morphology of antennule and leg 5. The copepodid stages of this species retain certain characteristics not only of Acartiidae but also of Pontellidae and Parapontellidae.     

Storage of embryonically transcribed poly(A) RNA and its utilization during metamorphosis of the hydroidHydractinia echinata

Summary The formation of tentacles and stolons during metamorphosis is severely disturbed if inhibitors of mRNA metabolism are applied during certain phases of development. The periods of sensitivity to -amanitin are late gastrulation and the disk stage of metamorphosis. A cordycepin sensitive phase exists during the first hour of metamorphosis. In all drug sensitive phases an enhanced poly(A) synthesis is found indicating increased mRNA metabolism in these stages. Pulse-chase experiments show that planula larvae store a poly(A)-rich RNA population sedimenting between 28–18s. These long living molecules are of embryonic origin, are located in RNP particles and are degraded during metamorphosis. The particles in question appear to be stored mainly in interstitial cells. In early metamorphosis no uridine is incorporated but labelled poly(A) is added to preexisting molecules.     

Requirement for protein synthesis for survival of unmodified bacteriophage T1 in a restricting host.

At high multiplication of infection, a substantial fraction of restricting cells (P1 lysogens) could be productively infected by unmodified coliphage T1 (T1.0) provided that protein synthesis was uninhibited during the first 5 min of infection. Successful infection under restricting conditions was accompanied by more genetic recombination than was seen under nonrestricting host, the recombination frequency declined for markers on T1.0 genomes; no effect was seen on recombination between markers on modified (T1.P) genomes. This suggested that recombination between unmodified genomes may be essential for their survival under conditions of host restriction. In a restricting host, genetic markers on T1.0 could recombine with T1.P even when the rescuing phage was added 6 min after T1.0 infection. However, even marker rescue recombination was diminished when protein synthesis was inhibited during early infection. Since DNA restriction is an early event, protein synthesis may be required soon after infection of a restricting host by T1.0 in order to preserve restriction-damaged DNA in a form that can participate in recombination. Experiments are also described that rule out some possibilities for the role of such a protein(s).     

Seasonal changes in the abundance of Microcrustacea in a small pond

Sampling methods, and methods used in extracting, identifying and counting the Microcrustacea in samples are outlined. Numbers of parthenogenetic females and young stages of Daphnia obtusa were highest from March/June to Sept./Nov., with a trough in July. Ephippial females were recorded during some or all of the months from March to Nov. with two numerical maxima, one in early summer and one in autumn. Males were present during the same period as ephippial females. Naupliar, copepodid and adult stages of Cyclops strenuus strenuus were recorded in every month; numbers of all three stages tended to be high from March to Nov. and low from Dec. to Feb. Females were present in smaller sumber than males. Females with eggs sacs occurred every month with two peaks of abundance in April–July and Oct. Adult Canthocamptus staphylinus were absent during the summer/early autumn from June/July/Aug. to Sept./Oct. Numbers were highest from Nov. to May/June with a peak in spring. Females carrying an egg sac were recorded each month with the highest proportion occurring sometime in the period Feb. to May. Possibly two peaks, one in early summer and one in autumn, of total numbers of Cyclocypris ovum were recorded. Comparisons with other available information on the life-cycle of the species are made. The effects on the species of a six-week period of drought in the autumn of 1969 are considered.     

Challenges of metamorphosis in invertebrate hosts: maintaining parasite resistance across life‐history stages

1. Insects lack the acquired immune system of vertebrates, but there is some evidence that insect immunity can be primed against an encountered pathogen to mitigate the intensity of future infections within a life stage. 2. Many invertebrates have multiple life‐history stages separated by complete metamorphosis, but different life stages can often be infected by the same pathogens, and the potential loss of immune priming during metamorphosis could therefore have detrimental effects on the host. Evidence that invertebrate immune priming can persist through metamorphosis is still missing, and consequently it is unclear how host–parasite interactions change across different life‐history stages in the context of infection history. 3. By experimentally manipulating the infection history of the flour beetle Tribolium confusum, we show that intestinal gregarine parasite infections during the larval stage reduced parasite load in adults, demonstrating that a host‐controlled mechanism for parasite resistance can persist through complete metamorphosis in insects. 4. Infections reduced larval developmental rates and increased host mortality but only during the crucial metamorphic stage, indicating that parasites impact multiple life stages. In general, our results demonstrate that invertebrates can show surprisingly robust immune priming despite dramatic physiological changes and protect hosts across completely different life‐history stages.     

Taxonomic studies on brackish copepods in Korean waters--II. Ontogeny and phylogeny of appendages in copepodid stages of Tortanus derjugini Smirnov, 1935 (Copepoda, Calanoida)

The development of all copepodid stages of a brackish calanoidcopepod Tortanus (Eutortanus) derjugini from western Korea isinvestigated. Homologies of segmentation and setation in adultcharacters are traced through all copepodid stages. The fourthand fifth pedigers are secondarily fused at the moult to theadult. Antennulary setation shows sexual dimorphism first atstage IV. The exopod of leg 1 is divided into two segments withoutaddition of any element at the moult to stage II, while an innerseta on the first segment is newly added at the moult to stageIII. This pattern differs from the basic pattern proposed forother copepods, in which the division of the exopod is precededby the formation of an element or both occur at the same time.In the fifth leg of the male some elements are suppressed duringthe final moult into the adult. The ontogenetic analyses indicatethat adult characters of some subgenera of Tortanus are expressedby their suppressions at the developmental phases. Suppressionof elements is important for the evolution of Tortanus.     

Patterns of Haemoproteus beckeri parasitism in the gray catbird (Dumatella carolinensis) during the breeding season

We determined the prevalence and intensity of blood parasites in breeding gray catbirds (Dumatella carolinensis) at Killbuck Wildlife Area in Wayne and Holmes Counties, Ohio (USA) from June through August 2000. Of 98 catbirds sampled, 40 (40.8%) had detectable infections of Haemoproteus beckeri. Overall prevalence of H. beckeri in this population is high relative to that reported in earlier blood parasite surveys of both breeding and migrant catbirds. Mean intensity of H. beckeri infection did not vary significantly between young and old birds or among sampling periods. We found no effect of age on prevalence or intensity of H. beckeri infection. Older birds were not more likely to be infected than younger birds, despite longer exposure to arthropod vectors. Prevalence varied significantly with season and was highest in June and lowest in August. This pattern also was observed in older birds sampled repeatedly. This seasonal variation may reflect both newly acquired infections and chronic infections relapsing in response to hormonal changes associated with breeding. Evidence of transmission was observed in the single hatching year bird that lacked detectable infection in early summer, but demonstrated a very high intensity infection in late summer. These observations provide supportive evidence that hematozoa infections are acquired on the breeding grounds during the first year of life and relapse during the breeding season in subsequent years.     

Effects of Plasmodium yoelii nigeriensis infection on Anopheles stephensl egg development and resorption

Abstract It has been shown previously that infection with Plasmodium yoelii nigeriensis reduces the number of eggs produced by female Anopheles stephensi. Here we examine the mechanism underlying fecundity reduction. Ovaries from infected and uninfected (control) female mosquitoes were examined12, 24 or 36 h after blood-feeding during the first gonotrophic cycle (replicated) or the second gonotrophic cycle (unreplicated). Follicular development was assessed according to Christophers' stages and the proportions of developing and resorbing follicles per ovary were determined. Resorption of some follicles commenced within 12h of blood-feeding, affecting significantly more follicles in the infected females: 1.1% v. 3.2%. The difference was greatest 36h after blood-feeding: 25% reduction (10 v. 35%) in the first cycle; 16% reduction (9 v. 25%) in the second gonotrophic cycle. The mean speed of oogenesis was also found to be significantly retarded in infected mosquitoes. During the second gonotrophic cycle, for example, only 92–94% of follicles reached stage III by 24 h and stage IV by 36 h in infected females, whereas all the developing follicles of uninfected females reached these stages more or less synchronously in the time specified.