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The susceptibility of various races of salmonids towards infections with the skin parasitic monogenean Gyrodactylus salaris Malmberg, 1957, differs markedly. Norwegian and Scottish salmon strains are known as extremely susceptible to infection, whereas Baltic salmon races such as the Neva strain (Russian origin) and the Indals river (Swedish origin) salmon have been characterized as relatively resistant. However, the status of the many other Baltic strains has remained unknown. The present study reports on the susceptibility of the Baltic salmon from the Swedish river Lule. It was shown that this strain is susceptible to infection but to a lesser extent than the Scottish salmon. Further studies showed that injection of immuno-suppressants (dexamethasone) greatly increased population growth of G. salaris on Scottish salmon but not on the Baltic salmon. Mucous cell density on fins differed between strains, and a general trend to decreased cell density on infected fish 8 wk post-infection, compared to uninfected fish, was observed. The largest decrease in mucous cell density following infection was seen in the most resistant fish. After administration of immuno-suppressants, this decrease in mucous cell density was inhibited in the Scottish salmon but not in the Baltic salmon. Thus, there seems to be a relationship between the fishes' ability to discard mucous cells and the ability to resist infections with Gyrodactylus salaris. Although the Lule salmon seems more susceptible to infection compared to previous reports on the Neva salmon, the results support the notion that Baltic salmon strains are generally more resistant than East Atlantic salmon.

Planarians are the free-living members (order Tricladida) of the phylum Platyhelminthes. They are triploblastic, acoelomate, unsegmented and located at the base of the Lophotrochozoa clade. Besides their huge regenerative capacity, planarians have simple eyes, considered similar to the prototypic eye suggested by Charles Darwin in his book 'On the Origin of Species'. The conserved genetic network that determines the initial steps of eye development across metazoans supports a monophyletic origin of the various eye types present in the animal kingdom. Here we summarise the pattern of expression of certain genes involved in the eye network that have been isolated in planarians, such as Otx, Pax-6, Six, Rax and opsin. We describe the effects of RNA interference-mediated loss of function on eye regeneration. Finally, we discuss the relevance of these findings for the evolution of the eye gene network.

The directly transmitted viviparous gyrodactylids have high species richness but low morphological and biological diversity, and many species are recorded from only a single host. They therefore constitute a guild of species ideal for studies of the evolutionary significance of host specificity. The group has the widest host range of any monogenean family, being found on 19 orders of bony fish. However, individual species range from narrowly specific (71% of 402 described species recorded from a single host) to extremely catholic (Gyrodactylus alviga recorded from 16 hosts). Gyrodactylid-host interactions extend from 60 mya (G. lotae, G. lucii) down to 150 years (G. derjavini on Oncorhynchus mykiss). Co-evolution with the host is comparatively rare within the gyrodactylids, but host switching or ecological transfer is common, and has been facilitated by the mixing of fish strains that followed glaciation. In this review, we consider the factors responsible for gyrodactylid specificity patterns, using examples from our work on salmonid gyrodactylids including G. salaris, responsible for major epidemics on wild Atlantic salmon (Salmo salar) in Norway since 1975, and G. thymalli from grayling and G. derjavini from trout.G. salaris has a wide host range with highest population growth rates on Norwegian salmon strains. However, growth rates are variable on both host strains and species, because of the multitude of micro- and macro-environmental factors influencing parasite mortality and fecundity. A better predictor of performance is the proportion of fishes of a strain which are innately resistant to the parasite, a measure which is negatively correlated with the time to peak infection in a host strain. Population growth rate is also negatively correlated with age of infection; the initial rate, therefore, predicts best the suitability of a fish as host for G. salaris. The host response to gyrodactylids appears to be the same mechanism in all salmonids with innate resistance as one end of a spectrum, but influenced by stress and probably under polygenic control. Hybrid experiments show that performance of G. salaris on a host is heritable, and usually intermediate between that of the parents. This host response mechanism, coupled with the initial parasite population growth on a fish, determines the host specificity, i.e. whether the fish will be susceptible, a responder or innately resistant. The use of population growth rate parameters allows comparison of different hosts as a resource for a gyrodactylid. In the case of G. salaris, East Atlantic and Baltic strains of Atlantic salmon are core hosts, but other salmonids can physiologically sustain infections for considerable periods, and may be important in parasite dispersal and transmission. A further group of non-salmonid fishes are unable to sustain G. salaris reproduction, but can act as transport hosts.Population growth parameters are very labile to stressors and environmental factors, particularly temperature and salinity, and also other aspects of host ecology and water quality. These factors may also influence the spectrum of hosts that can be infected under particular conditions, and probably favoured ecological transfer of gyrodactylids between host species in periglacial conditions. G. salaris may still be undergoing post-glacial range expansion (aided by anthropogenic spread) as shown by the increase in the species range over the last 25 years. The origin of G. salaris, G. teuchis and G. thymalli is discussed in relation to glacial refugiums during the last ice age.

Dendromonocotyle colorni sp. n. (Monogenea: Monocotylidae) is described from the dorsal skin surface of two specimens of Himantura uarnak (Forsskål) kept at the Eilat Underwater Observatory in Israel. Dendromonocotyle colorni is distinguished from the other eight species in the genus by the morphology of the terminal papillar sclerite on the haptor, the distal portion of the male copulatory organ and the morphology of the vagina. The development of the male copulatory organ is detailed for D. colorni and the adaptations of species of Dendromonocotyle to life on the dorsal skin surface of rays are discussed. Dendromonocotyle octodiscus Hargis, 1955 was identified from the dorsal skin surface of the southern stingray Dasyatis americana Hildebrand et Schroeder off Bimini, Bahamas and represents a new host record.

Reproductive success of Gvrodactylus salaris from River Lierelva was compared experimentally on 3 stocks of salmon (12.5 degrees C +/- 0.2). Isolated fish from 2 susceptible Norwegian stocks (Rivers Lier and Alta) and 1 resistant Baltic stock of salmon (River Neva) were infected with a single gravid worm in order to record the temporal sequence of births and age at death of individual parasites. Establishment success (proportion of worms surviving to give birth) was generally low and mortality high, but significantly fewer worms survived on Neva (45% with mean survival of 3.5 days) compared to Alta and Lier fish (60 % mean survival 7.9 and 5.2 days, respectively). There was a dramatic difference in parasite fecundity between the host stocks: only 2 births occurred on Neva fish compared to third and fourth births on both Alta and Lier hosts. The timing of the first birth was more variable on Neva hosts and was significantly extended (mean 2.3 days) relative to that on Alta and Neva fish (1.8 days). However, timing of the second birth did not vary on any of the 3 salmon stocks. Age-specific mortality and fecundity data are consistent with exponential population growth of G. salaris on Alta and Lier fish but eventual extinction on Neva hosts. This is the first demonstration that gyrodactylids maintained on different host stocks exhibit variations in fecundity, development and mortality, which may in turn account for the variable virulence noted when G. salaris infects different salmonid hosts.

The spermatozoon of the monopisthocotylean monogenean Pseudodactylogyrus sp. (a gill parasite of eels) has a single axoneme showing a 9 + '1' pattern, a nucleus and a mitochondrion, but has no cortical microtubules. This species thus provides a very simple model for the study of tubulin in the 9 + '1' axonemes of the Platyhelminthes, in contrast with digenean sperm which have a more complex spermatozoon with two such axonemes and cortical microtubules. Indirect immunofluorescence labelling of tubulin shows that the elongating spermatids, initially lying in all directions in the early stages, are arranged as parallel elements in further stages. The number of spermatids in an isogenic group could also be precisely counted and equals 32. Nuclear labelling with fluorescent dyes shows that the nuclei, first located in the common mass of the spermatids, later elongate and migrate into the growing spermatids, and that the nucleus is located in the central part of the mature spermatozoon, with the two extremities devoid of nucleus. Labelling with antibodies directed against acetylated, tyrosinated, and polyglutamylated tubulin gave positive results, thus indicating that these post-translational modifications of tubulin are present in the axoneme of spermatids and spermatozoa of monopisthocotylean monogeneans.

The epidermis of members of the major parasitic taxon Neodermata is distinctive among flatworms, being a syncytial, insunk, non-ciliated epidermis that develops through a wholesale replacement of larval epidermis at metamorphosis when the larva attacks a host. How it arose in evolution from what must have been a turbellarian-like ancestor is not immediately evident. While many turbellarian flatworms have also adopted a symbiotic way of life, the literature on ultrastructure of epidermis in these symbionts shows quite a variety of morphologies, many not so different from that of their free-living relatives. Various turbellarians do have syncytial or insunk epidermises or reduction of epidermal ciliation as is characteristic of the Neodermata, but co-occurrence in a single turbellarian of all features common to neodermatans has not been reported. Urastoma cyprinae, for example, which is ectosymbiotic on bivalves, has a ciliated cellular epidermis that is little different from what is known of epidermises of its free-living relatives. The endoparasitic Anoplodium hymanae, from the coelom of sea cucumbers, also bears a ciliated cellular epidermis, as is typical of many other rhabdocoels, but it shows marked phagocytic activity as well as incorporation of endosymbiotic bacteria. The closest similarity to neodermatan epidermis is that of the turbellarian Genostoma kozloffi, an ectosymbiont of the crustacean Nebalia: covering the bulk of the body is a non-ciliated syncytium with multiple branching connections to insunk nucleated portions, much as in epidermis of adult neodermatans and, on its ventral surface, is a field of ciliated cellular insunk epidermis resembling the epidermis of some larval neodermatans. Developmental clues to the origin of the neodermatan epidermis can be seen in turbellarian embryos. Before hatching, embryos of proseriate and triclad embryos go through 3 generations of epidermis, each replacing the next; 2 generations of epidermis are reported in the literature on rhabdocoel embryos. This process of replacement parallels the epidermal replacement that larval neodermatans undergo at metamorphosis. Ultrastructural study of developing acoel, polyclad and macrostomid embryos shows that they, too, have epidermal replacement and growth through immigration of deeper-lying cells, comparable to the processes seen in higher flatworms. Succession of distinct generations of epidermis in such animals as the proseriates, triclads and rhabdocoels is probably an adaptation to development of ectolecithal eggs, providing the means for the embryo to use yolk that resides in vitellocytes, outside its blastomeres. We propose that the Neodermata has taken advantage of this developmental mechanism, producing successive generations of epidermal cells even in its larval stages, to counter the defenses of hosts.

1. The phenomenon of asexual multiplication is rare in the animal kingdom. It occurs, however, in all main flatworm taxa. Flatworms are characterized by an extensive versatility, ranging from the different types of asexual multiplication to the different orthogonal plans for the nervous system. The role of the nervous system in the asexual multiplication taking place in flatworms is pointed out and discussed. 2. Immunocytochemical studies of the changes in the flatworm neuroanatomy show that the nervous system, particularly the main never cords, has a central role during asexual development. 3. Antibodies to different neuronal substances yield different immunoreactivity patterns and develop according to different time schedules. Serotoninergic nervous elements seem to have a leading role. 4. Substances produced by the nervous system influence fissioning and subsequent regeneration in free-living flatworms in the following ways. (a) A function as a wound hormone has been suggested for the neuropeptide RF-amide. (b) Mitogenic effects have been shown for several biogenic amines and neuropeptides. (c) Inhibitory roles are suggested for somatostatin and melatonin in connection with cell proliferation respective fissioning. 5. Growth factors have been observed both in free-living and parasitic flatworms. 6. Cells reactive to antibodies against epidermal growth factor increase in number in parallel with increases in mitotic activity in the gull tapeworm and occur in regions with high mitotic activity. A correlation between these two phenomena is suggested.

Polystome phylogeny is examined, with emphasis on the dimorphism of the frog parasite Polystoma integerrimum, which exists in a fully differentiated and a neotenic form, and the evolutionary development of exclusively neotenic genera. Protopolystoma, which infects the aquatic toad Xenopus, has essentially the same morphology as the neotenic (branchial) adult of P. integerrimum and is interpreted as a neotenic genus; however, it inhabits the bladder of its host, the infection site of the normal adult of the dimorphic species. The sphyranurid Sphyranura, ectoparasitic on the external gills of the mud puppy Necturus, resembles the two-sucker larva of P. integerrimum in possessing a single pair of haptorial suckers in place of the 6 suckers of adult polystomes, and is probably a neotenic parasite associated with a neotenic host. The neotenic animals in general are parasites of aquatic hosts, and the uterus is lost or reduced in these genera; by contrast, uterine function is greatly enhanced among polystomes infecting amphibians best adapted to terrestrial life.

Extracts of cercariae of the avian schistosome Trichobilharzia ocellata were analysed for the presence of ecdysteroids by radioimmunoassay, high-performance liquid chromatography monitoring fractions by radioimmunoassay, and gas chromatography/mass spectrometry (selected ion monitoring). Both free ecdysteroids and polar conjugated ecdysteroids were detected in the cercarial extracts. The free ecdysteroid fraction, as well as the hydrolysed polar conjugated ecdysteroid fraction, contained both ecdysone and 20-hydroxyecdysone in approximately equal amounts. The amount of ecdysteroids detected is comparable to those found in other platyhelminths. A possible role for the ecdysteroids in the development of the parasite and/or the interactions between the parasite and its intermediate host, the freshwater snail Lymnaea stagnalis, is discussed.

We analyzed the effects of infection with Trichobilharzia ocellata on hemocytes of its snail host, Lymnaea stagnalis, and correlated them with successive stages of parasite development. Circulating hemocytes were studied at 0, 2, 4, 6, and 8 weeks post exposure (p.e.) with respect to cell number, distribution of subpopulations (as characterized by morphology, determinants recognized by either of two lectins and a monoclonal antibody) and to proliferative, phagocytic and endogenous peroxidase activity. Infection results in a net elevated level of activity of circulating hemocytes at 2 weeks p.e., when mother sporocysts are present in the head-foot-mantle region, as well as at 4 weeks p.e., when daughter sporocysts are migrating to and growing in the digestive gland region. A lower level of activity was observed at 6 weeks p.e., when cercariae are differentiating within daughter sporocysts. A net activation was again found at 8 weeks p.e., when cercariae are escaping. So, infection with T. ocellata results in a net general activation of the internal defense system of L. stagnalis, during several stages of development of the parasite.

The development of immunoreactivity (IR) in the nervous system of asexually reproducing Microstomum lineare has been studied by a combination of simultaneous and double immunostaining with antisera to 5-HT and RF-amide, as well as with monoclonal antibodies to SCPB (molluscan small cardioactive peptide). Immunoreactivity appears in a distinct sequential order. 5-HT antigenicity in the postpharyngeal commissure indicates the initiation of the development of a new zooid. The development of a new brain and pharyngeal plexus always starts in connection to the parental nerve cords. Significantly different developmental patterns are observed for the IR to 5-HT and RF-amide, whereas IR to SCPB has the same localization as that to RF-amide, but appears both weaker and later during the development. Influences of the immunoreactive substances on the asexual reproduction and the feeding behaviour are discussed.

1. The occurrence and distribution of twelve dehydrogenases have been studied histochemically in two species of rhabdocoel turbellarians entosymbiotic in marine bivalves. 2. Both species (Paravortex scrobiculariae in Scrobicularia plana and P. cardii in Cerastoderma edule) possess dehydrogenases concerned with the three major energy-producing pathways. 3. P. scrobiculariae shows a much greater emphasis on glycolysis than does P. cardii. 4. It is suggested that the emphasis on glycolysis in P. scrobiculariae is an adaptation to regular variations in oxygen supply in its habitat; this adaptation facilitates feeding and associated migrations within the host which occur at times when the host is not submerged by the tide and when oxygen is likely to be in short supply. 5. Dehydrogenases associated with the pentose phosphate shunt are especially evident in the developing embryos of both species and are probably concerned with synthesis of nucleotides and nucleoproteins.

B. E. Bychovskij used the type of the adhesive structures and the formation of larvae as the basis of dividing Monogenea into groups. Using these principles he divided this class in two subclasses. Recent data confirm his chief statements.

A global study of the Metazoan parasites of Boops boops (L., 1758) have been made in the "golfe du Lion". Fourteen different species (eight Platyhelmintha, one Nematoda and five Crustacea) have been inventorized. Their respective localisation on the hosts, globals and specifics corresponding prevalences as well as their variations according to the size of the fish, and their abundance have been precised. Parasitics associations have also been examined.

The formaldehyde-glutaraldehyde-sucrose (FGS) method for in situ localization of catecholamines has been applied to the nervous system of the marine polyclad flatworm Notoplana acticola. This histochemical fluorescence technique revealed the presence of a small population of fluorescent cells within the brain. The number and positions of these neurons were constant in animals of the same size, but varied with the size of the worm. The brains of small animals (8 mm in length) were found to contain 20 fluorescent cells, whereas the largest animals studied (30 mm in length) were found to have 28 such cells. Various intermediate cell numbers were found in animals between these two sizes. The origin of the newly added fluorescent cells is uncertain. Peripheral fluorescence was found in association with the tentacular ocelli (eyespots) and interneurons within the ventral submuscular nerve plexus. The fluorescent spectrum from these cells measured in situ had a lambda max of 526 nm. Treatment with HCl shifts this peak to 530 nm. L-dopamine fluoresces with a similar peak emission before HCl treatment (525.5 nm) and shifts to the appropriate longer wavelength (530 nm) following acidification. This strongly suggests that the fluorescent substance in the neurons is dopaminergic in nature.

The evolution of tegument ultrastructures during development was studied in two Polystome species, Polystoma integerrimum and Polystoma pelobatis. It differs from Monogenea and other Platyhelminths in the presence of nuclei in the tegumentary syncytium of the oncomiracidium and their deferred elimination which occurs in the post-larva attached to the gills of the tadpole. This represents a delay in the loss of embryonic characteristics in Polystoma larvae which may be related to the possibility of neotenic development of these larvae. This delay allows us to follow naturally the considerable cytoplasmic changes which accompany the elimination of embryonic nuclei (disappearance of the ergastoplasm, golgi complexes and ribosomes, and of the vacuoles) and the transfer of control of this "enucleated" cytoplasm to nuclear information from tegumentary parenchymatic cells (appearance of new inclusions in the "annexed" cytoplasmic zone, maintenance of numerous organelles involved in the formation of these inclusions in the deep perinuclear region). The ultrastructual characteristics of ciliated cells and the tegumentary syncytium are discussed from the general point of view of the Platyhelminths and with respect to their adaptative function in the Polystomatidae. The originality of the Polystomatidae among the Monogenea is emphasized.