Description of two new actinosporean types from a brook of Fuji Mountain,Honshu, and from Chitose River,Hokkaido, Japan

Actinospore infection of oligochaetes living in the mud of 3 freshwater biotopes in Japan was studied. Using the cell-well plate method, a new aurantiactinomyxon type was found in 0.77% of the examined Tubifex tubifex oligochaete specimens from a brook near Yamanashi Prefectural Fisheries Experimental Station on Fuji Mountain. In 0.14% of Lumbriculus variagetus collected from Chitose River, near Chitose Salmon Hatchery, a new siedleckiella type was found, while at the same time 8.1% of the Lumbriculus spp. oligochaetes released triactinomyxons of Myxobolus arcticus. Of the examined Rhyacodrilus komarovi oligochaetes collected from the Mena River system, Hokkaido, 0.2, 0.6, 0.5 and 0.8% were infected with echinactinomyxon, neoactinomyxum and 2 types of triactinomyxon spores, respectively, and described in our previous paper. The oligochaetes released actinospores for several weeks. Actinospore infection showed high intensity in positive oligochaetes in the case of all the actinosporean types. Two of the actinospore types (aurantiactinomyxon and siedleckiella) presented here have not been previously described.     

Description of a new actinosporean type from South African freshwaters

Actinospore infection of oligochaetes collected from the mud of 2 freshwater biotopes in South Africa was studied. Using the 'cell-well plate method', a new aurantiactinomyxon type was found in 1.1% of the examined Branchiura sowerbyi oligochaete specimens from the Rietvlei River, north of Johannesburg, Gauteng, South Africa. In 1.5% of B. sowerbyi collected in a pond (Padda Dam), near the Rand Afrikaans University, Johannesburg, the same aurantiactinomyxon type was found. Infected oligochaetes were found only after collection and no actinosporean release was recorded in Branchiura specimens kept alive for several weeks. Actinospore infection showed a high intensity in oligochaetes in both positive cases. Until now, no actinosporean stages of myxosporeans have been described from South Africa. The aurantiactinospore type presented in this communication differs from the known types already described in the literature.     

Actinosporean infections in oligochaetes in a river system in southwest Ireland with descriptions of three new forms

Sediment samples were collected in April, June and August 1998 from a river in southwest Ireland. A total of 1,836 oligochaete specimens were collected from these sediments, and examined for actinosporean infections. Actinosporean-infected oligochaetes have been determined, symptoms and course of infection have been considered and the spores released from them have been described. Spore characteristics for each actinosporean were defined and the prevalence of infection in each oligochaete species was also determined. The overall prevalence of infection with actinosporeans within the oligochaete sample was approximately 2%. Seven types were distinguished, belonging to echinactinomyxon, aurantiactinomyxon and neoactinomyxum forms, from Tubifex ignotus, Lumbriculus variegatus and Limnodrilus hoffmeisteri, and compared with actinosporean types previously reported in the literature. Three of these forms have not been previously described, while the other 4 are morphologically and morphometrically similar to known forms. These actinosporeans may be stages in myxozoan life cycles.     

Description of raabeia,synactinomyxon and neoactinomyxum developing stages of myxosporeans (Myxozoa) infecting Isochaetides michaelseni Lastočkin (Tubificidae) in Lake Balaton and Kis-Balaton Water Reservoir,Hungary

Molecular and morphometric investigations were conducted on the actinosporean morphotypes of myxosporeans surveyed in oligochaetes of Lake Balaton and Kis-Balaton Water reservoir. Oligochaetes belonging to the species Isochaetides michaelseni Lasto?kin and Branchiura sowerbyi Beddard as well as to the genera Nais Dujardin, Dero Müller and Aeolosoma Ehrenberg were studied during an 18-month period. Actinosporeans were obtained exclusively from I. michaelseni (7,818 specimens) with very low prevalence (0.01–0.06%). Four new actinosporean morphotypes of the collective groups raabeia (2 types), synactinomyxon (1 type) and neoactinomyxum (1 type) were found and described, including the first synactinomyxon collective group from Hungarian biotopes and a new raabeia morphotype. Except for Synactinomyxon type 1, the 18S rDNA analysis revealed that the spores did not match any myxospore entity found in the GenBank.     

Occurrence of actinosporean stages of myxosporeans in an inflow brook of a salmon hatchery in the Mena River System,Hokkaido, Japan

Actinospore infection of oligochaetes living in the mud and on the roots of vegetation in an inflow brook of a Hokkaido salmon hatchery was studied within the framework of a Japanese-Hungarian research program. Two triactinomyxon types, 1 echinactinomyxon, and 1 neoactinomyxum type were isolated from the oligochaete Rhyacodrilus komarovi Timm, 1990 collected during the survey. The aurantiactinomyxons were recorded over a period of 3 mo starting from the day after oligochaete collection. The oligochaetes released actinospores for several weeks from the first day of the study. Spore excretion of individual oligochaetes was not synchronous. Of the oligochaetes examined, 0.7, 7, 3 and 3%, were infected with the echinactinomyxon, neoactinomyxum and the 2 types of triactinomyxon spores, respectively. Actinospore infection was intense in the positive oligochaetes in all 4 types. Of the 4 actinospore types presented here, 3 are described for the first time.     

Development of Myxobolus macrocapsularis (Myxosporea: Myxobolidae) in an oligochaete alternate host,Tubifex tubifex

The development of Myxobolus macrocapsularis Reuss, 1906, a myxosporean parasite of the gills of common bream Abramis brama L., was studied in experimentally infected oligochaetes. In 3 experiments uninfected Tubifex tubifex Muller and Limnodrilus hoffmeisteri (Claparéde) were exposed to mature myxospores of M. macrocapsularis. In all experiments, typical triactinospores developed in T. tubifex specimens but no infection was found in L. hoffmeisteri. Triactinospores were released from oligochaetes 66 to 99 d after initial exposure. At that time pansporocysts containing 8 triactinospores were located in the gut epithelium of experimental oligochaetes, but free actinosporean stages were also found in the gut lumen of the oligochaetes. Each triactinospore had 3 pyriform polar capsules and a barrel-shaped sporoplasm with 32 secondary cells. The spore body joined the 3 caudal projections with a stout style.     

Development of Myxobolus portucalensis Saraiva & Molnár, 1990 (Myxosporea: Myxobolidae) in the oligochaete Tubifex tubifex (Müller)

The extrapiscine development of Myxobolus portucalensis, a myxosporean parasite of the subcutaneous connective tissue of eel Anguilla anguilla L. was studied in the experimentally infected oligochaete Tubifex tubifex. After infecting parasite-free tubificids with mature spores of M. portucalensis, development of actinosporean stages was first observed with a light microscope 26 days after infection. Triactinospores of M. portucalensis emerged from the worms after 160 days of intra-oligochaete development. It was observed in histological sections that early pansporocysts were located in the gut epithelium of the experimental oligochaetes. Mature pansporocysts, each containing eight triactinospores, appeared 149 days after infection. After the rupture of pansporocysts, free actinospores were found in the lumen of the oligochaete gut. Released actinospores were floating in the water and showed a typical triactinomyxon form. Each triactinospore had three pyriform polar capsules, a sporoplasm with 32 secondary cells, a moderately long style and three slightly curved, sharply pointed caudal processes. The prevalence of infection in T. tubifex proved to be 52.5% (n = 40). No infection was found in Limnodrilus hoffmeisteri or in control Tubifex specimens     

Experimental detection of the actinospores of Myxobolus pseudodispar (Myxosporea: Myxobolidae) in oligochaete alternate hosts

The development of Myxobolus pseudodispar Gorbunova, 1936, an intracellular myxosporean muscle parasite of the roach Rutilus rutilus L., was studied in experimentally infected oligochaetes. In one experiment, uninfected Tubifex tubifex Müller and Limnodrilus hoffmeisteri (Claparéde) were exposed to mature spores of M. pseudodispar. Triactinomyxon spores developed both in T. tubifex and L. hoffmeisteri specimens. Triactinospores were first released from the oligochaetes 76 d after initial exposure. At that time, pansporocysts containing 8 triactinospores were located in the gut epithelium of experimentally infected oligochaetes, but free actinosporean stages were also found in their gut lumen. Each triactinospore had 3 pyriform polar capsules and an elongated cylindrical sporoplasm with 8 secondary cells. The spore body joined the 3 caudal projections with a relatively long style. One of the 3 caudal projections was shorter than the other two. The total length of the triactinospore was on average 206.5 microns.     

Hungactinomyxon,a new actinosporean type and collective group(Myxozoa) from <Emphasis Type="Italic">Branchiura sowerbyi</Emphasis> Beddard (Oligochaeta)

Actinosporeans are characterised by great morphological diversity. As it has been proved that the class Actinosporea is a synonym of the Myxosporea, actinosporean genera have only been regarded only as actinosporean collective groups. While most actinosporeans are released individually by their oligochaete hosts, members of the synactinomyxon, siedleckiella and antonactinomyxon collective groups are released as eight connected structural elements. These actinosporean types are differentiated by the type of unit and junction of the caudal processes. On the basis of these characteristics, a new actinosporean type, constructed from eight echinactinomyxon units, is described as hungactinomyxon. Adjacent units are joined by two of their three processes and form two interconnected cubes, each containing four echinactinomyxons. Molecular biological studies also suggest that this new actinosporean type differs from other actinosporean types built up from eight structural elements for which 18S rDNA sequences are available in GenBank.     

放射孢子虫在中国的首次发现

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Description of a new Neoactinomyxum type actinosporean from the oligochaete <Emphasis Type="Italic">Branchiura sowerbyi</Emphasis> Beddard

Actinosporean infection of oligochaetes living in the mud of a commercial gibel carp pond with myxosporean disease was studied. Six actinospore types were detected exclusively from the oligochaete Branchiura sowerbyi Beddard with very high prevalence (18%). Five out of the six types were identified as the same actinosporeans described in previous reports, the sixth actinosporean was identified as a new Neoactinomyxum type and described here based on morphological and molecular characterisation. Spore body of the actinospore was globular, much smaller than caudal processes. Three caudal processes were disc-like in apical view, hemispherical in side view, closer together and encircling the spore body. The number of sporoplasm cells was detected as eight in one specimen. The new actinosporean markedly differed from other Neoactinomyxum types in literature having much bigger caudal processes. DNA sequence analyses further confirmed the morphological identification, and revealed the actinosporean described here (KU641392) possessed less than 94% sequence similarity with myxozoans available in the GenBank database.     

The life cycle of Sphaerospora truttae (Myxozoa: Myxosporea) and some features of the biology of both the actinosporean and myxosporean stages

A previously undescribed echinactinomyxon type actinosporean was shown experimentally to be the alternate stage of Sphaerospora truttae. The echinactinomyxon type spores were found to be released from Lumbriculus variegatus and occasionally Tubifex tubifex. Overall infection prevalence of the echinactinomyxon was 0.14%. Peak release was in March-May each year. S. truttae spores developed in experimentally infected salmon 4.5 mo post-exposure to echinactinomyxon spores. Extrasporogonic stages of S. truttae first appeared in naturally infected salmon in early July and persisted for 8 to 10 wk. Correlation between peak release of echinactinomyxon spores and appearance of extrasporogonic stages of S. truttae is discussed.     

Actinosporean stages of Myxozoan parasites of oligochaetes from Nera River basin, Central Italy

The aim of this study was to check the occurrence of actinosporean stages of Myxozoan parasites of oligochaetes in the Nera River basin (Central Italy). The investigation was conducted near twelve trout farms (rainbow and brown trout) along the Nera River and in the littoral of the Piediluco Lake. Four actinosporean types, belonging to the collective groups aurantiactinomyxon, triactinomyxon, raabeia and echinactinomyxon, were detected in some of the biotopes investigated (Piediluco Lake and Sordo River). Differential diagnosis, carried out on a sub-sample of spores, showed a complete fitting with forms previously described in literature, with the exception of the triactinomyxon type.     

Prevalence and susceptibility of infection to Myxobolus cerebralis,and genetic differences among populations of Tubifex tubifex

The prevalence of infection and susceptibility of the aquatic oligochaete Tubifex tubifex to Myxobolus cerebralis, was examined in 2 studies on the upper Colorado River, Colorado, USA, where whirling disease occurs in wild trout populations. In the first study, the prevalence of infection ranged from 0.4 to 1.5%, as determined by counting the number of T. tubifex releasing triactinomyxons of M. cerebralis directly following their collection from the field. The susceptibility of those T. tubifex not releasing triactinomyxons was assessed by the number of these oligochaetes releasing triactinomyxons 3 mo following experimental exposures to spores of M. cerebralis. The prevalence of infection following experimental exposures of these T. tubifex ranged from 4.2 to 14.1%. In a second study, all T. tubifex collected at 2 different times directly from the 2 field sites in Colorado were exposed to spores of M. cerebralis. Individual oligochaetes representing those groups of T. tubifex releasing and those groups not releasing triactinomyxons at 3 mo were screened with molecular genetic markers. T. tubifex populations found at the 2 study sites consisted of 4 genetically distinct lineages that varied with respect to their susceptibility to experimental exposure to M. cerebralis. Lineages I and III contained the most oligochaetes susceptible to M. cerebralis and were the most prominent lineages at Windy Gap Reservoir, a site of high infectivity for wild rainbow trout on the upper Colorado River. In contrast, at the Breeze Bridge site which is below Windy Gap Reservoir and where M. cerebralis infections are less severe in wild trout, oligochaetes in lineages V and VI that are resistant to M. cerebralis were more prominent. These results suggest that certain habitats, such as Windy Gap Reservoir, are conducive to large and more homogenous populations of susceptible T. tubifex lineages that may serve as point sources of infection for M. cerebralis. Although not a direct objective of this study, there was no evidence of M. cerebralis infections among any oligochaetes other than those that would be classified as T. tubifex by standard morphological characteristics.     

Transmission of Thelohanellus hovorkai Achmerov, 1960 (Myxosporea: Myxozoa) to common carp Cyprinus carpio through the alternate oligochaete host

Thelohanellus hovorkai (Myxosporea: Myxozoa) was transmitted to common carp Cyprinus carpio by exposing fish to Aurantiactinomyxon spores collected from the oligochaete Branchiura sowerbyi. The morphological characteristics of the actinosporean stage are described in detail. B. sowerbyi were exposed to T. hovorkai spores isolated from the experimentally infected carp, and after 3 and 4 months the worms exhibited prevalences of the actinosporean stage at 19.47% (7/36) and 14.6% (6/41), respectively. Control, unexposed worms were negative for the actinosporean infection. This is the first report of an Aurantiactinomyxon transforming into a myxosporean belonging to the suborder Platysporina.     

Molecular phylogeny of tubificid oligochaetes with special emphasis on Tubifex tubifex (Tubificidae)

Tubifex tubifex is a cosmopolitan freshwater oligochaete whose presence has been studied as a health indicator of the aquatic environment and as a host for several myxozoan parasites of fish. Unfortunately, current morphological criteria used to distinguish Tubifex spp. (Tubificidae) are inadequate. We therefore developed mitochondrial 16S ribosomal DNA markers to examine phylogenetic relationships among aquatic oligochaetes and to distinguish species of Tubifex that might serve as hosts for a particular myxozoan parasite, Myxobolus cerebralis. Our phylogenetic analyses of oligochaetes based on a 378-bp segment yielded one most parsimonious tree with three major groups that corresponded to the families Lumbricidae, Sparganophilidae, and Tubificidae. T. tubifex and T. ignotus formed a monophyletic assemblage, and a sister relationship between the genera Tubifex and Limnodrilus was strongly supported. A second analysis of the relationship within the genus Tubifex identified six genetically distinct lineages of T. tubifex from North America and Europe that were separated by genetic distances comparable to those found for "well-defined" species of Limnodrilus. Therefore, the existence of several morphologically indistinguishable, thus cryptic, species of Tubifex in North America and Europe is suggested.     

The effect of cohabitation of Tubifex tubifex (Oligochaeta: Tubificidae) populations on infections to Myxobolus cerebralis (Myxozoa: Myxobolidae)

The competitive interactions between susceptible and resistant Tubifex tubifex (Oligochaeta: Tubificidae) exposed to Myxobolus cerebralis (Myxozoa: Myxobolidae) infections were investigated in two laboratory trials. Competition was assessed by the total parasite production over the course of the trials in mixed and pure cultures of M. cerebralis exposed worms, and by the genetic analyses of worms from the control and experimental groups at the beginning and end of the experiments. Mixed cultures of resistant and susceptible worms showed a 70% reduction in production of parasites released when compared with pure cultures of susceptible worms. In studies with laboratory and field-collected oligochaetes the mixed cultures at the end of the cohabitation experiments were dominated by resistant Tubifex from lineage V (HB strain) this strain of Tubifex has a competitive advantage over worms from other lineages. The results of this study suggest that certain species of Tubifex may be dead-end hosts to M. cerebralis by absorbing or inactivating the parasite and may also show greater survival compared to susceptible oligochaetes in certain whirling disease enzootic habitats.     

Molecular relationships and phylogeny in a community of myxosporeans and actinosporeans based on their 18S rDNA sequences

The community of myxosporeans and actinosporeans inhabiting a typical Scottish highland stream and the outflow area of an adjacent salmon hatchery was analysed on the basis of their 18S rDNA sequences. Nine myxosporeans belonging to the genera Sphaerospora, Chloromyxum, Zschokkella, Myxidium, Hoferellus and Myxobilatus were identified from mature spores in different organs of the fish species present. Twelve actinosporean types belonging to the collective groups of neoactinomyxum, aurantiactinomyxon, raabeia, echinactinomyxon and synactinomyxon were found to be released from oligochaete worms collected from sediments. Twenty of the 21 sequences obtained from these myxozoans are new entries to the myxozoan database, and the genera Chloromyxum, Hoferellus and Myxobilatus were entered for the first time. Study of the molecular relationships between the different taxa and with other myxozoan sequences available showed that the myxosporeans inhabiting the urinary system clearly cluster together, independently of host species or spore morphology. However, the sequences of the two Sphaerospora species encountered show considerable differences from other members of this group and all other freshwater myxosporeans, and they were found to occupy an ancestral marine position. Three actinosporeans, i.e. Neoactinomyxum eiseniellae, Aurantiactinomyxon pavinsis and Raabeia 'type 3' were found to represent alternate life cycle stages of Chloromyxum sp., Chloromyxum truttae and Myxidium truttae, respectively (approximately 1400 identical base pairs each). Three other actinosporeans encountered (two echinactinomyxon and one raabeia type) showed over 92% sequence identity with myxosporeans from GenBank, whereas all other actinosporeans formed a closely related group devoid of any known myxosporeans.     

First record of an actinosporean (Myxozoa) in a marine polychaete annelid

The marine polychaete Nereis (Hediste) diversicolor (Annelida) from shallow water in the Oresund, Denmark, was found to be infected with an actinosporean stage of a myxozoan parasite. The body length of the pyriform actinospore is 12-16 microm and its maximum width is 10-12 microm. The spore is triangular in apical view, with the 3 spherical polar capsules distally. The spore is without caudal processes. Eight spores develop in each pansporocyst. Free spores and pansporocysts were found in the musculature and parapodia but not in the intestine. The myxosporean stage in fish is unknown. This is the first record of an actinosporean stage in a marine polychaete, but because marine oligochaetes are rare, compared with polychaetes, the latter are believed to play an important role as invertebrate (alternate) hosts in marine myxozoan life cycles.