Placental Type Interactions and Evolutionary Trends of Development of Uterus in Cestodes

In Proteocephalus thymalli and P. torulosus, a contact of the placental type in uterus was shown to be formed at two different levels. At the first level an interaction occurs between outgrowths of uterine epithelium and thin capsule of embryos closely adjacent to uterine wall. The next level is formation of contact between neighboring egg capsules, which allows distributing nutrients among fetuses present in the uterine cavity. Placental interactions in Clestobothrium acheilognathi are limited in time and space. First, a relatively small number of eggs are involved in interaction of the placental type in the uterine sac, while uterine duct is filled with freely lying eggs. Second, the closest contact is observed in eggs with non-sclerosed egg shell. One of the main evolutionary tendencies in cestodes has been shown to be a modification of uterus for formation of close interrelations with embryonic membranes in the course of transition from the extrauterine to the intrauterine type of embryonic development. Uterus in parasites with a polylecital type of the egg is suggested to serve to the greater extent as a reservoir, whereas in cestodes with oligolecital eggs, uterus performs its direct function—supply of developing embryos with nutrients. As a result, modifications of uterine epithelium are formed: from the appearance of the placental type interactions formed repeatedly in phylogenetically distant groups of cestodes to formation of branched outgrowths separating the uterine space into units or disintegration to actively functioning uterine capsules.     

Ultrastructure of the reticulate uterus and specific features of matrotrophy in three species of higher cestodes (Cestoda,Cyclophyllidea)

The ultrastructure of the reticular uterus has been analyzed in pregravid and gravid proglottids of cyclophillid cestodes dwelling in water (Alcataenia dominicana and A. larina) and in terrestrial hosts (Arostrilepis tenuicirrosa). Cells of the medullary parenchyma surrounding the uterus are filled with lipid inclusions in all species investigated. The hypertrophic development of small excretory ducts that surround the uterus, contact the uterine epithelium, and penetrate the diverticula is characteristic of Alcataenia dominicana and Arostrilepis tenuicirrosa. A comparative analysis of the results and the data available for other cestode species allow for the assumption that the reticulate structure of the uterus, lipid accumulation, and contacts between the uterine epithelium and the excretory ducts are morphological and functional adaptations that enable matrotrophy and the attainment of maximal fecundity by cyclophillid cestodes.     

Morphology and ultrastructure of the uterus of Lineolepis scutigera (Dujardin, 1845) Karpenko, 1985 (Cestoda,Cyclophyllidea, Hymenolepididae) in formation of uterine capsules

The microanatomy and ultrastructure of the uterus in different stages of ontogenesis were studied in a cyclophyllidean cestode Lineolepis scutigera. In the early stages, developing embryos lie freely in the cavity of a single unbroken uterus, which is later fragmented into separate compartments. As a result, numerous spherical uterine capsules are formed; each of them contains one formed egg. Capsules represent a fragmented but actively functioning uterus. Muscular cells containing numerous lipid inclusions are located around them. In the final stages, placenta-like relationships are formed between eggs and the epithelium of capsules. A comparative morphofunctional analysis of uterine capsules in cestodes is presented. Attention is paid to the formation of close interactions between the uterine epithelium and developing eggs.     

Schistocephalus solidus: a molecular test of premature gamete exchange for fertilization in the intermediate host Gasterosteus aculeatus

We tested the hypothesis that the cestode Schistocephalus solidus is capable of premature gamete exchange as a plerocercoid in the last intermediate stickleback host. The existence of such a reproductive mode is suggested by the highly advanced gonadal development in the plerocercoid and the large fitness gain of outcrossing. In addition, eggs from selfing cestodes have a higher hatching rate when the cestode originated from a doubly infected stickleback than when it came from a singly infected fish. We hatched eggs from 10 singly breeding cestodes that originated from doubly infected sticklebacks with the prediction that some should be outcrossed and share alleles with both the breeding cestode and the second cestode in the coinfection if the hypothesis is correct. However, all of the 430 tested larvae matched only the alleles of the breeding cestode. It is therefore very unlikely that S. solidus engaged in gamete exchange in its fish host. We suggest an alternative hypothesis to explain the higher hatching rate of eggs produced by cestodes from doubly infected fish as compared to those from single infections.     

Structure of Reproductive System of the Amphicotyllide Cestode Eubothrium rugosum (Cestoda, Pseudophyllidae)

A tendency for increase both of numerical and of ecological factors of infestation in pseudophyllid cestodes is realized due to morphological peculiarities of the structure of their reproductive system. In the species E. rugosum with the closed bursiform uterus and intrauterine development of eggs, there are revealed the same degree of maturity of oocytes in ovary, the lack of sphincter or filtering cell at the base of the ovicapt infundibulum, and the presence of a large vitelline reservoir. By analogy with trematodes, an interconnection is traced between the degree of egg maturation in the uterus and morphology of uterus and vittelicles. The thickening of the egg membrane has been shown to occur not only by deposits of individual vitelline globules at the internal superficial egg layer, but also by binding of lipid droplets excreted from the surrounding parenchyma to the uterine tube along its entire length to the external egg surface. The male reproductive system of Eubothrium rugosum is characterized by the presence of a small cirrus pouch, by the cirrus supplied with microtrichia, which corresponds to the similar vagina structure, and by localization of prostate glands outside the cirrus pouch. Comparison is presented with other ultrastructural peculiarities of organization of the reproductive system in the earlier studied species of pseudophyllid cestodes, Diphyllobothrium latum.     

Hatching strategies in monogenean (platyhelminth) parasites that facilitate host infection

In parasites, environmental cues may influence hatching of eggs and enhance the success of infections. The two major endoparasitic groups of parasitic platyhelminths, cestodes (tapeworms) and digeneans (flukes), typically have high fecundity, infect more than one host species, and transmit trophically. Monogeneans are parasitic flatworms that are among the most host specific of all parasites. Most are ectoparasites with relatively low fecundity and direct life cycles tied to water. They infect a single host species, usually a fish, although some are endoparasites of amphibians and aquatic chelonian reptiles. Monogenean eggs have strong shells and mostly release ciliated larvae, which, against all odds, must find, identify, and infect a suitable specific host. Some monogeneans increase their chances of finding a host by greatly extending the hatching period (possible bet-hedging). Others respond to cues for hatching such as shadows, chemicals, mechanical disturbance, and osmotic changes, most of which may be generated by the host. Hatching may be rhythmical, larvae emerging at times when the host is more vulnerable to invasion, and this may be combined with responses to other environmental cues. Different monogenean species that infect the same host species may adopt different strategies of hatching, indicating that tactics may be more complex than first thought. Control of egg assembly and egg-laying, possibly by host hormones, has permitted colonization of frogs and toads by polystomatid monogeneans. Some monogeneans further improve the chances of infection by attaching eggs to the host or by retaining eggs on, or in, the body of the parasite. The latter adaptation has led ultimately to viviparity in gyrodactylid monogeneans.     

Some issues related to reproduction of Pacific cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis> in waters of the eastern coast of the northern Kuril Islands and the southern extremity of Kamchatka

The analysis of data collected in December 1996 and 1998 on the reproduction of cod Gadus macrocephalus in Pacific waters of the northern Kuril Islands and the southern part of Kamchatka was performed. It was shown that the individual absolute fecundity of the cod varies within 0.197–9.729 million eggs and the relative fecundity, in the range of 24-1386 eggs. The fecundity of 1000 mature females comprises 2179–2449 million eggs. The low individual fecundity of fish is related to pseudobranchial tumor growth. The main role in cod reproduction is played by females of two-three size groups characterized by the highest numbers of mature females. It is suggested that different fecundity within the range of Pacific cod is related to environmental conditions, in particular, water temperature in the spawning grounds during spawning rather than to the habitation latitude.     

Morphological Characterisation of <Emphasis Type="Italic">Andrya</Emphasis> Railliet, 1893, <Emphasis Type="Italic">Neandrya</Emphasis> n. g. and <Emphasis Type="Italic">Paranoplocephala</Emphasis> Lühe, 1910 (Cestoda: Anoplocephalidae) in Rodents and Lagomorphs

The taxonomic significance of the main morphological features of the 25 species allocated to Andrya Railliet, 1893 and Paranoplocephala Lühe, 1910 is re-evaluated in the light of the recent molecular phylogenetic hypotheses for anoplocephaline cestodes. The present analysis and the existing phylogenetic data suggest that the structure and complexity of the early uterus are not, as previously assumed, the main phylogenetic or systematic determinants for anoplocephaline cestodes. Instead, the position of the early uterus with respect to other organs, combined with the morphology of the female genitalia, appear to allow a fairly straightforward discrimination of the three genera recognised here, without contradicting current phylogenetic hypotheses. A new genus, Neandrya n. g., is proposed for N. cuniculi (Blanchard, 1891) n. comb. (previously in Andrya), amended diagnoses are provided for Andrya and Paranoplocephala and a diagnostic key to these three genera is presented.     

Distribution of zinc in parasitic helminths

The distribution of zinc in representative groups of parasitic helminths was determined by the use of the atomic absorption spectrophotometer. The results of these analyses have shown that growing flukes (smaller forms) with active oogenesis and spermatogenesis contained more zinc than old (large) or very old adults with an empty uterus and large lobulated testes. In cestodes, the neck region and immature proglottids showed more zinc concentration than mature and gravid proglottids and fully grown cyst walls. Similarly, the youngest endogenous daughter cysts of Echinococcus granulosus contained more zinc in their walls than those of larger/older forms. Zinc was concentrated more in nematode eggs than in adult females.     

Fine structure of the female reproductive system in Sobolevicanthus gracilis and Cloacotaenia megalops (Cestoda: Cyclophyllidea)

The fine structure of the female reproductive organs and ducts in two cyclophyllidean cestodes was studied by transmission electron microscopy. All the studied ducts (vitelline, fertilization and vaginal ducts, and oviducts) as well as the uterine and ootype walls have the luminal surface elevated to form apical cytoplasmic lamellae and are surrounded by bands of circular muscules, which are attached to the basement layer. The structure of the studied ducts epithelium corresponds to the pattern described in other ducts of the cestode genital system, namely a nucleate syncytial layer. An exception is the vagina Sobolevicanthus gracilis, in which the surface is lined with not numerous atypical microtriches. The comparison of our results and the data reported for other species is given. It is found out that in different species of cestodes forming oligolecital eggs, there are observed various uterine structures and diverse contacts between the eggs capsules and the uterine epithelium. The formation of special structures and contacts is the evolutionary way from the extrauterine type to more progressive intrauterine type of embryo development in cestodes.     

Egg maturation and oviposition strategy of a capital breeder,Cleorodes lichenaria,feeding on lichens at the larval stage

Abstract 1. Capital breeders rely solely on resources acquired before breeding, whereas income breeders may obtain considerable amounts of resources following the commencement of reproduction. In income breeders oviposition occurs over a longer time period with a small number of eggs ready to be laid at the start of adult life, whereas capital breeders reproduce more rapidly and contain higher numbers of mature eggs at eclosion relative to potential fecundity. 2. This study explored egg maturation and oviposition strategy of a nocturnal geometrid moth, Cleorodes lichenaria. It was predicted, based on the long larval period of C. lichenaria and the known biology of other geometrids, that this lichen‐feeding geometrid moth concentrates egg production at the beginning of its breeding period and that external nutrients have a limited role on its fecundity. 3. Approximately 46% of eggs were ready to be laid in newly hatched females and approximately 40% of their potential fecundity was realised during the first night, after which the number and the weight of eggs decreased steadily. Feeding or drinking did not increase the fecundity of females. Pupal mass correlated positively with realised fecundity and ovigeny index. 4. Reproduction of C. lichenaria is based solely on larval‐derived resources. It is suggested that females are able to resorb thoracic tissues to increase fecundity and that this ability is probably linked to relatively long lifespan and low ovigeny index of this species, characteristics rarely observed in other geometrid moths.     

Fecundity and egg output by Toxocara canis in the red fox, Vulpes vulpes

The role of the red fox Vulpes vulpes in the dissemination of eggs of Toxocara canis into the environment is considered with reference to female worm fecundity and egg output in the faeces of infected foxes collected from four localities in southern England. A significant positive correlation was found between female worm size and the number of eggs in the uterus but there was no significant relationship between T. canis worm numbers and egg output in fox faeces. Reliable estimates of worm burdens in foxes could not, therefore, be determined from faecal egg counts alone. The highest mean egg output of 2145.0 epg recorded from adult foxes indicated that fox cubs are not necessarily the main sources of environmental contamination with T. canis eggs. Saturated magnesium sulphate was found to be a more effective flotation solution than zinc sulphate and sodium chloride for recovering eggs from fox faecal samples.     

Reproductive cycling and fecundity estimation in the upland bully

The upland bully is a prolific and widespread member of New Zealand freshwater fish communities that may mature at age 1 (or earlier). It begins spawning in spring, laying as many as eight clutches of eggs over the spring and summer, at periods as brief as 13 days. Ovaries contain two modes of maturing oocytes, so that oogenesis must be continual during the spawning season. This poses difficulties in fecundity estimation, which were solved by captive rearing. The ovoid eggs are about 2 mm long; egg size declines with clutch number during the summer, but increases with fish size. Clutch size varied, in 70 spawnings, from 121 to 880 eggs, with total seasonal fecundity up to >5000 eggs. The production of multiple clutches greatly increases seasonal fecundity and may explain the success of this species in sometimes unreliable or harsh environments.     

Revision of the Progamotaenia zschokkei (Janicki, 1905) complex (Cestoda: Anoplocephalidae), with the description of six new species

A review of cestodes currently attributed to Progamotaenia zschokkei (Janicki, 1905) (Cestoda: Anoplocephalidae) suggests that six additional species are present: Pr. tenuis n. sp. found in Thylogale stigmatica (Gould) and T. thetis (Lesson), Pr. lomatosoma n. sp.from Macropus agilis (Gould), Pr. petrogale n. sp. from Petrogale spp. and Macropus dorsalis(Grey), Pr. fimbriata n.s p. from Lagorchestes conspicillatus Gould and L. hirsutus Gould, Pr. obesa n. sp. from Onychogalea fraenata (Gould) and O. unguifera (Gould) in north-eastern Australia and Pr. kellerae n. sp. from O. unguifera in north-western Australia. The new species are differentiated by a combination of characters: prominence of scolex lobes, number of lobes in the fimbriated velum, testis number and distribution, size of the cirrus-sac, morphology of the uterus and presence or absence of a pyriform apparatus in eggs from terminal segments. The morphological differences found support earlier molecular studies, in that cestodes within each host genus are distinct, with the exception that considerable morphological variation exists within Pr. petrogale. Principal components analysis suggested that most of the cestodes from Petrogale spp. belonged to Pr. petrogale. The status of specimens from Pe. lateralis is unclear. More detailed molecular studies are required to determine the significance of the morphological variation within this taxon. A key is provided to species within the complex and the related species Pr. villosa (Lewis, 1914).     

The evolution of the forms of hermaphroditism in Cyclophyllidea (Cestoda). 2. Morphofunctional causes of the formation of tapeworms having a protogynous type of the genital apparatus development

Morpho-biological causes of the formation of protogynous forms of Anoplocephalidae (Cyclophyllidea) parasitizing rodents have been studied. The reduction in the number of segments in the strobila of anoplocephalideans (Anoplocephaloides, Paranoplocephaloides) in comparison with related polymeric species of the taxon (Paranoplocephala) suggests that protogyny is resulted from the secondary oligomerization of strobila in the phylogenesis of these cestodes. It is evident that the ecological cause of the selection for a small number of segments in the strobila is a tough competition among large polymeric cestodes that results in the formation of small anoplocephalideans located in the back part of the small intestine (Paranoplocephaloides) and in the blind gut (Anoplocephaloides) of rodents. There is slowing down of strobilation rates, acceleration of genital apparatus development and somatic proglottid morphogenesis during the change into protogyny. Due to an earlier, regarding testicle morphogenesis, formation of the ovary and uterus, uterus segments are developing most of the time in the strobila. Thus, proterogyny is one of the universal mechanisms to extend the terms of hexacanth embryogenesis in the second oligomeric strobila of tapeworms.     

成虫取食对棉铃虫雌蛾繁殖的影响

试验对雌蛾设置５个食物处理,定量地研究了成虫取食对棉铃虫蛾繁殖和寿命的影响,成虫取食对雌蛾与寿命和主效量均显著影响,补充营养延长雌蛾寿命,提高产卵量,并且」补充营养对产卵量的影响比其对寿命的影响更大,分析表明被营养的作用在于提高雌蛾后期的存活率和产卵量,前期补充营养比后期补充营养具有更大的作用。雌雌全重及其蝮部干重受成廊和的自龄的双重影响,补充营养时雌蛾体重、腹部干重及其脂肪含量下降慢,补充营养可     

The occurrence of Eubothrium fragile (Cestoda: Pseudophyllidae) in twaite shad, Alosa fallax (Lacépède) in the River Severn

The occurrence, size and maturity changes of Eubothrium fragile have been studied in postlarvae, juveniles and adult twaìte shad, Alosa fallax , from several locations in the River Severn. Parasites were only found in adult shad and not in post-larvae or juveniles. No juvenile or recently acquired cestodes were identified as such, but adults were present in shad throughout the whole period of their spawning migration. A large proportion of the parasites were gravid upon arrival in the river and, although eggs were subsequently released into fresh water, there was no loss of cestodes from the fish. It was concluded that E. fragile is a marine species, that the parasites found in adult shad in fresh water were the residue of a marine life cycle and that the eggs released in fresh water were part of the parasite's natural reproductive wastage. The distribution and biology of E. fragile were discussed and it is considered that it is typical of the marine species of the genus.     

The function of multiple mating in oviposition and egg maturation in the seed beetle Callosobruchus maculatus

Abstract.  To comprehend the significance of multiple mating in female Callosobruchus maculatus , the material-benefits hypothesis is tested, as well as the potential function and mechanism of additional mating for females. The results show that longevity and lifetime fecundity are significantly higher in doubly-mated females, whereas females subject to an interrupted second copulation (which transfers no ejaculate) show resemblance with singly-mated females in lifetime fecundity and daily fecundity, supporting the material-benefits hypothesis. Female lifetime fecundity increases with ejaculate size in doubly-, but not singly-mated females. Doubly-mated females exhibit an immediate increase in fecundity on the day of remating. Moreover, dissection of ovaries after remating shows that the number of unlaid eggs is lower in doubly-mated females in comparison with singly-mated females, whereas the total number of unlaid and laid eggs shows no significant difference. This suggests that the second mating increases the oviposition rate, but not the egg maturation rate in female C. maculatus . Thus, ejaculate serves as a stimulus for oviposition, at least on the remating day. However, because doubly-mated females lay more eggs in total than singly-mated females, doubly-mated females appear to mature more eggs than the singly-mated counterparts. This is attributed to the 'indirect-driven' hypothesis, which states that the reduction in the number of mature eggs in the oviducts brought about by oviposition stimulates the maturation of oocytes in ovarioles to replenish the number of mature eggs.     

Genetic variation in male effects on female reproduction and the genetic covariance between the sexes

Abstract.— Males of many insect species increase the fecundity and/or egg size of their mates through the amount or composition of their nuptial gifts or ejaculate. The genetic bases of such male effects on fecundity or egg size are generally unknown, and thus their ability to evolve remains speculative. Likewise, the genetic relationship between male and female investment into reproduction in dioecious species, which is expected to be positive if effects on fecundity are controlled by at least some of the same genes in males and females, is also unknown. Males of the seed beetle Stator limbatus contribute large ejaculates to females during mating, and the amount of donated ejaculate is positively correlated with male body mass. Females mated to large males lay more eggs in their lifetime than females mated to small males. We describe an experiment in which we quantify genetic variation in the number of eggs sired by males (mated to a single female) and found that a significant proportion of the phenotypic variance in the number of eggs sired by males was explained by their genotype. Additionally, the number of eggs sired by a male was highly positively genetically correlated with his body mass. The between-sex genetic correlation, that is, the genetic correlation between the number of eggs sired by males and the number of eggs laid by females, was highly positive when eggs were laid on Acacia greggii seeds. This indicates that males that sire many eggs have sisters that lay many eggs. Thus, some of the genes that control male ejaculate size (or some other fecundity-enhancing factor) when expressed in males appear to control fecundity when expressed in females. We found no significant interaction between male and female genotype on fecundity.     

Time requirement for capacitation of boar spermatozoa assessed by their ability to penetrate the zona-free hamster egg.

Boar spermatozoa were preincubated for various times in the isolated uterus and oviduct from a maturing gilt and used to inseminate zona-free hamster eggs. The proportions of eggs penetrated and activated were increased, and the interval between insemination and sperm penetration was shortened when the spermatozoa were preincubated for 4--5.5 h instead of 2--.5 h. Overall penetration rates were higher and sperm penetration occurred about 1 h earlier when the eggs were inseminated with spermatozoa preincubated in the uterus than in the oviduct. It is concluded that the change in ability of boar spermatozoa to penetrate zona-free hamster eggs is due to capcitation which requires 4--4.5 h and 5--5.5 h of preincubation in the isolated uterus and oviduct, respectively.