Schistosomatoidea and Diplostomoidea
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See discussions, stats, and author profiles for this publication at: http://www.researchgate.net/publication/262931780 Schistosomatoidea and Diplostomoidea ARTICLE in ADVANCES IN EXPERIMENTAL MEDICINE AND BIOLOGY · JUNE 2014 Impact Factor: 1.96 · DOI: 10.1007/978-1-4939-0915-5_10 · Source: PubMed READS 57 3 AUTHORS, INCLUDING: Petr Horák Charles University in Prague 84 PUBLICATIONS 1,399 CITATIONS SEE PROFILE Libor Mikeš Charles University in Prague 14 PUBLICATIONS 47 CITATIONS SEE PROFILE Available from: Petr Horák Retrieved on: 06 November 2015 Chapter 10 Schistosomatoidea and Diplostomoidea Petr Horák , Libuše Kolářová , and Libor Mikeš 10.1 Introduction This chapter is focused on important nonhuman parasites of the order Diplostomida sensu Olson et al. [ 1 ]. Members of the superfamilies Schistosomatoidea (Schistosomatidae, Aporocotylidae, and Spirorchiidae) and Diplostomoidea (Diplostomidae and Strigeidae) will be characterized. All these fl ukes have indirect life cycles with cercariae having ability to penetrate body surfaces of vertebrate intermediate or defi nitive hosts. In some cases, invasions of accidental (noncompat- ible) vertebrate hosts (including humans) are also reported. Penetration of the host body and/or subsequent migration to the target tissues/organs frequently induce pathological changes in the tissues and, therefore, outbreaks of infections caused by these parasites in animal farming/breeding may lead to economical losses. 10.2 Schistosomatidae Members of the family Schistosomatidae are exceptional organisms among digenean trematodes: they are gonochoristic, with males and females mating in the blood vessels of defi nitive hosts. As for other trematodes, only some members of Didymozoidae are P. Horák (*) • L. Mikeš Department of Parasitology, Faculty of Science , Charles University in Prague , Viničná 7 , Prague 12844 , Czech Republic e-mail: [email protected]; [email protected] L. Kolářová Institute of Immunology and Microbiology, First Faculty of Medicine , Charles University in Prague and General University Hospital in Prague , Studničkova 7 , Prague 12800 , Czech Republic e-mail: [email protected] R. Toledo and B. Fried (eds.), Digenetic Trematodes, Advances in Experimental 331 Medicine and Biology 766, DOI 10.1007/978-1-4939-0915-5_10, © Springer Science+Business Media New York 2014 332 P. Horák et al. gonochoristic, and only adults of Aporocotylidae and Spirorchiidae inhabit blood circulation of their hosts. The family Schistosomatidae comprises 14 genera parasitizing mammalian and avian hosts. Besides the genus Schistosoma that has medical and veteri- nary importance (human and mammalian parasites), three genera (Bivitellobilharzia , Heterobilharzia , Schistosomatium ) infect mammals, and ten genera (Allobilharzia , Anserobilharzia, Austrobilharzia , Bilharziella , Dendritobilharzia , Gigantobilharzia , Jilinobilharzia , Macrobilharzia , Ornithobilharzia , Trichobilharzia ) cause avian diseases. 10.2.1 Bird Schistosomes 10.2.1.1 Life Cycle Bird schistosomes have a two-host life cycle; generally the eggs containing mature miracidia are released via feces from the infected host to the water environment where miracidia hatch in a while. In case of Trichobilharzia regenti , however, the miracidia hatch directly in the host nasal tissue, and leave the host upon contact with water. Miracidia are active swimmers and seek after appropriate intermediate snail host. In order to fi nd the host, miracidia possess receptors to be able to respond to physical factors (gravity, light) as well as chemical cues. Concerning the latter stim- uli, carbohydrates bound in miracidia-attracting glycoproteins (MAGs, miraxons) emitted by the snail hosts seem to serve for chemoorientation and host recognition by Trichobilharzia szidati (syn. Trichobilharzia ocellata ) [ 2 , 3 ]. Subsequent larval development of bird schistosomes, characterized by an asexual multiplication in intermediate hosts, proceeds in freshwater, brackish, or saltwater snails. The details of intramolluscan development are known for a lim- ited number of bird schistosomes. With regard to morphogenesis of intramollus- can larval stages, and also their infl uence on snail physiology and immunity, T. szidati seems to be the most studied model [ 4 ]. The parasite inhibits certain immune functions, such as phagocytosis, effi ciency of encapsulation and bacte- rial killing, and stimulates production of humoral defense molecules, e.g., mol- luscan defense molecule (MDM) and granularin. Moreover, the developing trematode larvae induce production of schistosomin, a molecule released proba- bly from snail hemocytes and connective tissue cells, and competing with snail hormones for receptor binding sites on the target reproductive tissues (inhibition of oviposition) [ 5 ]. Several weeks (depending on temperature) after exposure to miracidia, the snails start to liberate cercariae. These freely swimming larvae (Fig. 10.1a ) respond to some physical and chemical stimuli and exhibit specifi c behavior in the aquatic environ- ment. Trichobilharzia szidati recognizes substrates being warmer than the ambient water and attaches to the surfaces containing ceramides and cholesterol. As the infec- tion of a new host comprises several steps, the penetration itself is triggered by other chemical signals—unsaturated fatty acids for T. szidati and free sterols for Austrobilharzia terrigalensis [ 3 ]. Besides the action of cercarial muscles, secretions of cercarial penetra- tion glands containing histolytic enzymes play a crucial role in skin penetration. 10 Schistosomatoidea and Diplostomoidea 333 Fig. 10.1 ( a ) Ocellate furcocercaria of Trichobilharzia regenti (length of about 800 μm) from the intermediate snail host, Radix labiata. (Author: Dr. J. Bulantová); ( b ) Advanced schistosomulum of Trichobilharzia regenti isolated from the spinal cord of a duck 11 days post infection, and then cultivated 3 days in vitro. The estimated length of the fully mature worms is about 11 mm. (Author: Dr. J. Bulantová) Whereas some human schistosomes use serine peptidases (e.g., cercarial elastase) as the main penetration enzymes, in T. szidati and T. regenti (and also in Schistosoma japonicum ) cysteine peptidases (cathepsin B-like) from the penetration glands are supposed to play a role [ 6 ]. After the attachment, the body of cercariae enters the skin, whereas the tail is removed. Once in the host, cercarial body transforms to schistosomulum being able to evade host immune attack: (1) Thick glycocalyx on the surface is rapidly removed and a new surface double membrane of the tegument is formed. (2) The density of surface antigens drops to the level undetectable by specifi c antibodies and some lectin probes [ 7 ]. In birds, the adult schistosomes live either in visceral organs or nasal mucosa; therefore, we distinguish visceral and nasal schistosomes. These two groups also differ in migratory routes within the host body; this statement is however based on a few experimental models. After penetration of the host skin, visceral schistosomes enter blood vessels and use the blood circulation to reach the fi nal location; during the migration schistosomula can therefore be found in the lungs, liver, etc. On the other hand, nasal schistosomes search for peripheral nerves and use them to crawl to the spinal cord and brain. This temporary location is typical in the phase before reaching the nasal area of their avian hosts. Of course, depending on the infection dose, some migrating schistosomula may be found in ectopic locations (e.g., nasal T. regenti in the lungs). In addition to the infections of compatible defi nitive hosts, cercariae of some genera of bird schistosomes (e.g., Austrobilharzia , Bilharziella , Gigantobilharzia , Trichobilharzia ) have been confi rmed as the causative agent of human cercarial dermatitis. It means that the human skin possesses components that may be recog- nized by cercariae as signals for attachment and penetration of the accidental host. It is remarkable that the content of attractive fatty acids is even higher in the human skin when compared with the bird skin [ 3 , 8 ]. 334 P. Horák et al. 10.2.1.2 Occurrence Bird schistosomes can be found around the world [ 9 ]. Geographical latitude plays a minor role provided the intermediate snail and the defi nitive bird hosts are available for transmission. Therefore, bird schistosomes and cercarial dermatitis have been reported, e.g., from Iceland and Norway [ 10 , 11 ]. The prevalence rate in birds may differ with regard to local conditions, but in some cases it can be high, reaching even 90–100 % (e.g., the USA and New Zealand) [ 12 , 13 ] . W a t e r f o w l ( sensu lato ) seems to serve as dominant defi nitive host; nevertheless, other groups of birds (e.g., passerines) have also been reported [ 9 ]. On the other hand, the infections of snail intermediate hosts are less frequent. In many localities, the prevalence rate does not exceed 1 %. However, outbreaks with prevalences up to 50 % have also been recorded (e.g., Denmark and Russia) [ 14 , 15 ]. Predominantly, pulmonate snails of the families Lymnaeidae, Physidae, and Planorbidae are used for larval development of bird schistosomes. Nevertheless, other groups of snails may also be involved in the life cycles of some genera/species [ 9 ]. 10.2.1.3 Damage to the Defi nitive and Accidental Hosts As mentioned above, the infective larvae (cercariae) penetrate the skin of bird hosts; usually feet serve as the entry site, because the rest of the body covered