Gregarinasina

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Gregarines
A live specimen of a septate (or cephaline) gregarine showing the distinctive "head"-like section of the trophozoite containing the epimerite at its anterior end. Septate gregarines are intestinal parasites of arthropods.
Scientific classification
Domain: Eukaryota
Kingdom: Chromalveolata
Superphylum: Alveolata
Phylum: Apicomplexa
Class: Conoidasida
Subclass: Gregarinasina
Orders

Archigregarinorida
Eugregarinorida
Neogregarinorida

The gregarines are a group of Apicomplexan protozoa, classified as the Gregarinasina[1] or Gregarinia. The large (roughly half a millimeter) parasites inhabit the intestines of a large number of invertebrates. They are not found in any vertebrates. However, Gregarinasina is closely related to both Toxoplasma and Plasmodium, which cause toxoplasmosis and malaria, respectively. Both protists use protein complexes similar to those that are formed by the gregarines for gliding motility and invading target cells.[2][3] This makes them an excellent model for studying gliding motility with the goal of developing toxoplasmosis and malaria treatment options.

Life cycle[edit]

Gregarines occur in both aquatic and terrestrial environments. Although they are usually transmitted by the orofaecal route some are transmitted with the host's gametes during copulation (Monocystis).

In all species four or more sporozoites (the precise number depends on the species) equipped with an apical complex escape from the oocysts, a process called excystation, find their way to the appropriate body cavity and penetrate host cells in their immediate environment. The sporozoites emerge within the host cell, begin to feed and develop into larger trophozoites. In some species, the sporozoites and trophozoites are capable of asexual replication - a process called schizogony or merogony. Most species however appear to lack schizogony in their lifecycles.

In all species two mature trophozoites eventually pair up in a process known as syzygy and develop into gamonts. During syzygy gamont orientation differs between species (side to side, head to tail). A gametocyst wall forms around each pair of gamonts which then begin to divide into hundreds of gametes. Zygotes are produced by the fusion of two gametes and these in turn become surrounded by an oocyst wall. Within the oocyst meiosis occurs yielding the sporozoites. Hundreds of oocysts accumulate within each gametocyst and these are released via host's faeces or via host death and decay.

Lankesteria cystodytae are intestinal parasites of ascidians. They are examples of aseptate gregarines; these lack epimerites and instead possess attachment organelles known as mucrons

.

Taxonomy[edit]

The gregarines were recognised as a taxon by Grasse in 1953.[4] The three orders they are currently divided into were created by Levine et al in 1980.

There are currently approximately 250 genera and 1650 species known in this taxon. They are divided into three orders based on habitat, host range and trophozoite morphology.[5]

Archigregarines are found only in marine habitats. They possess intestinal trophozoites that are similar in morphology to the infective sporozoites. Phylogenetic analysis suggests that this group is paraphyletic and will need division.

There are generally four zoites in each spore in this group.

Eugregarines are found in marine, freshwater and terrestrial habitats. These species possess large trophozoites that are significantly different in morphology and behavior from the sporozoites. This taxon contains most of the known gregarine species. The intestinal eugregarines are separated into septate - suborder Septatina - and aseptate - suborder Aseptatina - depending on whether the trophozoite is superficially divided by a transverse septum. The aseptate species are mostly marine gregarines.

Urosporidians are aseptate eugregarines that infect the coelomic spaces of marine hosts. Unusually they tend to lack attachment structures and form gamont pairs that pulsate freely within the coelomic fluid.

Monocystids are aseptate eugregarines that infect the reproductive vesicles of terrestrial annelids. These latter species tend to branch closely with neogregarines and may need to be reclassified.

There are generally eight zoites in each spore in this group.

Neogregarines are found only in terrestrial hosts. These species have reduced trophozoites and tend to infect tissues other than the intestine.

There are generally eight zoites in each spore in this group.

The eugregarines and neogregarines differ in a number of respects. The neogregarines are in general more pathogenic to their hosts. The eugregarines multiply by sporogony and gametogony while the neogregarines have an additional schizogenic stage - merogony - within their hosts. Merogony may be intracellular or extracellular depending on the species.

DNA studies suggest that the archigregarines are ancestral to the others.[6]

Characteristics[edit]

  • Meiosis occurs in all species
  • Monoxenous - only one host in life cycle - almost all species
  • Mitochondria have tubular cristae and are often distributed near the cell periphery
  • Apical complex occurs in the sporozoite stage; lost in the trophozoite stage in eugregarines and neogregarines
  • Trophozoites have a large and conspicuous nucleus and nucleolus
  • Inhabit extracellular body cavities of invertebrates such as the intestines, coeloms and reproductive vesicles
  • Attachment to host via a mucron (aseptate gregarines) or an epimerite (septate gregarines); some gregarines (urosporidians) float freely within extracellular body cavities (coelom)

The parasites are relatively large spindle-shaped cells, compared to other apicomplexans and eukaryotes in general (some species are > 850 µm in length). Most gregarines have longitudinal epicytic folds (bundles of microtubules beneath the cell surface with nematode like bending behaviour): crenulations are instead found in the urosporidians.

Molecular biology[edit]

The gregarines are able to move and change direction along a surface through gliding motility without the use of cilia, flagella, or lamellipodia.[7] This is accomplished through the use of an actin and myosin complex.[8] The complexes require an actin cytoskeleton to perform their gliding motions.[9] In the proposed ‘capping’ model, an uncharacterized protein complex moves rearward, moving the parasites forward.[10]

History[edit]

The gregarines are among the oldest known parasites having been described by the physician Francesco Redi in 1684.[11]

References[edit]

  1. ^ Carreno RA, Martin DS, Barta JR (November 1999). "Cryptosporidium is more closely related to the gregarines than to coccidia as shown by phylogenetic analysis of apicomplexan parasites inferred using small-subunit ribosomal RNA gene sequences". Parasitol. Res. 85 (11): 899–904. doi:10.1007/s004360050655. PMID 10540950. 
  2. ^ Menard R (2001) Gliding motility and cell invasion by Apicomplexa: Insights from the Plasmodium sporozoite. Cell Microbiol 3: 63-73
  3. ^ Meissner M, Schluter D, Soldati D (2002). "Role of Toxoplasma gondii myosin a in powering parasite gliding and host cell invasion". Science 298: 837–841. 
  4. ^ Grasse PP (1953) Traite de Zoologie Anatomie, Systematique, Biologie. I. Protozoaires: Rhizopodes, Actinopodes, Sporozoaires, Cnidosporidies. Masson et Cie, Paris
  5. ^ Perkins FO, Barta JR, Clopton RE, Peirce MA, Upton SJ (2000) Phylum Apicomplexa. In: Lee JJ, Leedale GF, Bradbury P (ed.s) An illustrated guide to the Protozoa. 2nd ed. Society of Protozoologists. Lawrence KS Vol 1. pp.190-369
  6. ^ Leander BS (2008). "Marine gregarines: evolutionary prelude to the apicomplexan radiation?". Trends Parasitol 24 (2): 60–7. doi:10.1016/j.pt.2007.11.005. 
  7. ^ Walker, M M, Mackenzie C, Bainbridge SP, Orme C (1979) A study of the structure and gliding movement of Gregarina garnhami. J Protozool 26: 566-574
  8. ^ Heintzelman MB (2004) Actin and Myosin in Gregarina polymorpha. Cell Motil Cytoskeleton 58:83-95
  9. ^ Mitchison TJ, Cramer LP (1996). "Actin-based cell motility and cell locomotion". Cell 84: 371–379. 
  10. ^ Sibley LD, Hakansson S, Carruthers VB (1998). "Gliding motility: An efficient mechanism for cell penetration". Curr Biol 8: 12. 
  11. ^ Redi F (1984) Osservazioni intorno agli animali viventi, che si trovano negli animali viventi

External links[edit]