Rhoptry

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Tachyzoites of Toxoplasma gondii, transmission electron microscopy. Rhoptries: ro (click to enlarge) Parasite140105-fig3 Toxoplasmosis in a bar-shouldered dove - TEM of 2 tachyzoites.tif
Tachyzoites of Toxoplasma gondii , transmission electron microscopy. Rhoptries: ro (click to enlarge)

A rhoptry is a specialized secretory organelle. They are club-shaped organelles connected by thin necks to the extreme apical pole of the parasite. These organelles, like micronemes, are characteristic of the motile stages of Apicomplexa protozoans. They can vary in number and shape and contain numerous enzymes that are released during the process of host penetration. The proteins they contain are important in the interaction between the host and the parasite, including the formation of the parasitophorous vacuole (PV). [2] [3]

Contents

Characteristics

Rhoptries are one of the three characteristic secretory organelles present in all Apicomplexa along with micronemes and dense granules. [4] Rhoptries and micronemes are localized at the apical complex of the Apicomplexan organism, which suggests common ancestry of the members of the phylum and the evolution process they have experienced. [4] The name rhoptry indicates its shape as it comes from the Greek word for “club-shaped.” [5] These large membrane-bound organelles [6] are electron-dense and highly acidic [5] and have similar high density across those in Apicomplexan species. [7]

There is a variation in the number of rhoptries present in different species and during different developmental stages. For example, the tachyzoite stage of Toxoplasma gondii, which is found during the acute phase of toxoplasmosis, has 10 to 12 rhoptries, while the bradyzoite stage observed during the chronic phase of the infection has one to three rhoptries. [6] Plasmodium falciparum merozoites have two, the sporozoites have two to four, and the noninvasive ookinetes have none. [8] Meanwhile, Cryptosporidium sporozoites only have a single rhoptry. [6]

Structure and Content

Rhoptry mainly comprises two regions: the rhoptry neck and the rhoptry bulb. [6] Those two regions physically divide the parasites and have different features and materials. The rhoptry neck is an electron-dense duct [6] that narrowly extends at the anterior tip [9] and contains the rhoptry neck proteins (RONs), [6] which are named after where they localize in the parasite. [9] On the other hand, the rhoptry bulb is a larger, bulbous base that is electron-lucent and contains the rhoptry bulb proteins (ROPs) and membranous materials. [6] So far, eight rhoptry bulb proteins, ROP1 through ROP8, have been identified in T. gondii . [5] Those two classes of proteins, RONs and ROPs, follow the typical secretory pathway from the endoplasmic reticulum to the Golgi, then finally, where they are normally stored, the rhoptry. [9] They have critical functions in the host invasion and replication within the host of Apicomplexan parasites. [6] During the host invasion process, the proteins are secreted at different times at which they each function. RONs are exocytosed first because they contribute during the invasion. [8] ROPs follow afterward and perform a post-invasion role. [8]

Synthesis

De novo assembly of rhoptries occurs during cell replication. [4] They are first synthesized as pre-rhoptries, which are spherical-shaped, [4] trans-Golgi-derived vesicles. [9] Yet, how these immature rhoptries are formed is still unknown. [9] Pre-rhoptries elongate and mature into the functional rhoptries just before cytokinesis, which then move to the apexes of the parasites to localize to their normal position—the apical complexes. [4]

Functions

The three unique secretory organelles of Apicomplexamicronemes, rhoptries, and dense granules—release their contents by exocytosis at different stages of the host invasion as the process is regulated in time and space. [8] Microneme contents are secreted first to the apical end of the parasite when the parasite attaches to the host cell, followed by rhoptry as invasion proceeds, and then dense granules near post-invasion. [5] [8] The micronemal proteins secreted to the parasite’s surface direct the rhoptry proteins to the host cell by forming complexes together. [8] The rhoptry proteins then localize to different locations within the host cell, including the plasma membrane, the cytosol, the nucleus, the parasitophorous vacuole membrane (PVM), and the PV lumen. [8] The primary functions of rhoptries are to assist host invasion and to exploit host cellular functions for enhanced parasitism. [4] Still, the specific roles differ depending on where they localize within the host upon direct injection into the host cytoplasm and on the host species. [4] During the initial stage of host invasion, rhoptry contents help the parasite attach to the host, [4] and the rhoptry membranous material forms the PVM around the parasite entering the host cell to establish its protective intracellular protective compartment for successful development [8] by inducing invagination of its plasma membrane. [6] In Plasmodium , some rhoptry proteins localize to the PVM and promote the formation of the vacuole. [4] Apicomplexan parasites also utilize rhoptries to divert the host cell’s immune response. The host can come to favor the parasitic invasion if the rhoptry proteins manipulate the host’s actin cytoskeleton. [4] Furthermore, rhoptry proteins in Toxoplasma gondii can mistraffic the host’s immune factors for its virulence. [4] Another function of rhoptry proteins is nutrient import during the lytic cycle of Apicomplexa. [8]

Related Research Articles

<span class="mw-page-title-main">Apicomplexa</span> Phylum of parasitic alveolates

The Apicomplexa are organisms of a large phylum of mainly parasitic alveolates. Most possess a unique form of organelle structure that comprises a type of (non-photosynthetic) plastid called an apicoplast—with an apical complex membrane. The organelle's apical shape is an adaptation that the apicomplexan applies in penetrating a host cell.

<span class="mw-page-title-main">Toxoplasmosis</span> Protozoan parasitic disease

Toxoplasmosis is a parasitic disease caused by Toxoplasma gondii, an apicomplexan. Infections with toxoplasmosis are associated with a variety of neuropsychiatric and behavioral conditions. Occasionally, people may have a few weeks or months of mild, flu-like illness such as muscle aches and tender lymph nodes. In a small number of people, eye problems may develop. In those with a weak immune system, severe symptoms such as seizures and poor coordination may occur. If a woman becomes infected during pregnancy, a condition known as congenital toxoplasmosis may affect the child.

<i>Toxoplasma gondii</i> Type of protozoan parasite

Toxoplasma gondii is a parasitic protozoan that causes toxoplasmosis. Found worldwide, T. gondii is capable of infecting virtually all warm-blooded animals, but felids are the only known definitive hosts in which the parasite may undergo sexual reproduction.

<i>Plasmodium</i> Genus of parasitic protists that can cause malaria

Plasmodium is a genus of unicellular eukaryotes that are obligate parasites of vertebrates and insects. The life cycles of Plasmodium species involve development in a blood-feeding insect host which then injects parasites into a vertebrate host during a blood meal. Parasites grow within a vertebrate body tissue before entering the bloodstream to infect red blood cells. The ensuing destruction of host red blood cells can result in malaria. During this infection, some parasites are picked up by a blood-feeding insect, continuing the life cycle.

<span class="mw-page-title-main">Aconoidasida</span> Class of single-celled organisms

The Aconoidasida are a class of apicomplexan parasites created by Mehlhorn et al in 1980.

<span class="mw-page-title-main">Microneme</span>

Micronemes are secretory organelles, possessed by parasitic apicomplexans. Micronemes are located on the apical third of the protozoan body. They are surrounded by a typical unit membrane. On electron microscopy they have an electron-dense matrix due to the high protein content. They are specialized secretory organelles important for host-cell invasion and gliding motility.

<i>Plasmodium knowlesi</i> Species of single-celled organism

Plasmodium knowlesi is a parasite that causes malaria in humans and other primates. It is found throughout Southeast Asia, and is the most common cause of human malaria in Malaysia. Like other Plasmodium species, P. knowlesi has a life cycle that requires infection of both a mosquito and a warm-blooded host. While the natural warm-blooded hosts of P. knowlesi are likely various Old World monkeys, humans can be infected by P. knowlesi if they are fed upon by infected mosquitoes. P. knowlesi is a eukaryote in the phylum Apicomplexa, genus Plasmodium, and subgenus Plasmodium. It is most closely related to the human parasite Plasmodium vivax as well as other Plasmodium species that infect non-human primates.

<span class="mw-page-title-main">Gregarinasina</span> Subclass of protists

The gregarines are a group of Apicomplexan alveolates, classified as the Gregarinasina or Gregarinia. The large parasites inhabit the intestines of many invertebrates. They are not found in any vertebrates. Gregarines are 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 for invading target cells. This makes the gregarines excellent models for studying gliding motility, with the goal of developing treatment options for both toxoplasmosis and malaria. Thousands of different species of gregarine are expected to be found in insects, and 99% of these gregarine species still need to be described. Each insect species can be the host of multiple gregarine species. One of the most-studied gregarines is Gregarina garnhami. In general, gregarines are regarded as a very successful group of parasites, as their hosts are distributed over the entire planet.

Intracellular parasites are microparasites that are capable of growing and reproducing inside the cells of a host. They are also called intracellular pathogens.

An apicoplast is a derived non-photosynthetic plastid found in most Apicomplexa, including Toxoplasma gondii, and Plasmodium falciparum and other Plasmodium spp., but not in others such as Cryptosporidium. It originated from algae through secondary endosymbiosis; there is debate as to whether this was a green or red alga. The apicoplast is surrounded by four membranes within the outermost part of the endomembrane system. The apicoplast hosts important metabolic pathways like fatty acid synthesis, isoprenoid precursor synthesis and parts of the heme biosynthetic pathway.

Acidocalcisomes are rounded electron-dense acidic organelles, rich in calcium and polyphosphate and between 100 nm and 200 nm in diameter.

<i>Colpodella</i> Genus of single-celled organisms

Colpodella is a genus of alveolates comprising 5 species, and two further possible species: They share all the synapomorphies of apicomplexans, but are free-living, rather than parasitic. Many members of this genus were previously assigned to a different genus - Spiromonas.

Chromera velia, also known as a "chromerid", is a unicellular photosynthetic organism in the superphylum Alveolata. It is of interest in the study of apicomplexan parasites, specifically their evolution and accordingly, their unique vulnerabilities to drugs.

<span class="mw-page-title-main">Apicomplexan life cycle</span> Apicomplexa life cycle

Apicomplexans, a group of intracellular parasites, have life cycle stages that allow them to survive the wide variety of environments they are exposed to during their complex life cycle. Each stage in the life cycle of an apicomplexan organism is typified by a cellular variety with a distinct morphology and biochemistry.

<span class="mw-page-title-main">Morn repeat containing 1</span> Protein-coding gene in the species Homo sapiens

MORN1 containing repeat 1, also known as Morn1, is a protein that in humans is encoded by the MORN1 gene.

Goussia is a taxonomic genus, first described in 1896 by Labbé, containing parasitic protists which largely target fish and amphibians as their hosts. Members of this genus are homoxenous and often reside in the gastrointestinal tract of the host, however others may be found in organs such as the gallbladder or liver. The genera Goussia, as current phylogenies indicate, is part of the class Conoidasida, which is a subset of the parasitic phylum Apicomplexa; features of this phylum, such as a distinct apical complex containing specialized secretory organelles, an apical polar ring, and a conoid are all present within Goussia, and assist in the mechanical invasion of host tissue. The name Goussia is derived from the French word gousse, meaning pod. This name is based on the bi-valve sporocyst morphology which some Goussians display. Of the original 8 classified Goussians, 6 fit the “pod” morphology. As of this writing, the genera consists of 59 individual species.

<span class="mw-page-title-main">IRGs</span>

Immunity Related Guanosine Triphosphatases or IRGs are proteins activated as part of an early immune response. IRGs have been described in various mammals but are most well characterized in mice. IRG activation in most cases is induced by an immune response and leads to clearance of certain pathogens.

<span class="mw-page-title-main">Duffy binding proteins</span>

In molecular biology, Duffy binding proteins are found in Plasmodium. Plasmodium vivax and Plasmodium knowlesi merozoites invade Homo sapiens erythrocytes that express Duffy blood group surface determinants. The Duffy receptor family is localised in micronemes, an organelle found in all organisms of the phylum Apicomplexa.

Hammondia hammondi is a species of obligate heteroxenous parasitic alveolates of domestic cats. Intracellular cysts develop mainly in striated muscle. After the ingestion of cysts by cats, a multiplicative cycle precedes the development of gametocytes in the epithelium of the small intestine. Oocyst shedding persists for 10 to 28 days followed by immunity. Cysts in skeletal muscle measure between 100 and 340 μm in length and 40 and 95 μm in width. Some of the intermediate hosts develop low levels of antibody and some cross-immunity against Toxoplasma.

<span class="mw-page-title-main">Parasitophorous vacuole</span>

The parasitophorous vacuole (PV) is a structure produced by apicomplexan parasites in the cells of its host. The PV allows the parasite to develop while protected from the phagolysosomes of the host cell.

References

  1. Rigoulet, Jacques; Hennache, Alain; Lagourette, Pierre; George, Catherine; Longeart, Loïc; Le Net, Jean-Loïc; Dubey, Jitender P. (2014). "Toxoplasmosis in a bar-shouldered dove (Geopelia humeralis) from the Zoo of Clères, France". Parasite. 21: 62. doi:10.1051/parasite/2014062. ISSN   1776-1042. PMC   4236686 . PMID   25407506.
  2. Bradley, Peter J; Chris Ward; Stephen J. Cheng; David L. Alexander; Susan Coller; Graham H. Coombs; Joe Dan Dunn; David J. Ferguson; Sanya J. Sanderson; Jonathan M. Wastling; John C. Boothroyd (October 7, 2005). "Proteomic Analysis of Rhoptry Organelles Reveals Many Novel Constituents for Host-Parasite Interactions in Toxoplasma gondii". J. Biol. Chem. 280 (40): 34245–34258. doi: 10.1074/jbc.M504158200 . PMID   16002398.
  3. Richard, D; et al. (March 2009). "Identification of rhoptry trafficking determinants and evidence for a novel sorting mechanism in the malaria parasite Plasmodium falciparum". PLOS Pathogens. 5 (3): e1000328. doi: 10.1371/journal.ppat.1000328 . PMC   2648313 . PMID   19266084.
  4. 1 2 3 4 5 6 7 8 9 10 11 Sparvoli, Daniela; Lebrun, Maryse (July 2021). "Unraveling the Elusive Rhoptry Exocytic Mechanism of Apicomplexa". Trends in Parasitology. 37 (7): 622–637. doi:10.1016/j.pt.2021.04.011. ISSN   1471-4922. PMID   34045149.
  5. 1 2 3 4 Black, Michael W.; Boothroyd, John C. (September 2000). "Lytic Cycle of Toxoplasma gondii". Microbiology and Molecular Biology Reviews. 64 (3): 607–623. doi:10.1128/MMBR.64.3.607-623.2000. ISSN   1092-2172. PMC   99006 . PMID   10974128.
  6. 1 2 3 4 5 6 7 8 9 Ben Chaabene, Rouaa; Lentini, Gaëlle; Soldati-Favre, Dominique (March 2021). "Biogenesis and discharge of the rhoptries: Key organelles for entry and hijack of host cells by the Apicomplexa". Molecular Microbiology. 115 (3): 453–465. doi: 10.1111/mmi.14674 . ISSN   0950-382X. PMID   33368727.
  7. Sam-Yellowe, T. Y. (1996-08-01). "Rhoptry organelles of the apicomplexa: Their role in host cell invasion and intracellular survival". Parasitology Today. 12 (8): 308–316. doi:10.1016/0169-4758(96)10030-2. ISSN   0169-4758.
  8. 1 2 3 4 5 6 7 8 9 Cova, Marta Mendonça; Lamarque, Mauld H.; Lebrun, Maryse (2022-09-08). "How Apicomplexa Parasites Secrete and Build Their Invasion Machinery". Annual Review of Microbiology. 76 (1): 619–640. doi: 10.1146/annurev-micro-041320-021425 . ISSN   0066-4227.
  9. 1 2 3 4 5 Lebrun, Maryse; Carruthers, Vern B.; Cesbron-Delauw, Marie-France (2014-01-01), Weiss, Louis M.; Kim, Kami (eds.), "Chapter 12 - Toxoplasma Secretory Proteins and Their Roles in Cell Invasion and Intracellular Survival", Toxoplasma Gondii (Second Edition), Boston: Academic Press, pp. 389–453, doi:10.1016/b978-0-12-396481-6.00012-x, ISBN   978-0-12-396481-6 , retrieved 2023-11-27