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Recent Advances in the Field Trichomonas Vaginalis [Version 1 F1000Research 2016, 5(F1000 Faculty Rev):162 Last updated: 17 JUL 2019 REVIEW Recent Advances in the Trichomonas vaginalis Field [version 1; peer review: 2 approved] David Leitsch Institute of Parasitology, Vetsuisse Faculty of the University of Bern, University of Bern, Längassstrasse, Bern, 3012, Switzerland First published: 11 Feb 2016, 5(F1000 Faculty Rev):162 ( Open Peer Review v1 https://doi.org/10.12688/f1000research.7594.1) Latest published: 11 Feb 2016, 5(F1000 Faculty Rev):162 ( https://doi.org/10.12688/f1000research.7594.1) Reviewer Status Abstract Invited Reviewers The microaerophilic protist parasite Trichomonas vaginalis is occurring 1 2 globally and causes infections in the urogenital tract in humans, a condition termed trichomoniasis. In fact, trichomoniasis is the most prevalent version 1 non-viral sexually transmitted disease with more than 250 million people published infected every year. Although trichomoniasis is not life threatening in itself, it 11 Feb 2016 can be debilitating and increases the risk of adverse pregnancy outcomes, HIV infection, and, possibly, neoplasias in the prostate and the cervix. Apart from its role as a pathogen, T. vaginalis is also a fascinating organism with F1000 Faculty Reviews are written by members of a surprisingly large genome for a parasite, i.e. larger than 160 Mb, and a the prestigious F1000 Faculty. They are physiology adapted to its microaerophilic lifestyle. In particular, the commissioned and are peer reviewed before hydrogenosome, a mitochondria-derived organelle that produces publication to ensure that the final, published version hydrogen, has attracted much interest in the last few decades and rendered T. vaginalis a model organism for eukaryotic evolution. is comprehensive and accessible. The reviewers This review will give a succinct overview of the major advances in the T. who approved the final version are listed with their vaginalis field in the last few years. names and affiliations. Keywords Trichomonas vaginalis , trichomoniasis , non-viral sexually transmitted 1 Robert Hirt, Newcastle University, Newcastle disease , hydrogenosome , metronidazole upon Tyne, UK 2 Sven B. Gould, Heinrich-Heine-University Düsseldorf, Düsseldorf, Germany Any comments on the article can be found at the end of the article. Corresponding author: David Leitsch ([email protected]) Competing interests: The authors declare that they have no competing interests. Grant information: This work was supported by project J3492 of the Austrian science fund (FWF). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Copyright: © 2016 Leitsch D. This is an open access article distributed under the terms of the Creative Commons Attribution Licence, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. How to cite this article: Leitsch D. Recent Advances in the Trichomonas vaginalis Field [version 1; peer review: 2 approved] F1000Research 2016, 5(F1000 Faculty Rev):162 (https://doi.org/10.12688/f1000research.7594.1) First published: 11 Feb 2016, 5(F1000 Faculty Rev):162 (https://doi.org/10.12688/f1000research.7594.1) Page 1 of 7 F1000Research 2016, 5(F1000 Faculty Rev):162 Last updated: 17 JUL 2019 Introduction much lower, with 40 to 50 per 1000 men and women7. In total, Trichomonas vaginalis (Tv) is a globally occurring anaerobic/ 276 million infections with Tv are believed to occur worldwide microaerophilic protist parasite which colonizes the epithelium of and per annum1. These numbers are very high indeed, but it is the human urogenital tract. Although often asymptomatic, Tv infec- estimated that most Tv infections (up to 80%) are asymptomatic8. tions can cause inflammation in the cervix, the vagina, and the ure- Importantly, men are infected equally frequently, but 89% of tricho- thra. Based on estimates of the World Health Organization (WHO) moniasis cases are actually diagnosed in women because of their from 20081, trichomoniasis constitutes the most prevalent non-viral higher incidence of symptoms, which are sometimes severe and sexually transmitted disease (STD) worldwide, affecting more than debilitating. The main concern about Tv infections, however, is their 276 million people every year. Women and men are infected with predisposing nature for other diseases or sequelae2, and a number comparable frequency, but in men symptoms are normally mild and of new studies give justification to this concern. For example, Tv infections are cleared by the host’s immune system within weeks. was found to be associated with human papilloma virus infec- In women, however, Tv infections can persist for many years, and tions and cervical cytological abnormalities9. Moreover, in a meta- symptoms, mainly pruritus caused by inflammation and odorous analytical study, strong statistical evidence was presented for an vaginal discharge, can attain a severity which is debilitating. As association of an underlying Tv infection and preterm birth10. Most trichomoniasis is not a life-threatening disease, it was often belit- importantly, however, evidence for a predisposition for infection tled as a “nuisance infection” in the past. A large number of studies with HIV in Tv-infected individuals is mounting. In a meta-study from the last 20 years or so, however, have shown that underlying on 31 studies, it was concluded that the risk of HIV acquisition Tv infections increase the risk of adverse pregnancy outcomes and is increased 2- to 3-fold in Tv carriers11, and it was found that Tv contagion with HIV virus2. Given the fact that HIV and Tv are often infection increased the risk of HIV infection 2.5-fold in macaques, jointly epidemic in many parts of the world, this is an alarming find- which serve as a non-human primate model. Accordingly, it was ing. Moreover, a higher risk of developing prostate cancer due to Tv calculated that annual screening for Tv would save US$553 per infection has been suggested2. woman and lifetime in the prevention of new HIV infections to sus- ceptible male partners in the United States alone12. Before the development of the 5-nitroimidazole drug metroni- dazole in 1960, trichomoniasis was notoriously difficult to treat. In order to get a picture of the genetic diversity of Tv, a large-scale Nowadays, most patients can be successfully treated with metroni- study13 was conducted, subjecting 235 Tv isolates, collected from dazole or another more effective 5-nitroimidazole derivative, tini- all around the globe, to microsatellite genotyping14. Intriguingly, Tv dazole. However, resistance to 5-nitroimidazoles does occur and was found to group into two distinct clusters or “types”. Both types seems to be on the rise3. In addition, allergic reactions to nitroimi- occur worldwide with comparable frequency, although type 1 is dazoles have been reported and side effects of nitroimidazole presumably the older clade13. Interestingly, the presence of Tv virus treatment can be disturbing. (TVV) is unequally distributed within the two types, with about 70% of all type 1 isolates, but only 30% of type 2 isolates, carrying Apart from its role as a pathogen, Tv has attracted great interest the virus. Conversely, metronidazole resistance is far more preva- from geneticists, biochemists, and evolutionary biologists after the lent in type 2 isolates. discovery of the hydrogenosome4, a mitochondrion-like organelle which generates hydrogen. Due to its microaerophilic lifestyle, Tv Treatment does not have the ability to generate ATP by oxidative phosphoryla- Since 1960, metronidazole and other 5-nitroimidazoles, such as tion but depends on substrate-level phosphorylation. Originally, it tinidazole, have been the mainstay of Tv treatment3. 5-nitroimida- was assumed that the hydrogenosome is an ancestral form of the zoles have been reported to damage DNA, form adducts with pro- mitochondrion5, which kindled interest in Tv as an assumed archaic teins (partly with inhibiting consequences15), and cause oxidative eukaryote. This notion, however, has been thoroughly revised after damage in the cell by depleting thiol pools15. 5-nitroimidazoles are publication of the Tv genome in 20076. It is now apparent, based on in fact prodrugs, which have to be reduced at their nitro groups in phylogenetic studies, that the hydrogenosome constitutes a reduced order to become toxic. This reaction, however, takes place quan- form of fully developed mitochondria. Nevertheless, from the evo- titatively only in microaerophilic/anaerobic organisms and has lutionary and cell biologist’s point of view, the hydrogenosome been suggested to be catalyzed by several enzymes and factors remains an intriguing organelle, and the extraordinary size of the Tv such as ferredoxin16, nitroreductase17, and thioredoxin reductase15. genome, exceeding 160 Mb, will certainly provoke further research Resistance to 5-nitroimidazoles in clinical Tv isolates does occur, in the years to come. sometimes leading to extended and complex treatment regimens18. Clinical metronidazole resistance is based on decreased oxy- In this review, I will give a brief but comprehensive overview of the gen scavenging in the cell, leading to higher intracellular oxygen advances in the research on Tv from the last 5 years or so, spanning concentrations19. Accordingly, expression of flavin reductase 1, an from the epidemiology to the infection biology, treatment, and cell enzyme that uses flavin mononucleotide (FMN)
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