Drug Target for Pathogenic Amoebae

Drug Target for Pathogenic Amoebae Horizontal Gene Transfer of a Chlamydial tRNA-Guanine Transglycosylase Gene to Specific Algal and Protozoan Lineages: A Putative Drug Target for Pathogenic Amoebae Abstract: tRNA-guanine transglycosylases are found in all domains of life and mediate the base exchange of guanine with queuine in the anticodon loop of specific tRNAs. They are also known to regulate virulence in bacteria such as Shigella flexneri, which has prompted the development of drugs that inhibit the function of these enzymes. Here we report a group of tRNA-guanine transglycosylases in eukaryotes (algae and protozoa) which are more similar to their bacterial counterparts than previously characterized eukaryotic tRNA-guanine transglycosylases. In silico analysis of these bacterial-like tRNA-guanine transglycosylasesrevealed thatthe majority are predicted to be targeted to mitochondria, although some are likely to localize to chloroplasts, the secretory pathway or the cytosol. We provide evidence demonstrating that the gene encoding theseenzymes was acquired by these eukaryotic lineages via horizontal gene transfer which from the Chlamydiae. Given that the S. flexneri tRNA-guanine trans glycosylase can be targeted by drugs, we propose that the bacterial-like tRNA-guanine transglycosylases could potentiallybe targeted in a similar fashion in pathogenic amoebae that possess these enzymes such as Acanthamoeba castellanii. Keywords: mitochondria, tRNA-guanine transglycosylase, queuine tRNA-ribosyltransferase, horizontal gene transfer, tRNA, queuosine, Chlamydiae Abbreviations: TGTase: tRNA-guanine transglycosylase E-TGTase: Eukaryotic tRNA-guanine transglycosylase B-TGTase: Bacterial tRNA-guanine transglycosylase BL-TGTase: Bacterial-like tRNA-guanine transglycosylase HGT: Horizontal gene transfer Introduction Base modification of tRNAshas been implicated in tRNA structure, aminoacyl tRNA synthetase interaction andinfluencing codon-anticodon basepairing[1]. The function of the modification will depend on itstype and the position of the modified base. For example, most bases that are modified within the anticodon loop (positions 34-36) of tRNAsare important for accurate translation by facilitating interactions with their cognate codons in mRNAs [1]. One such modification that influences codon-anticodon basepairingis the incorporation of queuine within the anticodon loop. Queuosine is a modified guanosine analogue found in tRNAs from all three domains of life.Despite its wide phylogenetic distribution, queuosine is only found in a select group of tRNAs (tRNAHis, tRNAAsp, tRNATyr and tRNAAsn) [2].Reduced incorporation of queuosine in these tRNAs alters their codon recognition ability and has been linked to various cancers [3,4]. tRNA-guanine transglycosylases Queuosine modification of tRNA is mediated by tRNA-guanine transglycosylases (TGTases)(also known as queuine tRNA-ribosyltransferases). TGTases catalyze this modification via base exchange where the guanine at position 34 of the tRNA is post-transcriptionally removed and substituted with queuine or a queuine precursor [5].Eukaryotes are not capable of de novo queuine synthesis but acquire it through diet or their gastrointestinal microbiota [6].After its acquisition, the eukaryotic TGTase (E-TGTase) mediates the replacement of guanine with queuine in the anticodon loop. In contrast, queuosine modification of bacterial tRNA is more complex. Prokaryotesuse GTP-cyclohydrolase-like enzymes tosynthesizea queuine precursor(e.g. preQ1) from GTP. The bacterial TGTase (B-TGTase) then mediates the base exchange with guanine to incorporate preQ1, unlike E-TGTases that use queuine itself as the substrate.This incorporatedpreQ1 is then modified by S-adenosylmethionine tRNA ribosyltransferase to e poxyQ, which is further modified to form queuosine [6].In addition to tRNA modification, B-TGTasesplay a role in regulating the expression of bacterial genes.TGTase mutants (vacC) in the bacterium Shigella flexneri exhibit reduced expression of the virG and ipaBCDgenes, which encode virulence factors that facilitate the spread and invasion of the pathogen [7]. This is a result ofthe VacCTGTase beingrequired to modify a single base in virF mRNA, which encodes the transcriptional activator ofvirG and ipaBCD[8].Thus, B-TGTases can modify substrates otherthan tRNA and are important mediators of bacterial virulence. As a result, B-TGTases have served as a targetfor the development of drugs that interfere with their function [9].Here we report a new group of TGTases in eukaryotes that display significantly greater similarity to B-TGTases than E-TGTases. We hereby refer to these proteins as bacterial-like TGTases (BL-TGTases).In silico analysis identified 25 BL-TGTases in distinct protozoa n and algal lineages and the reason for their similarity to B-TGTases is explored in this article. Variation in the subcellular localization of bacterial-like tRNA-guanine transglycosylases To investigate the putative subcellular localization of BL-TGTases, three bioinformatic programs were utilized: Mitoprot [10], Predotar [11] and Target P [12]. The putativelocalizationfor each BL-TGTase was supportedby predictions from at least two of the three programs.Most BL-TGTases possess N-terminal mitochondrial targeting signals (Table 1), suggesting a role in modification of mitochondrial tRNAs.Interestingly, the BL-TGTases from Ostreococcus lucimarinus and Chondrus crispus were predicted to localize to mitochondria with one program (Predotar) but to the plastid with another (Target P). While it is possible that these two proteins may localize to both organelles, further experimentation is required to elucidate their subcellular locations. The BL-TGTase from the diatom Phaeodactylum tricornutum was predicted to localize to the endoplasmic reticulum (ER) of the secretory pathway, indicating it maymodify other substrates in this organelle. Bacterial-like tRNA-guanine transglycosylase genes originated from a Chlamydial gene acquired via horizontal gene transfer While the localization of BL-TGTases varied, all 25 of the proteins exhibited higher levels of amino acid similarity to B-TGTases despite their existence in eukaryotes. A Bayesian analysis of phylogeny using MrBayes [13] withBL-TGTases,B-TGTases and E-TGTasesconfirmedthis similarity(Figure 1).The BL-TGTases were most similar to TGTases from members of the Chlamydiae.In fact, the Chlamydial TGTases were more similar to BL-TGTases than other B-TGTases. Given that Chlamydiaeare bacteria, the topology of the tree in the present study is incongruent with the universal tree of life. Instead, this topology is consistent with a horizontal gene transfer (HGT) event. That is, the genes encoding BL-TGTases originated from a Chlamydial TGTase-encoding gene that was acquired via prokaryote-to-eukaryote HGT. In addition to the strong statistical support for the BL-TGTase-Chlamydial TGTase sister group, there are several other factors that support this notion. The Chlamydiaeare known to be major contributors of genes to several eukaryotic genomes via HGT [14,15]. This includes genes encoding tRNA modification enzymes such as the Chlamydial tRNA guanine methyltransferases found in protozoa, diatoms and algae[16,17] and Chlamydial tRNA genes in vascular plants [18].Similarly to the present study, sister groups were observed between the Chlamydial and the horizontally acquired eukaryotic genes in these cases. Lastly, the majority of eukaryotic lineages in which we identified BL-TGTases have previouslybeen reported to possess HGT-derived genes acquired from the Chlamydiae[16,19]. Thus, the notion that BL-TGTases resulted from the acquisition of a B-TGTase from the Chlamydiaevia HGT in eukaryotes is highlyplausible. Indirect acquisition of a Chlamydial tRNA-guanine transglycosylase in protozoa via anon-Chlamydialbacterial intermediate Interestingly, a B-TGTase sequence from the ÃŽ ´-proteobacterium ‘Candidatus Babela massiliensis’clustered with the BL-TGTases of protistsrather than the B-TGTases (Figure 1). Although the protozoan BL-TGTases displayed similarity to Chlamydial B-TGTases, the possibility of a HGT event from ‘Ca. B. massiliensis’to protistswas present. SinceChlamydiae and ÃŽ ´-proteobacteria are not closely related, the phylogeny of their B-TGTases was investigated. Interestingly, the ‘Ca. B. massiliensis’TGTase clustered with the Chlamydial TGTase clade rather than other ÃŽ ´-proteobacterial (Pelobacter, Geobacter, Myxococcus, Desulfobulbus) B-TGTases (Figure 2). ‘Ca.B. massiliensis’and members of the Chlamydiae are found as obligate intracellular symbionts of protists such as Acanthamoeba, Dictyosteliumand Naegleria [20,21]. The presence of both of these bacteria within the one eukaryotic cell would provide the ideal conditions for HGT between them. Therefore, it is likely that at least two independent HGT events have occurred:1) The Chlamydiae donated a TGTase-encoding gene to an ancestral ‘Ca.B. massiliensis’species; and 2) ‘Ca.B. massiliensis’then donated this gene to theAmoebozoa and Heterolobosea.How the BL-TGTase genes were acquired in the algal lineagesremains to be elucidated, but may have occurred via additional HGT events (either prokaryote-to-eukaryote or eukaryote-to-eukaryote). Bacterial-like tRNA-guanine transglycosylases as drug targets for pathogenic amoebae In addition to their role in queuosine modification of tRNAs, TGTases are important for S. flexneri virulence [7,8].As a result, studies have focused on the development of TGTase inhibitors that specifically target the S. flexneri B-TGTase to treat shigellosis,while the E-TGTases of the human host remain unaffected. Some of these inhibitors,such as lin-benzoguanine,function by occupying the binding site for preQ1[22,23].While most eukaryotic species that possess BL-TGTases are non-pathogenic, we identified a BL-TGTase in Acanthamoeba castellanii, the causative agent of amoebic keratitis and encephalitis.Naegleria gruberi, whichalso has a BL-TGTase, is non-pathogenic, but is closely related to Naegleria fowleri, the etiologic agent of primary amoebic meningoencephalitis, which may possess an unidentified BL-TGTase. Given the development of B-TGTase inhibitors has already been achieved, the BL-TGTasesin pathogenic eukaryotes could also potentially be targeted with the same drugs. Alter natively, new inhibitors could be developed following resolution of the BL-TGTase crystal structure. To confirm BL-TGTases as a putative drug target future research should attempt to characterize these proteins and determine if they have retained their prokaryotic functions and mechanism of action. Concluding remarks In this report, we have described a group of TGTases in algae and protozoa (BL-TGTases). Theseproteins are predicted to localize to various subcellular locations including mitochondria, chloroplasts, the ER and the cytosol, depending on the organism. Lastly, we showed that via multiple HGT events, BL-TGTases were originallyfrom bacteria of the Chlamydiae lineage. The bacterial origin of these proteins could be exploited in the development of drugs similar to those synthesized for the S. flexneri B-TGTase. Research into the identification and synthesis of BL-TGTase inhibitors may provide a novel treatment for infectious diseases which are caused by pathogenic amoebae that possess these proteins.

Coca-Cola India Case Study Analysis

Coca – India Case study Lakeesha Customer Inserts His/her Name Customer Inserts Grade Course Customer Inserts Tutor’s Name April 11, 2013. Introduction Coca-Cola is the leading brand of the world in soft drink category. It was founded by a pharmacist in Georgia known by the name of John Pemberton. He used to sell a potion for physical as well as mental disorders. However, the potion soon turned into world’s largest selling brand after carbonated water in addition to fountain water with other constituents were added to the brand.Furthermore, soon after the addition of carbonated water, Coca-Cola brand was given the honor of 1887 and in just year of two years, the brand began to spread its sales all over the world. However, the brand got under the wave of ten billion cases in the past decades, which made its growth come to sudden plunge and with lesser improvement in the sales. During the era of 2000, the brand had per annual sale growth of 0. 2 %, which is a deep loss for the company as back in 1980s, its sales growth, had been 5-7%.Furthermore, as it began its sales in India, the Coca-Cola began to see its rise again since the brand became a favorite of the one billion population of the country, which comprised of every rural or urban part of the community. Ethical Issues Many of the most important cases were filed against the Coca-Cola Company in the last few years. They have been the result of an allegation imposed on the company by its four employees in spring 1999. According to the lawsuit, the company has used discriminatory acts on its African-American employees just simply on the basis of their being black. Such diverse types of warning were being issued on the company for its discriminatory practices on the employees. Moreover, many of the employees there have been reported to experience an unethical discriminatory culture in accordance with their race and case. Additionally, another major case was filed against the company on 8 Jun e 1999 (Kochan, 1997). This case comprised of thirty-three children enrolled in Belgium school, which fell ill and were hospitalized.Then again, after few more Belgians went into the trap of this illness. The bottles had been tinned up in a Coco- Cola plant of Dunkirk, France. Moreover, the crisis intensified later after the quantity of people who were stricken with intestinal problems increased from eight to 250, all over the country. However, on June 29, a report was issued which claimed according to research by a toxicologist that the blame was on the carbon dioxide impurities found in Antwerp while on the other hand, there was fungicide in Dunkirk.In order to fix the crisis, many resolutions were passed by the company, which started free selling campaigns in the countries sending free vouchers. There beach parties, dancers were hired for concerts and advertisements were issued on televisions thanking the public for being loyal to the company. Hence, around 5. 5 million people we re influenced by the company to get its fame back. After this, a major crisis had to be dealt with by the company.On February 4, 2003, the Centre for Science and Environment filed a case against Coca- Cola (Analysis of Pesticide Residues in Soft Drinks, 2003). The CSE was led by a group of activists that claimed about the Kinley Bottled water to contain traces of pesticides. It comprised of ingredients like DDT and Malathion, which are highly toxic and can result in the destruction of human bodies. Let humans apart, these substances can even damage aquatic animal and plant life when disposed of in water bodies.These tests were taken from a sample of seventeen bottles that had been sold lately. So after the test was being conducted by Pollution Monitoring Laboratory, the results showed that these Coca-Cola products contained more than 30- 36 times of such toxic substance which was surpassing the global standards approved all over the world. These pesticides, according to research cou ld result in cancer, birth defects, and even damage to nervous as well as immune system of the body. Hence, it was proved that it was unethical to sell products, which could result in loss of human lives.The EEC standard was 0. 005 mg/l of pesticides while in case of Coca-Cola it was 0,018 mg/l, which was thirty six times more (Pendergrast, 1993). In order to resolve issue, the company could show results of its own samples so that employment of many of its workers could be saved. Most important ethical issue that could be assumed from the above mention is Gupta’s dilemma which showed with evidence that Coca- Cola had surpassed the limits of its pesticides concentration to more than forty five times. Alternative ResolutionsSome of the resolutions that the company could make was firstly to marketplace. This was meant that to use such type of substances in its products that were approved by the standards of market. This will build up a strong foothold of the company in the world of market. Moreover, the billions of stakeholders that work in the company could lose their employment if the company continued with unethical acts. This will eventually attract more investors to contribute to the business as a result of which more Coca- Cola brands could be opened.Moreover, an ethical and proper cultural environment should be given to the employees so that more talented people could come and give their services to the company. Hence, it can benefit themselves as well as the company. In addition, Coca-Cola Company can use its resources in the development of community by contributing in environmentally sustainable practices. Hence, this will result in a better profit making and market value of the company. Best Ethical ResolutionIn my opinion, by following the given standards approved by the market, Coca-Cola can get its market value back. This will eventually result in higher employment and lives of many people would be saved if the substances used were not injurio us to health. References: † Analysis of Pesticide Residues in Soft Drinks. † Centre for Science and Environment (CSE (2003). Kochan, Nicholas,. The World’s Greatest Brands: an International Review by Interbrand. New York University Press,, 1997. Pendergrast, Mark. For God, Country and Coca-Cola. Charles Scribners, 1993.

An Impartial View of College Essay Examples

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The Issue Of Gun Control Essay - 2091 Words

President Obama said at a press conference held in response to another mass shooting in Oregon on October 01, 2015 â€Å"The solution to such violence is obvious. It cannot be this easy for somebody who wants to inflict harm on other people to get his or her hands on a gun† (Simple Minded Gun Control). Despite how obvious the president said the solution is, the fact remains that gun control is still a controversial issue today. The reason why this has attracted so much attention is because not everyone is in favor of gun control and each side brings up excellent points about the issue. Research related to this issue strongly supports the claim that there SHOULD be more gun control laws. The first task I will complete in this research paper is clearing up any misconceptions about Gun Control and all the terminologies I will use. Followed by my three arguments that prove this position which are (1) Incidents like Sandy Hook or Oregon will be less likely to occur (2) It reduces t he high rates of accidental deaths and (3) As the years pass by and technology updates the laws should be up to date as well. Members from the National Rifles Association state that No law-abiding American should be forced to face evil with empty hands. I say I agree with President Obama there SHOULD be more gun control laws because it should not be as easy as it is for someone who wants to inflict harm on others to get their hands on a weapon. Allow me to first clear up anyShow MoreRelatedGun Issue And Gun Control1401 Words   |  6 PagesUniversity, Gun Politics has been a course I have aspired to take. While many enroll in such a course looking for an escape from the â€Å"collegiate liberal echo chamber† or as an outlet for their conservative agendas, I saw the class as an â€Å"entrance to the dark side.† My views on guns prior to the class were, I would call, polarized yet uninformed. In most of my discussions, I would cite the Australian 1996 Na tional Firearms Agreement as precedent for how American politicians should approach the gun issueRead MoreThe Issue Of Gun Control Essay1646 Words   |  7 Pagessomebody who wants to inflict harm on other people to get his or her hands on a gun.† (Simple Minded Gun Control). Gun control is a controversial issue worldwide. The reason why this has attracted so much attention is because not everyone is in favor of gun control and each side brings up excellent points about the issue. Research related to this issue strongly supports the claim that there SHOULD be more gun control laws. Three arguments that prove this position are (1) Incidents like Sandy HookRead MoreThe Issue Of Gun Control1489 Words   |  6 Pagessplit on the issue of gun control. We have seen many violent shootings and innocent people dying because of gun violence. Some Americans believe we need strict gun laws to protect our children and ourselves from these horrific tragedies. Other Americans believes it i s our rights as Americans to posses’ guns and we are entitled to that right in our constitution. There are also some Americans that are stuck in the middle and can see both sides but recent events have definitely caused this issue to be inRead MoreThe Issue Of Gun Control1031 Words   |  5 Pagesintegrate them in a way that will trigger a new viewpoint on the subject. Gun control is something that should be enforced across the world in order to save lives. In the United States there is a ratio of 88.8 guns per one hundred persons (GunPolicy.org). Those numbers award us with the highest total per capita number in the world. With that amount of firepower there needs to be an efficient way to regulate who exactly can purchase a gun. Since 1791 when the second amendment was ratified there have beenRead MoreThe Issue Of Gun Control1705 Words   |  7 Pagesaway without preface or reason. All someone needs to do is pull the trigger. In today’s world, guns are far too acce ssible to the people of society. We hear in the news, stories of mass shootings, homicides, and suicides; most of which are caused by the activation of a gun. A hot-button issue, gun control is one of the most debated topics in American politics. Should we, or should we not, be able to own guns? Although it infringes on the Second Amendment that provides the right to bear arms, this amendmentRead MoreThe Issue Of Gun Control951 Words   |  4 PagesGun control has been a big topic for the past decade in the united states. These debates will rise and fall time in and time out after something horrific happens in the state. Anti-Gun supporters do not realize that it is extremely difficult to regulate something in the states that is a big portion of our economy.Would stricter gun laws change anything? So far statistically It has been proven otherwise one must consider how a citizen would defend themselves when they are faced with terror. How willRead MoreThe Issue Of Gun Control929 Wor ds   |  4 PagesGun Control Getting arrested yesterday was not the highlight of my week. I was hungry; so I went to Wal-Mart get some charcoal, lighter fluid, and steak. I put the charcoal in the grill and the lighter fluid on the coals. I got a little lighter fluid on my arm, but I didn’t pay it much attention. I lit the match and threw it on the coals. I looked down and my sleeve was on fire. I was waving my arm in the air, trying to put it out. Then here they come the police pulled in and tackled me to theRead MoreThe Issue Of Gun Control1552 Words   |  7 PagesThe issue of gun is always controversial. Firearm can be used as defending or an assaulting weapon. In United States, firearm increase the rate of homicide, suicide and gun violence, which can harm and murder people. Moreover, taking away people’s gun will not work effectively because the murder and criminal will find another ways to get guns. Also, the black market will appear for the people who cannot get guns from regulated market s. In addition, if regulated guns are banned, murderers may useRead MoreThe Issue Of Gun Control1678 Words   |  7 PagesJanuary 2014 The 2nd Amendment Over the past few years, the issue of gun control has been widely discussed. You surely have heard the phrase, â€Å"Guns don’t kill people, people kill people† uttered and i wholeheartedly support this statement. It is important to treat guns responsibly so they do not end up in the wrong hands. I believe gun control violates our inalienable rights. Another piece of our liberty will surely be taken away if guns are banned, and socialism and totalitarianism will be rightRead MoreThe Issue Of Gun Control1208 Words   |  5 PagesWill Christopherson Traverson English 2 1 March 2017 Gun Control The United States has 88.8 guns per 100 people, or about 270,000,000 guns, which is the highest total and per capita number across the globe. The current public gun control debate in the United States seems to be placed on standby until it is sparked up by a major mass shooting. There were at least 126 mass shootings between January 2000 and July 2014.(pro). Opponents of more gun laws accuse supporters of using a horrific event to further