Peptidylarginine Deiminase and Neurodegenerative Diseases

Peptidylarginine Deiminase and Neurodegenerative Diseases Abbreviations AD: Alzheimers disease CNS: Central nervous system GFAP: Glial fibrillary acidic protein NFT: Neurofibrillary tangles MAP2: Microtubule-associated protein 2 MBP: Myelin basic protein Nef3: Neurofilament 3 PAD: Peptidylarginine deiminase RT-PCR: Reverse transcriptase-polymerase chain reaction SP: Senile plaque Introduction Neurodegenerative diseases, such as Alzheimers disease and Multiple Sclerosis, are a group of progressive conditions, which affect a persons learning abilities and the day-to-day routine management. Citrullination/deiminasion is a process describing the enzymatic involvement in conversion of arginine residues into citrulline in target proteins. Here, is a review, entailing the main features of the peptidylarginine deiminase (PAD) enzyme family, with a view on its history, presence and future in neurodegenerative disorders. Background The amino acid Citrulline was firstly mentioned in 1955 (Smith and Young), and the citrullination process of L-arginine conversion to L-citrulline was firstly described in 1958 (Rogers and Simmonds).Citrullination is a posttranslational process, and also results in a 1Da decrease in size, due to an hydrolytic reaction (Vossenaar et al., 2003; Gyorgy et al., 2006; van Venrooji and Pruijn, 2003). However, it was only in 1977 when peptidylarginine deiminases, the enzymes that are responsible to the citrulline modification, were firstly described (Rogers et al.). Alzheimers disease: GFAP, an astrocyte-specific marker protein, is involved in the progression of Alzheimers disease via PAD2 citrullination. The increase of PAD2 can be seen in the hippocampi of AD patients. GFAP and vimentin citrullination was also identified by using a 2D gel electrophoresis and MALDI-TOF mass spectrometry. Activation of PAD occurs upon a mass influx of Ca2+, and therefore an increase in citrullinated proteins (Keller et al., 2000; Maccioni et al., 2001). The PAD family consists of five known members (1-4 6), each is present in a different area in the human body. PAD2 and PAD4 area of activity is the central nervous system (CNS), although PAD2 in larger amounts than PAD4 (Kubilus and Baden, 1983; Watanabe et al., 1988; Terakawa et al., 1991). Yet, the role of the different PADs is still largely understood. PAD3 was found in the root sheaths of hair follicles, as part of the citrullination process of keratinisation (Kanno et al., 2000). PAD4 was found in human myeloid leukemia HL-60 cells, and later in peripheral blood granulocytes (Nakashima et al., 1999; Asaga et al., 2001). PAD2 and PAD4 are also present in the myelin sheath. In fact, hyper-citrullination of myelin basic protein (MBP) is now known to result in myelin sheath loss in multiple sclerosis patients (Moscarello et al., 1994; Wood et al., 2008; Musse et al., 2008). Moreover, PAD4 is also involved in histone deimination in brain tissues of such patients (Mastronardi et al., 2006). Following immunocytochemical studies, PAD2 have been detected in glial cells and astrocytes (Asaga and Ishigami, 2000; Vincent et al., 1992; Asaga and Ishigami, 2001), microglial cells (Vincent et al., 1992; Asaga et al, 2002), oligodendrocytes (Akiyama et al., 1999), and Schwann cells (Keilhoff et al., 2008). Furthermore, during hypoxic insult (Asaga and Ishigami, 2000) and during kainic acid administration (Asaga and Ishigami, 2001; Asaga et al., 2002), neurodegenerative regions shown to have PAD2 activity and deimination of various proteins. These findings suggest that PAD2 plays an important role in protein citrullination in neurodegeneration.  Senile plaques (SPs) and neurofibrillary tangles (NFTs) are the two main forms of protein aggregation, and therefore responsible for neuronal death in AD (Katzman, 1986; Smith, 1998). They are mainly found in the hippocampus and cerebral cortex, when levels of PAD2 were detected to be more than threefold higher in the hippocampus than in the cortex of rat brains (Asaga and Ishigami, 2000). A study by Ishigami et al. (2005) was set to evaluate the involvement of protein citrullination in AD patients brains, and identified two citrullinated proteins- GFAP and vimentin, by using a two-dimensional gel electrophoresis and MALDI-TOF mass spectrometry. It went on to identify also a citrullinated MBP, again in the hippocampus region of the AD patients. Multiple sclerosis: Higher rates of citrullinated MBP were found in the CNS of MS patients than in the CNS of healthy adults (Mastronardi et al., 2006; Nicholas and Whitaker, 2002; Raijmakers et al., 2005). However, similar levels of citrullination have been observed in children under the age of 4. An indication that citrullination is involved in the development of mature myelin (Moscarello et al., 1994; Wood et al., 1996). Due to known increased citrullination, PAD2 and PAD4 rates in myelin from normal-appearing white matter (NAWM) of seven MS patients were examined, and were found to be significantly higher compared to NAWM from six control individuals, using immunoblots with PAD2 and PAD4-specific antibodies (Wood et al., 2008). The anti-citrulline antibody- F95, was also used in the same manner, resulting in the same outcome (Nicholas et al., 2004). The study by Wood et al. (2008) also discovered that PAD2 can citrullinate 18 out of 19 arginine residues, whereas PAD4 can only citrullinate 15 out of 19 arginine residues localised in MBP. It has been found and reported that MPB deimination increases its susceptibility to degradation by proteinases, which are elevated around active plaques and in the cerebrospinal fluid (CSF) (Cuzner and Davison, 1973; Einstein et al., 1972; Richards and Cuzner, 1978). It has been also reported that macrophages and reactive astrocytes are able to produce the proteinase cathepsin D (Allen and McKeown, 1979; Prineas and Wright, 1978). Cathepsin D then produces peptides that contain the immuno-dominant epitopes of MBP, located in the CSF of MS patients (Whitaker, 1977; Whitaker and Granum, 1980). It was then discovered that different MBP strains, containing greater amounts of citrulline per mole of MBP than their counterparts were digested at a much faster rate by cathepsin D (Pritzker et al., 2000). An explanation for this discovery by Pritzker et al. can be explained by the three-dimensional atomic structure of the human MBP they created. The stru cture showed a clear correlation between the open conformation of the atomic structure and the higher rate of citrullination in-site, which allows cathepsin D a better access to Phe-Phe linkages in MBP, and so an increased digestion of citrullinated MBP. Current perspectives In addition to MBP, GFAP also have shown to be citrullinated in MS. In a study by Nicholas et al. (2004), GFAP citrullination rate in both the NAWM and lesions of patients with SPMS was compared with the white matter of control brain tissue, and was found to be in higher amounts. The findings were obtained using GFAP anti-citrulline antibodies (such as P95) and confocal microscopy. On an extended study by Nicholas et al. (2004), using dual staining wit GFAP anti-citrulline antibodies, it was found that lesions taken from nine MS patients were highly citrullinated compared to white matter taken from six healthy control individuals. Western blotting has also confirmed that the majority of the deimination occurred in GFAP. PAD4 is enable to translocate from the cytoplasm to the nucleus, due to its ability to carry a functional nuclear localisation signal (Nakashima et al., 2002). PAD4 modification can also affect nuclear proteins, histones H2A, H3 and H4, and nucleophosmin/B23 (Hagiwara et al. 2002).   Deimination of histones causes change in chromatin confirmation, and therefore gene transcription gets turned off (Shimoyama et al., 2010). In a study by Cuthbert et al. PAD4 was found to inhibit transcription of estrogen-responsive genes in MCF-7 cells. Levels of PAD4 in MCF-7 cells were shown to rise in response to estrogen, followed by deimination of the pS2 promoter by PAD4, which coincides with downregulation of this gene (Cuthbert et al. 2004). Wang et al. showed that incubation of purified substrates H3 and H4 with PAD4 in vitro generated citrullinated H3 and H4, which coincided with a dramatic reduction in H3 Arg 17 and H4 Arg 3 methylation (Wang et al. 2004). This effect was also mimicked when HL-60 granulocytes were incubated with PAD4 in vivo, in the presence of calcium ionophore. PAD4 was found to be elevated in NAWM from 17 patients with MS, compared to 9 control subjects (Mastronardi et al. 2006). This was shown by fractionation of these samples into membrane-containing, non-microsomal, and nuclear fractions, followed by quantitation of the amount of PAD 1-4 antibody binding. The nuclear fraction contained a 3.5-fold increase in the level of PAD4 in patients with MS compared to controls. Through western blot analysis using anti-PAD4 antibody, this was attributable to increased PAD4 in the MS NAWM. Using an antibody against citrullinated proteins, this increase in PAD4 was found to be accompanied by an increase in citrullinated proteins in brain tissue taken from patients with MS, whereby strong nuclear labeling in NAWM from MS patients was seen compared to controls (Mastronardi et al. 2006). This increase in PAD4 was also accompanied by an increase in nuclear histone H3 citrullination, as shown by immunostaining of MS and control tissue with an antibody against citrullinated protein, which revealed strong nuclear staining of cells in the MS white matter. These findings were confirmed by western blot analysis, which showed a gre at abundance of citrullinated H3 in MS NAWM, with only traces in white matter from controls. This citrullination of histones greatly affects the chromatin structure and function, as deimination of arginine residues of histones decreases their positive charge, which compromises its ability to interact with DNA and possibly resulting in apoptosis of affected cells (Moscarello et al. 2007; Wang et al. 2004). It is not known whether excess citrullination is a primary or a secondary event to the inflammatory process in MS or whether the regulation of PAD isoforms may be part of the genetic susceptibility to MS. Single-nucleotide polymorphisms in the PAD4 gene, also associated with the autoimmune disease rheumatoid arthritis, increase mRNA stability, suggesting that this could result in greater PAD4 protein expression and hence increased citrullination of proteins (Suzuki et al. 2003). Increased PAD2 protein expression in human astrocytes in vitro has also been reported in response to inc reased intracellular calcium levels when cells were subjected to elevated pressure or in response to hypoxia (Bhattacharya et al. 2006a, b; Sambandam et al. 2004). A number of pathological processes, including excitotoxicity, occur in the CNS of patients with MS, which would lead to raised intracellular calcium ions in neurons and glia (Shideman et al. 2006; Smith 2007). Large numbers of activated macrophages are present in inflammatory demyelinating sites within MS lesions. Since these cells contain PAD enzymes and there is increased cell death due to raised intracellular calcium ions, this would lead to activation of PAD enzymes when released from dying cells (Bhattacharya et al. 2006a). Thus myelin proteins may be citrullinated both intracellularly, during myelin degradation following phagocytosis, as well as extracellularly, following release of PAD enzymes from dying cells. In addition, significant hypomethylation of the PAD2 promoter has also been found to occur in MS NAWM compared to controls, which may lead to increased PAD2 expression and subsequent increase in citrullination, as hypomethylation leads to increased gene transcription (M astronardi et al. 2007). Future perspectives Conclusion In the last 60 years since it was firstly mentioned, the research area of PADs have made a tremendous leap and came about with many important discoveries regarding the mechanisms and pathologies concerning PADs. However, it still seems to be largely unknown and have many possible routes of research. Especially when considering the wide localisation of the PADs throughout the human body. Such areas of research may ask after the origins of the calcium influx to the brain tissues, and the timing of that physiological event; how may the rate of calcium influx affect the rate of neurodegeneration; what would prove to be the most effective PAD-inhibitor treatment, etc. However, looking at the milestones along the research of PADs, it does seem that the right questions are being asked, and that the available/emerging technologies are suitable for this research. References Smith, D. G. and Young, E. G. J. Biol. Chem., 217, 845 (1955). Rogers GE, Simmonds DH. Content of citrulline and other amino acids in a protein of hair follicles. Nature. 1958; 182(4629):186-7. [PubMed: 13566234] Vossenaar ER, Zendman AJ, van Venrooij WJ, Pruijn GJ. PAD, a growing family of citrullinating enzymes: genes, features and involvement in disease. Bioessays. 2003; 25(11):1106-18. [PubMed: 14579251] Gyorgy B, Tà ³th E, Tarcsa E, Falus A, Buzà ¡s EI. Citrullination: a posttranslational modification in health and disease. Int J Biochem Cell Biol. 2006; 38(10):1662-77. [PubMed: 16730216] Van Venrooij WJ, Pruijn GJ. Citrullination: a small change for a protein with great consequences for rheumatoid arthritis. Arthritis Res. 2000; 2(4):249-51. [PubMed: 11094435] Rogers GE, Harding HW, Llewellyn-Smith IJ. The origin of citrulline-containing proteins in the hair follicle and the chemical nature of trichohyalin, an intracellular precursor. Biochim Biophys Acta. 1977; 495(1):159-75. [PubMed: 410454] Keller, J.N., Hanni, K.B., Markesbery, W.R. Impaired proteasome function in Alzheimers disease. J Neurochem. 2000; 75:436-439. Maccioni R. B., Muà ±oz J. P. Barbeito L. The molecular bases of Alzheimers disease and other neurodegenerative disorders. Arch Med Res. 32, 367-381 (2001).   Ã‚  

Pre-Marital Sex & Role of Youth in Building a Nation

PREMARITAL SEX Premarital sex is sexual activity practiced by persons who are unmarried. Is it ok to have premarital sex? † That is a common question among teens and engaged couples. In your mind, you are probably weighing the pros and cons of premarital sex. On the positive side of the scale, there is acceptance from your peers, hope for pleasure, and the fulfillment of sexual desires. The negative side of the scale carries the weights of morals, fear of pregnancy or disease, and guilt. Premarital sex is serious moral issue confronting high school students.This isn’t to say that sexual temptations only existed in youth. But these days, being youth, they are more cut off from familial restrictions and parish supports, confronted with peer pressure that alleges, â€Å"Everyone is doing it,† and perhaps exposed to an irreligious environment. Premarital sexual relationship is an important subject – especially today. Young people are bombarded with the worldâ⠂¬â„¢s standards of morality, or immorality. The values and moral standards, which were endorsed by most Filipinos in years past, are now ridiculed and/or ignored by many.CONCLUSION Premarital sex has no moral grounds, it is against God, and it is unsafe physically and emotionally. Although sex is pleasurable, it is designed by God to be enjoyed by two married people. ROLE OF YOUTH IN BUILDING A NATION Youth† is the critical period in a person’s growth and development from the onset of adolescence towards the peak of mature, self-reliant and responsible adulthood comprising the considerable sector of the population from the age of fifteen (15) to thirty (30) years.Their attitudes, values, mind-set, and priorities will determine the development of the country in the future. As a human we have different kinds of role in the nation and one of this is to become an effective good leader, for us to have a progressive nation, we need to be a responsible person, because we canà ¢â‚¬â„¢t have a progressive nation if we’re not responsible enough, but we must begin to our selves.And in order for us to become a good leader, we must know first how to be a good follower, because we can never be a good leader if we don’t know how to be a good As a member of our community we can help building our nation. We can serve our nation by being united as we having one objections and one goal we may able to attain the works easily. We can do that by participating in any programs in our community. As a youth we can join what they called â€Å"SK† or Sanguniang Kabataan

The Code Of Discipline Of The Berbice Uprising - 1819 Words

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Social Influences on Behavior Essay - 1300 Words

Social Influences on Behavior Rhonda Hager PSY/300 June 25, 2012 Teresa Neal Social Influences on Behavior Introduction All humans’ behavior is affected by social influences to some extent. The level of influence will vary from person to person, depending on the several factors, such as self-esteem, their level of self-awareness, morals, and values (Velden, 2007). People do many things to ensure they are accepted by the people in their group and to keep from being ostracized by individuals around them (Kowalski amp; Westen, 2011). Individuals are usually guided by their own sense of what is right or wrong and will make logical decisions based on this. 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Kinship and Marriage in Gurung Society - 603 Words

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A Social Partnerships in the New Millenium Free Essays

Social partnership is very fashionable right now. It has become the new â€Å"buzzword† in Industrial Relations. The ‘new’ government, the TUC and the CBI are all promoting partnership at work, every week there is another conference held somewhere in the country on the topic of partnership. We will write a custom essay sample on A Social Partnerships in the New Millenium or any similar topic only for you Order Now There is also a huge amount being written about the issue, some Industrial Relations commentators have even hailed this as the most exciting thing to happen to UK industry in years. However, not every group completely agrees with this. For example, some groups believe that Social Partnership has had a very negative effect on British Trade Unions. At www.labournet.org.uk a group have named a bulletin board ‘Solidarity’. It is a page dedicated to ‘challenging social partnership’ and it’s apparent corrosive effects on union power. It is very typical of the types of arguments put forward by groups that are oh so against social partnerships. Stakeholders like the authors of solidarity believe that social partnerships tie unions to the coat tails of employers. What they want is a ‘new unionism’ which opposes subordination to the ‘global market’ and builds on the experience of the Liverpool Dockers. This group is completely opposed to any kind of privatisation and really wants more from unions than just a concern for working conditions. They want a break â€Å"..with the narrow work place concerns of ‘non-political’ trade unionism† and believe that unions should fight † for the interests of the working class and oppressed as a whole†. They also see a need for a repeal of all â€Å"anti-union† legislation and an end to state interference into unions, such as social partnership agreements. Taken from ‘Solidarity’, the journal that openly states it’s purpose is to challenge social partnerships in trade unions, here are some ‘facts’ that display how social partnerships have had a harmful effect on British trade unions: In Rover and the car industry, where the unions have identified the interests of workers with commercial success in the ‘global market’, they have been gravely weakened and meekly accepted massive job cuts. USDAW has reached a ‘Partnership Agreement’ with Tesco which takes away the right of the union members as a whole to vote on pay deals. The TUC has collaborated with a privatised utility in the Energy Industry and set up a company with them – Union Energy – thus effectively abandoning the fight for re-nationalisation. They are collaborating with a privatised utility which has decimated trade union members’ jobs. The unions have swallowed ‘Investors in People’ which identifies the interests of union members with the ‘business aims’ of private companies. The TUC and most unions have accepted that the increased competition of the ‘global market’ means unity with ‘our employers’ and competing with workers in other countries. Stakeholders like these aim to challenge social partnership by campaigning for complete independence of the unions from the employer. They reject globalisation and counterpoise to it a working class internationalism which recognises that workers have more in common with those in other countries than they do with ‘our own’ bosses. It is a very Marxist standpoint . They feel that for those in the unions who support their independence from the employers and the state, a systematic struggle against ‘social partnership’ in all its manifestations is a central task if they are to break the unions from the employers’ coat tails. They are insistent that this outlook has only been adopted by union leaders. It has though percolated down to many workplaces, reinforced by the fear of unemployment. This far left view contrasts greatly with those unions that actually see social partnership in a very positive, if slightly unclear light. Unions like FIET, The International Federation of commercial, clerical, professional, and technical employees see social partnership as a â€Å"new and challenging area of work for the union.† For unions that see the millennium as a time to accept social partnership, it will mean unions and companies learning to do things differently, rather than trying to campaign against this change. Some unions like FIET have accepted that we now have a government committed to promoting the partnership approach, and to ensuring that it becomes a permanent feature in the workplace. Unions like this have therefore accepted that social partnership is going to be around for a long time. I think that left wing reactions to social partnerships, like that of ‘Solidarity’ is very much a reactive one rather than considered. I think that those stakeholders set so strongly against this issue need to understand that the government’s political commitment to partnership goes far beyond the UK. We now have a new and positive approach to Europe and this social partnership model is at the heart of the EU’s approach to all Industrial Relations. Indeed Social Partnership does mean long term changes to the Industrial Relations scene in Europe and in Britain, all that unions need to work on is understanding what these changes will mean for their members and the companies that they work in. Looked at from this more positive, optimistic standpoint then social partnership has the potential to deliver some very real benefits: Potentially it means that employers will be working with the trade unions to bring about improvements in the quality of work. It means that employees, through their trade union, will be given a much greater say in how their company is run. Social partnership should give unions the opportunity to be consulted earlier, more often and on a wider range of issues than has happened before. Surely the growth in partnership at work can only be a welcome one, if not somewhat overdue? However, these encouraging and upbeat pointers can only be achieved if the unions get themselves involved in equal terms right from the beginning, otherwise employers could try to do things without the unions. As stakeholders like ‘Solidarity’ state I think that there is indeed a risk that social partnership may be used to try and undermine trade union organisation in the work place. Employers could potentially claim that they can have a partnership directly with their staff and do not require the union’s participation. If the unions are excluded in this way then not only could there be disastrous consequences for union members but social partnership will not work for employers either. Partnership can only be successful when employees as partners are properly supported and resourced through a dependable trade union. To illustrate this point further, let us consider what may happen if the retail sector tried to introduce partnership without trade union involvement : Most employees in this sector work part-time. There are increasingly complex and unsociable working hours in retailing which in turn means that most employees may have very little contact with anyone beyond their shift. In these circumstances employees have less of an opportunity to form a collective view. Simply meeting together can be extremely difficult in such organisations, let alone having the resources and confidence to develop their own ideas and concerns. Therefore it is clear that in most cases employees are unlikely to become equal and effective partners when they do not have the time or the means to generate their own agenda. A dependable and experienced trade union means that employees can enter into partnership properly supported and resourced. To achieve this, a real commitment is required from employers to supporting trade union membership, as the best way for their staff to be given a voice and more importantly the confidence to use it. Basically partnership needs partners. But how will social partnership actually change things in real terms for employees and the unions? By answering this question I will hopefully be able to assess if social partnership is a viable way forward in the new millennium. Firstly, unions need to take a positive approach, for them social partnerships should be about: Co-operation, not confrontation. Improving the quality of working life. Employers listening to and respecting unions and vice versa. Employees developing their own agenda through their trade union. If they agree on the above terms then hopefully it will mean: Moving away from the common practice of the union only talking to companies once a year. At the moment, even with companies with whom there exists a good relationship, unions often only talk to the organisation at the time of the annual wage negotiations. Many unions feel that the annual wage round sometimes becomes a little tired and predictable, with companies offering the lowest increase that they feel they can get away with. However, partnership is different. For the unions, it means talking to companies throughout the year. It means having a constant dialogue with employers about what is happening at all levels of the company. It should also mean that the union is consulted on a wider range of issues. Companies are used to contacting unions when they are implementing redundancies or sell offs. Mainly because there is a legal obligation on them to do this, but unions have not in the past been consulted on the overall direction of a company or about strategic decisions that may affect employees. A Social Partnership means that the union should be consulted on a far wider range of issues than they have been used to. It means earlier and better consultation sessions, theoretically no more being told about something when it has already been implemented. How to cite A Social Partnerships in the New Millenium, Essay examples

Protein Synthesis and Protein Degradation

Question: Discuss about the Protein Synthesis and Protein Degradation. Answer: Introduction: Protein turnover refers to the process by which protein synthesis and protein degradation is balanced. Selective process of protein turnover mediated by ubiquitin therefore refers to protein regulatory process that control degradation of some proteins in cell process. Ubiquitin is 76 amino acid regulatory protein that helps in controlling degradation of selected proteins during protein turnover. According to Tymoczko, Stryer Berg (2002) ubiquitination refers to the addition of ubiquitin to substrate that can signal degradation of the substrate. The following paper describe selective protein turnover of ubiquitin-mediated degradation of keratin in the intermediate filament protein. Degradation of keratin in the intermediate filament occurs through a series of steps that include activation and conjugation, Transesterification, ligation by ubiquitin ligase and Transfer to target protein via E3 to lysine residue intermediate filament protein. Firstly, ubiquitin is activated by ubiquitin activation enzyme E1 that is followed by conjugation of ubiquitin. Secondly, transesterification that involve transfer of ubiquitin to ubiquitin-enzyme E2. Thirdly, the E2 complex is then delivered to ubiquitin-ligase E3 substrate site in the intermediate filament proteins. Fourthly, the keratin protein is delivered via E3 to lysine residue in the intermediate filament protein (Harper Schulman, 2009). In the last step, ubiquitin binds to the lysine residue on the keratin through isopeptide bond. Ubiquitin therefore bind to target protein at different sites and to amino group at N-terminus. The overall result is cleavage of keratins bonds at different amino group and high energy is required in this process. The degradation of ubiquitin- substrate is done by 26S proteasome complex that at the end release reusable ubiquitin and free ubiquitin (Philpott Itzhaki, 2012). Reference Harper, W. Schulman A.(2009) Ubiquitin protein activation. Nature review molecular cell biology 12(5): p. 23-31 Philpott, A Itzhaki, L. (2012) Regulation and control of Proteins Degradation. Biology Open 1(13): p. 136-72 Tymoczko, J. Stryer L Berg, M. (2002) Biochemistry: Protein Turnover. New York : W H Freeman and company.