Role of mTOR in Pain

Role of mTOR in Pain Juliette Lanskey Introduction Pain perception protects the human body from damage, yet when the underlying mechanisms are disrupted pain can become a debilitating condition. There are almost 10 million Britains that suffer from pain on a daily basis affecting not only their personal wellbeing and quality of life but also the economy. Back pain is an example of chronic pain and back pain alone costs the NHS around  £5billion per annum and it was reported that 4.9 million days are lost per year to british businesses (http://www.britishpainsociety.org/media_faq.htm). Unfortunately, understanding the mechanisms that go awry leading to pain that is more harmful than beneficial is proving challenging. As a result, there is a deficit in treatments available to control chronic pain despite much research. There is therefore an urgent requirement to understand the mechanisms underlying pain perception in order for the development of therapeutics to reduce the sufferings of humans and the economy. This dissertation shall focus on a potential target, the mammalian target of rapamycin (mTOR), which recent research has highlighted as playing a significant role in chronic pain. Pain Pathways The processing of painful stimuli by the nervous system is termed nociception. Pain is nociception with additional psychological and emotional inputs. Noxious stimuli cause an action potential to run through a specialised set of neurons termed by Sherrington in 1906 as nociceptors; the pain neurons. Nociceptors have free nerve endings to detect noxious stimuli and can be classified into two main groups taking messages from the periphery to the central nervous system, called Aà ¯Ã‚ Ã‚ ¤ and C fibres. Aà ¯Ã‚ Ã‚ ¤ fibres are medium-diameter, mylinated neurons and this myelination and wider diameter allows rapid signal conduction. It is the Aà ¯Ã‚ Ã‚ ¤ fibres that transmit the first, sharp, localised pain of an injury whilst C fibres which are small-diameter, unmyelinated neurons transmit slow, diffuse, secondary pain 2009CELLULARANDMOLECULAR. Hence Aà ¯Ã‚ Ã‚ ¤ fibres are crucial in rapidly signalling an injury whilst the slow, burning pain from C fibres is important for protecti on during the healing period. These primary afferent nociceptors transfer messages from the periphery to the dorsal horn of the spinal chord. The dorsal horn is particularly important for processing and modulating noxious information. The dorsal horn is composed of six rexed laminae with transition zones approximately dividing different cell types. Indeed, nociceptors terminate in particular laminae. The majority of Aà ¯Ã‚ Ã‚ ¤ fibres terminate densely in lamina I, though some do also terminate in lamina V. C fibres mainly terminate in laminae I and II, although there are also a few C fibre terminations in lamina V. Thus the majority of neurons which terminate in the superficial dorsal horn specifically respond to noxious stimuli while neurons terminating deeper in the dorsal horn tend to respond to innocurous touch (large diameter, rapid conducting Aà ¯Ã‚ Ã‚ ¢ fibres transmit such innocuous information CELLULARANDMECHANMECHS2009). It is within the dorsal horn that nociceptors synapse onto central projection neurons which transmit the noxious information up to the brain. The main central pathways run up to the brain via the thalamus or brainstem and terminate in areas such as the periaqueductal grey and the parabrachial nucleus REF. There are also descending pain pathways originating in the periaqueductal grey, rostral ventral medulla and coeruleus which pass signals to the dorsal horn modulating nociception. There are also modulating circuits in the dorsal horn composing of exitatory and inhibitory interneurons contacting further neurons in the spinal cord (Fields2006thesis). Nociceptors have a pseudo-unipolar morphology allowing bidirectional signalling. This means that nociceptors are able to transmit action potentials antidromically from the the central nervous system to the nociceptor terminals (Dubin, 2010). A result of central processing is increased sensitivity of the area at and around a site of tissue damage or inflammation (PUBHUNT). A chemical soup of cytokines and growth factors is released at the site of injury and causes an increase in the sensitivity of a subset of surrounding nociceptors. This means that these nociceptors have a reduced threshold for noxious stimuli (thus will now respond to less intense stimuli than before) and also an increased response to noxious stimuli. This sensitisation of neurons at the site of injury is called primary hyperalgesia. mTOR inhibitors do not affect primary hyperalgesia and thus it is unlikely mTOR is involved, however there is another phenomenon called secondary hyperalgesia which mTOR does seem to be involved in. Secondary hyperalgesia is when a set of neurons not directly at the site of injury but in the surrounding, undamaged area undergo an increase in sensitivity due to central processing (pubhunt). Recent studies have demonstrated t hat the mTOR plays a role in creating this sensitivity. Acute pain is the pain that follows immediately after an injury to protect the body from further damage and aid the process of healing but when pain exists for more than 3 months it is defined as chronic pain (SITETHESISMerskey and Bogduk, 1994; Russo and Brose, 1998). This chronic pain does not protect the body but rather hinders the quality of life. The pathology of chronic pain often consists of decreased pain thresholds and increased response to stimsuli; the nociceptors are more sensitive. Moreover, whilst nociceptors are generally silent, firing action potentials only when stimulated (dubin2010), in chronic pain, there is an increased tendency for spontaneous activity (JULIUSANDBASBAUMTHESIS). Altogether, chronic pain leads to allodynia (pain from a normally non-noxious stimulus), hyperalgesia (heightened sensitivity to noxious stimuli) and spontaneous pain. mtor The mammalian target of rapamycin is a regulator a number of cellular processes including synaptic plasticity, protein synthesis and cellular metabolism (XONCUETALTHESIS). It is a molecule belonging to the kinase family and forms two complexes with raptor; mTOR complex 1 (mTORC1) and mTOR complex 2 (mTORC2). It is when part of these complexes that it administrates its cellular functions though much more is known about mTORC1 and so it is predominantly the role of mTORC1 in pain perception that this dissertation shall focus on. Signalling cascade There is a signalling cascade involving mTOR, the downstream targets of which lead to cellular activities resulting in the organisation of the cytoskeleton, the regulation of metabolism and cell survival (wullschlegerTHESIS). The signalling cascade is initiated by a signal such as a neurotransmitter acting on transmembrane receptors which activate phosphoinosital 3 kinase-AKT pathway. This results in the phosphorylation and thereby the activation of mTOR. Phosphorylated mTOR in turn phosphorylates the 4E-binding protein and in this phosphorylated state the 4E-binding protein is unable to bind and thus inhibit a protein called eIF4E. So when mTOR is activated it has the downstream affect of enabling eIF4E to associate with eIF4G, this is an essential step for initiating translation (TJ PRICE GERANTON). The fact that mTOR plays such a significant role in the regulation of translation is a hint of its importance in pain plasticity. Previously, it was thought by some that translation could only occur in the cell soma. However, others noted both the half-life of axonal proteins and the time it takes for a protein to travel down the length of the axon and concluded that the axoplasmic transport is too slow for protein synthesis only to occur in the cell soma HUNT. Indeed, following the discovery of ribosomes and ‘Golgi outposts’ in dendritic spines it is now believed that local protein synthesis at the sites of dendritic synapses plays a significant role in plasticity (2009REVIEW). Research demonstrates that chronic pain arises as a result of plastic changes that occur during persistent acute pain. During any pain there are noxious signals to the central nervous system enabling the pain to be perceived, if these signals persist it has been shown that this causes and maintains plastic changes that result in chronic pain. Indeed, it has been shown through advanced structural imaging methods that there are large scale alterations in the brain structure of sufferers of chronic pain CHRONICPAINPLASTICITY. There is relatively little research into the possibilities of targeting this pain plasticity to help patients cope with chronic pain in comparison with genetic studies. It is consequentially an exciting new avenue of exploration and the role of mTOR in pain plasticity is of particular interest. EARLIER As mTOR plays such a crucial role in cellular function it is unsurprising that mTOR dysfunction is believe to be involved in a number of maladies. The role of mTOR in cancer, diabetes and neurodegeneration is being explored and a number of mTOR inhibitors have already been tested for treating certain maladies. For example†¦..This has demonstrated that mTOR inhibitors are potential treatment regimes BUT THERE ARE SIDEEFFECTS MTOR THESIS MTOR SIGNALLING MTOR PLASTICITY see 2009reviewnociception and AMPKPG 6 is v good Dealing with pain MTOR AND RESEARCH thesis 2 Pain Pathways and Plasticity 3 The mammalian target of rapamycin 2007 Decreased Nociceptive Sensitization in Mice Lacking the Fragile X Mental Retardation Protein: Role of mGluR1/5 and mTOR – mTOR’s role in nociceptive synaptic plasticity through translation regulation; ‘ the mTOR inhibitor rapamycin inhibited formalin- and DHPG-induced nociception’ †¦. mTOR is a major regulator of protein synthesis for it controls the initiation of translation (PUBLISHEDHUNT2009). It is thought that by controlling protein translation it maintains the sensitivity of nociceptors following local injury. Targetting mTOR could reduce the secondary hyperalgesia that occurs from pain and thus help patients cope with pain†¦ 4 The mTOR signalling cascade Unicellular organisms that are sensitive to nutrient availability in their environment control translation via a rapamycin-sensitive translation pathway. This process is controlled by a protein kinase, TOR, which is blocked by rapamycin. Interestingly, neurons appear to have co-opted this evolutionarily conserved mechanism to control activity-dependent local translation. Mammalian TOR, or mTOR, is activated by neurotransmitter receptor signaling cascades and phosphorylates downstream factors that control translation. Hence, mTOR is intricately involved in synaptic plasticity in the CNS, a mechanism that is linked to its role in controlling translation in dendrites (Jaworskiet al., 2006). The major mechanism of mTOR-regulated translation is control of the initiation of cap-dependent translation (depicted in Fig 1) (Gingras et al., 2004). This occurs because one of the major targets of mTOR phosphorylation is the elongation associated factor 4E-BP (Gingraset al., 1999). 4E-BP binds c c ap-binding factor eIF4E and, when it is hypo-phosphorylated, inhibits the formation of the eIF4E/eIF4G elongation complex preventing translation. When 4E-BP is hyper-phosphorylated, 4E-BP dissociates from eIF4E allowing eIF4G binding and the initiation of cap-dependent translation. Recently a small molecule inhibitor of eIF4G binding to eIF4E was discovered (4EGI-1) and this molecule inhibits cap-dependent translation (Moerke et al., 2007). Hence, mTOR is crucial for regulating activity-dependent translation in neurons via its regulation of elongation factors (Bankoet al., 2006; Tanget al., 2002; Tsokaset al., 2007) and the mTOR pathway is amenable to specific pharmacological manipulation. 6 Experiments suggesting inhibiting mTOR could help control pain 2007 Decreased Nociceptive Sensitization in Mice Lacking the Fragile X Mental Retardation Protein: Role of mGluR1/5 and mTOR – mTOR’s role in nociceptive synaptic plasticity through translation regulation; ‘ the mTOR inhibitor rapamycin inhibited formalin- and DHPG-induced nociception’ NOT THAT RELEVANT – basically saying that because of mutation translation can’t happen properly†¦meaning mTOR has less control. However there is a subsection with rapamycin injections which does show decreased nociception with rapamycin 2011 – systemic inhibition of mTOR – mTOR regulation of nociceptive sensitivity; ‘ inhibiting the mTORC1 pathway systemically alleviated mechanical hypersensitivity in mouse models’ Good intro relating mTOR to chronic pain Local cutaneous intrathecal administration of rapamycin blocks activation of downstream targets of mTORC1 alleviating mechanical hypersensitivity – 21, 29 3 43 46 62 SHOULD PUT 1 OR 2 LOCAL EXPERIMENTS BEFORE THIS ONE If targeting mTORC1 signaling pathway has a potential thera- peutic application for controlling chronic pain, systemic rather than local administration (as has been used previously [21,29]) requires further investigation. Here we examined the effective- ness of temsirolimus (CCI-779), a clinically used rapamycin ester derivative, given systemically

Greek Inspired Architecture in Boston, MA

Greek Architecture in Boston My first discovery of classical architecture was made the moment I stepped outside my apartment, as my building has two Doric columns framing the front door. As I meandered over the hill and across the Common I realized exactly how prevalent classical architecture really is. Elements of classical architecture can be found on almost every building in the old residential neighborhoods and many commercial buildings feature columns and arches as well.I saw all three orders of columns, arches, and even a dome. The only architectural element I was unable to find was a iron vault. The Massachusetts State House, about half a block from my apartment, has many examples of classical architecture including a large gold dome, twelve Corinthian columns, and seven arches visible from outside. The State House, which houses the state legislature and the governor's offices, was built in 1798 by leading architect of the time Charles Bullfinch.The dome was originally made ou t of wood shingles but has since been sheathed in copper and covered in 23 karat gold. (http:// www. Satisfaction. Gob/formalities/mischievousness's. Asp) As far as I observed he arches and columns follow the rules of the classical orders except that the frieze above the columns is blank. I'm not sure if the frieze above Corinthian columns is traditionally blank or not. A pioneer, which signifies the importance of the logging industry in Massachusetts, tops the dome. Http://www. Satisfaction. Gob/formalities/ mischievousness's. Asp) I doubt this would be found on domes of the ancient world. Just as classical architecture was meant to convey power and grace in Ancient Greece and Rome, I'm sure Charles Bullfinch used elements are classical architecture n the design of the State House for the purpose of conveying the wealth, power, and beauty of the government and state of Massachusetts. Another building that incorporates classical elements is the Boston Museum of Fine Art.The current MFC building was built in 1909. There are rows of ionic columns and construction that looks like post-and-lintel. In the area that would be the frieze it says â€Å"Museum of Fine Arts† which could probably be loosely considered a continuous frieze. It was especially interesting to me to see the post-and-lintel type openings since most alluding seem to prefer the Roman's arches over the Ancient Greek post-and-lintel system. The Museum of Fine Art is another building that, due to its purpose, should convey an air of greatness.The last building I would like to use as an example is my apartment building, which was built in 1909. There are likely hundreds of residential apartment buildings in the area whose doors are flanked by a column on each side, particularly the apartments that were built in the early sass. The columns of my apartment building are of the Doric order but the abacus of each is rounded, not angular. (Strickland peg 15) There is a session but no cornice, pedimen t, or any other aspect of the Doric order. At night the columns are illuminated.It is clear by the emphasis the building management places on the columned entrance that they believe it to denote a sense of wealth, greatness, or class. While the State House and MFC certainly do this, I find my building does not quite succeed. It feels that without any other of the classical elements the columns are lacking. It is particularly interesting to me how long classical architecture has been in style. It as been used for centuries on everything from prominent government buildings to much more humble residences.Architecture that was once reserved for temples and important public spaces can now be found on everything from strip malls to pharmacies. An important difference in the use of the classical architectural elements is that Ancient Greeks built in their style partly for the logical purpose of creating a building that will stand. Today, even more so than in Ancient Rome, many aspects of t he architecture are purely ornamental. Http:// consistory. Schenectady. Org/temple-of-portions. HTML

The Basics of Samples of Autobiography Essay for College

The Basics of Samples of Autobiography Essay for College What You Need to Do About Samples of Autobiography Essay for College Before You Miss Your Chance Choosing our service, you will understand that studying can be simple if you gain from the help of capable experts. The primary goal of essay writing services is to assist you with issue and support at the same time you write your paper. After discovering our website, you will no longer will need to bother friends and family with these kinds of requests. Another benefit of our website is the quickness. The Upside to Samples of Autobiography Essay for College Additionally, an excellent autobiography sample can provide you some ideas of what things to write about or enable you to remember certain events from your life. All events are put in chronological order. Although autobiographies are usually summations of pivotal events in your life, the events you opt for may differ based on the class curriculum. 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It has often being found that autobiography essay is a little expository in nature and in some specific cases might be based on prejudice. Thus, when you plan your autobiography, make certain to concentrate on events that had significant effect on your life and totally ignore the tidbits. If you're suffering from low self-esteem, when you believe your life is meaningless, or you merely feel sad write an autobiography. You may as well start right with the story rather than putting a very long autobiography introduction. How to begin writing an autobiography is dependent upon what sort of it you are writing. If you don't understand how to compose an autobiography in the suitable way it's time to learn from the very best writing examples. Even if you're writing an official autobiography for work, you still must entice the reader with good text. Many are highly-reputed. There's no place more important to create a decent first impression than on paper. 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Life after Death, Reincarnation, Resurrection and...

Life after Death, Reincarnation, Resurrection and Immortality of the Soul Belief in life after death has taken many forms, some which are unique in particular religious belief systems, though; others can be found in more than one religion. For most religions, life after death is an article of faith. In Western religions, the belief is founded in scriptural evidence, but for all religions the belief in life after death is the same: life after death has been promised to humans by an all powerful[1] There are many views of life after death in particular which have been much adhered to and much discussed by philosophers. This essay will attempt to put forward some of the key ideas and arguments†¦show more content†¦And if human beings can exist without their bodies then the view that they can survive death seems a plausible one. We normally think of death as the end of a persons bodily life. But if people are distinct from their bodies, then the fact that their body dies does not entail that they die. This view that the body can survive as a disembodied state does not lack modern supporters. According to Richard Swimburne it is coherent to suppose that a person can exist without a body. Swimburne argues that if X can be without Y then X and Y are distinct. This view that a people are essentially other that their bodies is referred to dualism. There are a number of points, which seem to go in favour of this view on life after death. For one there is the fact that we often naturally talk as if our real selves were distinct from our bodies- as for instance, when we say we have bodies and as when we agree that we can be the same person over a number of years even though our bodily make- up has dramatically changed in the meantime. Other views for life after death, which fit into the motion of dualism, include the immortality of the soul and the reincarnation of a person. 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