Schwann Cells and Nerve Regeneration

Schwann Cells and Nerve Regeneration Yinghui Xu   Ã‚   Abstract The majority of the nervous system cells are glial cells, which have various properties and functions. Schwann cells are the principal glial cells of the peripheral nervous system(PNS). In normal nerves, they function to form insulating myelin sheath around axons and provide support to neurons. In injured nerves, however, they change their properties and switch function to that of supporting nerve regeneration. Understanding their ability in the switching process could lead to better treatment for those suffering from nerve injuries. This essay gives an overview of the structure and development of Schwann cells and the causes of nerve injuries. It also provides an explanation in the role that Schwann cells play in supporting nerve regeneration and how different approach could promote nerve regeneration. Introduction to Schwann cells Schwann cells are the most abundant glial cells in the PNS. They are a diverse group of cells originated from neural crest cells and they play an essential role in the PNS of both vertebrate and invertebrate. They can be subdivided by their morphology and anatomical location into four subsets, namely the myelinating Schwann cells, non-myelinating Schwann cells, satellite cells and perisynaptic Schwann cells (Armati Mathey, 2014).These different types of Schwann cells and their anatomical locations are shown in Figure 1. Figure 1 Locations of different types of Schwann cells in the PNS(Armati Mathey, 2007). (a) Satellite Schwann cells in the dorsal root ganglia. (b) Non myelinating Schwann cells in mixed peripheral nerve fibres. (c) Myelinating Schwann cells in mixed nerve fibres. (d)Perisynapic Schwann cells at the neuromuscular junction (NMJ). Each subset of Schwann cells has different functions. For example, perisynaptic Schwann cells are essential for modulating NMJ function whereas satellite Schwann cells have the ability to regulate the neuronal environment (Armati Mathey, 2007). Non-myelinating Schwann cells associate with several axons via a single layer of myelin. Myelinating Schwann cells are the most studied Schwann cell subset and they are most known for their function in speeding up the propagation of action potentials. Myelinating Schwann cells wrap around axons of neurons to form the insulating myelin sheath. Between the myelin sheaths are periodic gaps called nodes of Ranvier where action potentials occur. Action potentials jump from one node to the next node,in a process called saltatory conduction, which increases conduction velocity and allows signals to be propagated long distances without any degradation. In addition to their function in the propagation of action potentials, Schwann cells have many subtle yet essential functions, one of which is its role in nerve regeneration. This essay will focus on the characteristics of Schwann cells and how they respond to injury that allows the PNS to regenerate after damage. Development of Schwann cells The myelinating and nonmyelinating Schwann cells in the PNS are originated from the neural crest, which is a multipotent structure that also gives rise to other glial cells of the PNS. Schwann cell development occurs in well-defined developmental steps, which are strictly regulated by a number of signals. (Figure 1.2). During the first stage of Schwann cell development, neural crest cells produces Schwann cell precursors. These then generate the immature Schwann cells. At birth, the immature Schwann cells differentiate into either the myelinating or nonmyelinating Schwann cells. Figure 1.2 Main stages of Schwann cell development (Jessen, et al., 2015). Peripheral nerve injury Peripheral nerve injury is injury to peripheral nervous tissue. It can occur at any point along a peripheral nerve. Nerves can be damaged by a number of mechanisms such as compression, traction, or a cut. They can also get damaged by toxins, infection and physical agents such as freezing and electrical current (Murray, 2014). 2.1 Structure of peripheral nerves In order to understand the pathophysiology of peripheral nerve injury, it is important to know the anatomical structure of a peripheral nerve (Figure 2). Axon is made up of neurofilaments and microtubules that transport substances between the nerve cell body and the axon terminal. All peripheral nerve axons are associated with Schwann cells. In larger (myelinated) nerves, Schwann cells generate layers of myelin around the axon to form a sheath, whereas in smaller (unmyelinated) nerves a single Schwann cell associates with several axons via a single layer of myelin. The connective tissue (the endoneurium) that surrounds the axon-Schwann cell unit provides structural support to individual axons. Nerve fibres are further protected by a surrounding multipayered cellular tube (the perineurium) that surround nerve fascicles (Jessen, et al., 2015; Murray, 2014). Finally, the entire nerve trunks are protected by the outermost connective tissue (the epineurium) (Murray, 2014). Figure 2 Anatomical structure and main cellular components of a peripheral nerve (Jessen, et al., 2015). 2.2 Pathogenesis of peripheral nerve injury Regardless of cause, there is a limited range of responses to peripheral nerve injury. Wallerian degeneration. Wallerian degeneration occurs when there is a loss of axonal continuity by cut or crush. Communication is lost between the cell body and the segment of nerve distal to the site of injury. As a consequence, the distal nerve segment degenerate. The details of the initiation of this degeneration remain to be fully explained, but evidence shows that an increase in intraaxonal calcium activates proteases, which leads to the degradation of the axon cytoskeleton. Wallerian degeneration is also associated with the degeneration of the myelin sheath by lipases and proteases. Schwan cells (and later macrophages) are involved in further myelin degeneration. Schwann cells also proliferate and undergo morphological changes and are induced to generate molecules that will be required for nerve regeneration later on. Figure 2.2 Wallerian degeneration2.3 Types of peripheral nerve injury There are two main types of peripheral nerve injury (Burnett Zager, 2004). The first type is axonotmesis, in which axons are damaged but the connective tissue sheaths are preserved. This type of injury can be caused by compression, traction and laceration but it is mainly seen in crush injury (Murray, 2014). Axonal regeneration is very effective and function is restored in 3-4 week when modelled in rodents by nerve crush (Jessen, et al., 2015). The second and the more severe type of nerve injury, neurotmesis, involves damage to both the axon and connective tissue sheath. Severe traction, crush and laceration can lead to neurotmesis (Murray, 2014). In this case, functional recovery is generally poor. The Schwann cell injury response When nerve injury occurs, Schwann cells respond rapidly and undergoes a radical phenotypic change. The process has two major parts, one of which is the reversal of myelin differentiation. Molecules involved in forming Schwann cells before myelination are up-regulated while myelin associated genes are down-regulated (Jessen, et al., 2015). The other part of the response involves the appearance of phenotypes that are not associated with normal Schwann cells or immature Schwann cells. These cells have a repair supportive phenotype and contribute to the nerve regeneration process. Schwann cells function in multiple modes during nerve regeneration. There are four main functions that Schwann cells carry out in response to nerve injury (Niu, 2009). These functions include(1) activating macrophages and clearing debris; (2) forming the Bungner band; (3) secreting neurotropic factors; (4) secreting ECM and CAM. 3.1 Schwann cells secrete cytokines to attract macrophages When an axon is cut, the end still attached to the cell body is called the proximal segment, while the other end is called the distal segment. The process of nerve regeneration involves the formation of axonal sprouts at the proximal stump and the growth of axonal sprouts until they enter the distal stump. Efficient nerve regeneration needs a suitable microenvironment which is regulated by macrophages via their phagocytotic and clearing functions (Barrette, et al., 2008). Experiment by Huang et al. showed that Schwann cells can secrete macrophage migration inhibitory factor (MIF) when peripheral nerve injury occurs (Huang, et al., 2002). MIF is a pluripotent cytokine that functions in inflammatory reactions and immune responses. It also plays an important role in macrophage activation, attracting macrophages to clear away debris at the injury site and regulate the microenvironment to allow for efficient regeneration. Nishio et al. administered anti-MIF polyclonal antibody into regenerating rat sciatic nerves using the silicone chamber model. The results showed that the regeneration length of the nerve in the anti-MIF antibody-treated group was significantly shorter than that in the control group at weeks 2,4,6 after surgery (Nishio, et al., 2002). In addition, a large number of apoptotic Schwann cells were observed the anti-MIF antibody-treated nerves. These results suggest that MIF also contributes to the acceleration of peripheral nerve regeneration and the prevention of Schwann cell apoptosis (Nishio, et al., 2002). Besides MIF, other cytokines such as MCP-1 were also secreted by Schwann cells When peripheral nerve injury occurs, Schwann cells secrete MIF and other cytokines which activates macrophages. Invading macrophages also secrete cytokines which contribute to Schwann cell proliferation (Kubota Suzuki, 2000). Injury-induced Schwann cell proliferation plays an important role in axon regeneration as described in the following section. Schwann cells are also capable of phagocytosing myelin and they cooperate with macrophages to clear away myelin and debris (Jessen, et al., 2015), providing a good microenvironment for nerve regeneration to occur. 3.2 Schwann cells proliferate to form the Bungner band to guide axonal growth Schwann cells are highly plastic. The change has been characterised as de-differentiation (Chen, et al., 2007), but it is also seen by some groups as activation (Armstrong, et al., 2007). The two terms seem to be contradictory to each other, but this can have a simple explanation, which is that the change in Schwann cell phenotypes during Wallerian degeneration in fact involves both of these processes. The conversion of Schwann cells to repair Schwann cells involves both a loss- dedifferentiation and a gain-activation of phenotypes In the distal stump of injured nerves, denervated Schwann cells undergo a phenotypic change to form repair Schwann cells. These repair Schwann cells form Bungner bands which are regeneration tracks that provide support for injured nerves and guide regenerating axons to their targets. 3.3 Schwann cells secrete neurotropic factors (NTF) to enhance nerve regeneration NTFs are peptides or small proteins that support the growth and differentiation of neurons. The level of their expression is elevated in Schwann cells during nerve injury. Because of their strong function in promoting neuronal growth, they may be used to prevent the impairment of function or death of neurons during nerve injury (Niu, 2009). Most NTFs produce their effects by signalling through receptor tyrosine kinases (Malenka, et al., 2009). NTFs can be divided by their cellular mechanisms into three families, namely the neurotrophin family, the CNTF family and GDNF family (Henderson, 1996) 3.3.1 Neurotrophin family The neurotrophin family includes nerve growth factor (NGF), brain-derived neurotrophic factor (BDNF) and neurotrophin-3/4/5/6/7 (NT-3/4/5/6/7). There are two receptor types for neurotrophins. The first type is called the tropomyosin receptor kinanse (Trk) which has an high affinity to different neurotrophins. TrkA is the receptor for NGF, Trk B can be the receptor for BGNF, NT-4 and NT-3, and TrkC is activated only by NT-3. The other type of is called p75 neurotrophin receptor which has a low affinity and can bind with all neurotrophins (Niu, 2009).. NGF is the first discovered neurotrophin. It is critical for the proliferation and survival of neurons, especially after nerve injury. Schwann cells express an elevated level of NGF during nerve injury, which promotes axonal regeneration and accelerates the process of nerve repair (Niu, 2009). Recombinant NGF have been produced in laboratory and it is regarded as a feasible therapy for neurodegeneration of the CNS and PNS (Colangelo, et al., 2005). BDNF help to support the survival of existing neurons and promote the growth and differentiation of new neurons (Huang Reichardt, 2001). Schwann cells that are genetically modified to overly express BDNF significantly improve the survival of spinal gangalion neuron (Pettingil, et al., 2008). Takano et al. developed an in vitro model for retinal explants and showed that BDNF greatly promoted the regeneration of neurites from retinal ganglion cells in a damaged retina. A considerabe number of neurites were observed within 24 hour in the group containing BDNF while only a small number of neurites were observed after 3 days in the control group (Takano, et al., 2002). Other neutrophins use different ways to accelerate nerve repair when damage occurs. Using NT-3 alone or in combination with other NTFs can promote nerve regeneration in the injured spinal cord (Taylor, et al., 2006; Arvanian, et al., 2006). Research by English et al. showed that neutotrophin-4/5 is required for the early growth of regenerating axons in peripheral nerves (English, et al., 2005). 3.3.2 Ciliary neurotrophic factor (CNTF) family The CNTF family includes CNTF, leukemia inhibitory factor (LIF) and interleukin-6 (IL-6). LIF and IL-6 can act directly on neurons to promote axonal regeneration (Jessen, et al., 2015). The most prominent function of the CNTF family is its prevention of motor neuron degeneration, which suggests that it can be used as a therapeutic treatment for human degenerative motor neuron diseases (Sendtner, et al., 1992). 3.3.3 GDNF family GDNF is a small protein that supports the survival of many types of neurons including motorneutrons. It can also prevent apoptosis of motor neurons caused by axotomy. 3.4 Schwann cells produce extracellular matrix proteins (ECM) and cell adhesion molecules (CAM) The endoneurium surrounding peripheral axons contain a large amount of ECM secreted by Schwann cells. ECM is rich in collagen and contains glycoproteins such as fibronectin and laminin (Niu, 2009). ECM regulates key aspects of Schwann cell development including the formation and function of myelin (Court, et al., 2006). In addition, they mediate Schwann cell proliferation and axon growth (Armstrong, et al., 2007; Webber Zochodne, 2010). CAMs secreted by Schwann cells are involved in peripheral nerve repair. They play an important role in axon growth and formation of nerve bundles. Study from Lavdas et al showed that Schwann cells have the ability to migrate in the CNS, promoting myelin regeneration and making it possible for remyelination in the CNS (Lavdas, et al., 2006) . Experiment from Park et al. showed that the attachment and proliferation of Schwann cells are affected by special CAMs. It is therefore important to consider choosing optimal CAMs for tissue-engineered nerve regeneration (Park, et al., 2008). Summary Schwann cells use the above four ways to function during peripheral nerve repair. Many years of research have already proved that Schwann cells can secrete a large amount of molecules to support neurons and accelerate the damage repair process. However, further research is needed to show whether As Although the PNS is able to regenerate, much research still needs to done to maximise regeneration potential.      Ã‚  

Essay on ”Race, Class, Violence and Denial: Mass Murder and the Pathologies of Privilege” by Tim Wise Essay

Often when racial inequality and discrimination is being discussed, we get to think of terms such as â€Å"white privilege† and American history with the Civil Rights Act in 1964. But we think of it, mainly as history. And that, according to Tim Wise, an anti-racism activist and American writer, is the biggest self-deception of the modern American world. Throughout an article posted on his own webpage, concerning school shootings, Tim Wise discusses the general American attitude towards this relatively new phenomenon in American society. With the use of especially pathos Wise argues that the most concerning thing about these events is how society is handling them afterwards. The problem is, according to Wise, that white people tell themselves ‘white lies’, and therefore never think that such actions could be taking place in their communities. He claims that there’s a reason why this happens in the outwardly ordinary societies. It’s because the people, trying to maintain at certain surface of innocence, refuse to see the signs of trouble, even when it’s going on before their very eyes. He wants us to take responsibility and stop excluding some parts of society from the â€Å"danger-zone† just because they look bucolic and normal on the outside and almost in the form of a provocative scolding he explains what damages these ‘white lies’ can do – not only to people of color, but to white people as well. Tim Wise wrote this article 3 days after the massacre at the Sandy Hook Elementary School in Newton, Connecticut, where the 20-year-old Adam Lanza shot 20 pupils and 6 staff members. Without any further explanation Wise starts out his article, and from the very beginning he makes sure that the reader is attentive: â€Å"The senselessness alone would have been sufficient†(p.8 l. 7). We immediately sense the dramatic tone just as we also quite quickly understand the author’s profound personal concern about this issue. The title indicates what is to come; yet it is not very specific and it does not deal with school shootings as an issue at all. It gives us an idea of the author’s personal fight against racism which is also reflected in the article and probably all of his writing. Wise uses heavy vocabulary in the title and he continues to do so in the rest of the article. His ethos is strengthened by his use of language. The language is rather difficult, and the fact that he leaves out almost all explanatory references, â€Å"For the right, it’s the culture of poverty, or perhaps some specific aspect of â€Å"black culture†(†¦)(p.11.l138) shows that he expects his readers to have a certain amount of knowledge about politics and society. Even though the language contains many complex sentences and has many intellectual references, it still maintains a natural and easy-to-read flow. As mentioned before, he uses pathos a lot in his argumentation. When attempting to persuade the reader, he is mainly focusing his arguments on emotions. But he is no pleaser, though. He tries to speak to our guilty conscience, and it becomes clear which reaction he wants. â€Å"There is dysfunction and pathology and general awfulness where some of the beautiful people too reside? Yes precious, yes indeed.†(p.9 ll.47-48) His tone is extremely provocative and ironic, but we never doubt the seriousness of this matter to him. Actually this only indicates the importance of the subject and the author’s character appears very passionate. Maybe to the extent that he fails to remain objective – but then again, was that ever his purpose? His use of irony gives him more options too. It makes it easier for the reader to accept some of his quite exaggerated statements. The fact that the article is published on Wise’s own webpage called: www.timwise.org is quite crucial to the way it appears. The author has created the webpage for himself and the purpose of it is to be the platform for his views on different matters. The ones who visit his webpage and read his articles have this in mind from the beginning. Also they must be interested in him in some way or another. His main claim, that the school shootings is a result of the surroundings, works absolutely fine. When he refers to the general weapon availability in the United States, he backs up his ground with facts for the first time in his article, â€Å"(†¦)we should know by now that with 280 million guns in circulation, they can’t all be tucked into the waistbands of young black men who reside somewhere else(†¦)(s.9 ll.61-62) He is surprised that people act surprised. Also, he criticizes white people for thinking that the only ones who can’t control weapons are the poor and colored ones. He uses the topic â€Å"School shootings† to raise a debate about a still ongoing racial discrimination in the US. It seems intelligent of him to choose this topic, since the horror of it is something almost everyone more or less agree on, since it is something everyone wants to end. He claims that the only way to ever set a stop to this madness is for the communities to take responsibility. All these arguments connect in a fine way, but his use of rebuttal does almost not exist. If the article had been for at newspaper or anything else with more critical readers of more dissimilar opinions, he would need that to seem less subjective. Wise has a clear purpose with this article. And at most points, he succeeds in persuading the reader. He wants us to wake up. He wants society to wake up from its conscious sleep and take some responsibility. So, who are the enemies? And who can be the heroes? According to Tim Wise we as a society plays both of the parts.

Magical Answers to Essay Writing Topics for Class 3 Disclosed

Magical Answers to Essay Writing Topics for Class 3 Disclosed Top Essay Writing Topics for Class 3 Choices An argumentative essay requires you to choose a topic and have a position on it. Moral argumentative essay topics are a few of the simplest to get carried away with. Choosing topics for argumentative essays is critical for your general success. Recent argumentative essay topics that are related to society is going to do. So you don't actually need to get a topic! Try out another topic and do the very same 5-minute writing test till you locate a topic you know it is easy to write on. If you get to select your own topic, that's wonderful. The very best topic for your essay is one which is aligned with your subject of study. Application essays about challenges reveal how you respond to difficulty to individuals who are really interested in how you are going to deal with the subsequent four years by yourself. 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Identifying an issue and proposing one or more solutions ought to be an important component in your essay. Don't neglect to explain why the predicament is significant to you! State issue and explain why a solution should be figured out. Sure, with this kind of a huge selection of topics to select from, picking just one may be challenging. Writing about nuclear weapons is always a great idea. Very often it becomes tough to choose one particular topic either due to the many ideas in the student's head , or due to their complete absence. Another reason is to observe how well students argue on various views and demonstrate understanding of the studied subject. Whispered Essay Writing Topics for Class 3 Secrets Remember you could make funny argumentative essays if you do a few things. It is preferable to grow up in a family with a great deal of brothers and sisters. Preferably, it needs to be something that you're an expert in. Opt for a distinctive topic that others may not think of, and whatever you select, make certain you know a lot about it! Whatever They Told You About Essay Writing Topics for Class 3 Is Dead Wrong...And Here's Why The college essay is among the main facets of your college application. A college essay topic may or might not be freely dependent on the student based on the course and the professor. Explain your commitments, and you're going to be the sort of student colleges find immensely attractive. Students are accustomed to the simple fact which th eir professors give them with the assignment's topic. The Do's and Don'ts of Essay Writing Topics for Class 3 You may find there's a compelling argument for learning another language after all! Needless to say, you can select any topic, nobody could ever know that you're describing experience that you never actually had, but don't forget that it is always simpler to tell the truth than to invent lies. Some can require research while some may require the usage of the five senses and figurative language. You have to be able to use persuasive language. Top Choices of Essay Writing Topics for Class 3 While you're just beginning to compose essays, you shouldn't struggle attempting to produce something to discuss. Naturally, you have to begin writing. Luckily, you have our top-class group of writers who can assist students and teach them how to pick the most suitable essay direction. Normally, the teachers or professors assign the topics independently. Essay Writing Topics for Class 3: the Ultimate Convenience! You should think about a task to locate a theme not an issue but an opportunity and even a benefit. Write about a problem you have or wish to address. The ideal thing is to choose a topic that will provide you with an opportunity to present new ideas or a fresh perspective on a problem. For instance, you can pick a topic for elementary, middle, or higher school. The Truth About Essay Writing Topics for Class 3 You want to totally explore the subject and ensure it is personal. If it's necessary to compose your whole essay in 1 day, do your very best to give yourself breaks so you don't burn out. There are several steps that you should take as a way to write a great essay. Don't do all your writing on the very same moment.

US Military Essay - 1053 Words

Many people are concerned about the U.S. military because of how many risks come with joining it. However, the U.S. military also explains the benefits that one could receive. Joining the military is a decision that should not be taken lightly. One must understand all aspects of the military. In order for a voter to be fully informed about joining the US military, one must consider both the risks, like mental illnesses and commitment and benefits, like education, and experience. 1. Risks of Joining the Military: Mental Illnesses Suffering from a mental illness is a common effect of joining the military that many people aren’t fully aware of. Specific mental illnesses one can suffer from are Posttraumatic Stress Disorder (PTSD) and†¦show more content†¦PTSD, depression, and the lack of treatment should all be taken into consideration when thinking about the military. Depression in soldiers can be caused by the length of deployment because they don’t see their families or friends for months. Commitment Being in the military takes dedication as well as devotion because it is a big commitment that one must follow through with. Rod Powers, a writer, states, â€Å"Before you join, recognize the fact that a stint in the military is not civilian employment. Its not just like having a regular job. You cant just get up and quit anytime you want to (Powers). One must be patient because this is a responsibility Powers also states, â€Å"Along with not being able to quit, there is also the factor of being deployed for months. Soldiers may find themselves spending up to seven months out of every year deployed† (Powers). A soldier must be able to handle being separated from family and friends for a good amount of time. This separation can cause stress and anxiety between soldiers and their families. 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