Analysis of Peripherin-2 Cone Mutation V268I

Analysis of Peripherin-2 Cone Mutation V268I Introduction: 1.1 Composition of Retina: Eye an organ which is known for its sensory function Vision, serves as a basis for perception of the world outside. Three layers of tissues enclose the eye, outermost layer sclera, middle layer choroid and inner layer retina. Central macular region of the retina called as fovea consist of photoreceptor cells, where maximum visual acuity is possible due to minimal light scattering. Iris found in region between the cornea and lens acts like an aperture controlling the amount of light that could enter and pass through retina (Siegel and Sapru.2011). Retina consists of following layers: The outer most primary epithelium layer with pigmented melanin cells absorbs uncaptured light preventing reflection back to rest of retina. The later parts of retina layers involved in sensing and processing of light stimulus are the layers with photoreceptor cells- light sensitive region with rod and cone cells, external limiting membrane, outer nuclear layer, outer plexiform layer, inner nuclear layer, inner plexiform layer and inner most layer with ganglion cell axon fibers. Optically as light when imaged onto retinal photoreceptor layer, light passing through this layer with rods and cones converts light stimuli to electric signal through horizontal cells, further processed through bipolar, amacrine before they reach ganglion cells. From the axons of Central Nervous System (CNS) ganglion cells information is passed in the form of action potential to optic disc and optic nerve signal is received by brain’s visual center. (Ayoub, 2008). 1.2 Composition of Photoreceptors: Retina is made up of two different types of photo receptors- Rods and cones, the ratio of these photoreceptors vary in different species depending on its habitat being diurnal or nocturnal, in human’s rod cells are in majority with cone cells in a ratio of 20:1 (Mustafi et al., 2009). Rods and cones consist of outer segment, inner segment and synaptic terminal. Towards outer surface retina outer segments are located, these segments contain membranous discs arranged in the form of a stack, and these regions are involved in photo transduction by detecting photons and converting them into electrical signals. Inner segment consist of nucleus, both segments are integrated by a stalk containing microtubules. The synaptic terminal carries out synaptic contact with other cells (Siegel and Sapru. 2011). Retinal pigment epithelial cells phagocytize the aged disks which are shed from distal end when new a new disk is added to the base (Young. 1967). Fig: 1 Structure of rod and cone cell. In rods outer segment disc membrane is separated by plasma membrane, disks consist of enclosing lipid bilayer compartment called as disk lumen or intra distal space. Lamellar region of the disk is joined by curved rim region, lamellar region consist of photoreceptor protein, Rhodopsin – an integral membrane protein, and curved rim regions consist of filaments extending to cytoplasmic regions of adjoining disks maintaining structure of outer segment. Rim regions also contain membrane proteins Rim-1, a high molecular weight protein and two other lower molecular weight proteins Peripherin/rds and Rom-1. In cones discs are not completely formed, they are adjoined and are continuous with plasma membrane. (Molday. 1994). These two proteins Peripherin/rds and Rom-1 are homologous in structure, peripherin/rds is expressed both in rods and cones while Rom-1 is only found in rods, they can interact to form dimers and tetramers. (Elizabeth. 2005) Cones are known to be responsible for vision during daylight and mediate color vision, while rods do not. Axial structure of cones makes them capture light efficiently during day, with fast responsive membrane potential with less integration time. There are three types of cones L, M, S cones which are specific to a region of visible spectrum to which they are found to be sensitive. L cones are sensitive to ÊÅ ½max~ 553-565, M cones are sensitive to ÊÅ ½max~ 530-537, S cones are sensitive to ÊÅ ½max~ 415-430, frequency of photons. (Mustafi et al., 2009). Rods are specialized to work under dim light conditions, rhodopsin is the most abundant protein found in the rods constituting about 85 % of total protein in rod outer segments. (Elizabeth. 2005). Circuit transmitting information to ganglion cells is different in rods and cones, each rod bipolar cell is connected to many rod cells, and many rod bipolar cells are connected by amacrine cell. More convergence make rods better detector of light but reduces spatial resolution. In cones it’s less convergent as each ganglion cell receives input from only one cone bipolar cell, which is connected to single cone cell (Mustafi et al., 2009). Signal transduction in Photoreceptors: The signaling cascade in photoreceptors involves from light absorption till signal generation by membrane polarization, resulting in controlling the rate of release of neurotransmitter through synaptic terminal. In Dark light condition gyanylate cyclases (GCs) actively produce high Cyclic guanosine monophosphate (cGMP) levels, There by the cGMP sensitive cyclic nucleotide-gated (CNG) channel is in its open state and gives rise to the influx of Na+ and Ca+2. Rods and cones are depolarized to ~35 to 45 mV, where outer segment membrane channels are open permeable to Na+ by effluxing k+ balancing cations. This constitutes a circuit known as â€Å"Dark or circulating current†. In this condition with the channel opened for influx of ca+2 initiates the release of neurotransmitter from synaptic terminal. In light conditions cation channel in outer membrane is closed, thereby membrane hyperpolarizes towards equilibrium potential for k+, causing halt in neurotransmitter release (Jindrova. 1998). When photon is absorbed by Rhodopsin- a 40 kDa protein belonging to G protein coupled receptors, consist of a protein opsin, a single polypeptide with seven transmembrane helical segments forming three cytoplasmic loops and a chromophore 11-cis retinal, which binds to lysine, 296 residue on opsin’s seventh helix. In rods and cones photon absorption leads to isomerization of 11-cis retinal form to all trans retinal confirmation and in the surrounding opsin repositioning of cytoplasmic loops occur which in turn activates the protein to Metarhodopsin II (R*) state, which stimulates the heterotrimeric G protein transducin (Hargrave et al.1993; Scot and John . 2008). Preceding from here all the steps occur in dark state, Transducin stimulates cGMP phosphodiesterase (PDE6) to hydrolyze cGMP to 5’GMP by dissociation of ÃŽ ³ subunit from PDE ÃŽ ±ÃŽ ² subunits, resulting a vast increase of 5’GMP and decrease in cGMP concentration leading to the closing of the CNG channel. As a consequence, the outer membrane hyperpolarises to ~-70 mV and release of neurotransmitter is halted to bipolar cells, finally light signal information is sent by termination of transmitter release to the brain. For regeneration of dark current, the Rod cell needs to restore back the concentration of cGMP which is regulated by concentration of Ca+2 and by gyanylate cyclases (GC), Ca+2 molecules are bound by gyanylate activating proteins. In light conditions, where the CNG channels are closed and Ca+2 levels are low, the low level of calcium activates gyanylate activating proteins, which in turn stimulates gyanylate cyclases leading to production of cGMP. Increased level of cGMP causes to open CNG channel and dark current is restored (Jindrova. 1998). Additional mechanisms are found to be involved to restore the molecules from active to inactive state. Inactivation of Rh* by phosphorylation, catalyzed by rhodopsin kinase (Chenet al.1999). Phosphorylated rhodopsin is blocked by the protein arrestin by binding to it preventing activation of transducin, resulting in breakdown of activated rhodopsin (Xu et al,. 1997). The all trans retinal disassociates from opsin, diffuses to cytosol transported to outersegment then into pigmental epithelium where it is reconverted to 11- cis retinal, then the recycled 11 -cis retinal is transported back to outer segments. Peripherin- 2: This membrane protein is named on basis of localization studies done in 1987, which revealed that this protein was found localized around the periphery region of the outer segments, disks in rods and lamellae or disks in cones. (Molday et al,. 1987). In humans peripherin/rds is encoded by cDNA Peripherin/rds with open reading frame of 346 amino acids, weighing 39.3 KDa (Travis et al., 1989). Peripherin is an integral membrane glycoprotein having four trans membrane domains. When photoreceptor cells fail to develop in outer segments it results in retinal disorder named retinal degeneration slow (rds). Amino acid sequence of peripherin in bovine photoreceptor cell is 92.5% identical to protein encoded by rds gene in wild type mouse. Localization of these proteins was also found to be in rod outer segment (ROS) membranes, implying the role of peripherin-2 as a cause in retinal disorder. (Connell et al, .1991). Peripherin protein is also known and addressed with many other names like pho toreceptor peripherin, peripherin-2, peripherin/rds, rds/peripherin and rds. The two dimensional structure proposed consisted of four membrane spanning domains, with its N and C terminals located intracellular in extradiscal space, two extra cellular loops D1 and D2 ( EC1 and EC2) are localized with each containing a N-linked glycan in intradiscal space. The D2 loop among four species mouse, rat, human and bovine is found to be 92% identical, predicted that evolutionary drift in this loop could be a reason for its involvement in protein -protein interactions. (Connell and Molday, 1990; Travis et al., 1991). Fig: 2 Structure of Peripherin-2. D2 loop in intradiskal region is found to be key in protein folding and tetrameric subunit formation, changes in amino acids in this loop resulted in either protein misfolding or reduction in sediment coefficient of protein or even both, seven cysteine residues found in this loop to be important for intramolecular disulphide bonds, one of the cysteine residue is responsible for polymerization of tetramers (Loewen and Molday, 2000). When seven conserved cysteine residues were replaced they showed abnormalities, C214S linked mutant for adRP was not normally folding and interacting with rom-1 protein to form tetramer. While in C150S mutant failed to form intra molecular disulphide bonds, these results revealed that cysteine residues were crucial for folding of protein and subunit assembly, failure of these functions are linked to retinal disorders like adRP. (Goldberg et al., 1997). Human rom-1 protein with 351 amino acids exhibiting 35% identity to peripherin-2, hydropathy profiles revealed that rom-1 and peripherin-2 have a similar topology with four transmembrane domains, one of differences noted in the both the proteins were consensus sequence for N-linked glycosylation is absent in the rom-1 (Molday. 1994). Peripherin homologous protein rom-1 forms disulfide-linked dimers with each other (Bascom et al.,1992). Peripherin-2/rom-1 core native complex was found to exist as a tetramer, membrane curvature was found to induce with tetramer formation when they were reconstructed in lipid vesicles (Kevany et al,. 2013). Peripherin-2 associates with itself and also with its homologue rom-1 in forming homo and heterotetramers core, these tetramers when linked intramolecularly together by disulphide bonding forms octamers and higher order oligomers (Loewen and Molday. 2000). A highly conserved region within the C-terminal domain of peripherin/rds was found to be import ant for membrane fusion, it forms a complex with melanoregulin (MREG) onto the last five residues of the C-terminus (Gln341-Gly346), membrane fusion is important in organelle biogenesis, disc morphogenesis and disc shedding. (Boesze-Battaglia et al., 2007). Peripherin/rds plays a major role in rod and cone outer segment morphogenesis. In mice, absence of peripherin/rds leads to develop normal photoreceptor inner segments but they fail to form outer segments, these outer segments undergo apoptosis (Nir and Papermaster. 1986). Peripherin- 2 is found to have differential roles in rod and cone cells with respect to their binding partners. Peripherin 2 found to be binding with the Glutamic acid rich protein (GARP) portion of ÃŽ ² subunit of rod CNG channel. This interaction is predicted to anchorage disc rim and rod plasma membrane, but in cone CNG channel such interaction was not observed, as GARP is not expressed in cone CNG channel or in other from. (Conley et al,.2010). Peripherin-2 links CNGB1a CNG channel subunit to rhodopsin, these three forms a complex localizing its contact between disk rims and plasma membrane regions. FRET experiments revealed that transmembrane -4 (TM4) of periperin-2 is key for rhodopsin interaction. In peripherin-2, G266D mutation specific to TM4 region, eliminated the specific binding capacity of peripherin-2 with rhodopsin (Becirovic et al,. 2014). Peripherin-2 being a multifunctional protein and mutations in peripherin/RDS gene results in a broad spectrum of retinal disorders like macular dystrophies, cone and cone-rod dystrophies and retinitis pigmentosa. Studying Peripherin/RDS mutations and protein structure elucidates pathophysiological mechanisms underlying these retinal disorders for effective therapeutic intervention (C.J.F. Boon et al,. 2008). Zscan29: At present, very less information is known and available regarding the function, structure and localization of this protein Zscan29. mRNA specific for this protein were found to be expressed in retina through RT- PCR experiments from mice different tissues. In mice 5 different isoforms of this protein were found, with longest isoform having 869 amino acids (aa), and other isoforms were of 834 aa, 548 aa,265 aa,206 aa. The isoform with 265 aa was found to interact with the C- terminus of CNGB1a sub unit of CNG channel. In long isoforms with 869aa and 834aa, a SCAN domain, a DUF2 domain, 2 GT-1 domains and 6 zinc finger domains were found. The scan domain is predicted to be a DNA binding domain and involved in transcription regulation, as this domain in other proteins were also found to be associated as transcription factors. DUF-2 Domain function is unclear and yet to be known, Two GT-1 domains of 79 aa each, were predicted to play a major role in light sensing mechanisms by binding GT trihelix transcription factors. (___citation__). In Arabidopsis GT-1 is predicted to respond to light signals via calcium dependent phosphorylaton, after GT-1 binding to GT cis element of light inducing gene (Bauer et al). Fig: 3 Isoforms of Zscan29 protein with its domains. Aims of this work: Analysis of Peripherin-2 cone mutation V268I: Verification of the interaction of WT peripherin-2 and peripherin-2 with mutation at V268I position with S-opsin, M-opsin and Rhodopsin via co immunoprecipitation from HEK293 cells. Interaction and localization of Peripherin-2, V268I mutation in mouse retina. Analysis of Peripherin-2 mutations at position P210: Verification of the interaction of peripherin-2 protein with mutations at P210 position with wild type Peripherin-2 and its homolog Rom-1 via coimmunoprecipitation from HEK293 cells. In vitro imaging of HEK293 cells expressing peripherin-2 protein with mutation at P210 position. Expression and localization of Peripherin-2, P210L and P210R mutations in mouse retina. Affinity purification of Anti-Zscan29 antibody and checking antibody efficiency.

Cosmic Creation Myth

Cosmic Creation Myth across Culture Paper Kristin Sheffield HUM 105 For decades myths have been told about the development of the world. Each culture has their version of how life began. These myths are centered on creation; how the earth was created, how the sky, land or the sea was created, and how humans and animals were created. These myths give cultures the security of belief. Different cultures believe differently, but these beliefs help certain cultures to continue to face problems or issues, worship, and create happiness and peace. The Enuma ElishApsu, the father of Sweet Waters  and Tiamat, Mother of Salt Waters created 3 unruly children. The children gods were just being children, but unfortunately they were disrupting Tiamat’s peace. She grew to hate their behavior and asked Apsu to deal with them. When he tried they just ignored him. Due to their unwillingness to obey their parents Apsu's resolution was to destroy them. Tiamat was very upset by this resolution. S he told Apsu that his resolution was evil and they needed to be more understanding. Her pleas were ignored by Apsu. Apsu and Tiamat’s creations soon realized Apsu's evil plans to destroy them.At first they cried, and then they succumbed to their fate. However somehow they had befriended Ea, the wisest God, who eventually killed Apsu and made his co-conspirator his slave. This is a Babylonian Myth. This myth later became the reason for a national holiday in which they â€Å"emphasized the reestablishment of order†, (Rosenberg. 2006). The Creation of the Universe Ife Olorun, the god with the greatest knowledge, had a son, Obatala, who wanted to create land where Olokun, the goddess of endless waters and wild marshes ruled. Obatala went to his older brother for advice on how to proceed.His brother's gift was one of prophecy. He advised his brother and sent him on his way. Obatala did everything he said he'd do. He created land in the middle of marshy water; he then create d plants. But he wanted more and more. While drunk on wine, he started playing with clay. He asked Olorun to â€Å"breathe life into them†, (Rosenberg. 2006). Once he sobered he realized they were imperfect. He decided never to drink that type of wine again and vowed to protect all the humans that suffered because he was drunk. He â€Å"became the protector of all those who are born deformed† (Rosenberg. 006). This myth came from Africa; the Yoruba people. They showed more emotion in their stories; Love, jealousy, sympathy and anger. As with many myths, the creators of these myths were neither male nor female. They were groups of people. The Enuma Elish myth was from the Babylonians and also the Assyrians as well. For The Creation of Universe Ife, this myth came from the Yoruba people of Africa. Together these myths share similarities about suffering and appreciation for what they have. In these myths something vital has to be restored in order to move forward.With the Enuma Elish it was the balance between good and evil. With The Creation of Universe Ife he became the protector of imperfect people. Both The Enuma Elish and The Creation of Universe Ife had mainly water as the major element. In The Enuma Elish both parent gods were from Salt or Sweet waters. Olokun, in The Creation of Universe Ife was god of mash and wild waters. In both myths anything that disrupted peace made the waters â€Å"surge back and forth† (Rosenberg. 2006). Another similarity was humans were created. In The Enuma Elisa Ea killed Kingu by severing his blood vessels.The first humans came from the blood. In The Creation of Universe Ife, Olorun â€Å"breathed life† into the clay for Obatala creating humans. The difference was Obatala wanted companions, but Ea’s purpose was to have the humans â€Å"serve the gods† (Rosenberg. 2006). These myths were designed to understand the unexplainable. Different cultures explained them differently. Each cultu re had god and goddesses. Stories of these gods and goddesses explained things to humans such as how we (humans) came to be; how the earth formed; why humans look differently from other humans or animals; etc.Different cultures believe how things came to be differently. It depends on their faith, religion, and beliefs as to what they will believe. In all the stories that are being told, it would only take a person with influence to announce the story is true in order for other people to start believing. Once other people start believing, the proof is optional. Reference Rosenberg, D. (2006). World Mythology: An anthology of great myths and epics (3rd ed. ). Chicago, IL: McGraw-Hill.

The Untold Story on Order College Research Paper Online That You Really Need to Read or Be Left Out

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Wild and Rebellious Adolescence - 1601 Words

The movie thirteen is a raw psychodrama directed by Catherine Hardwicke is based on the life of a young teenage girl, Tracy Freeland whom catapults from pre adolescence/childhood to a wild and rebellious thirteen year old girl. Filmed in Los Angeles, Tracey and her mother’s relationship are put to the test when she befriends Evie. Evie is a popular girl from junior high school who introduces her to the world of sex, drugs and self-mutilation. We see a physical and psychological change in Tracey almost overnight, as her kinship with Evie transforms into a toxic relationship. Tracey’s early traumatic life experiences, manifests into a co-morbidity of depression and borderline personality disorder that affects her ability to cope with an†¦show more content†¦Tracey has recurring flashbacks about walking in on Bradley smoking crack cocaine in their kitchen, and almost overdosing. The trauma of his presence in their house can be seen as a significant moment affectin g in her socialisation in the film. Tracey turns from an innocence girl, too a rebellious and dangerous teenager as her home becomes a toxic nightmare. Tracey meets Evie in her high school, she begins to idealise Evie; Evie has the popularity, and the confidence that Tracey wants. Tracey’s family environment and growing psychological issues impact her ability to have a scene of self; this is shown when Tracey conforms to become Evie. Their friendship grows rapidly; once Evie moves into the Freeland house the two girls go into a downward spiral of hard drugs, sex, lies, piercings and petty crime. Evie manipulates Tracey into false sense of security. She gets the attention she craves from Evie; slowly Tracey shuns Mel completely out of her life. The influence of Evie on the family dynamics is major factor influencing Tracey’s resentment towards her mother. Evie and Tracey become attached to one and other, as their friendship becomes toxic a nightmare for the entire Freeland family. Mel noticed the influence, Evie had in Tracey’s attitude but she became manipulated by Evie also. Evie was portrayed as a compulsive liar. She tells Mel, that she was sexually abused by her uncle at a youngShow MoreRelatedWild Swans Analysis2040 Words   |  6 Pagesï » ¿Year 11 Short Story Unit Writing Task How does â€Å"Wild Swans† explore the central character’s experience of adolescence? In Alice Munro’s short narrative Wild Swans, the female adolescent narrator, Rose, travels alone on a train ride to Toronto. On the train she is sexually harassed by a minister and through Rose’s reactions to the harassment, readers are given a vivid image of how Rose experiences adolescence. Throughout the story, Rose experiences fear and confusion about growing up, promptingRead MoreThe Crucible And Conversion By Arthur Milller1094 Words   |  5 Pagesdespite the fact they are based centuries apart. The time of adolescence in a woman’s life is an extremely influential period when one either stands out as a dominant leader, or falls in line as a follower. Though The Crucible and Conversion are based over three hundred years apart, both focus on similar motifs regarding female adolescence including rebellious lust, and the desire for attention and one’s individual identity. The theme of rebellious lust is prevalent in both The Crucible and ConversionRead MoreThe Crucible By Arthur Milller1316 Words   |  6 Pagesof teenage girls who face similar problems. The time of adolescence in a woman’s life is an extremely influential period when one either stands out as a dominant leader, or falls in line as a follower. 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