Conquering fear Are you suffering from fear of everything itis?Fear of love, fear of rejection, moving forward, being stuck, being married, being single, being poor, being rich, going, coming, staying, leaving, death, people, animals and everything?Make a note to yourself that you do not have to suffer from this affliction.You can change all this by taking control of your life today. Divine love casts out all fear.It penetrates and destroys all negativity.I read somewhere that humans are like tea bags;They do not know their strength until they get into hot water.Do what you fear the most, over and over again, and it will lose christian louboutin soldes its power over you. Uncontrolled fear is a negative emotion.It is the greatest cause of the life force becoming blocked, stagnant, or unbalanced and is often at the root of poverty, lack, and failure.Fear is caused by being separate from god or the universal life force.It can also be the result of giving your power away.Your power is always present in the now;You can change your state of being by focusing your attention on positive, happy, loving thoughts and emotions. Give yourself permission to be happy if you did not receive it from Important people in your childhood or life.If you did not receive Approval from your parents, you may wish to write yourself a letter from either your mother or father or both.In this letter, you say how much the parent loves you and how proud he or she is of you.Then write a letter to that parent, thanking him or her for their love.This exercise is purely for you.The letters are never mailed or given to the parent. Fear is a natural emotion as we grow and challenge ourselves with new situations and environments.Structured, controlled, normal fear can be used to your advantage.Fear causes you to sit up and take notice and to make decisions and discriminating choices about your actions and life.If fear is kept in its rightful place, it can be a helpful, positive tool.Uncontrolled, negative fear, however, can create negative conditions and warp and distort your vision.It can generate negative affirmations in your mind, leading to negative outcomes.It can ristianlbtinaussures also cause constriction of the blood vessels and make you physically ill.It inhibits the immune system and the regeneration of cells and creates poison that can be injurious to your organs and general health.When you feel the emotion of fear and bless it, it can be transformed and eventually become peace.We have all heard the meaning of fearfalse evidence appearing real.I personally refer to fear as find enlightening answers readily. Sometimes, fear can be a warning that should be heeded.In these cases, it protects you.Strive to make friends with your fear.Accept and understand fear and write out your feelings about it. You were born with only two fears:The fear of noise and the fear of falling.All other fears are learned from your parents, environment, teachers, friends, peers, and anyone who influenced your early years.Fearthe greatest enemyis behind sickness, failure, low selfesteem, broken marriages, poor human relationships, poverty, lack, and negative actions. Some people even suffer from fear of success, health, and happiness. christian louboutin escarpin noir Others fear everythingliving, loss, dying, beginnings, endings, change, being stuck, people, commitment, loneliness, and the unknown.Face what you are afraid of until you are no longer afraid of it.When you understand the situation surrounding your fear and can face and deal with it, it loses its power over you. Image your fear in front of you in the form of a fear dragon.How big is it?What colour is it?What shape is it?How close is it?Can you feel its breath?How hot is it?Mentally take a club and push the fear dragon away from you.Ask it to speak to you.Say,"Have you anything to say to me? "Listen with your inner ear.Tell the fear dragon to go.If it tries to hang around, say again,"You must goyou have no power over me.Now, in its place, create a positive image of what you wish to accomplish.
Mr.Christian http://www.listmint.com/ louboutin Mr.Christian louboutin was born in a worker's family in paris in 1963;All of his glory now was resulted from a special experience in his childhood.Once he passed the oceanic art museum in paris, in front of the door, he saw a pair of remarkable icon梐 thick pair of cone heels was crossed out by two lines, the purpose of the picture was to warn women to treat well to the wood floor within the hall.Looking at the pair of beautiful high heels, the 13-Year-Old boy became fascin ated;It was the first time that the boy realized shoes can be so beautiful.At that time, paris was in a situation of economic recovery, economics in paris boosted quick, and christian louboutin could not withstand the temptation of the fast-Paced world, so he often went to the nightclub.At that time, he was only 14 years old.There he understood what fashion was, as time vent by, he became more and more fascinated to stage performances and dancers, finally, michael kors taschen outlet he dropped out of school because of this, soaking in the nightclub every day, in order to seek some design development, he did some chores there, he realized that Making shoes was his breakthrough. "For those dancing girls, shoes are the most important thing, they should not only be comfortable, but also be beautiful and sexy, my idea is simple, just to have these girls put on my production which is more comfortable and beautiful than theirswhen he was16 years old, christian louboutin made the first pair of shoes in his life, although he made his best effort to promote sales of his shoes, those dancers did not believe him, then he realized that he must undergo a professional training.Then, in 1981, he first apprenticed in the follies bergeres at a friend's referral, and then he learned shoe technology in a prestigious brand company--Charles jourdan, where he made up his deficiencies in the process of shoes-Making.Christian louboutin soon emerged in the industry because of his talent, creativity and self-Confidence.However, at that time, he did not want to join in any corporation, nor did he want to create his own brand.Fledgling christian louboutin had finally developed his own brand in 1992, he produced colorful high heels, exotic, which were identified by the media as he best out of the mainstream?As soon as they were presented, and they received a great concern.Many people thought that the"Red tread?Were symbols of his shoes, which was not true.At the beginning he did not want to wipe the soles red, but every time when he designed shoes, the logo was his headache.And the red color was originally came from his female assistant red nail polish.He thought the red tread would be a big success.Soon, the red tread were popular all around the world, many royal families, especially large stars joined to christian louboutin, which made the shoes famous instantly.Looking through the list of his customers, we can see names like monaco princess caroline, cameron diaz, nicole kidman, sarah jesscia parker, and jennifer lopez.Red soles seem to have fascination, which touches fine women hearts;A woman wearing christian louboutin acts more confident and sexy in front of men.
Alveolar p55 tnf receptor attenuates ventilator btsbydrdreoutletuk Mechanics and blood gases and protein levels in lavage fluid.Was used to determine leucocyte recruitment and Model, while antitnf antibody was ineffective.In the lps+Vili model, p55 blockade prevented deteriorations in respiratory1 on alveolar macrophages, and interleukin 6 and monocyte chemotactic protein 1 levels in lavage fluid. (Ali/ards), although this frequently results in the development of ventilatorinduced lung injury(Vili).1 While the initiating trigger is unique, the final pathophysiology of VILI is indistinguishable from that of ALI of other Aetiologiesthat is, characterised by pulmonary oedema and inflammation.2 3 VILI may be particularly amenable to early pharmacological intervention as its iatrogenic nature means that onset occurs In a predictable manner and within the hospital setting. The proinflammatory cytokine tumour necrosis factor(Tnf)Has been consistently implicated in the pathogenesis of ali/vili, Both clinically and in experimental models.36 Clinical trials beats by dre solo hd of antiTNF therapy in patients with sepsis/ALI710 have unfortunately shown little or no benefits, and in some cases may have led to increased morbidity.The reasons behind Such a lack of benefit are unclear, but are likely to include issues relating to the design of the agents used and also to The complicated biology of tnf signalling.Tnf signals through two cell surface receptors, p55 and p75, and recent evidence Suggests that signalling through these individual receptors may have differentialeven opposingroles in a number of pathological Conditions.1113 We previously found that biologically active TNF is upregulated within the alveolar space early during the development of Vili.14 Moreover, using TNF receptor knockout mice, we found that the two receptors of TNF play opposing roles during VILI:P55 Receptor knockout mice were protected while p75 receptor knockout animals were more susceptible to pulmonary oedema.15 cheap beats by dre solo hd In light of these data, we propose that 'selective' inhibition of 'p55 TNF receptor' signalling, specifically within the Alveolar space, would be clinically much more effective than total tnf inhibition to ameliorate the deleterious effects ofconventional antibody technologies are not optimal for this purpose for a variety of reasons, perhaps the most important of Which being that monoclonal antibodies to this and, in general, other cell surface receptors have a tendency to induce receptor Crosslinking, thereby activating rather than inhibiting signalling.16 17 To investigate the possibility of pharmacological inhibition of p55 signalling, we therefore used a novel IgG fragment known As a domain antibody(Glaxosmithkline, stevenage, uk)That was selected to specifically bind and inhibit mouse p55 tnf receptor.
Command and the website service christian louboutin soldes modules The cSm was designed and built by north american aviaTion at its downey, california plant.The iniTial contract required the delivery of 11 nonflying mockup modEls for configuraTion analysis, 15 boilerplate test vehicles, and 11 flight spacecraft.The flight vehicles evolved into block i, those built without a forward docking facility for use in earth orbit for tesTing and flight operaTions training, and block ii, those with the docking equipment for full lunar operaTions.By early in 1964, the concept of in flight maintenance(Ifm), in which it was envisaged that the crew would carry out repairs to malfuncTioning equipment during flight, was abandoned.Carrying spares and tools involved too high a weight penalty and experience had showed that events during flight emergencies would probably occur and develop into an unmanageable situaTion too rapidly for repairs by the crew to be carried out.Instead, nasa required the main contractors to design in reliability and switchable redundancy whereby malfuncTioning components could be bypassed or isolated during flight and a secondary system used instead.Also by 1964, naa studies showed that the problems inherent in recovery of the spacecraft over land were very nearly insurmountable.ImpacTing the ground without the use of retro rockets was almost certainly going to injure the crew.In april 1964 the decision to recover theCommand moduleby parachute with a splashdown in the sea and recovery by us naval vessEls was accepted and approved by nasa.DifficulTies and delays were experienced by both naa and grumman with the ever changing requirements of the mission concept as it evolved.Naa design for the block ii spacecraft was being hindered by grumman's delay in settling on a final design for the lm and in parTicular for the docking arrangement between the two spacecraft.For the first four months of 1964 grumman led a mission study together with nasa, naa and mit to invesTigate the exact requirements of the hardware from lift off to splashdown.This produced the design reference mission(Drm), a three volume document specifying the exact requirements by which contractors would design the hardware.Unresolved problems with the design of both block i and block ii spacecraft brought about a change in north american's design team when paup was replaced by dale myers as north american's apollo program manager.Naa relaTionship with nasa became strained at Times, especially during the first years of the spacecraft's development.Reports of poor workmanship and safety pracTices troubled nasa project officials and the situaTion came to a head in 1965 when nasa apollo program manager, general sam phillips undertook a review and wrote to north american poinTing out various shortcomings in their procedures.Improvements were made, but on 27 january, 1967, the first block i, manned flight, apollo spacecraft caught fire on its launchpad killing its crew gus grissom, edward white and roger chaffee.The inquiry that followed was unable to pinpoint the exact cause of the fire but its report highlighted deficiencies in north american's design, workmanship and quality control.The apollo 1 fire forced north american to carry out a complete revision of theCommand moduledesign removing as many sources of flammable material as possible and incorporaTing fireproofing features to electrical wiring and connecTions, all of which delayed the start of producTion.Harrison storms stood down from the apollo project, as did nasa's joseph shea who felt a personal responsibility for the disaster.Storms was replaced by william d bergan and shea by george low from nasa headquarters, who took a demoTion to step into the posiTion of aspo manager.Bergan replaced key north american staff and introduced a system of individual teams for each spacecraft.This pracTice was also adopted by the lm's manufacturer, grumman who also were forced to review the lm's design.Command module(Cm)The cm was the real heart of the apollo spacecraft as it contained the three man crew, navigaTion and environmental systems for the overall flight and was the only part of the complete stack to return intact to earth.It was conical in shape with a blunt, convex base, measuring 12 feet 10 inches in diameter and 11 feet high and weighed 12, 500 pounds when unloadedthe body of the spacecraft was constructed in a double layer separated by a thermal insulaTion layer.An inner pressure shell was formed from a lightweight, double skinned, alloy matrix and an outer skin, made from a honeycomb steel alloy doubled up as a heat shield and micrometeorite protecTive layer.The cm shell was separated into three main compartments, the forward, crew and aft compartments and the distribuTion of the craft's weight was carefully calculated to provide an offset to its centre of gravity.This induced a natural stabilising force during reentry and provided aerodynamic lift.AtTitude control by the onboard computer to make adjustments to its flight trajectory was obtained through the rcs and made use of the craft's lifTing body characterisTics.Two individual rcs systems were uTilised by the cSm.The system uTilised in theService modulewas manufactured by the marquardt company of california, and incorporated Small 100 pound thruster rockets, clustered in groups of four(Quads), in two separate subsystems which were spaced equidistant around the circumference of the craft. TheCommand moduleused a separate system designed by Rocketdyne, which, of necessity, was flush fitTing with the spacecraft's outer surface to remove protruding equipment that would be damaged by the aerodynamic forces generated during reentry.The two systems, when used in various combinaTions with each other controlled the pitch, roll and yaw of the combined cSm and to make minor adjustments to the crafts speed.Both rcs systems used the same hypergolic fuEls and pressurisaTion as the Sps engine.Forward compartmentthe forward compartment surrounded the forward docking tunnel and contained the earth landing system(Els).This equipment, for use during the earth reentry phase, was protected by a heatshield, which was jetTisoned after passing through the thermal interface of reentry.The els provided the deployment of two drogue parachutes of 16.5 feet diameter by pyrotechnic mortars at about 24, 000 feet alTitude to stabilise and slow the latter part of the descent.At 10, 000 feet the drogues would be discarded and three main descent parachutes deployed by mortars to complete the descent to splashdown.Three inflatable bags were also housed within the forward compartment to right the spacecraft if it overturned at splashdown.Stable 1 was the desirable upright floaTing condiTion(Apex up)And stable 2 the inverted posiTion.Two reacTion control motors were also housed in the forward compartment to provide pitch atTitude control at the forward end of the spacecraft.Crew compartmentthe interior crew compartment provided a living space of 210 cubic feet, Similar to the capacity of a large family saloon car and was the only part of the cm to be pressurised with an internal atmosphere.During take off the crew occupied three couches situated side by side and facing the main control and instrument panel.The main control panel carried the majority of the flight controls, switches, circuit breakers, warning lights and alarms required to control and monitor the spacecraft's performance during major manoeuvres.After take off the central couch could be collapsed to provide more room to move about the interior and gain access to the lower equipment bay situated centrally under the couches.Two hatches allowed access to the crew compartment.The main side hatch, twenty nine by thirty four inches, was used for crew ingress and egress prior to and after the mission.The second, forward hatch, at the apex of the cm cone, allowed access to the thirty inch diameter docking tunnel, docking probe and drogue and to the lunar module when docked.Valves within the hatches enabled the crew to vent atmosphere to space or to equalise internal pressures between spacecraft.The crew were provided with five windows in the cm.One nine inches diameter in the side hatch, two windows, thirteen inches square either side of the hatch, used for observaTion and a further two triangular windows, angled to face forward, thirteen by eight inches, used during docking.All windows were constructed with triple panes with the outer pane nearly threequarters of an inch thick.Each pane was treated to filter infrared and ultraviolet light and could withstand temperatures of 2, 800 degrees fahrenheit.Removal of the centre couch provided access to the lower equipment bay and the guidance navigaTion and control system(Gncs)With its three main subsystems, the apollo guidance computer(Agc), the InerTial Measurement Unit(Imu)And the opTical alignment system(Ops).The opTical system was centrally placed under the centre couch posiTion.The crew were provided with two dsky's, one in the lower equipment bay for inputs from the ops and a second housed in the main instrument panel.CommunicaTion equipment, also housed adjacent to the agc provided voice, television and telemetry communicaTion with the manned space flight network(Msfn)And between command and lunar modules when separated during the landing and rendezvous manoeuvers.CommunicaTion with the msfn was through a high gain, sband antenna, mounted at the aft end of theService moduleand consisted of four, 31 inch diameter dishes mounted on a folding arm which was deployed after launch.Alignment of the antenna was controlled by the agc through the communicaTion system.The crews vital signs were monitored through the biomed monitoring system.A harness with sensors taped to the astronauts skin supplied heart rate, respiraTion and ekg informaTion via a conTinuously linked informaTion channel.The environmental control system(Ecs)Situated in the left hand equipment bay, below the left hand couch, monitored and controlled the cm's internal atmosphere, pressure and temperature and controlled the temperature regulaTion of internal electronic equipment.Manufactured by the hamilton standard co, it also controlled water producTion through the service module's fuel cells for cooling electronic equipment and supplied hot and cold potable water for the crew's consumpTion.Up to launch the internal pressure was maintained just above atmospheric, with a mixture of oxygen and nitrogen.During launch this pressure was bled away to be replaced by an internal atmosphere of pure oxygen at 5 psi and maintained at 75 degrees fahrenheit.The ecs also monitored the condiTion of the internal atmosphere and filtered out carbon dioxide from the crew's exhaled breath by the use of replaceable lithium dioxide christian louboutin platforms online filters.Control switches and circuit breakers occupied the bulkheads to either side of the couches.Bays and cupboards around the walls of the crew compartment and under the couches also housed all the equipment and supplies that would be necessary for the needs and comfort of the crew.Lockers on each side of the lower bay housed food, clothing, cameras, medical kit, personal hygiene kit and survival gear.These bays also provided storage space for the sample return containers(Src)That would house the moon rock samples on the return journey.Food packages consisted mainly of dehydrated food in bags that could be reacTivated by the addiTion of hot or cold water from the ecs water spigot in the left lower equipment bay.Individual meals of over 70 different items were supplied for the mission duraTion, each idenTified for day and crew member.A further snack pantry was provided to supplement the meals and prevent the raiding of meal combinaTions for favoured items.Waste disposal of urine and water was through a vent to space in the right lower equipment bay, while solid waste products were stowed, in defecaTion bags, in one of the cupboard spaces after treatment.Aft compartmentthe aft compartment and side walls, were divided radially into twenty four bays to house the various systems and consumable for the flight including ten marquardt reacTion control engines with their propellant tanks and helium tanks for pressurisaTion.Water tanks and five, silver zinc oxide batteries for power to supply the cm after jetTisoning the Sm were also stowed here.Three of the batteries powered to the cm's electronic equipment during reentry and two iniTiated the pyrotechnics used for cm/Sm separaTion and parachute deployment.An external umbilical harness connected theService moduleto the aft compartment of the cm to provide services between the two during the mission.On jetTisoning the Sm just prior to reentry, a sequencer deadfaced all electrical circuits between the two modules and closed oxygen, water and other supply lines.SeparaTion of the umbilical was achieved by a pyrotechnic powered guilloTine which cut all interconnecTing links between the two modules and retracted the umbilical arm clear of the cm.Further explosive bolts severed three stainless steel straps which secured the two modules together.The rear, blunt face of the spacecraft was covered by the all important, reentry heat shield.Its funcTion was to safeguard the crew from the 5000 degrees fahrenheit temperature generated by fricTion through the earth's atmosphere during the thermal interface(Ti).The apollo heatshield was a new design differing from the previous gemini type.Manufactured by the avco corporation of massachusetts, the heatshield consisted of a cover varying in thickness from two and half inches at the centre, to half an inch at the outer periphery.It was constructed from a brazed stainless steel honeycomb matrix whose 400, 000 cells were filled with a fibre glass and phenolic resin ablation material.During reentry the ablation material heated and burned off carrying the heat away from the spacecraft.Service module(Sm)TheService module(Sm), also built by North American Aviation with the CM, provided the motive and electrical power, oxygen and communication facility with earth.Cylindrical in shape, it measured 12 feet 10 inches in diameter and 24 feet 7 inches long and was attached to the cm during the flight.Constructed from one inch thick alloy honeycomb panels and partitioned off by milled aluminium radial beams internally into six longitudinal bays around a central tubular core, it weighed over 51, 000 pounds.The main service propulsion system(Sps)Engine was housed on a thrust structure at the rear of the central core and two helium tanks to pressurise the fuel system took up space at the forward end.Fuel tanks for the sps engine took up four of the bays while oxygen and hydrogen tanks and three fuel cells to provide the spacecraft's main water and electricity supply occupied a fifth.Power cells developed and manufactured by the pratt and whitney aircraft division provided electricity by combining oxygen and hydrogen which also produced water as a byproduct.The water was passed forward through the umbilical connection to the cm's environmental system to be used for cooling electronic systems and as potable water for the crew's consumption.The remaining sixth bay, the scientific instrument module(Sim)Bay, was left empty until later apollo flights when it carried remote sensing equipment for lunar data collection from orbit.The sps, a restartable, constant thrust, rocket engine, manufactured by aerojet general, provided 20, 500 pounds of thrust and was gimballed for directional thrust to provide the orbital and speed changes and mid course corrections to the spacecraft's trajectory.The engine was to be used for all large velocity changes of the spacecraft after the sivb third stage had been jettisoned.This would include braking the craft into a lunar orbit and when the time came, pushing it out of that orbit again into the homeward trajectory.The engine's firing time and alignment was controlled by the stabilisation and control system(Scs), a subsystem of the GNCS in the command module.
Heel height an economic indicator Inside the 1920s, lowheeled flapper footwear gave method to highheel pumps and platforms during the good depression. Within the 1970s oil crisis, platforms came back en vogue because the lowheeled sandals on the late 1960s have been cast aside. Images:Images:Images:In the 1990s, the low, thick heels in the"Grunge"Period have been replaced by"Sex and the city"Inspired stilettos just because the dotcom bubble burst. "Usually, in an financial downturn, heels go up and stay up as shoppers turn to a much more flamboyant fashions as a suggests of fantasy and escape,"Says trevor davis, a customer item professional with 'international business enterprise machines' worldwide organization services unit, within a press release. Ibm carried out a study of social media posts and is predicting that women's heel heights, that are presently in nosebleed territory, are poised to come back down michael kors taschen online to earth. Possibly the shift signals a modify within the economic outlook, or perhaps it's a sign of resignation. "This time, some thing distinctive is happening perhaps a mood of longterm austerity is evolving among buyers sparking a wish to decrease ostentation in iaelorstasen every day settings,"Davis says. Ibm came to its conclusion by using special application to sort through social media posts about footwear trends.A number of separate sorts have been created, initially broadly, and then narrowing down to bloggers that are passionate about footwear and have massive followings.The outcome is a single you won't locate inside the shoe retailer.Right now, shops are nevertheless carrying skyhigh heels, however the shift to flatter shoes is around the way. In actual fact, discussions about escalating heel height peaked toward the finish of 2009, after which declined following that.Involving 2008 and 2009, one of the most influential bloggers wrote regularly about heels that ranged in height from 5 to eight inches, but by 2011 they had been writing about the return of the kitten heel plus the fantastic flat from designers jimmy choo and christian louboutin. This isn't to say that the skyhigh heels are gone.Alternatively, davis says, these soaring heels are discussed as glamwear and not for the office or possibly a buying trip. Based on ibm, this study also underscores that companies and retailers can not feel that they're the ones in handle of exactly where the trends head.Bloggers have turn out to be influential tastemakers in their own suitable.
Alpha inhibits adipogenesis via beats by dre outlet uk a beta chpdrebtshdphones CorrESponDEnCE:Jk SEtHi, DEpArtmEnt oF CliniCAl BioCHEmiStry, univErSity oF CAmBriDGE, ADDEnBrookE'S HoSpitAl, Box 232, HillS roAD, CAmBriDGE CB2 2qr, uk.InDEED, impAirED ADipoGEnESiS mAy ContriButE to tHE pAtHoGEnESiS oF oBESityASSoCiAtED ConDitionS inCluDinG Insulin rESiStAnCE, HypErlipiDAEmiAAndtypE ii DiABEtES.ADipoGEnESiS involvES tHE ACtivAtion oF A CASCADE oF trAnSCription FACtorS, wHiCH CoorDinAtE tHE ExprESSion oF GEnES rESponSiBlE For ADipoCytE FunCtion.1 ADipoGEniC Stimuli inDuCE tHE rApiD ExprESSion oF tHE trAnSCription FACtorS CCAAT/EnHAnCErBinDinG protEin(C/EBp)AnD C/EBp, wHiCH StimulAtE A trAnSiEnt StAGE oF mitotiC ClonAl ExpAnSion DurinG EArly ADipoGEnESiS.1 C/EBPAnD C/EBpAlSo promotE tHE ExprESSion oF tHE mAStEr ADipoGEniC trAnSCription FACtorS C/EBPAndpEroxiSomE proliFErAtorACtivAtED rECEptorGAmmA(PpAr).ToGEtHEr tHESE ACt CoopErAtivEly to rEGulAtE tHE ExprESSion oF GEnES ASSoCiAtED witH tErminAl DiFFErEntiAtion.1 LoCAlAndEnDoCrinE FACtorS mAy rEGulAtE ADipoGEnESiS By moDulAtinG tHE ACtivity oF tHESE trAnSCriptionAl EvEntS.2In ContrASt, inHiBition oF ADipoGEnESiS By TNF mAy BE morE rElEvAnt to pAtHoloGiCAl ConDitionS.ADipoGEnESiS iS oFtEn impAirED in DiSorDErS ASSoCiAtED witH ElEvAtED proDuCtion oF tnF, SuCH AS in CACHExiA, SEpSiSAndin oBESityrElAtED Insulin rESiStAnCE.7 THE lAttEr iS BASED on tHE HypotHESiS tHAt rEStriCtion oF ADipoSE tiSSuE ExpAnDABility mAy ContriButE to oBESityrElAtED HypErlipiDAEmiAAndSuBSEquEntly lEADinG to lipotoxiCityinDuCED Insulin rESiStAnCE.8 THE AntiADipoGEniC ACtionS oF TNF HAvE BEEn primArily AttriButED to SiGnAlS trAnSDuCED By TNF rECEptor1(TnFr1), 4, 9 tHrouGH wHiCH TNF StimulAtES numErouS SiGnAllinGPAtHwAyS.THESE inCluDE tHE ACtivAtion oF SpHinGomyElinASES, CASpASES, jnk, Erk1/2AndnuClEAr FACtorkAppA B(NFB).EACH oF tHESE HAvE BEEn impliCAtED in tHE nEGAtivE rEGulAtion oF ADipoGEnESiS.9 AltHouGH it HAS BEEn SuGGEStED tHAt ACtivAtion oF NFB iS nECESSAry For TNFinDuCED AntiADipoGEnESiS, 10, 11 tHiS ConFliCtS witH tHE rECEnt FinDinG tHAt NFB ACtivity inCrEASES DurinG ADipoGEnESiS.12 HEnCE, tHE nAturE oF tHE FinAl AntiADipoGEniC SiGnAllinGPAtHwAy(S)InDuCED By tnFr1 rEmAIns inComplEtEly unDErstooD.HErE, wE ConFirm tHAt tnF inHiBits ADipoGEnEsis By prEvEntinG tHE EArly inDuCtion oF ppArAndC/EBp.HowEvEr, tHis EFFECt CoinCiDEs witH EnHAnCED ExprEssion oF sEvErAl AntiADipoGEniC GEnEs tHAt ArE Also tArGEts oF tHE wnt/CAtEnin/tCEll FACtor 4(TCF4)PAtHwAy. THErEForE, wE invEStiGAtED wHEtHEr TNF inHiBitS ADipoGEnESiS By ACtivAtinG tHE CAtEnin/TCF4PAtHwAy.Our rEsults sHow tHAt tnF EnHAnCEs tHE trAnsiEnt CAtEnin/tCF4 trAnsCriptionAl ACtivity DurinG EArly AntiADipoGEnEsis. BloCkinG tHiSPAtHwAy viA StABlE knoCkDown oF CAtEnin or ovErExprESSion oF DominAntnEGAtivE TCF4 (DntCF4)AttEnuAtEs tnFinDuCED AntiADipoGEnEsis. THErEForE, wE propoSE tHAt TNF inHiBitS ADipoGEnESiS viA A CAtEnin/TCF4 DEpEnDEntPAtHwAy.Top oF pAGErEsultssHorttErm ExposurE to tnF is suFFiCiEnt to inHiBit ADipoGEnEsis in 3t3l1 CEllsto iDEntiFy tHE EArliEst ADipoGEniC EvEnts tHAt ArE impAirED By tnF wE First invEstiGAtED tHE impACt oF limitinG tnF ExposurE to tHE First 48 H oF 3t3l1 ADipoGEnEsis.FiGurE 1A sHows tHAt tnF mEDiAtEs tHE sAmE AntiADipoGEniC EFFECts wHEtHEr it is prEsEnt For tHE First 48 H oF ADipoGEnEsis or tHrouGHout tHE 8DAy ADipoGEniC proGrAmmE.THE 48 H tnF trEAtmEnt siGniFiCAntly prEvEntED tHE ExprEssion oF Ap2 tHAt oCCurrED DurinG normAl ADipoGEnEsis(FiGurE 1B).THis is ConsistEnt witH tHE prEviously rEportED EFFECts on F442A prEADipoCytEs13AndsuGGEsts tHAt tHis tnF ACtion is not CElltypE spECiFiCAndAntiADipoGEniC EFFECts oCCur EArly DurinG tHE inDuCtion pHAsE oF tHE ADipoGEnEsis.ConFluEnt 3t3l1 prEADipoCytEs wErE inDuCED to DiFFErEntiAtE As DEsCriBED in mAtEriAlsAndmEtHoDs.CElls wErE trEAtED witH or witHout tnF(2 nG/ml)EitHEr tHrouGHout tHE DiFFErEntiAtion proCEDurE(Full)Or For tHE First 48 H AFtEr inDuCtion(48 H). (A)At 8 DAysPostinDuCtion, tHE ExtEnt oF ADipoGEnEsis wAs AssEssED By stAininG For lipiD ACCumulAtion witH oil rED o.THE DAtA ArE rEprEsEntAtivEOF FivEinDEpEnDEnt ExpErimEnts. (B 3t3l1 CElls wErE inDuCED to DiFFErEntiAtE witH or witHout tnF(2 nG/ml For tHE First 48 H)AnD totAl rnA wAs ExtrACtED At 0, 2, 4, 8, 12, 16, 24, 48, 96, 144And192 HPostinDuCtion.TrAnsCript lEvEls oF Ap2(B), PPAR2(C), C/EBP(D)AnD C/EBp(E)WErE DEtErminED By usinG rEAltimE pCrAndnormAlisED to 18s rrnA.OF tHrEE inDEpEnDEnt ExpErimEnts.StAtistiCAl siGniFiCAnCE is inDiCAtED As Follows:CompArED to Control trEAtmEnt At tHE sAmE timE point(PppFull FiGurEAndlEGEnD(240K)To invEstiGAtE FurtHEr tHE tEmporAl CHArACtEristiCs oF tnFinDuCED AntiADipoGEnEsis, wE AnAlysED ExprEssion oF sEvErAl kEy ADipoGEniC trAnsCription FACtors.In Control CElls, ADipoGEniC inDuCtion siGniFiCAntly inCrEAsED tHE ExprEssion oF ppAr2 From 8 HPostinDuCtion onwArDs,AndinCrEAsED C/EBp ExprEssion wAs DEtECtED witHin 4 HPostinDuCtion(FiGurE 1CAndD).THus tHE inCrEAsED ExprEssion oF tHEsE kEy ADipoGEniC trAnsCription FACtors oCCurs EArliEr tHAn HAs BEEn prEviously rEportED. 14, 15 TNF trEAtmEnt ComplEtEly prEvEntED tHE inDuCtion oF BotH PpAr2AnD C/EBp(FiGurE 1CAndD).ADDitionAlly, tnF supprEssED tHE ExprEssion oF C/EBp to BElow prEinDuCtion lEvEls, rEDuCinG it By ApproximAtEly 60% At 8 H postmDi(FiGurE 1D).In ContrAst, tnF DiD not AltEr tHE ExprEssion oF C/EBp, An inDuCEr oF C/EBpAndppAr(FiGurE 1E).THEsE DAtA sHow tHAt tnF mEDiAtEs its AntiADipoGEniC EFFECts EArly in tHE ADipoGEniC proGrAmmE.TnF EnHAnCEs tHE trAnsiEnt ExprEssion oF AntiADipoGEniC mEDiAtors DurinG tHE EArly stAGEs oF AntiADipoGEnEsisto invEstiGAtE How tnF supprEssEs C/EBpAndppAr2 ExprEssion, wE AnAlysED tHE ExprEssion oF trAnsCription FACtors known to inHiBit tHEsE ADipoGEniC GEnEs.GAtA2AndGAtA3 supprEss ppAr By DirECtly BinDinG to its promotEr,AndForCEDExprEssion oF GAtA2or GAtA3 inHiBits ADipoGEnEsis.16 In Control CElls, BotH GATA2AndGATA3 mRNAs wErE DownrEGulAtED witHin 2 HPostinDuCtion.HowEvEr, nEitHEr oF tHEsE mrnAs sHowED AltErED ExprEssion in tnFtrEAtED CElls(FiGurE 2AAndB).SimilArly, tnF stimulAtEs tHE sustAinED ExprEssion oF CmyC DurinG AntiADipoGEnEsis in tA1 prEADipoCytEs17AndForCED ExprEssion oF CmyC prEvEnts ADipoGEnEsis By inHiBitinG tHE ExprEssion oF C/EBp.18 ADDitionAlly, BotH CyClin D1AnD ppArinHiBit ADipoGEnEsis By supprEssinG PPAR.19, 20 FiGurEs 2C sHow tHAt tHEMrna expression oF CmyC, CyClin D1AnD PPAR wAs siGniFiCAntly uprEGulAtED FollowinG ADipoGEniC inDuCtion,AndtHis ExprEssion wAs FurtHEr EnHAnCED By TNF trEAtmEnt.At lAtEr timEpoints(4 DAys), only CyClin D1 ExprEssion rEmAinED ElEvAtED in TNFtrEAtED CElls(FiGurE 2D).RnA sAmplEs wErE CollECtED DurinG 3t3l1 ADipoGEnEsisAndAntiADipoGEnEsis As DEsCriBED in FiGurE 1.ExprEssion oF GAtA2(A), GATA3(B), CmyC(C), CyClin D1(D)AnD ppAr(E)WAs DEtErminED By usinG rEAltimE pCrAndnormAlisED to 18s rrnA.GEnE ExprEssionAndstAtistiCAlly siGniFiCAnt DiFFErEnCEs ArE rEportED As DEsCriBED in FiGurE 1Full FiGurEAndlEGEnD(112K)TNF ACtivAtES CAtEnin/TCF SiGnAllinG DurinG AntiADipoGEnESiSIntriGuinGly, CmyC, CyClin D1AnD PpAr ArE tArGEt GEnES oF tHE CAnoniCAl Wnt SiGnAllinGPAtHwAy, wHiCH itSElF HAS BEEn SHown to inHiBit ADipoGEnESiS.21, 22, 23 WE nExt invEstiGAtED Wnt10BMrna expression.UnDEr Control ConDitions, tHErE wAs A trAnsiEnt pEAk in wnt10B mrnA lEvEls, wHiCH suBsEquEntly DroppED to BElow prEADipoCytE lEvEls(FiGurE 3A).In ContrAst, wnt10B lEvEls DurinG tnFinDuCED AntiADipoGEnEsis wErE nEitHEr ElEvAtED nor mAintAinED in tHE First 48 H oF tHE proGrAmmE.InstEAD, wnt10B trAnsCript lEvEls wErE siGniFiCAntly supprEssED By tnF trEAtmEntAndonly BEGAn to rEturn to normAl AFtEr tnF wAs rEmovED.ExprEssion oF wnt10B(A)WAs DEtErminED As DEsCriBED in FiGurEs 1And2. (BAndC)3T3L1 prEADipoCytEs wErE Also trAnsFECtED witH tHE TCF rEportEr TopFlAsH As DEsCriBED in MAtEriAlsAndMEtHoDs.THrEE DAys posttrAnsFECtion, CElls wErE inDuCED to DiFFErEntiAtE in tHE ABsEnCE or prEsEnCE oF tnF(2 nG/ml).LuCiFErAsE ACtivity wAs AssAyED 12 H(B)Or 48 H(C)PostinDuCtion.OF FivE(B)Or Four(C)InDEpEnDEnt ExpErimEnts, witH EACH ExpErimEnt CArriED out in DupliCAtE. (D, EAndF)ConFluEnt 3t3l1 prEADipoCytEs wErE inDuCED to DiFFErEntiAtE witH or witHout tnF(For tHE First 48 H AFtEr inDuCtion)AndCytoSoliC protEin wAS ExtrACtED At tHE inDiCAtED timESPostinDuCtion. ExprESSion oF CAtEnin, AP2,AnD ppArwAS tHEn AnAlySED By WEStErn BlottinG.TimE 0 H protEIns wErE ExtrACtED BEForE DiFFErEntiAtion wAs inDuCED.THE p85 suBunit oF pi3k wAs usED As A loADinG Control.OF tHrEE inDEpEnDEnt ExpErimEnts. (G)CAtEninMrna expression DurinG ADipoGEnEsisAndtnFinDuCED AntiADipoGEnEsis.TotAl rnA wAs ExtrACtED As DEsCriBED in FiGurE 1. (H)3T3L1 prEADipoCytEs ExprEssinG CAtEnin s45A or An Empty vECtor wErE inDuCED to DiFFErEntiAtE witH MDI. THE ExtEnt oF ADipoGEnESiS At 8 DAySPostinDuCtion wAS ASSESSED By StAininG For lipiD ACCumulAtion witH oil rED OFull FiGurEAndlEGEnD (243K)WE nExt invEStiGAtED wHEtHEr tHE TNFinDuCED uprEGulAtion oF CmyC, CyClin D1AnD PpAr rEprESEntED A morE SpECiFiC EFFECt oF TNF on CAtEnin/TCF trAnSCriptionAl ACtivity uSinG A TCFDEpEnDEnt luCiFErASE rEportEr, TOPFLASH. TNF EnHAnCED TOPFLASHDEpEnDEnt luCiFErASE ACtivity At 12 HPostinDuCtion oF ADipoGEnESiS (FiGurE 3B) But not At 48 HPostinDuCtion (FiGurE 3C).THis is ConsistEnt witH tHE EArly But trAnsiEnt EnHAnCEmEnt oF wnttArGEt GEnEs DurinG AntiADipoGEnEsis(FiGurE 2C to invEstiGAtE wHEtHEr tnF mEDiAtEs its EFFECts on tCFDEpEnDEnt GEnE ExprEssion viA stABilisAtion oF CAtEnin, CytosoliC protEin lEvEls wErE ExAminED. AS SHown in FiGurE 3D, TNF DiD not AFFECt tHE lEvElS oF CytoSoliC CAtEnin DurinG tHE FirSt 48 HPostinDuCtion.HowEvEr, At lAtEr timE points(>48 H), CAtEnin protEin lEvEls wErE mAintAinED in TNF trEAtED CElls(FiGurE 3EAndF).In ContrAst, tnF DiD not AltEr tHE ExprEssion oF CAtEnin mrnA DurinG AntiADipoGEnEsis(FiGurE 3G).HowEvEr, tHE ConstitutivE ExprEssion oF A stABlE CAtEnin mutAnt(CAtEnin s45A)24 Also inHiBitED ADipoGEnEsis(FiGurE 3H).THis suGGEsts tHAt tnFinDuCED stABilisAtion oF CAtEnin mAy ContriButE to tHE AntiADipoGEniC ACtions oF tnF. CAtEnin knoCkDown EnHAnCES 3T3L1 ADipoGEnESiS AnDAttEnuAtEs tnFinDuCED AntiADipoGEnEsisTo ADDrESS tHE rolE oF CAtEnin in TNFinDuCED AntiADipoGEnESiS, wE GEnErAtED A 3T3L1 CEll linE in wHiCH ExprESSion oF CAtEnin iS StABly knoCkED Down By ECtopiC ExprESSion oF A SHort HAirpin RNA (SHrnA)SpECiFiC to CAtEnin.THE ExprEssion oF CAtEnin mrnAAndprotEin wAs siGniFiCAntly rEDuCED in tHEsE CElls(FiGurE 4AAndB).THE ADipoGEniC potEntiAl oF tHis CEll linE AppEArED similAr to Control prEADipoCytEs, witH BotH DiFFErEntiAtinG ComplEtEly in rEsponsE to Full inDuCtion CoCktAilAndwitH no morpHoloGiCAl DiFFErEnCEs AppArEnt At DAy 8(FiGurE 4C).HowEvEr, wHEn suBjECtED to suBmAximAl inDuCtion, tHE CAtEnin knoCkDown prEADipoCytEs ExHiBitED EnHAnCED lipoGEnEsis(FiGurE 4C)AndConSiStEntly ExprESSED HiGHEr lEvElS oF PpAr2AnD C/EBp(FiGurE 4DAndE).SurprisinGly, CAtEnin knoCkDown CElls Also ExprEssED morE ppAr2 mrnA wHEn inDuCED witH Full mDi(FiGurE 4D).THEsE rEsults inDiCAtE tHAt knoCkDown oF CAtEnin EnHAnCEs ADipoGEnEsis in 3t3l1 prEADipoCytEs. FurtHErmorE, tHEy SuGGESt tHAt CAtEnin mAy normAlly limit PpAr2AnD C/EBpExprESSion in mAturE 3T3L1 ADipoCytES.3T3L1 prEADipoCytEs wErE trAnsFECtED witH vECtors EnCoDinG SHrnAs AGAInst FirEFly luCiFErAsE(Control)Or murinE CAtEnin(SHrnA). (A) TotAl RNA wAS ExtrACtED From ConFluEnt ControlAndCAtEnin SHrnA 3T3L1 prEADipoCytESAndExprESSion oF CAtEnin mRNAWAs DEtErminED By usinG rEAltimE pCrAndnormAlisED to 18s rrnA. (B)CytosoliC protEin was ExtraCtED From ConFluEnt 3t3l1 prEaDipoCytEs ExprEssinG Control or CatEnin sHrnaAndExprEssion oF CatEnin protEin was analysED by wEstErn blottinG.P85 was usED as a loaDinG Control.THE rEsult sHown is rEprEsEntativE oF Four inDEpEnDEnt ExpErimEnts. (C ControlAndCatEnin sHrna CElls wErE inDuCED to DiFFErEntiatE witH sErum only(CosmiC CalF sErum CCs), Insulin(Ins), IBMX or MDI. THE ExtEnt oF ADipoGEnESiS At 8 DAySPostinDuCtion wAS ASSESSED By oil rED O StAininG (C)Or ExtraCtion oF total rna FollowED by rEaltimE pCr analysis oF ppar2(D)AnD C/EBp(E)Mrna expression. (F ControlAndCAtEnin SHrnA CEllS wErE inDuCED to DiFFErEntiAtE witH MDI in tHE prESEnCE oF tHE inDiCAtED ConCEntrAtionS oF TNF, wHiCH wErE mAintAinED in tHE ADipoGEniC mEDiA For tHE FirSt 48 HPostinDuCtion only. THE ExtEnt oF ADipoGEnESiS At 8 DAySPostinDuCtion wAS ASSESSED By oil rED O StAininG (F)Or ExtraCtion oF total rna FollowED by rEaltimE pCr analysis oF http://www.teamskyuser.co.uk/beats-by-dr-dre-pro.html ppar2(G)AnD C/EBp(H)Mrna expression.OF tHrEE inDEpEnDEnt ExpErimEnts.The results in(C)And(F) ArE rEprESEntAtivEOF tHrEE inDEpEnDEnt ExpErimEnts. For EACH inDuCtion trEAtmEnt, StAtiStiCAlly SiGniFiCAnt DiFFErEnCES BEtwEEn ControlAndCAtEnin knoCkDown CEllS ArE rEportED AS FollowS: PppFull FiGurEAndlEGEnD(353K)THE rEquirEmEnt For CAtEnin in TNFinDuCED AntiADipoGEnESiS wAS invEStiGAtED By inDuCinG ControlAndCAtEnin knoCkDown CEllS to DiFFErEntiAtE witH MDI in tHE prESEnCE oF A rAnGE oF ConCEntrAtionS oF TNF. FiGurE 4F SHowS tHAt 2 nG/ml TNF ComplEtEly prEvEntED lipiD ACCumulAtion in BotH ControlAndCAtEnin knoCkDown CEllS.However, 1ng/ml tnf was less effective at inhibiting lipid accumulation in catenin knockdown cells. AnAlySiS oF PpAr2AnD C/EBpmRNA ExprESSion ConFirmED tHAt TNF inHiBitED tHEir ExprESSion in A ConCEntrAtionDEpEnDEnt mAnnEr (FiGurE 4GAndH).However, in catenin knockdown cells, the concentrationdependent expression of c/ebp(Figure 4h)Was rightshifted.This indicates that tnf may regulate c/ebp expression in a catenindependent manner.Catenin knockdown promotes apoptosis during tnfinduced antiadipogenesisa striking observation made during our experiments was that catenin knockdown preadipocytes were more sensitive to tnfinduced cell death during adipogenic treatment. THiS oBSErvAtion wAS ConFirmED By SuBSEquEnt ASSESSmEnt oF ADHErEntAndviABlE CEll numBErS (At 24 h).In control cells, tnf had no significant effect on cell viability during antiadipogenesis(FiGurE 5AAndB).In contrast, in catenin knockdown cells, tnf significantly reduced the number of viable cells(FiGurE 5AAndB), suggesting that TNFinduced cytotoxicity may be enhanced in the absence of catenin.Consistent with this, tnfinduced dna fragmentation was markedly enhanced in the catenin knockdown cells(FiGurE 5CAndD).
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