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How are the nation's top quarterbacks progressing? Each week, ESPN RISE will track the performances of the 12 QBs who attended the 2009 Elite 11 quarterback event.
(Results based on games through Sept. 20)
Bishop Carroll (Wichita, Kan.)
College: Oklahoma
Bell provided a little of everything in getting Bishop Carroll off to a 3-0 start. He opened the game with a one-yard TD run, and in the 35-12 victory over South went 26-of-39 for 281 yards with two touchdown passes.
Next Game: Sept 25 vs. North (Wichita, Kan.)
Story Link
Joe Boisture
Saline (Saline, Mich.)
College: Michigan State
It was another tough outing for the future Spartan in a 40-7 loss to Bedford. Boisture connected for a 92-yard touchdown on the first play, but Saline was unable to build any momentum. He finished 7-of-22 for 162 yards.
Next Game: Sept. 25 vs. Monroe
Story Link
Robert Bolden
St. Mary's (Orchard Lake, Mich.)
College: Penn State
Contained in a tough loss two weeks ago, Bolden helped St. Mary's rebound with a victory over FAB 50 ranked De La Salle Collegiate. He ran for 89 yards and didn't need to provide much through the air in the team's first win.
Next Game: Sept. 26 vs. Detroit Jesuit
Story Link
Tyler Bray
Kingsburg (Kingsburg, Calif.)
College: Tennessee
Bray put together another stellar performance, throwing for three scores and 338 yards. The future Volunteer has Kingsburg off to a 2-0 start and will look to keep things going this weekend against Corcoran.
Next Game: Sept. 25 vs. Corcoran
Barry Brunetti
Memphis University School (Memphis, Tenn.)
College: West Virginia
In another crucial out-of-state matchup, Brunetti led Memphis University by land and air on the way to a 20-15 victory over Olive Branch (Miss). The dual-threat QB rushed for 71 yards and threw for 136 and two TDs.
Next Game: Sept. 25 vs. Memphis East
Story Link
Devin Gardner
Inkster (Inkster, Mich.)
College: Michigan
Gardner ran for a key score and two-point conversion as the Vikings got their first victory, 14-7 at perennial Cleveland-area power St. Edward.
Next Game: Sept. 25 at Highland Park
Jake Heaps
Skyline (Sammamish, Wash.)
College: BYU
Heaps was a little off his game in a 28-25 loss to Oaks Christian and fellow Elite 11 QB Nick Montana. Heaps was picked off three times and finished 13-of-31 for 256 yards and a touchdown.
Next Game: Oct. 2 at Newport (Bellevue, Wash.)
Story Link
Austin Hinder
Steamboat Springs (Steamboat Springs, Colo.)
College: Cal
It was the defense that stole the show in a 49-6 victory over Montezuma Cortez. The future Golden Bear chipped in two touchdown passes and 146 yards. He also ran for a score.
Next Game: Sept. 25 at Eagle Valley (Gypsum, Colo.)
Story Link
Nick Montana
Oaks Christian (Westlake Village, Calif.)
College: Washington
Montana helped Skyline outlast Oaks Christian 28-25 in a head-to-head Elite 11 QB matchup versus Jake Heaps, completing 21-of-34 attempts for 261 yards and two scores.
Next Game: Oct. 2 at Venice
Story Link
Jesse Scroggins
Lakewood (Lakewood, Calif.)
College: USC
Scroggins was on point again, throwing for three TDs and getting Lakewood off to a 2-1 start. In a 35-7 victory over Mayfair, the USC commit was 18-of-24 for 243 yards without an interception.
Next Game: Sept. 25 vs. Verbum Dei (Los Angeles)
Story Link
Phillip Sims
Oscar Smith (Chesapeake, Va.)
College: Alabama
Oscar Smith stayed undefeated and Sims was able to overcome a poor start in a 42-0 win over Kings Fork. Despite being intercepted for the first time this year, Sims was good for 219 yards and TD on 17-of-27 passing.
Next Game: Sept. 25 at Grassfield (Chesapeake, Va.)
Story Link
Chandler Whitmer
Downers Grove South (Downers Grove, Ill.)
College: Illinois
Whitmer was able to rally DGS after a mistake-filled first half. He hit 11-of-19 passes for 141 yards and two TDs while leading his team to a 37-22 win over Downers Grove North.
Next Game: Sept. 25 at Willowbrook (Villa Park, Ill.)


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Kinky hot devil Lana S sucks a strong cock ardently
Description:Blond head in red corset and stockings is surely a devil. This blondie is so horny and spoiled that she's an expert in pleasing a hot human's cock. Kinky and ardent girlie sucks the tool with delight. Besides this devil assistant desires to experience a real poking from behind.
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Ged Tutors in Harleysville, PA
Find Private & Affordable GED Tutoring in the Harleysville Area!
Temple University - Bachelors, Public Health , Temple University - Current Grad Student, Public Health - Epidemiology
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Recent Posts
Tag Archives: politics
See Politics Post Art
See politics, post art. Make sure that comma is in there because otherwise it sounds like an invitation to watch politics create art. The phrase came to mind one day when I realized that my artist friends were more likely … Continue reading
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Super Bowl Ads 2017 Sell Success Carefully
The game is over. As usual, I skipped the game and watched the ads. I'll assume some team won the game and some team lost the game. Congratulations. Sorry to hear that. The ads are entertaining, but I watch them … Continue reading
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Presidents Of Countries And Companies
Okay. I give in. I'll write about the President, and I'll try to bring it back around to personal finance because that's the purpose of this blog. So much weird stuff has happened since Inauguration Day that it takes a … Continue reading
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Pay Bills Do Laundry
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Adapting To Singularities
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We had the time of our Lives. The memory of our magical time at my hen weekend at Dar Jaguar with my best friends will be with us all forever… We lost ourselves in a stylish dream of 1001 Nights. Everything was organised beautifully for us and was the perfect mixture of indulgence, relaxation, culture, adventure and fun.
Hello Girls!
I am so excited to share our stylish, sassy and sophisticated Girls' Getaways with you!
In my work as a red carpet and wedding dress designer in London, Brides tell me all the time that they can't find anywhere original or distinctive to celebrate their hen with their best girlfriends. Especially if they don't want it to involve stretch white limos, pink bunny ears and sweaty nightclubs.
Well that's no longer true…
"Sex and the City" inspired and reminded us of the fun and importance of precious girlfriend-time and I wanted to create something really special at Dar Jaguar in Marrakech to allow for just this… The perfect, exotic, beautiful place to explore and adventure, shop til' you drop and relax with your best gals.
Whether you are wanting to be seduced by the souks , enthralled by exotic new experiences like camel riding, belly dancing and excursions to the desert, or you just want to be spoiled and indulged with a sensual massage by our candlelit rose filled fountain, we will create the PERFECT HEN CELEBRATION just for you. And I haven't even mentioned our stunning "1001 Nights Banquet" on the rooftop terrace…
For these reasons and more, Dar Jaguar has established a sparkling reputation for designing and hosting some of the most spectacular hen party packages in Marrakech. The riad is the perfect size, location and layout for Hen Party Accommodation, groups or friends wishing to celebrate.
But don't take our word for it, have a look at the gorgeous photos, our "MoroccanRoll" film and all the lovely testimonials…
Sound good? You can have this little jewel all to yourselves with four members of staff, for only € 90 per person per night based on a group of ten friends.
We look forward to hearing from you soon and creating the best celebration imaginable!
With love and fragrance,
Lucia Silver x
Exclusive Use of Dar Jaguar for a long weekend (or more)
Accommodation for 10 Friends in five luxury rooms (more accommodation can be arranged if required)
Four Professional Staff at your personal service, including a talented cook and night security
A delicious, fresh Breakfast every morning on the rooftop terrace
A Private Cooking Lesson with a shopping trip and a 3 course
An exotic Full Day Excursion out of Marrakech to the mountains, waterfalls, the desert or the seaside
Camel Rides and sand dune buggying!
A Private Professional Guided Tour of the best of the souks and the finest cultural and historical sites
An hour's Full Body Massage or Facial
A Private Group Belly Dancing Lesson
A Private Group Yoga Lesson
Henna Tattooing for those wanting to return with an exotic imprint…
Evenings organised to get the best out of the Nightlife and Marrakech party scene
Our renowned "1001 Nights Banquet" with Magical Dar Jaguar Entertainments
For further details regarding your Hen Weekend in Marrakech, please email us at firstname.lastname@example.com.
"We had the time of our Lives. The memory of our magical time at my hen weekend at Dar Jaguar with my best friends will be with us all forever… We lost ourselves in a stylish dream of 1001 Nights. Everything was organised beautifully for us and was the perfect mixture of indulgence, relaxation, culture, adventure and fun."
In August, my friends and I were invited over by the Dar Jaguar riad, owned by British-born designer Lucia Silver, to experience their hen weekend package. She not only offered us a fabulous place to stay, but put on an incredible itinerary to keep us entertained and inspired throughout our trip. When you visit a place more than once it's very easy to end up doing the exact same things the second (or in my case, third!) time around so it was awesome to be presented with some different options.
Lucia is a design obsessive and the riad is decorated in her own imitable and eclectic style. Leopard print wallpaper, four poster beds, golden bathtubs, deep black walls, a bubbling fountain full of rose petals, and a private plunge pool in the middle of the courtyard…! It really is like something out of your most vivid dreams!
…Before heading out to the Palmeraie for something we were all super excited about – camel riding! It was hilarious trying to get all those camels in a line to get a decent photo but so much fun! No trip to Marrekech is complete without a camel ride. On longer visits you can even arrange to do them out in the actual desert, which is a few hours drive from the city.
I was grateful to have Dar Jaguar to head back to at the end of the night though. Lucia even treated us all to hour long full body massages before bed. Wow!
Thank you to Lucia and everyone at Dar Jaguar for putting on such a spectacular few days. If you are looking for a unique hen weekend or honeymoon spot then you must check them out. As well as providing accommodation, they can put on bespoke itineraries for you too, just like we did.
Kat Williams, Rock N Roll Bride
Dar Jaguar, a divine (and divinely intimate) riad in the heart of the souk. Dar Jaguar is owned by Lucia Silver, a couture fashion designer from Notting Hill, and the very definition of a British eccentric. As soon as we met, I knew we were kindred spirits, and this was proven to me over and over again as the week wore on.
Yet another reason we loved staying at Dar Jaguar? We had the whole place to ourselves. Spoiled rotten. Scented oil massages in the courtyard, next to the fountain which was full of roses and floating candles…
Our belly-dancing lesson was absolutely one of the highlights of the trip for me. I really enjoyed trying something unfamiliar and getting out of my own way.
I have to say an ENORMOUS thank you to Dar Jaguar and Lucia for organising such a stellar trip, and I recommend her beautiful riad with my whole heart. I cannot think of a more wonderful place to host a hen weekend or bachelorette party, to have a Marrakech honeymoon, or just escape on a romantic getaway with your beloved… Lucia is a sweetheart with wonderful attention to detail, and you will feel like you are staying in your very own home.
Gala Darling
Never again will I consider spending a fortune on an expensive hen weekend in the cold English countryside. Lucia's Girls Getaway at the stunning Dar Jaguar was the experience of a Lifetime. The ten of us will never forget it – perfect sunshine, THE most beautiful house, brilliant staff, delicious food and the perfect "Sex and the City" itinerary – all arranged for us!
* Photography by Shell de Mar


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The South East has seen shake-ups and departures in senior management at both Lester Aldridge and ASB Law.
Lester Aldridge elected head of the dispute resolution group Michael Giddins as its new managing partner on 19 June.
Giddins beat rival candidates Rachel Lapworth, the firm's head of real estate, and corporate partner Tim Ford to the post.
Head of fast-track Karen Savage, who had announced her intention to stand at the end of last month, withdrew from the race.
The election took place following the decision last month by incumbent Roger Woolley to return to his property practice after six years at the helm. Giddins will take over in November for a three-year term, following a five-month handover period.
Eyebrows were raised across the region last month after ASB overhauled its management structure following the departure of chief executive officer Christopher Honeyman Brown earlier this year (The Lawyer, 29 May).
The non-lawyer chief executive joined the firm in 2001, after ASB was formed by the three-way merger of Argles & Court, Burstows and Stonehams. Honeyman Brown advised on the merger in his role at chartered accountants Horwath Clark Whitehill.
Honeyman Brown oversaw a stormy period in which 11 jobs were cut across ASB's offices and three private client partners left for London firm Goodman Derrick. As first reported by The Lawyer (20 February), Honeyman Brown left earlier this year to pursue "other interests", and has been unavailable for comment since.
In the wake of his departure, ASB lawyers voted to dispense with the chief executive post altogether and instead introduce a new role of managing partner. They also voted to hire a new finance director, who will form a new management team with the senior partner and MP.
Head of commercial litigation Andrew Clinton was elected to the managing partner post for a two-year term on 24 May, while Russell Bell was re-elected as senior partner for a further three years.
The firm expects to appoint a new finance director in the next few months, and had reduced the candidates to a shortlist of five, according to Bell.


text: http://www.flickr.com/photos/38221599@N05/sets/72157626682782189/ I just noticed all 3 of my rackets all have cosmetic issues on them, one obviously worse than the others. Im pretty confident of the answers, but I want to make sure none of them would qualify as serious damage. Specifically, -is the large chip at the throat of the racket between the grommets (the 3 pictures) just of the protective coating and paint, not into the twin tube or anything internal? is there anything more than cosmetic damage right now/in the future caused from it? -is the bubbling by the grommet in one pictures just of the protective coating, not paint? again, is there anything more than cosmetic damage now/in the future caused from it? -will the grommet that is closed in and/or the grommet that has "exploded" because the knot is inside it damage the strings?


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Law-breaking biker accosts reporter filming him, gets what he deserves
Reporter Stanley Roberts and a camera crew went down to the Golden Gate Bridge recently to film "People Behaving Badly" — which could be retitled "Drivers Acting Stupid."
His first instance is a car stopping in the middle of the highway to snap a picture. But what happens next is the crazy thing…
(via Reddit)
Read more: http://twentytwowords.com/law-breaking-biker-accosts-reporter-filming-him-gets-what-he-deserves/
4 responses to "Law-breaking biker accosts reporter filming him, gets what he deserves"
This is a segment on our news in the Bay Area about "People Behaving Badly". It's usually traffic stuff, bike riders being rude, people walking their dogs in places they shouldn't be, etc, etc. This just exploded because they happened to be filming a righteous buffoon who wanted a fight. Usually it's just Stanley commenting on bad drivers that drive us all insane.
"The point", MH, is that the guy on the bike felt the need to be all "I'm a rebel and I'm going to show this TV guy how tough I am" and came over and made a big stink about being videotaped (which is perfectly legal because he's in a public place with no expectation of privacy) – perhaps not realizing that he was in an area that had a lot of security cameras. His harangue apparently went on long enough to attract the attention of other authorities; like the reporter said at the end, if he'd just ridden on nothing would've come of it. Instead, he got busted for all the outstanding things the police wanted to talk to him about…
Sorry if I don't get the spirit of the piece… Sounds like grade school hall monitor or something. What's the point?
Cuz everyone just loves a rat.
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Owned since new?
Owned for how long?
4yrs year(s)
Buy this make again?
Price and equipment6.0/10
A base model with power mirrors & air/con; not a bad deal.
Under the bonnet8.0/10
oil plug & filter at the front with no obstacle's.
How it drives8.0/10
On the firm side with good feed back.
Comfort and practicality8.0/10
Heaps of rear leg room & large boot.
Servicing and running costs8.0/10
The computer was not to advanced;so I could service it,& didnt need a mechanic to stop the service sign on the dash display coming on.
Aesthetics and styling8.0/10
Smooth lines with the front air dam giving it a sporty look on a base model.
Safety and security6.0/10
Remote key was good; & I liked the rear centre lap sash seat belt.
Quality and reliability6.0/10
When things went wrong I found them easy to work out.In the 4yrs I had the VR;engine trans & diff had no major problems.
Overall verdict6.0/10
I sold the VR to upgrade;but wish I kept it as the 2nd car because the live axle gave it,good tyre wear (2mm even tread on all 4 tyres).Boot space was huge,due to spare wheel being to one side,& upright & Level steel floor meant I could carry large heavy loads.Oil changes were easy; no need to jack up car to get,oil tray under neath.Turning cirle was small for a large car.Overall a good towing & family car.


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It redensifies, revitalizes and nourishes the epidermis. With a redensifying and deep moisturizing effect, it relaxes the skin, boosts the natural "collagen capital" providing firmness, elasticity and tone to the skin. Revitalizes and smoothens micro wrinkles. Creamy beauty mask for face with nourishing and revitalizing action formulated with Plant stem cells, Collagen, Vitamins, oils and natural extracts such as Litchi fruit that counteracts tiredness and oxidative stress.


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The grand saga of an American ranching family continues in The Last Ranch, the final, mesmerizing book of New York Times best seller Michael McGarrity's gripping and richly authentic American West trilogy.
When Matthew Kerney returns to his ranch in the beautiful San Andres Mountains after serving in Sicily during World War II, he must not only fight to recover physically and emotionally from a devastating war injury, but he must also battle attempts by the US Army to seize control of his land for expanded weapons testing.
Forced off public grazing lands, banned from gathering his cattle on high mountain pastures, and confronted by military police guarding a high security army post on the northern reaches of the range, Matt finds himself at the center of a heavy-handed government land-grab. The reasons behind this surge of secrecy and control become clear when Matt witnesses the boiling, blinding explosion of the first atom bomb at Trinity Site.
As he struggles with an aging, stove-up father no longer able to carry a heavy load at the ranch, an ex-convict intent on killing him, and a failing relationship with a woman he dearly loves, Matt must draw upon all his mental and physical resources to keep his world-and the people in it - from collapsing.
Following the New York Times best-selling Hard Country and its sequel Backlands, The Last Ranch enthralls with the deeply rich, sometimes heartbreaking Kerney family saga as it steps brilliantly into the mid-20th-century world of the new American West.
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- Allen Anthony
A Great Series
- Sally Filler


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Sonus Festival by Time Warp!20/04/2017
Mark your calendars for August 20-24th this year, because we're going to Pag Island, Croatia!
That's right, it's the five-year anniversary of Sonus Festival, which means more international plauyers from around the world playing tunes of house and techno! This is happening at the Zcre Beach site, where ravers will get to party in paradise!
Huge names are coming to the event, like Richie Hawtin, Ricardo Villalobos, and Tale of Us! Time Warp, the organizers, even announced their phase two lineup, with names like Solomun, Sven Vath, and Adriatique!
I'm sure you all want to get tickets, especially because Croatia is absolutely beautiful! Check out Sonus Festival's website to see how! www.sonus-festival.com/tickets
Then, let us know what you love most about the summer festival season!


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BELOIT, Wis. -- It is with great sadness that we share the news that Ken Hendricks, founder, chairman and chief executive officer of ABC Supply, passed away early Friday morning, December 21. He suffered a massive head injury as a result of a fall at a construction site at his home Thursday evening and died in surgery. He was 66 years old.
This is an enormous tragedy and a great loss to the family, associates, and the community. Right now, we are focused on working through the initial impact of this tragedy with primary focus on the Hendricks family, as we keep them in our hearts, thoughts and prayers.
Funeral services have been set for noon on Friday, Dec. 28, at First Lutheran Church, Janesville, with Rev. James Melvin officiating. Burial will be in Oak Hill Cemetery.
Viewing will be on Thursday, Dec. 27 at the Eclipse Center (Old Beloit Mall) from 3:00 p.m. until the time of a memorial service at 7:00 p.m. and again on Friday at First Lutheran Church from 11:00 a.m. until the time of the funeral service.
In lieu of flowers memorials are suggested to Beloit Regional Hospice. Schneider Apfel Schneider & Schneider Funeral Home is handling arrangements for the family. The on-line obituary and registry can be viewed at www.schneiderfuneraldirectors.com.


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Wired News: Race for Next Space Prize Ignites: "Five to orbit is a significant number; it's the number required to win the next big space prize. America's Space Prize is a $50 million purse established last year by Las Vegas hotelier and, yes, space entrepreneur, Robert Bigelow. Bigelow will award the money to the first U.S. company to build, without government funding, a spaceship that can send five people into orbit twice within 60 days. Bigelow has more than an academic interest in commercial spaceflight; through his Bigelow Aerospace, he's expanding his real estate empire off-planet with the first commercial space stations. While he can launch his stations on existing unmanned commercial rockets, he needs an orbital passenger vehicle to succeed in his venture."
It looks like people and companies are getting into position for the next big economic bang. This time it will be space ventures competing to get off the ground. A number of successful billionaires are running this space rush to orbit. It shouldn't be too much longer for the Moon, Near Earth Objects, and Mars to be within reach.


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Win 7 x64 Home Premium (and x86 VirtualBox VM)/Win10
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latest Windows critical update issue
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When I install the latest Windows update (Cumulative Security Update for Internet Explorer 9 for Windows 7 for x64-based Systems ) and then reboot, it blocks IE9 and makes it freeze while loading, so that I am unable to access the internet. When I restore to a point before the update installation,...
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Can't update to latest USB 3 Driver
Still having doubts my WD Passport is performing at it's optimum on my USB 3 connection. So I went to the ASUS website and downloaded what they say is the latest USB 3 driver for my Laptop's motherboard. Keep getting an error 16 :( and the installation is terminated. Have contacted ASUS...
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EVE: the Kauffman Center's Iconic New Year's Eve event
EVE is your chance to experience the Kauffman Center like never before. Celebrate New Year's Eve in iconic style, enjoying great food and unlimited drink, eclectic and inspiring performances, and so much more!
We are thrilled to give you a small preview of what you can expect from the performers at EVE:
It's going to be one incredible party! We can't wait to see you!
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Barrier dysfunction is pivotal to the pathogenesis of inflammatory bowel diseases (IBD) and collagenous colitis. Glucocorticoids restore barrier function in Crohn's disease, but whether this reflects attenuated inflammation or an epithelial-specific action has not yet been addressed. Using filter-grown Caco-2 monolayers as an in vitro model of the intestinal epithelial barrier, we observed that glucocorticoids induced a time- and dose-dependent increase in transepithelial electrical resistance (TEER) in a glucocorticoid receptor-dependent manner without altering flux of larger solutes or changing principal tight junction architecture. This was accompanied by reduced paracellular cation flux, reduced expression of the pore-forming tight junction component claudin-2, and upregulation of the sealing tight junction protein claudin-4. In contrast, expression of occludin, claudin-1, -7, or -8 was not altered. Dexamethasone increased expression and activity of MAPK phosphatase-1 and inhibition of this phosphatase prevented the glucocorticoid-induced changes in TEER and claudin expression, whereas inhibiting p38 or MEK1/2 was not sufficient to replicate the glucocorticoid effects. Upon exposure to IFN-γ, TNF-α, or IL-1β, TEERs declined in dexamethasone-treated cells but remained consistently higher than in cells not receiving glucocorticoids. Treatment with IFN/TNF resulted in an upregulation of claudin-2 that was significantly attenuated by dexamethasone, whereas increased claudin-2 expression upon IL-1β stimulation was not affected by glucocorticoids. Taken together, barrier augmentation might represent a previously unrecognized mechanism of action, potentially contributing to the therapeutic efficacy of glucocorticoids in IBD and collagenous colitis.
- tight junction
- inflammatory bowel disease
- collagenous colitis
- IL-1β
- claudin
the intestinal epithelium forms the largest mucosal surface of the human body, separating the highly antigenic environment of the intestinal lumen from the milieu intérieur. Integrity of this barrier is of pivotal importance for homeostasis and compromised in a variety of pathological conditions, most prominently inflammatory bowel diseases (IBD) (7, 39, 44). Importantly, IBD-associated barrier dysfunction does not merely reflect the inflammatory epithelial destruction during phases with high disease activity, as increased intestinal permeability precedes clinical relapse in patients with inactive disease (56). Moreover, several studies reported structural and functional barrier abnormalities in IBD during remission and in macroscopically normal mucosa (30, 54). As, in addition, several animal models with targeted barrier defects show either overt inflammation or increased susceptibility to experimental colitis (19, 22, 43), sound evidence suggests a prominent role for barrier dysfunction as a cofactor in IBD pathogenesis.
At the molecular level, IBD-associated barrier dysfunction is linked to increased expression of myosin light chain kinase (MLCK) and elevated levels of phosphorylated MLC (5). In addition, changes in numerous tight junction components have been reported in IBD, with upregulation of the pore-forming transmembrane protein claudin-2 being the most consistent finding both in Crohn's disease (CD) and ulcerative colitis. Conversely, expression of other tight junction proteins with putative sealing properties, such as occludin, claudin-4, -5, and -8, was reduced in some but not all studies (18, 32, 52, 59). Increased paracellular permeability ensuing from these changes has been proposed to result in unrestrained delivery of luminal material to the lamina propria, where an inflammatory response is initiated. Subsequent secretion of proinflammatory cytokines, such as TNF-α, IFN-γ, and IL-13, causes additional damage to the intestinal barrier, forming a vicious circle of barrier dysfunction and inflammation (44). Increased paracellular permeability furthermore contributes to the pathogenesis of diarrhea via a leak-flux mechanism resulting from the passive back leak of ions and water into the intestinal lumen (39), and, of note, the existence of such mechanism has also been proposed in collagenous colitis, a condition with macroscopically intact mucosa but reduced expression of occludin and claudin-4 (9).
Glucocorticoids constitute potent anti-inflammatory agents and are first-line therapeutics for the induction of remission in moderate to severe IBD (4). They effectively control symptoms in collagenous colitis (27), but detrimental side effects prohibit their long-term use. Glucocorticoid therapy restores increased intestinal permeability in CD (55), an observation commonly interpreted to reflect the overall attenuation of inflammation. However, studies in various endothelial and nonintestinal epithelial cell lines provided evidence for the ability of glucocorticoids to stimulate tight junction sealing in the absence of inflammatory stimuli (14, 16, 36, 60). A recent report furthermore described upregulation of multiple tight junction proteins by corticosteroids in immature enterocytes as part of their well-known ability to promote intestinal maturation (23). The aim of our study, therefore, was to investigate whether glucocorticoids regulate barrier characteristics of the intestinal epithelium in the absence of immunological factors using an in vitro model. Our results demonstrate for the first time a direct barrier-stimulating effect of these compounds that is accompanied by changes in ion permeability and claudin expression patterns. Moreover, we provide evidence for a novel signaling pathway involving MAPK phosphatase-1 (MKP-1) to control tight junction permeability in intestinal epithelial cells (IECs), pointing to a previously unrecognized mode of action potentially contributing to the therapeutic efficacy of glucocorticoids in the treatment of IBD and microscopic colitis.
Cell culture.
Caco-2 cells were grown in DMEM containing 10% FBS, 1% nonessential amino acids, 10 mM HEPES, and 1% penicillin-streptomycin and subcultured every 3–4 days before reaching confluency. Cell culture media and supplements were obtained from Life Technologies (Darmstadt, Germany), and cells from passages 10–35 were used for all experiments. IFN-γ, TNF-α, and IL-1β were obtained from Cell Signaling Technology (Danvers, MA) and applied to the basal compartment of the Transwell system after a 35-day differentiation period.
Electrophysiological and permeability measurements.
For measurement of transepithelial electrical resistance (TEER), 1 × 105 cells were seeded on polycarbonate Transwell inserts with a diameter of 12 mm and a pore size of 0.4 μm (Corning Costar, Cambridge, MA). TEERs were recorded using the Millicell system (Millipore, Billerica, MA), and resistances obtained over a blank filter were subtracted from the obtained values. For assessment of Lucifer Yellow or dextran permeability (both obtained from Sigma, Taufkirchen, Germany), Transwell inserts were washed with HBSS followed by apical application of the tracer compounds. After 2 h of gentle agitation at 37°C, basal tracer concentration was measured using a fluorescence plate reader (Perkin Elmer, Rodgau, Germany). Dilution potentials were measured as previously described (8) and used to determine relative sodium-to-chloride permeability according to the Goldmann-Hodgkin-Katz equation.
Cell proliferation assay.
Cell proliferation was determined using the XTT assay (Roche Diagnostics, Mannheim, Germany) according to the manufacturer's instructions.
RNA isolation and quantitative RT-PCR.
RNA was prepared with Trizol (Life Technologies), and, following removal of residual genomic DNA (TURBO DNA-free, Life Technologies), cDNA was constructed using the iScript kit (Bio-Rad Laboratories, Munich, Germany). Quantitative RT-PCRs were carried out on an Mx3005P light cycler (Stratagene, La Jolla, CA) using SybrGreen and EvaGreen master mixes from Fermentas (St. Leon-Rot, Germany) and Bio-Rad as well as primers listed in Table 1. All reactions were carried out in technical triplicates, and a standard curve derived from pooled samples was used to determine primer efficiency and to ensure assay linearity. Reactions lacking cDNA and samples in which reverse transcriptase was omitted during cDNA synthesis served as negative controls.
Reporter gene assays.
The plasmid pGL3/prom.Cl4 harboring the human claudin-4 promoter has been described previously (13) and was kindly provided by Prof. Jean-Francois Beaulieu (Department of Anatomy and Cell Biology, University of Sherbrooke, Canada). Caco-2 cells were stably cotransfected with the reporter plasmid and a linear hygromycin marker (Clontech, Palo Alto, CA) for selection using the ExGen system (Fermentas). Luciferase activity was measured using an assay system from Promega (Mannheim, Germany) and normalized to total protein content as determined by a modified Bradford assay (Roth, Karlsruhe, Germany).
Western blotting.
Cells were lysed in urea buffer supplemented with protease and phosphatase inhibitors as previously described (15). Proteins were separated using tricine SDS-PAGE, and Western Blotting was performed according to standard procedures. Primary antibodies were obtained from Life Technologies [zonula occludens (ZO)-1, occludin, claudin-1, -2, -4, -5, -7, and -8], Cell Signaling Technology (p38, phospho-p38, p44/42, phospho-p44/42), and Santa Cruz Biotechnology [β-actin, MKP-1, and glucocorticoid receptor (GR); Santa Cruz, CA]; horseradish peroxidase-coupled secondary antibodies were from DAKO (Hamburg, Germany). Stripping of Western blot membranes was performed for 30 min at 50°C in a buffer containing 2% SDS and 0.7% β-mercaptoethanol. Densitometric quantification was performed using the AlphaEase software (Alpha Innotech San Leandro, CA) or ImageJ (U. S. National Institutes of Health, Bethesda, MD).
Immunofluorescence was performed on Transwell inserts fixed in methanol at −20°C. After being blocked in PBS containing 3% BSA and 0.2% Triton X-100, filters were incubated with primary antibodies used at a dilution of 1:100 (claudin-2) or 1:200 (ZO-1, occludin, claudin-4) overnight. After repeated washing, fluorophore-coupled secondary antibodies (Life Technologies) were applied for 1 h at room temperature, and cells were washed again in PBS/Triton X-100. Mounting was performed using the ProLong antifade kit (Life Technologies), and images were collected using the AxioVision software on an Axio M1 imager equipped for fluorescence (Carl Zeiss, Jena, Germany).
Electrophysiological and densitometry data were analyzed in SPSS (IBM, Armonk, NY) employing Student's two-tailed t-test for comparisons of two groups and one-way ANOVA with Dunnet and Bonferroni correction for multiple testing when comparing more than two groups. Quantitative RT-PCR data were analyzed using the REST software tool (31). Graphs were prepared using GraphPad Prism (GraphPad Software, San Diego, CA); error bars depict the SE unless otherwise noted.
Glucocorticoids increase transepithelial electrical resistance in a GR-dependent manner.
To test whether glucocorticoid hormones modulate barrier characteristics in IECs, Caco-2 monolayers were treated with dexamethasone, and TEER was recorded to assess paracellular ion permeability. TEER remained relatively constant within the first 3 wk after being seeded both in control and dexamethasone-treated cells, and no differences between groups were observed. However, once Caco-2 cells had undergone complete differentiation, a process known to take up to 21 days in culture (38), cells treated with dexamethasone (0.1–10.0 μM) displayed a dose-dependent increase in TEER that continued to rise until reaching a plateau ∼35 days after being seeded (Fig. 1A). In fully differentiated control cells, resistances also slowly increased with time but were consistently significantly lower than in steroid-stimulated cells. Significant increases in TEER were also observed upon treatment with hydrocortisone (relative TEER after 35 days compared with cells treated with vehicle 1.53 ± 0.10, n = 4, P < 0.01) and budesonide (relative TEER 1.46 ± 0.11, n = 4, P < 0.05, Fig. 1B). This effect depended on activation of the GR, as cotreatment with the GR antagonist RU-486 (10 μM) abrogated the dexamethasone-induced increase in TEER (Fig. 1C). Of note, Western blotting and quantitative RT-PCR revealed upregulation of the GR during Caco-2 differentiation both on the protein and RNA level (Fig. 1, D and E), providing an explanation for the steroid unresponsiveness of these cells before terminal differentiation.
Because it has been established that paracellular permeability toward ions and larger solutes can be regulated independently (40), passage of the fluorescent tracer Lucifer Yellow (LY, molecular weight ∼450 kDa, median diameter 9.9 Å) across a monolayer of fully differentiated Caco-2 cells was quantified. Although dexamethasone significantly increased TEER, permeability of LY was not changed (Fig. 1F). Similar results were obtained with fluorescence-labeled dextran of 4 kDa (data not shown).
Immunofluorescence against the tight junction transmembrane protein occludin and the adapter molecule ZO-1 revealed membranous colocalization of these proteins, and no differences in cell morphology or the subcellular distribution patterns of these proteins were observed between dexamethasone-treated cells and controls (Fig. 2). Similarly, barrier-modulating effects of dexamethasone were not associated with increased proliferation, as the number of viable cells was in fact slightly reduced upon glucocorticoid stimulation as evidenced by the XTT assay (relative number of viable cells compared with cells treated with vehicle alone 0.91 ± 0.01, n = 30, P < 0.001). Taken together, these data suggest that glucocorticoids selectively modulate ion permeability without affecting the passage of larger solutes or changing principal tight junction architecture.
Dexamethasone selectively modulates expression of claudin-2 and -4 without altering their subcellular distribution.
Paracellular ion permeability is determined by the expression pattern of tight junction transmembrane proteins, in particular those belonging to the claudin family (40, 45). We therefore investigated whether dexamethasone altered expression of these molecules. Our analysis focused on claudin-1, -2, -4, -5, -7, -8, and occludin, as changes in the expression of these proteins have been previously reported in IBD (18, 32, 52, 59) or collagenous colitis (9). As demonstrated in Fig. 3, dexamethasone induced a significant reduction in claudin-2 as assessed by Western blotting (relative abundance compared with cells treated with vehicle 0.37 ± 0.07, n = 4, P < 0.005), whereas claudin-4 was significantly increased in these cells (relative abundance 1.91 ± 0.13, n = 4, P < 0.01). In contrast, occludin, claudin-1, -7, and -8 displayed no significant changes upon dexamethasone stimulation, whereas claudin-5 was not detectable in fully differentiated Caco-2 cells.
Consistent with this, quantitative RT-PCR revealed downregulation of claudin-2 (relative expression in cells treated with dexamethasone compared with cells treated with vehicle 0.78 ± 0.05, n = 6, P < 0.01), as well as increased expression of claudin-4 (relative expression 1.25 ± 0.05, n = 6, P < 0.01, Fig. 3C), indicating that dexamethasone at least partly acted on the transcriptional level. This was confirmed in Caco-2 cells stably transfected with a luciferase reporter for the human claudin-4 promoter. Dexamethasone treatment of these cells significantly increased luciferase activity in two independent clones after 30 days (relative reporter activity compared with cells treated with vehicle 1.48 ± 0.09, n = 5, P < 0.01), whereas reporter activation was not changed in cells stably transfected with the empty vector (data not shown). Again, terminal differentiation was required, as no increase in promoter activity was observed before this point (Fig. 3D).
As claudin-2 has been shown to form cation-selective paracellular pores (2, 46, 58), whereas claudin-4 restricts sodium permeability in Madin-Darby canine kidney cells (48), we next investigated whether altered expression of these proteins was associated with changes in ion selectivity. Indeed, paracellular cation flux was reduced in Caco-2 cells treated with dexamethasone as reflected by a highly significant decrease in sodium-to-chloride permeability (PNa+/PCl− 3.21 ± 0.21 vs. 5.32 ± 0.16, n = 3, P < 0.001). This was antagonized by RU-486 (PNa+/PCl− 5.50 ± 0.35 and 5.66 ± 0.07 with RU-486 or a combination of dexamethasone and RU-486, respectively), confirming a role for the GR in the dexamethasone-induced alteration of paracellular ion selectivity (Fig. 3E).
We next assessed the impact of dexamethasone on claudin-2 and -4 trafficking. Indirect immunofluorescence revealed membranous colocalization, with ZO-1 as the predominant staining pattern for claudin-2 although focal cell contacts lacking claudin-2 were frequently found in fully differentiated control cells (Fig. 4A, arrows). Dexamethasone induced an almost complete loss of claudin-2 from the vast majority of cell contacts; however, rarely focal junctions with preserved claudin-2 staining remained detectable (Fig. 4A, arrowhead). Claudin-4 predominantly colocalized with occludin and displayed a more homogenous membrane staining that appeared more intense in dexamethasone-treated cells, indicating that increased abundance of this protein as observed by Western blotting indeed reflected a net gain of junctional claudin-4 (Fig. 4B).
Of note, dexamethasone-induced alterations in claudin expression appeared to intensify changes observed during spontaneous Caco-2 differentiation, as time course analyses revealed a profound reduction of claudin-2 abundance, as well as increased claudin-4 expression in these cells following terminal differentiation (Fig. 4C). Immunofluorescence furthermore demonstrated that this was accompanied by a progressive focal loss of claudin-2 staining from single ZO-1-positive junctions over time (Fig. 4D), whereas the majority of claudin-4 remained homogenously distributed along tight junction strands with Caco-2 differentiation (Fig. 4E).
Barrier effects of dexamethasone depend on MKP-1.
The MAPK phosphatase MKP-1 is an important mediator of the anti-inflammatory effects exerted by glucocorticoids (12). Quantitative RT-PCR revealed a highly significant upregulation of MKP-1 in fully differentiated Caco-2 cells treated with dexamethasone (relative expression compared with cells treated with vehicle 2.66 ± 0.09, n = 6, P < 0.0001) that coincided with the observed increase in TEER (Fig. 5A). Western Blotting confirmed this observation on the protein level (Fig. 5B). As MKP-1 targets the MAPKs p38 and p44/42 (50), we furthermore assessed activation of these pathways. Figure 5C demonstrates that dexamethasone reduced the phosphorylation of both MAPKs without altering the total amount of these proteins, suggesting that increased expression of MKP-1 indeed results in increased enzymatic activity.
We next sought to determine the functional relevance of these findings by employing the MKP-1 inhibitor triptolide (11). As shown in Fig. 5D, triptolide dose dependently antagonized the effect of dexamethasone on paracellular ion permeability with 10 nM of the inhibitor being sufficient to completely prevent the steroid-induced increase in TEER (relative TEERs compared with controls 2.2 ± 0.13, 1.4 ± 0.87, and 1.04 ± 0.066, with dexamethasone and 0, 5, and 10 nM triptolide, respectively). Similarly, coadministration of sanguinarine (10 μM), a structurally unrelated MKP-1 inhibitor with an IC50 of 10 μM (49), resulted in a highly significant reduction in TEER compared with cells treated with dexamethasone alone (relative TEER 1.38 ± 0.10). In line with this, Western blotting revealed that both inhibitors antagonized the steroid-induced changes in claudin-2 and -4 expression (Fig. 5E). Pharmacological inhibition of p38 and p44/42 either alone or in combination using the specific inhibitor compounds BIRB796 (500 nM), U0126 (10 μM), or SB203580 (10 μM; data not shown), however, failed to increase TEER in Caco-2 cells (Fig. 5F), suggesting that blocking of these downstream MAPKs is not sufficient to recapitulate the effects of glucocorticoids on paracellular permeability.
Impact of dexamethasone on the decline in TEER and changes in claudin expression induced by IL-1β or a combination of IFN-γ and TNF-α.
Proinflammatory cytokines play a pivotal role in mediating barrier dysfunction in IBD (44). To assess the impact of dexamethasone on the decline in TEER under these conditions, Caco-2 cells were allowed to differentiate for 35 days in the presence or absence of dexamethasone (10 μM) followed by stimulation with a combination of IFN-γ (10 nM) and TNF-α (5 nM) or IL-1β (10 nM). As shown in Fig. 6A, both of these treatments resulted in a highly significant decline in TEER within 48 h (mean TEER 324 ± 11 Ω/cm2 without cytokines vs. 223 ± 9 Ω/cm2 and 264 ± 10 Ω/cm2 with IFN/TNF and IL-1β, respectively, n = 11, P < 0.001). TEER values were consistently significantly higher in cells treated with dexamethasone (mean TEER 496 ± 10 Ω/cm2 without cytokines vs. 368 ± 12 Ω/cm2 and 401 ± 14 Ω/cm2 with IFN/TNF and IL-1β, respectively, n = 11, P < 0.001 vs. cells receiving these cytokines in the absence of dexamethasone); however, the relative decline in TEER upon cytokine stimulation was similar between groups both with IFN/TNF (relative TEER compared with baseline 0.69 ± 0.025 and 0.74 ± 0.015 for vehicle and dexamethasone, respectively, n = 11, P = 0.14) and IL-1β (0.82 ± 0.02 vs. 0.81 ± 0.02, n = 11, P = 0.59).
Treatment with IL-1β resulted in a highly significant upregulation of claudin-2 after 48 h (Fig. 6B), and, although claudin-2 expression was significantly lower in dexamethasone-treated cells compared with controls, its relative increase in abundance was similar between both groups (2.01 ± 0.11 and 1.80 ± 0.22 in control and dexamethasone-treated cells, respectively, n = 3, P = 0.44, Fig. 6D). Combined treatment with IFN and TNF also increased claudin-2 expression, although to a lesser extent, in control cells (relative protein abundance compared with cells without cytokines 1.60 ± 0.12, n = 3, P < 0.01), which was significantly attenuated by dexamethasone (relative protein abundance 1.13 ± 0.12, n = 3, P = 0.37 vs. dexamethasone-treated cells without cytokines and P < 0.05 vs. control cells with IFN/TNF, Fig. 6D).
In contrast to our observations with claudin-2, IL-1β did not affect claudin-4 expression either in controls or dexamethasone-treated cells (Fig. 6C), whereas combined treatment with IFN/TNF slightly increased claudin-4 abundance in both groups (relative protein abundance 1.45 ± 0.12 and 1.40 ± 0.12 for controls and dexamethasone-treated cells, respectively, n = 3, P < 0.05 vs. cells without cytokines and P = 0.79 for controls vs. dexamethasone-treated cells, Fig. 6D).
Glucocorticoids are the mainstay for inducing remission in IBD and the treatment of collagenous colitis (4, 27). Barrier dysfunction constitutes an important factor in the pathogenesis of these conditions (39, 44), and it has been noticed that steroids restore increased intestinal permeability in patients suffering from CD (55). By demonstrating that glucocorticoids time and dose dependently reduce paracellular ion permeability in Caco-2 cells, we provide novel evidence that this observation might not be solely the result of an overall attenuation of inflammation but could reflect an epithelial-specific steroid effect.
Within recent years, evidence for a differential regulation of paracellular permeability toward ions and larger solutes has emerged. According to this concept, ions traverse the intercellular space via pores formed by claudin proteins, whereas flux of solutes larger than 4 Å occurs via the so-called leak pathway formed by temporary breaks in the tight junction strands (3, 40). Our observation that dexamethasone increased TEER without affecting LY or dextran permeability suggests that the main effect of glucocorticoids in IECs is to reduce flux via the pore pathway. In line with this, Caco-2 cells displayed an altered claudin expression pattern upon dexamethasone stimulation, whereas expression of MLCK and MLC phosphorylation, two major determinants of leak dysfunction in IBD (44), remained unchanged (unpublished observations). This is in contrast to numerous studies demonstrating that glucocorticoids reduce both pore and leak permeability in endothelial and mammary epithelial cells (14, 16, 36). Thus it appears that, depending on the biological context, corticosteroids may act via multiple mechanisms to modulate barrier characteristics. This is further emphasized by the observation that glucocorticoid stimulation of IECs did not alter expression or localization of occludin or ZO-1, as reported in other cell types (14, 36, 41).
Our finding that reduced claudin-2 expression paralleled the increase in TEER is in good agreement with numerous studies demonstrating that this protein increases paracellular conductance by forming pores within the tight junction strands (2, 17, 47). Conversely, claudin-4 forms a sealing tight junction component whose overexpression has been associated with increased TEER in various cell types (46, 48). Therefore, both downregulation of claudin-2 and increased claudin-4 expression constitute plausible mechanisms to account for the increase in TEER observed in our model. Similarly, as previous studies demonstrated claudin-2 to facilitate cation-flux, whereas claudin-4 restricts sodium permeability (2, 46, 48, 58), the reduction in sodium-to-chloride permeability found upon dexamethasone treatment is consistent with changes in claudin expression. Time-course experiments revealed both downregulation of claudin-2 and upregulation of claudin-4 in spontaneously differentiating Caco-2 cells in culture, supporting the idea that the promotion of intestinal maturation by glucocorticoids might not be restricted to immature enterocytes (23). Of note, numerous studies reported overexpression of claudin-2 (32, 52, 59) and downregulation or redistribution of claudin-4 in IBD (18, 29) although evidence for the latter is less unequivocal with other studies failing to replicate these findings (32, 59). Claudin-4 downregulation has also been reported in collagenous colitis (9), raising the possibility that reversal of altered claudin expression might contribute to barrier protection by glucocorticoids in IBD or microscopic colitis.
Although the approach employed in our study enabled us to specifically investigate glucocorticoid effects on IECs, data from animal models demonstrated that, in vivo, corticosteroids also participate in the stress-induced breakdown of the intestinal barrier (26). As this was abrogated by eradicating the gut microflora (42), it appears that glucocorticoids might also have detrimental effects on barrier function by changing characteristics of the intestinal microbiome. Dissecting the molecular pathways mediating barrier protection in IECs is therefore crucial for developing strategies to reverse barrier dysfunction in IBD and microscopic colitis.
In this respect, our study revealed two signaling intermediates involved in the barrier-modulating steroid effects. First, activation of the GR was required, as both increased TEER and altered ion selectivity were antagonized by RU-486. Following the conception that Caco-2 differentiation resembles changes of enterocytes along the crypt-villus axis (37), our observation that the GR was induced in Caco-2 cells with time suggests that the primary glucocorticoid targets in IECs are located apically along this axis. This is supported by in vivo evidence demonstrating increasing GR expression from the crypt to the villus in the small intestine and from the basal to the apical parts of colonic crypts (28, 33). Importantly, claudin-2 preferentially localizes to small and large intestinal crypts in normal mucosa (34), whereas expression in surface colonocytes has been frequently reported in IBD (32, 52). This implies the possibility that glucocorticoids might selectively act on pathologically altered claudin-2 expression patterns in IBD while having less impact on normal mucosa although further colocalization studies will be needed to confirm this hypothesis in vivo.
Apart from the GR, our study identified the MAPK phosphatase MKP-1 as a novel regulator of barrier function in IECs. Although MKP-1 has been attributed an important role in mediating the anti-inflammatory effects of glucocorticoid hormones (12), its impact on tight junction permeability has not been investigated up to now. A recent report, however, demonstrated that knockout of MKP-1 accelerated colitis development in the IL-10 knockout model (25), suggesting a potential role for this molecule in IBD pathogenesis. It has been established that MKP-1 acts by dephosphorylating ERK, p38, and JNK (50). In line with this, we found that dexamethasone reduced phosphorylation of ERK and p38, whereas we were not able to detect significant amounts of activated JNK in Caco-2 cells not exposed to proinflammatory stimuli (unpublished observations). However, whereas changes in TEER and claudin expression were reversed by MKP-1 inhibition, blockade of p38 and ERK either alone or in combination failed to recapitulate the barrier-modulating effects of dexamethasone. Thus, in line with the evolving perception of nonclassical MKP substrates (20, 21), our data might point to MAPK-independent signaling pathways regulating tight junction permeability downstream of MKP-1 in IECs.
Numerous studies have implicated proinflammatory cytokines such as IFN, TNF, or IL-1β in the barrier defect observed in IBD (10, 44). In vitro, these cytokines primarily induce a leak-type barrier defect that is mediated via activation of MLCK and interference with the junctional actomyosin ring (1, 61). This rapid and reversible leak-type barrier loss occurs independently of the claudin expression pattern, and, in line with our notion that glucocorticoids reduce permeability of the pore pathway without affecting leak flux, treatment with dexamethasone was not sufficient to prevent the decline in TEER induced by IFN/TNF or IL-1β. These findings contrast with a previous report demonstrating that glucocorticoids abrogate the increase in IEC tight junction permeability upon stimulation with TNF by inhibiting the upregulation of MLCK (6). The reason for this discrepancy is presently unclear but might relate to different culture conditions requiring different cytokine-stimulation protocols as in the aforementioned study. TNF alone was sufficient to reduce TEER in Caco-2 cells, whereas we and others observed the need for concurrent or sequential stimulation with IFN in this cell line (15, 51). Moreover, it could be speculated that prolonged exposure of IECs to glucocorticoids as in our experimental setting might lead to adaptive changes not observed with a short-term glucocorticoid protocol. Changes of the claudin expression pattern on the other hand are commonly regarded as a means to control the long-term characteristics of paracellular pore permeability (44), and hence it appears conceivable that reduced expression of claudin-2 and increased claudin-4 abundance resulted in significantly higher TEERs upon IFN/TNF and IL-1β stimulation in dexamethasone-treated cells. Generally, the role of proinflammatory cytokines in regulating claudin expression is not well understood and appears to be highly dependent on the cell type and the experimental protocol used (15, 24, 32, 53). In this regard, overexpression of claudin-2 has been most consistently observed upon stimulation of IECs with IL-13 (18, 32, 53). However, as opposed to HT29 or T84 cells, IL-13 failed to elicit any effect on barrier function in the Caco-2 line in our hands, which is in line with a previous report (35) and precluded us from testing the impact of dexamethasone on this phenomenon. However, we here report the novel finding that IL-1β also induces claudin-2 expression in IECs as previously described for rat hepatocytes (57). Whereas the functional significance of this observation remains to be established, it is noteworthy that the relative increase in claudin-2 expression upon IL-1β stimulation was similar in glucocorticoid-treated cells and controls and that its induction by IFN/TNF was significantly attenuated by dexamethasone, suggesting that the signaling pathways regulating basal, IFN/TNF-, and IL-1β-mediated claudin-2 expression differ in their sensitivity to glucocorticoid modulation.
Taken together, by describing an epithelial-specific barrier modulating effect of glucocorticoids, our data suggest a novel previously unrecognized mode of action potentially contributing to the efficacy of these compounds in the treatment of IBD and microscopic colitis. Further study of the molecular mechanisms involved may contribute to the identification and development of pharmacological strategies to restore impaired intestinal barrier function that lack the detrimental side effects of conventional steroids.
A. Fischer is supported by the "Friedrich C. Luft" Clinical Scientist Pilot Program funded by Volkswagen Foundation and Charité Foundation. This project received additional support from Charité Universitätsmedizin Berlin.
No conflicts of interest, financial or otherwise, are declared by the authors.
Author contributions: A.F. conception and design of research; A.F., M.G., and F.W. performed experiments; A.F., M.G., and U.-F.P. analyzed data; A.F., M.G., and U.-F.P. interpreted results of experiments; A.F. prepared figures; A.F. drafted manuscript; A.F., M.G., F.W., U.-F.P., B.W., D.C.B., and F.T. approved final version of manuscript; M.G., U.-F.P., B.W., D.C.B., and F.T. edited and revised manuscript.
The authors thank Prof. Jean-Francois Beaulieu for providing materials and Cornelia Tanneberger for excellent technical assistance.
- Copyright © 2014 the American Physiological Society


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The folks over at Hydro 74 have a track record of producing some of the most wonderful vector themes. They have really outdone themselves here with a special Chinese propaganda set that we just love.
"Here is a set that I was going to put together for myself, however, they didn't turn out as i would of hoped, thus i'l have to do some super clean versions. I've always enjoyed Soviet & Chinese propaganda. It's rich in history as well as rigid structures. It's a shame that Obey has pretty much killed it for anyone wanting to develop it, but then again, they make some damn good shirts and posters!"
*Source: Hydro 74


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Vigilante Women's Sun Valley Fleece II Jacket - Grey
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- Our model wears a size 8 and is 173cm tall


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Maybe not as you look at it, but I made the same mistake when I had some Thai written on my arm, it looks right to me, upside down to everyone else.
http://www. bonnersmusic.co.uk/ file_store/ 14615.jpg this is the original pic, it looks upside down from my eye view!
wow, you can play guitar with your eyes closed
You remind me of a pullthrough
How very shaggable.
Here's a link to this photo. Just copy and paste!
Here is direct link of this photo
Copy and paste the code below (HTML):
Copy and paste the code below (BBCode):
[URL=https://www.arrse.co.uk/community/gallery/photos/me-playing-guitar.7097/]Me playing Guitar[/URL] by [URL=https://www.arrse.co.uk/community/gallery/authors/punk_trooper.45379/]Punk_trooper[/URL] on Army Rumour Service
Separate names with a comma.


text: I've checked with the big art stores in Vancouver, Opus and Loomis. Neither of them carry Arches Platine. A place in Toronto has it but why the heck should I have to order all the way from there... Does anyone know if this paper exists in Vancouver or surrounding areas?


text:
- Attorney Resource Center
- Assisted Housing Preservation
- HUD Multifamily Housing Preservation
- RHS/RD Housing Preservation
- Basic Guides to Preservation Advocacy
- Public Housing
- Section 3 Program
- Section 8 Housing Choice Vouchers
- Domestic Violence and Housing
- Foreclosure Crisis
- Language Access to Housing (LEP)
- Reasonable Accommodation for Persons w/ Disabilities
- Utility Allowances in Federally Assisted Housing
- Low Income Housing Tax Credit
- Reentry
- Resident Engagement
- Choice Neighborhoods Initiative
- Rental Assistance Demonstration (RAD)
- Affirmatively Furthering Fair Housing (AFFH)
- Assisted Housing Preservation
- Publications
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- Housing Justice Network
- Help for Tenants, Homeowners, & Homeless
- Support NHLP
- About NHLP
March 26, 2012: NHLP Advocacy Leads to New HUD Guidance on Protecting Tenants at Foreclosure Act
Typically, the Federal Housing Administration (FHA) requires a foreclosed FHA-insured property to be vacant when it is conveyed to FHA after foreclosure. HUD provides a form notice for the mortgage lender to notify current occupants of the upcoming foreclosure sale with the statement that "HUD generally requires that there be no one living in properties for which it accepts ownership unless certain conditions are met."
When the Protecting Tenants at Foreclosure Act (PTFA), which keeps leases in place after foreclosure and grants tenants the right to a 90-day notice, was enacted in 2009, advocates grew concerned that the omission of PTFA protections from notice would lead tenants to vacate the property prematurely, in the process unwittingly give up their rights under the PTFA. For the last two years, the National Housing Law Project, as part of a coalition of housing advocates, has urged HUD to revise the notice to ensure tenants are aware their PTFA rights.
Largely reflecting NHLP's concerns, the guidance revised the FHA notice to inform tenants that they have rights under the PTFA to remain in the property after foreclosure. HUD also advised mortgagees that the notice must also alert tenants to additional state and local protections if applicable. Mortgagees are required to use the updated "notice to occupants of pending acquisition" no later than July 1, 2012.


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Earrings are one of those female accessories that can never be ignored. Now earrings are a little hard to pull off as they can make or even break your entire look. If you are a busy woman and you don't have time to put on some makeup then take help from the earrings to cover it up and look amazing. Have a look at some of the classic earrings you should try this season without hesitation.
Simple Studs:
Let's start our list with the simplest but cutest earrings that anyone can wear, the studs. Go for the stoned studs or simply pick the ones you like with your dress. Make sure your pull your hair off your ear to show what you are wearing.
Classic Hoops:
Hoops are not everyone's favorite but if you are among those women who like grabbing attention in a simple and stylish way then hoops should be your first choice. Let your hoops do the talking if your dress is simple and you are not wearing any makeup.
Tribal is the new emerging trend right now but if you think that tribal fashion will not suit you then start off with the tribal earrings. They are getting common among a lot of celebrities as they add a natural feel to any outfit.
You might be thinking that florals can only be worn in spring. This is not true when it comes to earrings as a pair of cute floral earrings can be worn any season of the year. Take a bold step and try the ornate ones to make the statement.
Statement Danglers:
Dangling earrings are very stylish but in order to turn people's heads as you walk by, try statement making ones. You can pick different styles and patterns to look cool such as the hash earrings or asterisk earrings.
Stone Earrings:
A lot of girls love to wear stones and if you are among the ones, then show it off by wearing stylish earrings with stones. There are a lot of cute stone earrings so you are not really out of options with this one. Wearing stoned earrings will quickly amp up your looks and will make you look rich and classy.
These are some cute earring choices you should be wearing this season. Have a look at some more beautiful examples of earrings mentioned below.


text: Clippesby Hall Camping Touring and Holiday Cottages Norfolk Broads. the seaside town Great Yarmouth, we're right at the heart of England's finest county. Se Clippesby Hall i Norfolk, England. Clippesby Hall, a multi award-winning camping and caravan park in the Norfolk Broads National Park with eight different. Châu Âu›; Vương quốc Anh (UK)›; England ›; East Anglia ›; Norfolk ›; Clippesby ›; Clippesby Nhà nghỉ›; Ảnh về Clippesby Hall.


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Results 1 to 19 of 19
May 21st, 2010, 10:04 AM #1Datafeeds = Duplicate Content Penalties?
I need a collective ruling on this one. I was talking with a new merchant yesterday and he vehemently objected to creating a datafeed because he was absolutely positive a datafeed would penalize his site in the Google. I hit the BS button and told him that I've never heard of this before, since there are thousands of merchants profiting from datafeeds.
But, I don't have all the answers so I'm asking for opinions.
May 21st, 2010, 10:05 AM #2
Google's pretty good about knowing whose content is the original one. Especially if all the affiliate sites are linking back to the merchant, with the original content.
That's my take on it, but maybe someone who has worked with a merchant has a different experience.
May 21st, 2010, 10:17 AM #3
If this was true then if you did a search on "Flirty Aprons" your affiliates could rank higher than the merchant. As it turns out the merchant ranks higher and #1 spot, above of amazon even. I also see that "Flirty Aprons" has a product feed. So, I think esnagel is right, Google knows who the original merchant is but I dunno about the other Search Engines.
I once had a merchant who did not want to give out datafeed to everyone, only select affiliates and I think his reason was the same. I didn't promote them heavily and did not pursue to ask for the feed but this merchant no longer has an affiliate program. Don't know if it's related though.
May 21st, 2010, 12:18 PM #4
- Join Date
- January 18th, 2005
- Posts
- 2,973
I agree with others who say that merchants should NOT worry about a duplicate content penalty arising from their affiliates' use of datafeed content.
However, I am aware that for some merchants, the product descriptions in their datafeeds are different from the product descriptions on their web sites, and in some cases the product titles are also different. Usually the differences involve "promotional" language (the datafeed version is shorter and more objective, while the merchant's product description on the site may add more flowery language and subjective comments).
May 21st, 2010, 04:11 PM #5
As well, there is the opportunity to "mix 'n match" merchants, for example, in PopShops, AvantLink & Golden Can. While merchants may not like this, it's one way to create unique shops that don't have duplicate content.Renée
Pay no attention to that woman behind the curtain. -Wizardress of Oz
May 21st, 2010, 04:37 PM #6
Google is smarter than that. I'm no SEO expert, but this strikes me as a bit of a "sky-is-falling" outlook.
With PopShops, he can arrange for the datafeed to be flagged Private, appearing only to those publishers given permission to see it. This will limit the exposure that he perceives to be a risk.Be at peace,
Lass Group Mall - Free parking first hour or with any purchase.
May 21st, 2010, 05:08 PM #7
It wouldn't be too hard to provide 1,000+ examples of merchants who distribute datafeeds to thousands of affiliates, but whose sites have not been negatively impacted by their affiliate's use of the feeds.CUSTOM BANNERS by GRAPHICS CANDY ~ Banner Sets and Website Graphics ~ Professional design, reasonable rates
DESIGNER DOG CHECKS ~ We double-dog dare ya to write one!
May 24th, 2010, 02:36 PM #8
If I talk to this guy again, I'll pass along all this good information. Thank you for the help.
June 8th, 2010, 07:52 PM #9
- Join Date
- December 20th, 2005
- Posts
- 413
June 9th, 2010, 07:53 AM #10
You can do the same with one merchant, depending on what's in their datafeed. If you can select items from different categories (but still related to your page theme), you'll have a unique page, as those items won't appear together on the merchant's site or most likely, an affiliate's site either.
Place some unique copy above/below the datafeed for good measure.Renée
Pay no attention to that woman behind the curtain. -Wizardress of Oz
June 9th, 2010, 11:31 AM #11
I think you guys may be overthinking the definition of "duplicate content", too. If you have a page that is completely blank except for the datafeed content, and so does another affiliate, that is clearly duplicate content. The more "other stuff" you add to the page, the less duplicate content problems you'll have.
Google doesn't look at a paragraph of descriptive text and automatically ding you for duplicate content - if it's on a page that provides other value to the viewer: more paragraphs of text, other images, anything that makes that page unique from the other 1000 affiliates that are using that one paragraph of descriptive text.Daniel M. Clark
Tech Manager
Greg Hoffman Consulting
June 9th, 2010, 01:19 PM #12
If anyone was going to be hit for duplicate content penalities it would be the POD companies like CafePress and Zazzle. Every single product page for a particular type of product has the exact same text description unless the "shopkeeper" has added a custom line or two.
Where merchants get the idea that this is a problem is from the SEO firms they hire that have no real clue about what they are doing and see the affiliate channel as competition and ask for crazy stuff to make the merchant think they are doing their job. SEO companies tend to ask for additions of variables to affiliate urls, they scream about duplicate content from the datafeeds, and what ever else they can find to blame on affiliates. They are usually the ones that come up with the "don't let affiliates use your name on pages" etc.Deborah Carney
TeamLoxly.com BookGoodies.com ABCsPlus.com
June 9th, 2010, 08:48 PM #13
- Join Date
- December 20th, 2005
- Posts
- 413
Renee and Daniel - thanks!
Time to put a few sites through the blender...[URL=http://themoneyforums.com]The Money Forums[/URL]
June 9th, 2010, 10:35 PM #14
As others have said, Google knows who owns the authoritative content. There is nothing for any merchant to worry about in terms of providing a datafeed for Affiliates..
In fact, I'll go so far as to say there was never a duplicate content "penalty" that ever existed from Google.
June 9th, 2010, 11:07 PM #15
I wouldn't want to derail this thread regarding datafeeds, but that's a fascinating suggestion, if that's what you're suggesting?
I would love to hear more.Generate more fake news.
June 9th, 2010, 11:17 PM #16
Yes that's what I'm suggesting. And nothing more to add, just that I don't buy it. You can get filtered to the deepest depths of the results for thin or duplicate content...and I suspect that's what some would refer to as a "penalty"
Google figured out a long time ago how to credit the true and authority source for content, so IMO retailers have nothing to worry about in terms of Affiliate program product feeds.
June 10th, 2010, 11:33 AM #17Renée
Pay no attention to that woman behind the curtain. -Wizardress of Oz
June 10th, 2010, 11:53 AM #18
Yes, you can dare say most In fact how about "no experience whatsoever with merchants, the affiliate marketing model or SEO."
June 10th, 2010, 12:10 PM #19
My comments were actually based on actual discussions with merchants and what their so called SEO firms requested or told them. No speculation on my part at all. Just jaw dropping requests from merchants (that I no longer manage because they never did "get it") that I replied with "no you can't do that" and "no that is not right".... etc.Deborah Carney
TeamLoxly.com BookGoodies.com ABCsPlus.com
Similar Threads
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Results 1 to 4 of 4
01-31-10, 23:49 #1Registered User
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Unanswered: Punctuated text added to list box is split
I need your help in solving why punctuated text gets split and some of it lost when added to a list box. In this instance, two lines are added to the list box instead of one. The split is at the position of the comma. The comma and semi-colon are omitted, all text following the semi-colon does not get into the list box. The code:
Dim txtString As String txtString = "The cabin looked deserted, and half covered with snow; and it would soon be dark." Me.lstScript.AddItem txtString
02-01-10, 06:12 #2King of Understatement
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txtString = """The cabin looked deserted, and half covered with snow; and it would soon be dark."""Testimonial:
pootle flump
ur codings are working excelent.
02-01-10, 13:04 #3Purveyor of Discontent
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pootle's example will take care of you. Google up "ms access escape quotes" if you want to know why.oh yeah... documentation... I have heard of that.
*** What Do You Want In The MS Access Forum? ***
02-01-10, 14:31 #4Registered User
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Thank you pootle flump for your response. I am using Access 2003 and maybe that causes your solution of using quotes like this: """text; more text""" to split the text when using AddItem with a list box, as it did for me without the extra quotes. It appears that the Access AddItem method treats the punctuated text as delimited strings separated by commas or semi-colons.
The solution I am using is a temporary table to hold strings of text. This table is the row source for the list box, and now the text appears intact in the list box.


text: Administrative Support Monday, October 09, 2017 Frederick, Maryland Temporary US$ 13.00 - US$ 14.00 per hour Answer the telephone with the utmost professionalism, urgency, screening, and prioritizing calls ? Email, mail and fax distribution of documents as needed ? Prepare and maintain departmental reports ? Schedule departmental meetings and taking ... see job description


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|25 Jun 2016
Windows 7 Ultimate x64
Babylon dictionary interface problem (Flag icon has gone)
Hello everyone !
I have a problem with my application "Babylon dictionary".
the little flag icon beside any dictionary name has disappeared and there is just a point instead of a flag distinguishing every dictionary.
Here is a screen shot :
Any idea what might be the problem ?
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Windows 7 Boot Problem at Win7 Flag
I have 2 HD's with OS Win7 on both(dual boot). About 3 weeks ago my PSU (Cooler Master Real Power Pro1250-Watt Power Supply - SLI Ready, 80PLUC, 135mm Fan) started making a noise around the fan area. I noticed a lot of dust in the vent and blow it out and the noise went away a couple of days later....
|Hardware & Devices
Windows Live Mail dictionary problem
I have made a mistake spell-checking an e-mail message. I typed "iin" meaning to type "in" -- and then, in spell-check, I accidentally told the dictionary to "Add" the word "iin" (rather than "change" the word to "in"). I guess I have a "heavy finger" because I mistakenly type "iin" a lot, and my...
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delete icon of babylon from firefox
hi guys. i installed babylon.but when i open firefox the icon of it always show n the top right of browser(take a look at picture below).it creats problem for me to add tab.how can i remove that?
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Action Center Flag System Icon cannot be turned on
Taskbar Properties/Notification area/Customize/Turn system icons on or off/Action Center is Off and grayed out. I actually want it showing all the time. Restore default icon behaviors and checking Always show all icons and notifications on the taskbar does not bring it back either. It is...
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There is a problem with the interface in Windows 7.
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What in windows controls the dictionary function of the built in spell checker and what does it effect and how do I configure it? Here's why... The letter " I " as a standalone letter should Never be lowercase in the English language (unless it is by itself as part of a string of letters for a...
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Pumpkin Mousse Cake
- 1 12.3-ounce package Mori-Nu tofu
- 2 c. canned solid-pack pumpkin
- 1 tsp. cinnamon
- ½ tsp. pumpkin pie spice
- 1/8 tsp. sea salt
- ½ c. pure maple syrup
- 1 14.4-ounce box graham crackers
- 1/3 c. pecan pieces
- 1 recipe carob glaze
Carob Glaze:
- ¼ c. pure maple syrup
- ¼ c. carob chips
- In a blender, process the Mori-Nu tofu, pumpkin, spices, sea salt and pure maple syrup until smooth.
- Spray a 12 x 9 inch pan with non-stick cooking spray and layer the ingredients in the order given.
- When spreading the pumpkin mixture on top of the graham crackers make sure to spread it to the edges of pan. Layer graham crackers first, then pumpkin mixture. Repeat twice. Sprinkle with pecan pieces.
- To make glaze: Heat the pure maple syrup in a small saucepan and cook until the syrup is white and creamy. Add the carob chips, stirring quickly until they are melted.
- Remove immediately from heat and drizzle on top of the pecan pieces on the pumpkin cake. Chill in refrigerator several hours or overnight. Cut into 2-inch squares.


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Firstly - it's my birthday! So Happy 41st Birthday to me :) Secondly - that would also mean it's my blog's birthday too - where has two years gone? It's turned into a bit of a monster, this blogging lark... and I wouldn't be without it for the world.
So rather than focus on my birthday - I can do that in real life and blog about it later - for this post I thought it was about time that ALL the extra specially loyal followers I have (you know who you are) were properly recognised. I had a think about this, and a nickname naturally came to me one day when messaging someone on Twitter... it's a rather lovely term of endearment that suits you lot perfectly because I think you're all so great.
So if you're a fan of Not Dressed As Lamb you get to call yourself a special name:
That's right, Lady Gaga has her Little Monsters, Glee fans are Gleeks, One Direction fans are called Directioners (I'm not going to mention that little singing boy from Canada) - and you, my dear readers, are my Little Lammies. All just for fun of course, but why not?
And as it's my two-year bloggiversary, and as I want to say a huge thank you to everyone for their amazing support, I thought I should highlight some of the most loyal readers by giving out the first ever Little Lammie Awards to the following twelve bloggers. I'm truly sorry I can't feature everyone, but I tried to pick just 10 - there were 12 finally. So in no particular order (I love all my Little Lammies equally), here they are.
Rachel - Rachel the Hat
Greetje - No Fear of Fashion
I'm a Little Lammie Button
I hope you'll check out these lovely ladies and that they'll all be proud to call themselves a Little Lammie! Their support - and YOUR support - is greatly appreciated. If you Tweet about this post, please include the #littlelammies hashtag, and not only that, I've done you a button if you want to continue the fun!

(You'll be able to resize the widget to any size you choose - you'll see the size is pretty big at 300x300. Simply change the HTML's width and height to the pixel size of your choice.)
The Little Lammies List
There's more! If you visit the page that the button will link to and get yourself on my new Little Lammies List by leaving me a comment (on that page) with your first name, blog name and URL, I'll add you to the list - the 12 Little Lammies above are already on there of course, I'd love to be able to give a little link love back as a thank you for your support!
Go to this page and leave your blog details: http://www.notdressedaslamb.com/p/little-lammies.html
So that's it - another year over. I think the next one will be even more exciting - DO. NOT. MISS. Thursday's post. A new project is underway (continuing on from my Best Features post), and I want everyone reading my blog to be part of it.
I couldn't be more excited! Plus it's my birthday! Aaaargh!!!!!
*her head explodes*


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Baby, Girls " Santa" Applique White Top & Stripe Ruffle Trousers Outfit (1)
Crafted from a soft cotton and elastane blend, so she"ll be in complete comfort.
-96% cotton, 4% elastan
-machine wash at 30 degrees.
-extra quality.
Customer Reviews
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Oct 25 11:39 a.m.
Article by: ARA Staff
The latest on iPhone running apps Read More
Oct 17 10:24 a.m.
Article by: ARA Staff
"At this point, unless you are likely to have a serious deficiency in overall nutrient intake -- or unless you have a specific indication for taking a multivitamin, such as pregnancy or the intent of pregnancy -- use of a multivitamin seems ill advised." Read More
Oct 17 10:10 a.m.
Article by: ARA Staff
Fauja Singh, 100, becomes the oldest person to complete a marathon. Read More
Oct 12 2:39 p.m.
Article by: New York Times
"Carlos was also dismissive of sprinting's current sensation, Usain Bolt of Jamaica, saying, 'I don't look at him.'" Read More
Oct 05 1:05 p.m.
Article by: ARA Staff
"A second survey, conducted by researchers at Loyola University Medical Center and published in June in the British Journal of Sports Medicine, reached the starker conclusion that almost half of runners may be drinking too much during their races." Read More
Oct 05 12:47 p.m.
Article by: ARA Staff
"Replacement operations increased 100% over the past 10 years and are expected to rise 500% by 2030 , according to the American Academy of Orthopedic Surgeons." Read More
Sep 29 12:20 p.m.
Article by: ARA Staff
"Some parts of their brains showed more activity than others, but in each of the samples, the brain cells held newborn mitochondria. There was no comparable activity in brain cells from the sedentary mice." Read More
Sep 27 3:42 p.m.
Article by: ARA Staff
"'Age for me is just a number'," Mr. Gebrselassie said in a recent interview" Read More
Sep 23 7:33 p.m.
Article by: ARA Staff
Diet not working out? Unrealistic Expectations? Sure, but its not your fault, says a new study. Read More
Sep 21 4:20 p.m.
Article by: ARA Staff
A new iPhone app motivates runners by chasing them with Zombies. Read More