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Plan your formal dance around this popular 19th century novel and movie about the southern US. This guide contains Gone with the Wind prom theme ideas.
I need some ideas for a Gone With the Wind prom theme!
Please help.
you could have a area for pictures with some huge columns like they have on the front of the plantation
homes with a sign that said Tara ,or a fake stairwell .you could make the columns out of cardboard . you could have fans for the girls and tussie mussies for the tables . i guess the carriage and horses would be to much though
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Climate variability, oceanography, bowhead whale distribution, and Iñupiat subsistence whaling near Barrow, Alaska
Ashjian, Carin J.
Braund, Stephen R.
Campbell, Robert G.
George, John C.
Kruse, Jack
Maslowski, Wieslaw
Moore, Sue E.
Nicolson, Craig R.
Okkonen, Stephen R.
Sherr, Barry F.
Sherr, Evelyn B.
Spitz, Yvette H.
MetadataShow full item record
Citable URIhttp://hdl.handle.net/1912/5127
KeywordBowhead whale; Plankton; Oceanography; Beaufort Sea; Subsistence whaling
The annual migration of bowhead whales (Balaena mysticetus) past Barrow, Alaska, has provided subsistence hunting to Iñupiat for centuries. Bowheads recurrently feed on aggregations of zooplankton prey near Barrow in autumn. The mechanisms that form these aggregations, and the associations between whales and oceanography, were investigated using field sampling, retrospective analysis, and traditional knowledge interviews. Oceanographic and aerial surveys were conducted near Barrow during August and September in 2005 and 2006. Multiple water masses were observed, and close coupling between water mass type and biological characteristics was noted. Short-term variability in hydrography was associated with changes in wind speed and direction that profoundly affected plankton taxonomic composition. Aggregations of ca. 50–100 bowhead whales were observed in early September of both years at locations consistent with traditional knowledge. Retrospective analyses of records for 1984–2004 also showed that annual aggregations of whales near Barrow were associated with wind speed and direction. Euphausiids and copepods appear to be upwelled onto the Beaufort Sea shelf during Eor SEwinds. A favorable feeding environment is produced when these plankton are retained and concentrated on the shelf by the prevailing westward Beaufort Sea shelf currents that converge with the Alaska Coastal Current flowing to the northeast along the eastern edge of Barrow Canyon.
Author Posting. © Arctic Institute of North America, 2010. This article is posted here by permission of Arctic Institute of North America for personal use, not for redistribution. The definitive version was published in Arctic 63 (2010): 179-194.
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Coherent patterns in bacterial growth, growth efficiency, and leucine metabolism along a northeastern Pacific inshore-offshore transect del Giorgio, Paul A.; Condon, Robert H.; Bouvier, Thierry; Longnecker, Krista; Bouvier, Corinne; Sherr, Evelyn B.; Gasol, Josep M. (Association for the Sciences of Limnology and Oceanography, 2011-01)We investigated the patterns in bacterial growth, production, respiration, growth efficiency (BGE), and bacterial leucine respiration and C-to-leucine yield (i.e., conversion factor [CF]) along a transect off the coast of ...
Ecological characteristics of core-use areas used by Bering–Chukchi–Beaufort (BCB) bowhead whales, 2006–2012 Citta, John J.; Quakenbush, Lori T.; Okkonen, Stephen R.; Druckenmiller, Matthew L.; Maslowski, Wieslaw; Clement-Kinney, Jaclyn L.; George, John C.; Brower, Harry; Small, Robert J.; Ashjian, Carin J.; Harwood, Lois A.; Heide-Jørgensen, Mads Peter (Elsevier, 2015-09-10)The Bering–Chukchi–Beaufort (BCB) population of bowhead whales (Balaena mysticetus) ranges across the seasonally ice-covered waters of the Bering, Chukchi, and Beaufort seas. We used locations from 54 bowhead whales, ...
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An Insider's View
Housing rebound helps furniture stock values
You cannot read anything without seeing the growing momentum in the housing sector. Just recently, we saw that new home sales reached a five-year high, pending home sales a six-year high and homebuilder confidence a seven-year high.
On a year-to-year basis, housing permits are up 24.8%, starts up 28.6%, housing completions up 20.8%, new single family home sales up 29%, existing home sales up 12.9% and the inventory of homes for sale is down 21.5% - because the demand is exceeding the supply. Hopefully, higher home prices will bring more homes into the market.
The headlines of 2008-2011 are rarely seen with bank foreclosures way down and the equity in most homes is rising, making homeowners feel wealthier with fewer "underwater" on their mortgages.
The usual bears on Wall Street now finally agree the housing recovery is real and sustainable. Professional investors, outside corporations, banks and lenders, private equity group and others are wondering how they can participate in this growth.
The first moves are to invest in the obvious categories like homebuilders, makers of housing components and their retailers. These have all gotten the attention of Wall Street and others as you might expect.
In these categories, there are a lot of large, multibillion-dollar market cap stocks that quickly become the favorites of the larger mutual funds and professional investors who dominate the U.S stock markets. The market caps of homebuilders Toll Bros., Lennar and D. R. Horton are $5.4 billion, $6.4 billion and $6.3 billion. Other large cap investments are Mohawk, Masco and Leggett & Platt at $8.3 billion, $7 billion and $4.3 billion. In home furnishings, some of our largest are Tempur-Sealy, Aaron's, Mattress Firm, Select Comfort and La-Z-Boy at $2.2 billion, $2.1 billion, $1.3 billion, $1.2 billion and $1.1 billion, respectively.
Most of our large funds and investors require a billion dollar market cap before they can make an investment for liquidity reasons. These days, only "small cap" funds can invest in companies with a market cap of less than $1 billion. That would include Ethan Allen, Haverty, Hooker, Flexsteel and Bassett at $851 million, $588 million, $181 million, $180 million and $175 million as of today.
The real story, however, is how much our values have grown. In just the last year, the furniture stocks are up an average of 29.1%. If you had invested $1,000 in furniture stocks in March 2009, you would have $4,006 today.
The net of this is that your business is worth more today and there are more buyers, investors and lenders than we have seen since 2006.
As hard as it is to believe, you are prettier and more attractive today than you have been in many years!


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» Voted By Fresno Bee Readers As The Best Place To Work In Central California
London Properties is a fantastic, unbelievable place to work! Whether you decide to become a Real Estate sales associate or join our highly-skilled support staff, you'll find yourself with the best people, the finest resources and the most positive energizing environment anywhere!
» Voted By Fresno Bee People's Choice Readers As Fresno's #1 Real Estate Company
Click to enlarge
Click to enlarge
Year after year, we sell more homes in our area than any other firm. Our sales volume now ranks us in the top 1% of all Real Estate companies nationwide. In fact, in the past 12 months we have closed over one transaction every 2.9 hours, 24 hours a day, 7 days a week. In 2015, we out-produced over 99.5% of all Real Estate companies in America. Closer to home in 2015, London Properties Associates averaged over 2.43 million dollars in closed volume per sales associate.
In 2015, of all luxury residential properties and land sales in the Valley over $750,000 London Properties participated in more luxury sales than any other broker. In fact, of these sales, London Properties represents more clients that our next two closest competitors combined. Over the years, our V.I.P. clients have found it reassuring to know that London Properties has participated in more luxury home sales than anyone in the valley.
Your new career in Real Estate can give you higher income, more independence, personal growth, more free time, and prestige. The decision you make today becomes your future. For more information, give me a call on my direct private line at 436-4080 or email me.
» Licensing And Training Courses:
We'll guide and assist you through the Real Estate licensing process and even provide post-licensing sales training. Our FasTrac system can put you on the way to a secure new future in just a few weeks.
» Start With The Best
Where you start your new career makes a difference to your success! Because London Properties leads by example, our success makes your success easier. We are ranked in the top 1% of all Real Estate firms nationwide. Our 40 years of outstanding production and stellar reputation make people want to do business with us and because of us, they'll want to do business with you!
» We Make It Easy
London Properties has helped more people start a new career in Real Estate than any other valley Broker. We make the process simple and easy. In Real Estate, bigger is better, and for you means more leads, more marketing support and better results online. London Properties is the Valley's #1 Real Estate Company because we provide our agents the ongoing training, career development, marketing, atmosphere, example and support they need to be successful. Our FasTrac Licensing program will help you secure your license faster so you can start making money sooner.
» Licensing And Training Courses
If you're new to the business, we make it easier than ever before to jumpstart your Real Estate career. If you're an established agent in need of continuing education, we can help you too!
Hi, my name is Wanda Fries.
In my previous career, I was a registered nurse. My career move to London Properties has more than doubled my income. Without a doubt, the best move I've ever made. Want to know how Real Estate could change your life? Call Barbara Gallagher. Her direct line is 436-4080. Or email her at email@example.com.
Bringing Your World To You
London Properties can make your life easier by bringing the world to you. Our climate-controlled, emergency-powered and securitized central computer system instantly delivers your transaction data, forms, emails, communications, marketing systems, and everything you need to do business, directly to your home, your cell or any location anywhere in the world. Now that's wired!
Now You're At One Address
We'll provide you with your own free private email account, allowing you to send mass emails to all 600 associates, managers, branch offices, and support staff with one easy click. We'll even forward all your existing Gmail, Hotmail, AOL, Facebook, and Twitter accounts to one location. No more checking multiple accounts. Everything you need to do business will now go directly to your office, your home, your cell, or any location in the world. We're here for you anytime, anywhere in the world.
Supporting You
Losing money because there's not enough time? Details and paperwork bringing you down? We'll provide you a full-time personal assistant or transaction coordinator on a file-by-file basis to do the detail stuff. Your files will be better organized, your sales will close faster and, best of all, you'll be free to make that next sale. We're here for you anytime, anywhere in the world.
Marketing You
Need help designing and implementing a personal marketing program? London Properties provides our sales associates free counseling from an independent marketing consultant who will design your own personal career-builder program. We'll even produce your new custom marketing materials at discounted prices. Need dozens of FREE postcards to mail with your name and number? We got that! We'll even address and post them for you. Hit up Reliance Marketing, London's in-house, full-service marketing department. Save time - visit reliancemktg.com and design and order your materials online with our custom software. Now you can control your own mailing campaigns from home on the fly. We're here for you anytime, anywhere in the world.
Providing Client Info 24/7
Need easy access to all of your transaction information at home? We got that! With RealTrak, London's proprietary transaction management system, all your important info is available to you 24/7 from anywhere in the world. RealTrak will automatically update your client contact and database and it will keep you in their mind by emailing your marketing material weekly, monthly, or whenever you want. We're here for you anytime, anywhere in the world.
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Need to send an urgent mass email regarding important needs, wants, status changes, price reduction or whatever to all 600 of your colleagues? We got that! All London offices and all our associates are connected through the same email server. You can instantly communicate to the entire company no matter where you are - at the office, at home, or on the road via your mobile device. Need someone to cover your open house? No problem! One quick message will get you a response. We're here for you anytime, anywhere in the world.
Info On The Fly
Need to answer a client inquiry, find the sales price or provide detailed listing data to a client when you're nowhere near your office? We got you covered! With mobilelp.com, simply enter the property address on your cell and you're good to go. Complete MLS information, including photos, will appear like magic. Any listing, any broker, any property, anytime, instantly on your cell. Mobilelp.com has data for every MLS from Merced to Hanford including Oakhurst and mountain communities. We're here for you anytime, anywhere in the world.
Getting It Online
Need to write up a sale or counter at 11 p.m., but no forms are available? We've got you covered 24/7. Simply go to LondonNet, our custom intranet site. You'll find all CAR ZipForms, as well as every disclosure, London Forms, and all the other documents you'll ever need. Print right to your office, home, or anywhere in the world. We're here for you anytime, anywhere in the world.
Bringing The Future To You
Feel overwhelmed keeping up-to-date with all the Real Estate tech stuff? Our full-time tech savant sends out weekly tech tips, attends technology conferences, and visits London offices weekly to demonstrate new tools to make sure you have access to every tech tool available. We'll provide regular workshops on how to use social networks like Facebook and LinkedIn, document signing, and all the new apps. Our tech training is FREE to London associates. We're here for you anytime, anywhere in the world.
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Your computer too slow, crashed, or bugged? We're here to help. Bring your ailing system to one of our in-house computer techs and you'll be back up and running in no time. You know that every London office provides free wireless internet connection, but we'll even install your own wireless connect system at your home. We're the only Real Estate firm in Central California with our own in-house tech team. We're here for you anytime, anywhere in the world.
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Need to keep your clients posted on new listings? We got that! At londonproperties.com you can sign them up for Find-It-First updates. They'll get all the new listings in their mailbox each morning. Not just from one MLS, but from all four major Valley MLS's. Every listing, every broker, every MLS, one easy search. We're here for you anytime, anywhere in the world.
Making You #1
Want a bigger presence online? We have our own in-house programmer to help you design a kickass website. We'll connect you with MLS search capability throughout the Valley, links to any community information, email to your cell and more features than you can imagine. We even provide you a FREE individual website for every one of your listings. We've made londonproperties.com the number one Real Estate site in Central California. More traffic than any other Realtor®. We can do the same for you. We're here for you anytime, anywhere in the world.
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We can all learn from each other, but you'll never find a Real Estate company where you can learn as much as you can at London Properties. Our FasTrac live training isn't just for folks new to the industry. FasTrac is taught by our two Brokers, our managers and our top-producing Realtors®. Experienced London agents frequently attend to get fresh ideas and inspirations. In addition we offer over 45 online classes that you can take from home or anywhere in the world. Get your Real Estate license. Renew your license or simply log on for career development ideas. Each London office has weekly sales meeting, legal updates, and tech training. You will receive tech tips, legal tips, and other important stuff daily via our company wide email intranet. No Real Estate company anywhere gives you more training. Any questions? We're here for you anytime, anywhere in the world.
We Make It Happen
We're a big family and, unlike most big families, our size does not mean you get less attention. You'll actually get more. We divide our associates into squads of 10 or less. Not only will you have more fun playing on a team and challenging other squads but you'll receive more care and encouragement. Your squad leader will be meeting with you weekly to check your commitments and your progress. They're like your mentor, and are available to answer questions and provide constant motivation. In addition, all nine of our Sales Managers and two Brokers are available by cell 24/7 at London Properties. You're never alone. We're here for you anytime, anywhere in the world.
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Why User Experience Design Matters To A Business
Most companies these days have to design something for their customers or users. It can be anything from a menu at a restaurant to the options at a bank ATM machine. While developers, programmers, coders, and IT personnel might be full of ideas and suggestions about how something should look or operate, you should ultimately let the clients that spend money with your business decide on how things operate anytime your business has a technological point of contact with them.
This is called user experience design, and it simply means that all design elements are ultimately focused not just on function, stability, and security, but the actual experience the user goes through when using your interface. This matters not just in touchscreens in brick and mortar stores, but also through your website and your smartphone app, which more and more businesses have on top of their online content. It even matters to some degree in how you set up your auto prompter and menu options on your phone system.
People want things to be convenient, easy, and even enjoyable when possible. When your existing clients feel so comfortable with your designs that they find dealing with your business to be a seamless part of their day, they are not likely to leave you for a competitor, sometimes even when another business might actually be cheaper for them!
Whenever it is time for an overhaul of any customer interface, don't just let your IT department whip something up and then roll it out all over the country. Let managers give it a test drive first. It's probably most efficient for a company to hire an app development company to avoid mistakes and a ton of wasted time. Do small scale tests in limited markets. Pull together focus groups. Most especially, let any of your current employees who are in customer facing roles try it out. They can really tell you whether or not your existing clientele are going to like it or not.
Tips on great user experience design from the pros
Regardless of how old a system is, always have an option to get feedback and survey data from clients and customers on their user experience. Any comments or suggestions that come up repeatedly need to be addressed, regardless of whether or not your IT people actually care or take it to heart. Don't be hard on them though. Engineers sitting in cubicle farms rarely know at heart that people shopping your business are eventually their paycheck. Your customer service folks don't ever forget that though.
How to choose the right UX company for the project
There are so many websites, toll-free numbers, and apps out there these days that consumers flock to the ones that save their time, go easy on their brains, or answer their questions and problems. Be an easy part of a good day, your customer will trust your business and stick with you for a long while. These kind of habitual customers are great repeat business and might stick with you long after alternatives come to the market that might even be better for them. Anytime your company works on software, an app, or a website that customers will engage, make sure that user experience design is the philosophy of the day.


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By now we all know the Sharks' story this season. Started out the shortened campaign with a blistering 7-0 record. Stumbled to a 5-9-6 record since. As of gametime on Wednesday, they sit eighth in the West.
And with this once-proud franchise barely hanging on to a playoff spot for a second season in a row, Sharks GM Doug Wilson is reportedly in "full panic mode," which includes shopping some of the team's underachieving players. Here's a nugget from Bruce Garrioch's report:
The San Jose Sharks GM isn't pleased with the club's inconsistency and he's open to all options. The club's UFA's — LW Ryane Clowe, C Michael Handzus and D Douglas Murray — are available and sources say Wilson "is in full panic mode." Murray is being "shopped hard" but one team backed off because of concerns about his conditioning. League executives believe Wilson is frustrated but he often stirs up discussions then doesn't deal. The Habs, Bruins and Penguins have interest in Clowe.
Although, as Fear the Fin points out, when it comes to these hot rumors, Garrioch is about as credible as a McDonald's salad. So, take this nugget with a grain of salt. Even still, this gossip was just too sexy to not share with you.
H/T Pro Hockey Talk via Fear the Fin


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Skye Maree
Roxy was fantastic right from the very start. I was setting up a new business so I needed everything! We started with a logo, then business cards, website, tax invoices, quotes, letterhead and swing tags. Roxy was really great to work with, she really listened to the concept I was wanted to create, took on board all my ideas and made them come to life on the screen. She always gave me plenty of different designs and colours to choose from. She was very professional and a pleasure to work with. I would reccommend her services to anyone that wants to start a business and get themself set up or wants to upgrade their image to one that is funky, modern and stands out from the rest.
Shane King – Markwell Constructions
From the very first phone call to Rocko Designs to the launch of our website, Rocko Designs DELIVERED. Rocko Designs provided great customer service and advice. Our website receives fantastic compliments. We recommend you phone Rocko Designs to discuss your future website
Murray Damro – Damro Electrical
On starting my electrical business I always knew it was going to hard to be noticed especially on the Gold Coast, you just have to take a drive up the M1 in the morning and notice all the electrical vans heading off to work. So I had heard about Roxy's design talent and contacted her. She was great, we able to sit down together and come up with a design that definitely stands out from the crowd, I personally have had so many great comments about my business cards, web site and van signage that she had designed. My business is starting to take off and I definitely have got Roxy to thank for getting me out there and noticed. I have no hesitation in recommending her for any of your future design needs.
Mandy Bell – The Freedom State
We have been working with Rocko Designs on both of our websites. We found Roxy particularly patient, especially since she had to work her way around a site written in Argentinean! We now have Rocko Designs perform all our website updating, and her advice on how to better market our sites have been invaluable. We now receive a report each month telling us how many people have visited the site, from what search engine and where they are from! We have used 2 other Web design companies before Rocko, and have received more information from Rocko Designs than both other companies put together
Julio de las Heras
Action Rent-a-Car has been in business for 11 years and over that time I have dealt with quite a few web designers, but always felt that I had to do most of the thinking when it came to design and themes. Roxy took control and with a simple brief has just finished our second website and I am realy pleased with the outcome, but more importantly the great feedback we are getting from our clients, even only a week after after it's launch. As we are a growing company with franchises coming on board I am glad to say that I will be using Roxy's expertise for all our future Web requirements.
Raymond Aoude – Abaa Car Rentals
Rocko Designs worked on our web page as if it were her own. We have had nothing but praise from our customers in regards to the quality and material on our website. The work and effort that was put into the set-up of the web site was huge. We continue to receive compliments regularly from our many satisfied hirers about our web site. I would strongly recommend using rocko for your next web design.
Brooke McDonald – Intergreat
I recently worked with Roxy on a logo design and splash page for my business and I found her approach to her work refreshing, creative and catchy! Roxy was succinct in delivering what I asked and punctual adhering to promised timelines – even if it was late at night! Roxy's committed approach meant my expectations were met. I highly recommend Roxy and Rocko Designs as an excellent creative resource for whatever your business needs.
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Being based in Singapore and needing a global presence that would be relevant in Asia, Australia and the Middle East, an innovative website was imperative for Little Red Wagon and as a constantly evolving brand, a significant challenge. Despite the fact that the company is based in Singapore and all of the briefing had to take place over the phone and via Skype and email, Rocko Designs was able to over deliver with a website that exceeded out expectations. Her intuitive needs analysis and collaborative style brought our vision for littleredwagon.com.sg to life and despite having never had the opportunity to see our product in person she was able to effectively capture the essence of our brand. She manages to combine a highly creative flair with a professional integrity that results in both vibrant, interactive sites and the ability to meet deadlines. In addition to creating the site, her attention to detail and real time response to changing retail demands, we have been thrilled with the ongoing, proactive support and e-marketing skills that Roxy has been able to offer – her drive to see our site and brand succeed is as strong as ours. I wouldn't hesitate to recommend Rocko Designs in any capacity and look forward to working with Roxy for many years to come.
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The U.S. Fish and Wildlife Service announced today that it will list the imperiled Gunnison sage-grouse as "threatened" under the Endangered Species Act (ESA), a move needed to prevent the bird's extinction.
A coalition of wolf conservation groups, environmental organizations and a retired federal wolf biologist sued the U.S. Fish and Wildlife Service today for repeated failures over the last 38 years to develop a valid recovery plan for the imperiled Mexican gray wolf, one of the most endangered mammals in North America. With only 83 individuals and five breeding pairs in the wild at last report, Mexican gray wolves remain at serious risk of extinction. The recovery plan, a blueprint for rebuilding an endangered species' population to sustainable levels, is necessary to ensure the lobos' survival and is legally required under the Endangered Species Act.
Twenty one imperiled species of sharks and rays were given added international protections at a recent multi-nation meeting on conservation of migratory species held in Quito, Ecuador November 4-9, 2014. The species include the great hammerhead, scalloped hammerhead, thresher sharks, silky shark, devil rays, reef manta ray and sawfishes. The additions represent a record for the number of shark and ray species listed by the United Nations Environment Programme's Convention on the Conservation of Migratory Species of Wild Animals (CMS).
A new poll conducted by Tulchin Research for Defenders of Wildlife found that a majority of registered voters throughout Colorado, Nevada, Wyoming and Montana support efforts to conserve greater sage-grouse across the West. In the poll, majorities had a favorable opinion of the grouse and supported efforts to protect its sagebrush habitat. The poll also showed that a majority would support listing the bird under the Endangered Species Act (ESA) if current conservation planning efforts fail to protect the species from possible extinction.
Florida voters have passed Amendment 1, which designates billions of dollars for natural resources, wildlife and habitat conservation.
Defenders of Wildlife and the Patagonia Area Resource Alliance today asked a federal court to hold the United States Forest Service and United States Fish and Wildlife Service accountable for their unlawful approval of the "Sunnyside" mineral exploration drilling project in southern Arizona's Coronado National Forest, in the Alum Gulch area of the Patagonia Mountains. The groups say the approval of the Canadian mining company Regal Resources' Sunnyside Project violates environmental laws and poses a potential threat to endangered species and the safety of drinking water for local residents. The Sunnyside Project involves drilling multiple exploratory holes up to 6,500 feet deep in one of the most biologically diverse mountain ranges in Arizona. The extensive drilling and construction would run 24/7 for months on end, and the total project operations could last up to 3 years.
Defenders of Wildlife today announced it has taken legal action against the U.S. Fish and Wildlife Service (the Service) for the agency's decision not to list the wolverine under the Endangered Species Act (ESA).
On Sunday, thousands of sheep will trail through the City of Ketchum. They're the feature attraction of the annual Trailing of the Sheep festival and are on the way to their winter grazing location. These sheep are different than most in one regard: they are part of a program that uses nonlethal methods to protect livestock from wolf predation.
Contaminants from a mine spill in Cananea, Sonora earlier this summer have likely reached the San Pedro River flowing into Arizona. And with recent storms, old copper and silver mine sites near Patagonia are leaking bright red contaminants into local streams. These toxic reminders of our mining history have the potential to wreak havoc on local water supplies and wildlife in the Coronado National Forest, one of the most biologically diverse areas in the world.
Today the U.S. Fish and Wildlife Service (the Service) announced it would provide $900,000 in grants as part of the Wolf Livestock Demonstration Project Grant Program. This program authorizes the Service to administer grants annually for five years to livestock producers in the states of Arizona, Idaho, Michigan, Minnesota, Montana, New Mexico, Oregon, Washington, Wisconsin and Wyoming willing to implement non-lethal deterrents to keep livestock away from wolves.


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Plasmid deficiency in urogenital isolates of Chlamydia trachomatis reduces infectivity and virulence in a mouse model
We hypothesized that the plasmid of urogenital isolates of Chlamydia trachomatis would modulate infectivity and virulence in a mouse model. To test this hypothesis, we infected female mice in the respiratory or urogenital tract with graded doses of a human urogenital isolate of C. trachomatis, serovar F, possessing the cognate plasmid. For comparison, we inoculated mice with a plasmid-free serovar F isolate. Following urogenital inoculation, the plasmid-free isolate displayed significantly reduced infectivity compared with the wild-type strain with the latter yielding a 17-fold lower infectious dose to yield 50% infection. When inoculated via the respiratory tract, the plasmid-free isolate exhibited reduced infectivity and virulence (as measured by weight change) when compared to the wild-type isolate. Further, differences in infectivity, but not in virulence were observed in a C. trachomatis, serovar E isolate with a deletion within the plasmid coding sequence 1 when compared to a serovar E isolate with no mutations in the plasmid. We conclude that plasmid loss reduces virulence and infectivity in this mouse model. These findings further support a role for the chlamydial plasmid in infectivity and virulence in vivo.
- Chlamydia
- plasmid
- mouse
- virulence
- infection
- genome
Chlamydial plasmids are all small [c. 7.5 kb, 8 coding sequences (CDSs)], highly conserved (< 1% variation within the species), nonconjugative, and nonintegrative, and their role in C. trachomatis pathobiology remains to be fully explored (Thomas et al., 1997; Seth-Smith et al., 2009). A long-standing observation has been that naturally occurring plasmid-deficient C. trachomatis are exceedingly rare, implying a significant, but noncritical role in fitness and transmission in vivo.
In vivo studies of the C. trachomatis plasmid have only recently begun and have thus far employed either the plasmid-free human C. trachomatis lymphogranuloma venereum (LGV, serovar L2, strain 25667R) or a plasmid-free trachoma isolate (serovar A, strain A2497P-) where the plasmid has been cured by novobiocin treatment (Carlson et al., 2008; Olivares-Zavaleta et al., 2010; Kari et al., 2011). In ocular infection of lower primates, plasmid-free C. trachomatis strain A2497P- was found to infect, but was significantly attenuated for shedding of viable chlamydiae and inducing inflammatory responses (Kari et al., 2011). In mice inoculated in the lower urogenital tract with a naturally occurring plasmid-free isolate of C. trachomatis LGV, serovar L2, reduced infectivity was seen (Carlson et al., 2008; Olivares-Zavaleta et al., 2010). It is important to note that in these studies, the plasmid-free C. trachomatis strains, while attenuated for infectivity in the mouse urogenital tract (L2, 25667R) or in primate eyes (A, A2497P-), nonetheless were able to elicit immune responses that protected against virulent and fully infectious matched plasmid-containing wild-type strains. This has led some to assert that plasmid-free strains may be effective attenuated vaccines (Olivares-Zavaleta et al., 2010; Kari et al., 2011). Similar results have been reported in the mouse model of urogenital infection with the mouse pathogen, C. muridarum (O'Connell et al., 2007). It is also important to note that none of the aforementioned studies have utilized urogenital isolates of C. trachomatis, and thus, it could be hypothesized that the plasmid of urogenital isolates possesses different qualities and would behave differently in an animal model when compared to the others thus far examined.
In the present study, we have assessed for the first time the infectivity and virulence of naturally occurring plasmid-free and a cocirculating plasmid-containing (hereafter, wild-type) urogenital isolates of C. trachomatis, serovar F, as well as the Swedish new variant C. trachomatis (nvCT; serovar E, with a mutation in CDS1) in a mouse model of urogenital or respiratory infection.
Materials and methods
Chlamydia trachomatis strains used in this study
Four urogenital isolates of C. trachomatis were used in this study; C. trachomatis Sweden 3 and 4 (hereafter, E/SW3 and F/SW4, respectively) were isolated in Malmö, Southern Sweden, in 2002 and are serovar E and F, respectively, representing plasmid-containing wild-type cervical isolates (Seth-Smith et al., 2009). Chlamydia trachomatis Sweden 2 (E/SW2), representing the Swedish nvCT (plasmid-containing serovar E, with a 377-base pair deletion in CDS1), was isolated from a male urethra in Malmö in 2006 (Seth-Smith et al., 2009; Unemo et al., 2010). Chlamydia trachomatis Sweden, serovar F, plasmid-free (F/SWFP−) is a cervical isolate from Malmö, Sweden, in 1995 (Persson et al., 1996; Wang et al., 2012). All isolates were cultured only in McCoy cells with minimal passage number.
Genome sequencing
Extraction of DNA was performed on strain F/SWFP− as previously described (Seth-Smith et al., 2009). DNA was sequenced on the Illumina GAII platform using 54-bp paired-end reads, resulting in a mean genome coverage of 1527 ×. A subset of Illumina reads was assembled using Velvet, version 1.0.12 (Zerbino & Birney, 2008), producing contiguous nucleotide sequences (contigs), which were scaffolded against the genome of strain F/SW4 (EMBL accession no. HE601804) (Harris et al., 2012), using abacas (Assefa et al., 2009). The two rRNA operons assembled into a single contigs, which were manually corrected, as were contig boundaries where possible. Manual insertion of capillary-sequenced hctB and tarp gene sequences was followed by self-mapping with all the sequence reads to confirm the assembled sequence and resulted in an improved high-quality draft sequence (Chain et al., 2009). After this process, the assembly of strain F/SWFP− comprised three contigs. Annotated features were transferred from the closely related strain F/SW4 using annotations_update.pl (https://github.com/sanger-pathogens/annotations_update) on the basis of blastn similarity using the default cut-off of 90% and manually curated using Artemis (Rutherford et al., 2000). Mapping of sequence data to reference genomes was carried out using smalt (http://www.sanger.ac.uk/resources/software/smalt/). Genome comparisons were visualized using the Artemis Comparison Tool (ACT) (Carver et al., 2005). The genome sequence and annotation have been deposited in the EMBL database under the accession number HE605380, and the read data are in SRA under the accession number ERS001400.
We selected an outbred strain of mice (Swiss Webster, SW) for these experiments. Five- to six-week-old female SW mice were obtained from Harlan Sprague Dawley (Indianapolis, IN) and housed with commercial rodent chow and water ad libitum with a 12 : 12 light/dark cycle. All mice were subject to a 7–10-day facility acclimation period prior to inclusion in the study. All studies were approved by the Midwestern University Institute Animal Care and Use Committee (MWU File # 1726).
Infection of mice and assessment of infection by culture
For urogenital infection, mice were pretreated with 2.5 mg of medroxyprogesterone acetate (DepoProvera, P4; Greenstone LLC, Peapack, New Jersey) and 7 days later were inoculated intravaginally (i.vag.) with log10-graded doses of McCoy cell-grown C. trachomatis as described elsewhere (Ramsey et al., 1999). To verify and track urogenital infection, cervical–vaginal swabs were collected in all mice at day 4, 7, 10, and 14 postinfection in each experiment. Chlamydia trachomatis was isolated inclusions visualized by indirect immunofluorescence and quantified as inclusion-forming units (IFU) in HeLa 229 cultures as previously described (Ramsey et al., 1999).
For respiratory infection, mice were anesthetized via inhalation of 4–5% isoflurane and placed in dorsal recumbency, and log10-graded doses of the strains were inoculated dropwise into the nasal nares exactly as described elsewhere (Ramsey et al., 2009). Mice were weighed daily, and weight gain or loss from baseline weight was recorded as a measure of morbidity. It should be noted that none of the inoculated mice exhibited any overt morbidity from the infection, and there were no mortalities following infection. Hence, lethal dose that induces mortality in 50% of the inoculated mice (LD50) was not determined.
Antibody responses
For both urogenital and respiratory inoculations, we have found it useful to assess seroconversion postinfection as a means to assess infection at levels below the level of detection of our culture methods (e.g. at low dose infection). For this purpose, blood was collected at day 0, just prior to infection, and 35 or more days postinoculation, and total plasma anti- C. trachomatis immunoglobulin G (IgG) levels were determined by enzyme-linked immunosorbent assay (ELISA) as described previously (Ramsey et al., 1989, 2009). An antibody titer ≥ 40 was considered positive for the determination of seroconversion. Immunoglobulin G antibody responses were also compared as a measure of elicitation of adaptive immunity and as an indirect measure of the degree of infection (greater infectious burden correlating to a greater antibody response). For quantitative comparison of antibody responses, titers were converted to log10, and the arithmetic mean of all seropositive animals was determined as described elsewhere (Ramsey et al., 1988).
Infection course data, as assessed by shedding of viable organisms (lower genital tract) and weight change over time (respiratory tract), were compared by analysis of the variance (anova, treatment group, days) with repeated measures. Results for ELISA and IFU counts at individual time points were compared using a two-tailed unpaired t-test. The incidence of viable chlamydial shedding and seroconversion was assessed using Fisher's exact test. Infectious dose that yielded a 50% of mice infected (ID50) was estimated by the method of Read and Muench (Reed & Muench, 1938).
Plasmid deficiency in a C. trachomatis urogenital isolate correlates with reduced infectivity in the mouse urogenital tract
In our first experiment, we intravaginally inoculated groups of 10 mice with log10-graded doses of either F/SW4 or F/SWFP− C. trachomatis and monitored outcomes of infection. Table 1 shows the results of the study. In both wild-type and plasmid-free inoculated groups, low numbers of viable organisms were shed intermittently through day 10 postinfection, but no animals in either group remained culture positive by day 14. The wild-type F/SW4 isolate had an estimated ID50 of 3.2 × 105 IFU. We were unable to infect 50% of the mice with the plasmid-free isolate using our highest dose of 106 IFU – despite a second experimental attempt using 7 additional mice (data compiled in Table 1). Thus, we assert only that the ID50 of the F/SWFP− isolate (as determined by culture) to be in excess of 106 IFU. In addition, at the 106 IFU dose, the day 4 and day 7, IFU isolation attempts indicated a greater infectious burden in animals inoculated with the F/SW4 isolate (P = 0.01 and P =P 0.03, respectively, by two-tailed unpaired t-test). As we have previously observed, seroconversion provides a more sensitive method of determining infection below the level of detection of culture (Ramsey et al., 2009). Thus, we were able to use seroconversion to achieve an ID50 for both groups with the plasmid-free isolate also displaying reduced infectivity (ID50 ~ 17-fold higher) by this measure. Antibody titers were not statistically different between the two groups. Cumulatively, these data show that while the mouse urogenital tract is not a highly permissive environment for urogenital isolates of C. trachomatis, plasmid deficiency renders the pathogen even less fit when compared to a wild-type serovar-matched isolate in this model.
↵P values were determined by comparing strains at each dose for either the ratio of culture positive to culture negative or the ratio of seroconverters by a two-tailed Fisher's exact test. Geometric mean antibody titers were compared by a two-tailed Student's t-test.
↵Number of mice culture positive of the total mice inoculated at the given dose with either F/SW4 or F/SWFP− strains. Culture detection of infection was attempted on swabs collected at 4, 7, and 10 days postinfection. Range of IFU counts (considering all mice and time points postinfection) is shown in parentheses.
↵Number of mice of the total tested displaying seroconversion for plasma IgG antibody (day 35 postinfection) against serovar F antigen. The mean geometric antibody titer ± the standard deviation of mice that seroconverted following inoculation is provided in parentheses (titers ranged from 40 to 5120).
↵ID50 is the calculated dose that will achieve 50% infection rate. Data are given in IFUs as calculated by the method of Reed and Muench (Reed & Muench, 1938).
Plasmid deficiency in a C. trachomatis urogenital isolate correlates with reduced infectivity and morbidity in the mouse respiratory tract
Unlike C. muridarum, which naturally ascends into the upper urogenital tract and induces pathogenic host responses, C. trachomatis isolates are less likely to ascend and are less virulent for the urogenital tract in the mouse model. Induction of chronic oviduct damage by C. trachomatis in the mouse requires direct inoculation into the upper genital tract of mice via surgical exposure of the ovarian bursa or oviducts (Tuffrey et al., 1986a, b). However, weight change following respiratory infection is a common means to generally assess human pathogen virulence in rodent models (Fox, et al. Ed., 2007) and has been used to assess chlamydial virulence in the mouse (Ramsey et al., 2009; He et al., 2011). For this reason, we sought to assess the in vivo infectivity and morbidity of F/SWFP− and wild-type F/SW4 C. trachomatis via intranasal inoculation of mice.
Table 2 shows that statistically significant differences in infectivity were observed between the groups at the lowest dose where 4 of 10 animals seroconverted when inoculated with 103 IFU of the F/SWFP− (estimated ID50 = 1292 IFU), but all 10 animals seroconverted when inoculated with plasmid-carrying F/SW4 (ID50 < 1000) at the same dose. Statistical significance was also achieved when assessing differences in antibody levels in the groups inoculated with 106 IFU of the wild-type strain evoking higher antibody levels in those that seroconverted. While not consistent across all doses, these data indicate enhanced infectivity in the presence of the plasmid.
↵P values were determined by comparing strains at each dose for either the ratio of seroconverters by a two-tailed Fisher's exact test or geometric mean antibody titers by a two-tailed Student's t-test.
↵Number of moribund mice of the total inoculated at the given dose with either F/SW4 strain or F/SWFP− strain through a c. 20-day monitoring period.
↵Number of mice of the total tested displaying seroconversion for plasma IgG antibody (day 35 postinfection) against serovar F (SW4) antigen. The mean geometric antibody titer ± the standard deviation of mice that seroconverted following inoculation is provided in parentheses (titer range = 20–10 240).
↵Calculated infectious dose that will achieve 50% infection rate (seroconversion) as calculated by the method of Reed and Muench (Reed & Muench, 1938).
Figure 1 shows that following an initial loss in weight in all groups (likely due to the stress of anesthesia and dosing procedures), all groups gradually recovered initial body weight and began to gain weight above baseline day 0 weights. Overall, a distinct trend was observed in the 103, 104, and 105 groups where mice inoculated with the wild-type F/SW4 strain gained less weight over time than those inoculated with identical doses of the F/SWFP− isolate. While this did not achieve statistical significance at the 103 dose – likely due to low number of the F/SWFP− isolate-infected mice with only 4 mice in this group seroconverting – the difference in weight gain in the 104 and 105 groups was highly significant. This indicates that the F/SW4 isolate caused greater morbidity than did the plasmid-deficient F/SWFP− isolate. Interestingly, at the highest intranasal dose (106), no differences in weight gain over time between the wild-type and plasmid-free infected groups were observed, although both groups gained less weight than identical groups at lower doses. This suggests that higher doses compensate for the loss of the plasmid when comparing weight changes observed in the lower dose groups. Lastly, it is notable that neither the plasmid-free C. trachomatis F/SWFP− nor the F/SW4 isolate induced any mortalities over the 21-day monitoring period, indicating low virulence of human urogenital C. trachomatis isolates in this nonhuman host.
A naturally occurring plasmid mutation in CDS1 results in reduced infectivity without affecting virulence
In 2006, the Swedish nvCT that contained a 377-bp deletion mutation in CDS1 was isolated in Sweden (Ripa & Nilsson, 2006). One serovar E isolate from this outbreak, Sweden 2 (E/SW2), like all nvCT strains, was undetectable by two of the major nucleic acid-based diagnostic test targeting CDS1. Because this mutant pathogen amplified rapidly in the Swedish population, it was hypothesized that evolutionary pressures exerted by screen-and-treat public health programs using the diagnostic test had selected the mutant. However, it could not be ruled out that the degree of spread could also have been attributed to selective biological fitness advantages incurred as a result of plasmid deficiency. This is why, we sought to test the hypothesis that mutation in the plasmid CDS1 observed in E/SW2 enhanced in vivo fitness when compared to a closely matched and cocirculating serovar E (E/SW3) isolate that carried a wild-type intact plasmid.
Table 3 shows the results of comparing infectivity of these two isolates in the urogenital tract following intravaginal inoculation. When assessed by culture, neither the wild-type nor plasmid mutant showed a difference in ID50 or chlamydial shedding. However, there were slight differences in the rate of seroconversion with c. twofold higher ID50 in the nvCT isolate E/SW2 and significantly increased antibody responses in the E/SW3 isolate when compared to the mutant at the 105 and 106 doses. Table 4 shows the results of comparing infectivity of the E/SW2 and E/SW3 isolates in the respiratory tract following intranasal inoculation. As seen with urogenital infection, the ID50 was increased in the plasmid mutants over the wild-type isolate. However, an accurate ID50 for the E/SW3 infected group could not be obtained because, even at the lowest dose of 103 IFU, all mice seroconverted, whereas at the identical dose of the plasmid mutant E/SW2, only 4 of 10 seroconverted. Additionally, as seen with urogenital infection, intranasal infection with the wild-type isolate stimulated higher antibody responses at the 104 and 106 doses, while the difference noted at the 105 dose did not quite achieve statistical significance. Weight change post-intra-nasal infection was also monitored in this group, but there were no remarkable differences in weight loss or reduction in weight gain over time as was seen following intranasal infection with the mutant-isolate in the previous experiments (data not shown). From these data, we conclude the mutation that occurred in the E/SW2 isolate had a modest effect on in vivo fitness of the isolate in the mouse host, and this effect was to reduce fitness, not to enhance it.
↵P values were determined by comparing strains at each dose for either the ratio of culture positive to culture negative or the ratio of seroconverters by a two-tailed Fisher's exact test. Geometric mean antibody titers were compared by a two-sided Student's t-test.
↵Number of mice culture positive of the total mice inoculated at the given dose with either E/SW3 or E/SW2 strain. Culture detection of infection was attempted on swabs collected at 4, 7, and 10 days postinfection. Range of IFU counts (considering all mice and time points postinfection) are shown in parentheses.
↵Number of mice of the total tested displaying seroconversion for plasma IgG antibody (day 35 postinfection) against serovar E antigen. The mean geometric antibody titer ± the standard deviation of mice that seroconverted following inoculation is provided in parentheses (titers ranged from 40 to 5120).
↵ID50 is the calculated dose that will achieve 50% infection rate. Data are given in IFUs as calculated by the method of Reed and Muench (Reed & Muench, 1938).
↵P values were determined by comparing strains at each dose for either the ratio of seroconverters by a two-tailed Fisher's exact test or geometric mean antibody titers by a two-tailed Student's t-test.
↵Number of moribund mice of the total inoculated at the given dose with either E/SW3 (plasmid-intact parental) strain or E/SW2 (plasmid mutant) strain through a c. 20-day monitoring period.
↵Number of mice of the total tested displaying seroconversion for plasma IgG antibody (day 35 postinfection) against serovar E antigen. The mean geometric antibody titer ± the standard deviation of mice that seroconverted following inoculation is provided in parentheses (titer range = 40–10 240).
↵Calculated infectious dose that will achieve 50% infection rate (seroconversion) as calculated by the method of Reed and Muench (Reed & Muench, 1938).
Genome comparisons
The genomes of nvCT serovar E strains E/SW2 and contemporaneous E/SW3 and E/SW4 have been described and published elsewhere (Harris et al., 2012). When comparing nvCT strain E/SW2 and E/SW3, we previously showed that there were a total of 229 SNPs separating these two strains, 158 of which are found in a 30-kb region spanning genes SW2 1391 (CTL-393) to SW2 1601 (CTL-417) (Harris et al., 2012). This SNP-dense region was shown to be a region of recombination and encompasses genes located alongside the plasticity zone or PZ. The PZ is a region of high variation both in terms of recombination and functional gene loss.
As a comparator to F/SW4 for our in vivo studies, the genome of strain F/SWFP− was sequenced and assembled. The F/SWFP− genome was 1 042 732 bp in length with no evidence of plasmid, either as an extrachromosomal element or integrated into the chromosome. There were no wholesale gene gains or losses. In addition to the lack of plasmid, the genome of F/SWFP− also differed from that of strain F/SW4 by 178 SNPs listed in Supporting Information, Table S2. The majority of these were found clustered in two regions likely to be explained by recombination. 31 SNPs between bases 167 543 and 171 733 of the F/SW4 genome, 26 being located within gene F/SW-41481 predicted to encode an integral membrane protein and a single SNP in F/SW-41491 – a FAD-dependent monooxygenase. Notably, this region overlaps with the putative recombined regions in nvCT serovar E strain E/SW2 (Harris et al., 2012). The second region of high SNP density contained 92 SNPs found between bases 710 211 and 725 729, affecting a 15-kb region encompassing 14 genes, F/SW4 6341–6471. The affected genes fall into several functional classes including core functions as well as more those orthologous to putative virulence genes. One example is the mviN gene, which has been shown to play a role in virulence in Francisella tularensis infections in mice (Ulland et al., 2010). Analysis of the pseudogenes within F/SWFP− did not indicate any unique pseudogenes compared with other sequenced strains of serotype E and F (F/SW4, E/SW2, E/SW3; Table S3).
For many years, the chlamydial plasmid was referred to as the 'cryptic plasmid of Chlamydia'. This descriptor referenced the unknown function of the plasmid that is carried by nearly all C. trachomatis ocular and urogenital isolates. However, it is arguably no longer accurate to refer it as 'cryptic'. For example, we now know the criticality of the plasmid in eliciting pathogenic immune responses in several model systems as well as in conveying certain key in vitro phenotypes (e.g. glycogen accumulation) and in controlling the expression of certain genes on the Chlamydia chromosome. Overall, studies of infection with both wild-type and plasmid-free C. trachomatis and C. muridarum strains in mouse urogenital infection indicate that the plasmid modulates infectivity and may be essential for pathogenic immune responses (O'Connell et al., 2007; Carlson et al., 2008; Olivares-Zavaleta et al., 2010; Russell et al., 2011). It is all but certain that either C. muridarum plasmid gene products or products of plasmid responsive loci on the bacterial chromosome engage Toll-like receptor-2 (TLR-2) causing deleterious responses in mouse urogenital infections with C. muridarum (O'Connell et al., 2011). Similarly, in ocular infection of lower primates, plasmid-free C. trachomatis was found to infect, but was significantly attenuated for inducing inflammatory responses (Kari et al., 2011). Although the plasmid seems essential for certain measures of pathogenicity in mouse urogenital tract infection with C. muridarum, in the mouse respiratory tract, deficiency of the plasmid in C. muridarum seems to have opposite effects, with increased pathological responses when the plasmid is not present (He et al., 2011). Likewise, it is important to note that plasmid-free C. psittaci intraperitoneal (i.p.) infections of mice or plasmid-free C. caviae urogenital infection in guinea pigs results in different outcomes (Miyairi et al., 2011; Frazer et al., 2012). In both models, the plasmid was dispensable regarding attenuation of virulence, and in the C. psittaci i.p. mouse model, a handful of chromosomal single nucleotide polymorphisms was left to account for virulence differences between wild-type and plasmid-free strains (Miyairi et al., 2011). Interestingly, the important human and zoonotic pathogen, C. pneumoniae, does not normally carry a plasmid nor does its chromosome contain any genes related to chlamydial plasmids (Thomas et al., 1997). It would be convenient if all observations of chlamydial strains and infections in various models were consistent, but one must remember that these differences likely reflect different chlamydial species and/or infection at a different anatomical site and thus adaption of the pathogen and its plasmid to each host and anatomical niche. Based on these observations, one could readily assert that each Chlamydia–host interaction is unique, and the role of the plasmid in these interactions is concomitantly variable.
In the present study, we have specifically assessed for the first time, the infectivity and virulence of naturally occurring wild-type and plasmid-free urogenital isolates of C. trachomatis, serovar F. Mice inoculated with a plasmid-free urogenital isolate of C. trachomatis displayed higher ID50 by both urogenital and respiratory routes of inoculation and reduced weight gain over time. There were no wholesale gene deletions or substitutions in these isolates although several SNPs were observed – especially in selected regions of the chromosome. While we cannot rule out a contributory role of genes affected by SNPs in these isolates, our data are consistent overall with studies in other model systems that support a critical role for the plasmid in both infectivity and virulence in this model.
With regard to nvCT E/SW2, it was originally speculated that the mutation of the plasmid in nvCT (E/SW2) was responsible for amplification in the Swedish regional population due to either a diagnostic or biological selective advantage (Ripa & Nilsson, 2006); Unemo & Clarke, 2011). Further, according to one large clinical study, women infected with nvCT were less likely to exhibit symptomatic urethral infection and lower abdominal pain (Bjartling et al., 2009). Transmission was rare among those with more international sexual networks such as heterosexuals in higher ages and men who have sex with men (Unemo & Clarke, 2011). Subsequently, no outstanding in vitro differences in nvCT were discerned by microscopy, growth kinetics, cell tropism, antimicrobial susceptibility, or inclusion glycogen inclusion when compared to similar wild-type isolates including, a serovar E C. trachomatis predominating in Sweden prior to the isolation of the mutant (Unemo et al., 2010; Unemo & Clarke, 2011). Likewise, in these studies, the genomic signature compared with other isolates was unremarkable with no wholesale gene deletions or substitutions. These results indicated that preponderance of this isolate in many regions of Sweden was likely due to a diagnostic selective advantage. Yet, fitness had heretofore not been directly tested in vivo.
In the present study, mice infected with nvCT E/SW2 displayed some moderately reduced parameters of infectivity, but measures of virulence in the respiratory tract were more equivocal. Overall, in vivo fitness appears reduced, and thus these data do not support the mutant's expansion as a result of in vivo fitness and thus would more likely support its expansion as a result of positive selective pressure from diagnostic test avoidance.
Some caution should be exercised in interpreting our present data. First of all, we carefully selected serovar-matched isolates that were predominating in the human population at or before the time the F/SWFP− or plasmid mutant E/SW2 was initially isolated (Unemo & Clarke, 2011). Genome sequencing has indicated that while the chromosomes of the comparator strains differ on the account of putative recombinations and SNPs, there were no other whole gene differences. However, this was not unexpected (Tables S1 and S2 (Unemo et al., 2010). This analysis also showed how apparently closely matched, cocirculating strains carry a number of SNPs, some in genes putatively associated with virulence. It is not possible to predict whether any of these SNPs contributed directly to the differences we have observed in the mouse model. However, our observations are quite consistent with the aforementioned in vivo observations of plasmid-free strains in human disease (Bjartling et al., 2009; Harris et al., 2012) and in other animal models (O'Connell et al., 2007; Carlson et al., 2008; Olivares-Zavaleta et al., 2010; Russell et al., 2011). Thus, we feel the most logical inference is to attribute the differences seen to presence and function of the plasmid.
Secondly, C. trachomatis, unlike C. muridarum, is not a natural pathogen of the mouse. Consequently, C. trachomatis infection induces neither the degree of infection burden nor the pathology that is observed following C. muridarum urogenital or respiratory infections (Rank, 2007). Thus, one should be mindful of these host and pathogen differences when interpreting our data. Nonetheless, it is pleasing that our findings largely support and agree with those of others investigating the chlamydial plasmid in mouse urogenital tract infections with LGV strains either C. muridarum or C. trachomatis (O'Connell et al., 2007; Carlson et al., 2008; Olivares-Zavaleta et al., 2010; Russell et al., 2011). In the context of what has already been reported by others regarding the chlamydial plasmid and in vivo fitness, our results add to the growing body of evidence that in many hosts and at specific anatomical sites, the plasmid plays a role in modulating infectivity and virulence, but this role is not universal. Now that a means to genetically manipulate chlamydiae and selectively mutate the plasmid has been elaborated and the function of the plasmid can be dissected and studied in greater detail (Wang et al., 2011; Song et al., 2013).
Supporting Information
Table S1. Single nucleotide polymorphisms (SNPs) in E/SW2 compared to E/SW3.
Table S2. Single nucleotide polymorphisms (SNPs) in F/SWFP compared to F/SW4.
Table S3. Pseudogenes in the genomes of the strains studied.
This work was supported in part by Midwestern University intramural funds and by Wellcome Trust Grant Nos. 091296 (I.N.C.) and 098051 (N.R.T. and H.M.B.S.-S.).
In this paper further evidence of the important role of the chlamydial plasmid in virulence is provided. In both the respiratory and urogenital tract sites, the plasmid-containing version of Chlamydia trachomatis was more virulent than the non-plasmid strain, when analysed in the mouse model. While this has previously been shown with the mouse strain of Chlamydia, this is the first report showing similar virulence characteristics of the human C. trachomatis strains.
- © 2013 Federation of European Microbiological Societies.
This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs licence (http://creativecommons.org/licenses/by-nc-nd/3.0/) which permits non-commercial reproduction and distribution of the work, in any medium, provided the original work is not altered or transformed in any way, and that the work is properly cited. For commercial re-use, please contact email@example.com


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Dr. Raub graduated from the University of Toledo College of Medicine in 1990. He works in Cincinnati, OH and 4 other locations and specializes in Allergy & Immunology. Dr. Raub is affiliated with Bethesda North Hospital, Good Samaritan Hospital, Mercy Hospital Anderson and Select Specialty Hospital.
Allergy & Immunology
Medical School: University of Toledo College of Medicine; Graduated 1990
Please verify insurance information directly with your doctor's office as it may change frequently.
*This physician has multiple office locations. See Office Information for details.
8000 5 Mile Rd Ste 315
Cincinnati, OH 45230
8000 5 Mile Rd Ste 315
Cincinnati, OH 45230
40 N Grand Ave Ste 101
Fort Thomas, KY 41075
89 Carothers Rd
Newport, KY 41071
89 Carothers Rd
Newport, KY 41071


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By Lori NewmanAugust 18, 2011
SAN ANTONIO -- As temperatures continue to soar over 100 degrees, San Antonio's sizzling summer heat is affecting people's ability to work, exercise and play the way they normally do.
"Heat-related illness can be avoided if people conduct outdoor activities during the coolest parts of the day," said Capt. Andrew Patterson, chief of Environmental Health Services at Brooke Army Medical Center. "Wear loose woven, light-colored clothing and stay hydrated."
According to National Weather Service forecast data, the hottest part of the day is generally from 4 to 7 p.m. in Texas.
"People also need to be aware of the symptoms of heat-related illnesses and what to do when they occur," Patterson said.
The Centers for Disease Control and Prevention website (http://www.cdc.gov) describes the symptoms and recommended treatment for the following heat-related illnesses:
• Heat Cramps " Symptoms include muscle cramps, pain or spasms in the abdomen, arms or legs. People who experience these symptoms should stop the activity and sit in a cool place. Drink clear juice or a sports beverage or drink water with food. Do not return to strenuous activity for a few hours after the cramps subside. People with heart problems, on a low-sodium diet or having cramps for more than one hour should seek medical attention.
• Heat Exhaustion " Symptoms include rapid heartbeat, heavy sweating, extreme weakness or fatigue, dizziness, nausea or vomiting, irritability, fast shallow breathing and slightly elevated body temperature. People who suffer heat exhaustion should rest in a cool area, drink plenty of water and take a cool shower, bath or sponge bath.
• Heat Stroke " Symptoms include high body temperature, confusion, loss of coordination, hot, dry skin or profuse sweating, throbbing headache and seizures or coma. To treatment for this serious heat-related illness move the person to a cool, shaded area; remove their excess clothing and apply cool water to their body. If they are able, have them slowly sip water.
"Cooling is the first priority and can reduce mortality," Patterson said.
An important sign of a life-threatening heat-related illness is mental confusion.
To assess this, ask the person questions such as their name, what month and year it is, where they are and what they were doing before becoming ill. If the person cannot answer, call for an emergency medical evacuation or ambulance immediately.
Staying hydrated is important, however hourly fluid intake should not exceed one and a half quarts per hour or 12 quarts per day.
Symptoms of over hydration or water intoxication are confusion, weakness and vomiting.
People who have over-hydrated should restrict their fluid intake and eat something salty such as tomato juice, pretzels or a dill pickle, explained Patterson. "But, medical attention is a must!"
Other less severe heat-related problems include dehydration, heat edema (swelling of the extremities), heat rash and sunburn.
Current Fort Sam Houston weather conditions and the Work/Rest and Water Consumption Table are available at http://samhouston.army.mil/weatherUpdate/FSHWeatherConditions.doc.
The U.S. Army Public Health Command website also offers several useful resource materials on heat injury prevention at http://usaphcapps.amedd.army.mil/hioshoppingcart/.


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No Evidence Anyone Involved in Berezovsky's Death, Police Say
There is no evidence to suggest any third party involvement in the death of Russian national Boris Berezovsky, whose body has been found at his U.K. home, police said.
Berezovsky, 67, was found dead yesterday at the property in Ascot, near London, Thames Valley Police said in a statement on its website, adding that the death remains "unexplained."
"We are at the early stages of the investigation and we are retaining an open mind as we progress," Detective Chief Inspector Kevin Brown, deputy senior investigating officer in the case, said in a statement today.
According to the police, an employee of Berezovsky said he called the ambulance service after becoming concerned for the Russian's welfare, having not seen him since late on March 22. The employee said that he forced open the bathroom door which was locked from the inside and discovered the body on the floor.
"The investigation team are building a picture of the last days of Mr Berezovsky's life, speaking to close friends and family to gain a better understanding of his state of mind," Brown said. "It would be wrong to speculate on the cause of death until the post mortem has been carried out. We do not have any evidence at this stage to suggest third party involvement."


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Barry Saunders and Simon Le-Chalmers, who owned the franchise's for Viva Soccer Gold Coast North and Viva Soccer Northern NSW, are both now, through a mutual agreement with Viva Soccer, running these regional areas as independent coaches allowing viva Soccer to concentrate on Metro areas.
Barry and Simon believe that with coach and parent support, we set small bite-size goals for young players with a developmental focus on trying to improve performance gradually and in small increments. When progress is swift, accept it and reassess the goals, then move on to the next challenge.
Through our programs we provide a safe, fun and structured environment where children can learn at their own pace from our foundation grassroots programs up to aspiring young talented players. All our coaches are FFA qualified, have current working with children checks and all coaches that run their own program must hold a current First Aid certificate.
Latest News:
All our programs are designed and support the initiatives set out by Football Federation Australia with the key coaching points being followed:
- Football is being played
- Football is being learned and therefore taught
- Football is being experienced and enjoyed
- Players understand the football purpose of the exercise
- Players recognise the game-related intention
- Players are challenged to improve both individually and as a group
Hello!! Toby and I are so excited about SIBA Soccer, I just wish we were still in Australia to be able to come along and join in. I really think you guys need to set up in NZ… WE MISS YOU! Toby still talks about you guys all the time. The best start to soccer he could have dreamt of!! To anyone out there reading this and thinking of joining SIBA Soccer for the first time just go for it!! You won't be disappointed, its so much fun!! Toby learnt so much and just loved Simon, Sticki and team. Once again WE MISS YOU!! Sami and Toby xx
Kingcliff Camp April 2013
Thank you so much for a great 3 days! Cullen just loved it – he cant wait to join you on a Tuesday afternoon at Kingscliff.
Thank you Bazza for a fantastic fun Soccer camp, both Joe and Jacob absolutely loved it. They were very disappointed that today was the last day and would have been happy to be with you guys all week. Will definitely be back! Also Lewis and George were great with the kids too
Murwillumbah FC Camp, April 2013
SIBA soccer is the best thing to do because we learn so many skills and have the best time ever Conor age seven
I love SIBA soccer because it helps me get better at soccer and I learn lots – from Niamh McCarthy age 5
Darcy was happy all day after that first day with Baz at Siba Soccer and that continued all week. We couldn't get the smile off his face. Barry and his team made Darcy feel very welcome and took away any fears he had from being one of the youngest there. They really took him under their wing which was great. I would recommend these guys to anyone.
Fantastic day down at Murwillumbah today with coach Bazza providing his SIBA Soccer inflatable pitch for all the kids to play whilst the parents took in the PL fixture between MFC and Mudgeeraba. Many thanks to coach Baz and SIBA Soccer from all at Murwillumbah Football Club.
To Coach Simon,
My son, Jesse-Jaymes,(aged 6) participated in the first SIBA SOCCER program last week and absolutely loved it. The coaches were both friendly and professional in their delivery of the program. There was a great atmosphere where I believe the children felt safe and respected as the coaches were instantly calling them all by their first names (and the parents too).
Jesse-Jaymes finished the 3-day program with new soccer skills, new friends and a fuelled passion to persue the game of soccer.
Thankyou SIBA SOCCER, we will definately be back for more !!!!
Thank you Simon for a fantastic soccer camp, Riley had a ball. It was so well organised and professionally run. Riley has benefited from your excellent teaching and coaching skills and it is great for the kids to be around such positive role models. We are looking forward to more SIBA Soccer activities to participate in!
Hi Soccer Simon,
Thank you for the Great SIBA Holiday Programme Slayter had a absolute ball. Meeting new kids and improving his soccer skills.
The program is definitely worth while and keeps you kids active and healthy. Thanks once again to you guys. Dannii & Slayter
Yay! There is my boy in blue. He had such a great time and is already talking about the next SIBA Soccer clinic. He has picked up some great soccer skills and had fun doung so. I love how encouraging your coaches are with all the kids. Thank you. See you at the next one
Thank you coach Simon and Vinnie for organizing and running such a great program for the kids..
They has a blast.
Their looking forward to the next one.
Great job.
Well done fellas, our little bloke Mav loved it. It all helps to get him prepared for his first soccer season with Coomera Colts U/6's. He'd had 3 seasons with Little Kickers so he's now looking forward to stepping it up a touch, cheers.
My son Hayden went to the soccer camp at Coomera and had a wonderful time. He learnt heaps of skills and made new friends. He wishes it was still going. Will look forward to the possibility of monday skill training. Will have his hand up for that.
Lachlan had an awesome 3 days, especially today in the rain and said he will be back next school holidays!! Thank you so much Simon and Vinnie 🙂


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DJ-producer Illich Mujica revives his Techtonics affair in a big way—he's bringing in Germany's Marc Romboy, the maker and player of emotive electronic rhythms (including that great 2011 remix of Terrence Parker's "Love's Got Me High") who heads up the Systematic label. Moth is the studio team of Luis Padron and Mujica himself, and Wender is one of the folks behind the fun Rinsed affair.
|Event phone:
|Event website:


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followed your advice and sold some more shares (about 10% of what I have left) to buy some AAPL at 395.
See you guys at profits, suckers.
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