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Frank Morring, Jr.
Senior Editor, Space,
Aviation Week & Space Technology
Frank is a Senior Editor and has been a journalist for 40 years, specializing in aerospace for over 20 years. Frank joined Aviation Week in 1989 as a defense/space reporter and senior space technology editor. In 2007 he was named deputy managing editor/space, responsible for coordinating space coverage across all bureaus and publications.
Frank began his career working for his hometown daily in Huntsville, Alabama, and moved to Washington in 1979 as correspondent for the Birmingham Post-Herald. He later covered the Cold War Pentagon for Scripps-Howard News Service.
Frank received a degree in comparative literature from Dartmouth College. He has Goddard and Von Braun media awards from the National Space Club, and a Neal Award.


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Pressed into the starting lineup by Maymon's ongoing knee problems, 6-foot-9 senior post Kenny Hall produced 14 points and 8 rebounds in helping the Vols to a 75-68 victory. Hall, who missed the last nine games of 2011-12 due to a disciplinary suspension, hit 6 of 9 shots from the field and blocked 2 shots versus the Bulldogs in perhaps the finest all-around game of his college career.
"One thing Kenny's done is he's doing a good job of bouncing back and being a good leader," head coach Cuonzo Martin said on his post-game show. "He's not very vocal as a leader but he's accepting his role, going inside, going outside, shooting his shot from the perimeter and making his free throws.
"Kenny Hall has really grown up. I'm so proud of his approach to the game, his ability to play as hard as he can play, but also stepping in for a guy like Jeronne Maymon and really rebounding the basketball. I think Kenny's doing a great job."
Though hindered by fouls (5) and turnovers (3), Vol teammate Jarnell Stokes produced a double-double of 14 points and 11 rebounds. The 6-foot-8, 270-pound sophomore raked six of his rebounds off the offensive glass.
Junior point guard Trae Golden also scored 14 points for the Vols, adding 5 assists and just 2 turnovers. Josh Richardson and Jordan McRae contributed a dozen points each to the Vols' balanced attack. Richardson did a little of everything, going 4 of 7 from the field, 1 of 2 from 3-point range and 3 of 3 from the foul line, grabbing 5 rebounds and making 2 steals. McRae hit 4 of 6 shots, including 2 of 2 from beyond the arc, and contributed a pair of assists.
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Jeremy Atkinson scored 18 points, D.J. Cunningham and Jon Nwannunu 12 each for Asheville, now 1-2 on the season and 0-9 all-time against the Vols.
Tennessee struggled out of the gate and trailed 19-20 with 8:36 left in the first half. Golden hit a 6-footer in transition to open an 11-2 run that McRae closed with a 3-pointer, giving Tennessee a 30-22 lead. The Vols widened the gap to 38-28 by the break.
The Big Orange opened the second half with a 12-5 spurt that produced a 17-point lead (50-33) with 15:45 remaining. But, as happened in the opener against Kennesaw State, the Vols lost focus and squandered much of their advantage. UNC Asheville kept chipping away and pulled within six points (74-68) with 15.4 seconds left. Richardson hit a free throw with 12.1 seconds remaining to secure the final margin.
"I feel like we've got to do a lot better job of closing out," Hall said, adding: "I think we just have to do a better job of keeping our poise out there on the floor, especially when we get a nice lead. We can't get too comfortable out there because a lot of better teams is going to take advantage of that."
Martin praised the Bulldogs for fighting back when Tennessee appeared ready to salt the game away.
"When we got up 15, 17 points, those guys stayed the course, found ways to make plays," the Vol coach said. "I don't think we were as sharp in about a five-minute span defensively, and they made the plays to make it a ball game."
Tennessee was solid offensively, shooting 50 percent (26 of 52) from the field and 50 percent from 3-point range (4 of 8). The Vols were a little sloppy at times, however, committing 14 turnovers.
Tennessee's defense was not at all sharp. It allowed the Bulldogs to shoot 50 percent from the field (23 of 46) and it forced just 11 turnovers. After being outrebounded by Kennesaw in Game 1, however, the Vols bounced back to dominate the backboards in Game 2 by a 37-25 margin.
Martin was pleased with the backboard work, noting: "I thought our guys did a tremendous holding those guys to three offensive rebounds, as much as they get the ball around the rim."
OKLAHOMA STATE PREVIEW: The Cowboys improved to 2-0 with their defeat of Akron, thanks to a superior all-around performance by Marcus Smart. In addition to contributing 16 points, he led Oklahoma State in rebounds (9), steals (4) and assists (3). Le'Bryan Nash led all scorers for the Cowboys with 18 points, while Markel Brown chipped in 17.... Oklahoma State shot just 37.5 percent from the field (21 of 56) but limited Akron go 30 percent (21 of 70).... Oklahoma State's Philip Jurick was signed by Bruce Pearl out of Chattanooga's East Ridge High School and redshirted as a Tennessee freshman in 2008-09 before transferring. The 6-foot-10 Jurick produced 2 points, 8 rebound and 2 blocked shots before fouling out in a 20-minute relief stint versus Akron.

subset: commoncrawl


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Google Docs is a free, Web-based word processor, spreadsheet, presentation, form, and data storage service offered by Google. It allows users to create and edit documents online while collaborating in real-time with other users. Google Docs combines the features of Writely and Spreadsheets with a presentation program incorporating technology designed by Tonic Systems.
iGoSyncDocs is a Google Docs desktop application, which is a small, easy to use, Java based application specially designed to offer an easy way to access and synchronize files to Google Docs across multiple computers.
Some features of iGoSyncDocs:
1. Access Google Docs Services from desktop application
2. Download or upload document via Google Docs Services
3. Share Document with friends, Google groups
4. Create new Document, Spreadsheet & Presentations
5. View Hidden, Started documents
6. Rename, trash or delete documents
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text:
Other operators sometimes give the exudate escape while making the grooves in what is now known as 'Smith's Operation.'
The microscopic characters of the exudate are pretty uniform.
The Gyropeltes, however, regards this exudate as delicious food and rapidly removes and devours it.
The poisonous substances produced by bacteria are diluted and washed away from the part by the exudate.
Autopsy: Both guinea-pigs showed marked congestion and a moderate degree of exudate in the peritoneum.
The exudate that fills the air cells and minute bronchial branches undergoes disintegration and softening when healing commences.
Upon standing this exudate separated into two layers, a reddish yellow serum above, and a grayish yellow partly solid layer below.
Superficial wounds require no special drain, as some exudate will find exit between the stitches and the rest will be absorbed.
A clear watery fluid moistening the surface of serous membranes or exudate resulting from inflammation of any of those membranes.
The nuclei of the bodies recognized in the exudate are evidently in a state of division and consequent multiplication.
exudate ex·u·date (ěks'yu-dāt')
A fluid that has exuded out of a tissue or its capillaries due to injury or inflammation.


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Jacqueline Buttke, APNP, is a certified nurse practitioner with a special interest in neurology, focusing on headache, neck and back pain. Jacqueline received her Master of Science degree in nursing from Concordia University in Mequon and her Bachelor of Science degree in nursing from the University of Wisconsin-Stevens Point. She is a member of the National Association for Orthopedic Nurses and the Wisconsin Nurses Association. She was born and raised in Wisconsin and enjoys the great outdoors: canoeing, biking, skiing, and camping. She also has a passion for ballroom dancing.
Ministry Health Care


text:
CANADA STOCKS-TSX hits seven-week high as resources gain
TORONTO, July 22 (Reuters) - Canada's main stock index opened higher on Monday to hit a seven-week high as gains in prices of commodities such as gold and oil drove gains in shares of natural resource companies. The Toronto Stock Exchange's S&P/TSX composite index was up 50.52 points, or 0.40 percent, at 12,735.65 shortly after the open.
© Thomson Reuters 2017 All rights reserved.


text:
Emily Perron is an adjunct faculty member in the Graduate School of Education.
Emily Perron
Professional Title: Adjunct Professor
Areas of Academic Focus and Expertise: Mathematics, secondary education
Area of Work and Concentration at Lesley: Mathematics in education
Representative List of Recent Courses Taught: CMATH 6110: Functions and Algebra II; CMATH 6113: Probability: The Mathematics of Uncertainty
Education: BA, MAT, Rhode Island College; MEd, Lesley University
Emily became an adjunct instructor for Lesley in 2011. She teaches in the Masters of Education program. She has also facilitated a number of math education courses since 2010, including Functions and Algebra II (CMATH 6110) and Geometry and Measurement II (CMATH 6112).She is a full-time math teacher at Woonsocket Middle School in Woonsocket, RI and has been there for seven years. She has taught grades 6 through 8 in the middle school and has worked with students from resource through accelerated. Emily has taught two after-school programs including "7th Grade Academic Lab," in which she worked with students failing in their second year of Grade 7 and helped to promote them to eighth grade, and "Algebra in the Real World," where she is currently working with seventh and eighth grade students to build their knowledge of basic algebra before they enter the high school. She has also collaborated with teachers from two other RI districts to write units of study for the Common Core State Standards with the help of the Dana Center at the University of Texas. Prior to teaching in Woonsocket, Emily taught a basic mathematics concepts course at Rhode Island College.Emily earned her first master's in secondary education from Rhode Island College and her second master's in math education from Lesley University. Her undergraduate work at Rhode Island College was in mathematics. She has a certificate in secondary education (7-12) with a concentration in mathematics, a middle school endorsement (5-8), and an English as a Second Language (ESL) endorsement. Emily is a lifelong learner and believes that we all learn something new every day.
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text:
Well 3 things come to mind here.
1. Makes little sense using sudo when you should be root.
2. If you can't find the package by the name you give then try searching for the package using apt. Guides on google abound. Try "search packages using apt".
3. Use google to look for alternate ways to install something. Try "install rfkill"


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It's Mike's birthday, and everyone is calling him the grandpa of the group since he is the oldest. A very enjoyable episode of My Boys and probably one of the best of the season (There's really not that much to choose from) but yeah, it was still a great episode. It had a surprise ending, funny one liners, and a great cast that interacted great.
Kenny warning Stephanie about the movie producer was a bit too predictable for my liking but it still was entertaining. At the time I didn't even know Kenny & Stephanie were dating until the end, that caught me by surprise since I barely started watching this show. We have something very eventful happening at the end which sort of sets the rest of the season in motion. Bobby losing his family fortune, he gets everything taken away and has to stay with PJ. Then we get a bit of a surprise. We think Mike is turning mid-30 status. In the end, we find out he's 40. He really is the grandpa of the group. Funny enjoyable episode.


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The Importance Of Article Marketing
Posted on January 21, 2011
Let's face it; we're not all fantastic writers, not to mention the amount of time it takes to write a simple 400 word article. The majority of entrepreneurs will hire copywriters or ghostwriters to write for them instead of taking the time. However, in your marketing ventures, don't forget about article marketing, an interesting way to promote your business.
What is article marketing?
Article marketing is when you write about topics in your expertise and then put those articles on popular article websites such as HubPages.
Who should use article marketing?
Anyone who has a business to promote, should be writing them. You want to be known as the expert in your field, right? Well, what better way to do that than to write about it and spread it across the web.
People are in love with reading articles in their favorite niche. The more original articles you post, the more you'll be noticed, and inevitably the more sales you'll generate from it.
How should I write them?
Make them as informative as possible. It's been said that you should give away all of your knowledge in order to gain the trust of others and I completely agree with that. Even if you give away everything you know, when someone needs help in your niche, you will be the first one they go to since you were kind enough to give away such valuable information. Best of all, make the articles original! No two articles should be the same. Try to stay away from being too "advertise-y", but I do suggest hinting to your brand in there as much as possible.
Where should I post them?
The best websites to post your original content are:
- Hubpages
- Squidoo
- Ezinearticles
- Knol
- Associated Content
There might be other niche blog sites that you'll want to consider posting your articles to, but those above are the main ones to focus on, and get the most hits. Plus, you'll be able to add links and a detailed profile.
You'll notice I said 'original content' quite a bit above. That's because Google loves original content. Even if you need to change around the paragraphs and words, before posting the same article to other sites, try to make them as original as possible.
Once you've posted your article, don't just run away and never return. Comment on other similar niche articles within the website to get the word out that you're an active member and an authority in your field. Also be sure to promote your article through popular social networking sites.
How do you use article marketing to your benefit?
56 Replies to "The Importance Of Article Marketing"
January 21, 2011 (3:21 pm)Reply
Hey Morgan,
Exactly right! AM is crazy powerful for building backlinks and traffic but it can only really be done if you take the time to create content as if you were posting to your blog.
You want to be able to get people to click through and you can only really do that by delivering very valuable information.
I've got some big plans for AM this year; now it's time to write!
January 21, 2011 (4:46 pm)Reply
Absolutely, very good points! Backlinks are so powerful within article marketing.
Good luck with your AM strategy!
Dennis Edell@Direct Sales Marketing
January 22, 2011 (2:24 pm)Reply
Since when?
January 22, 2011 (4:01 pm)Reply
Since Demand Studios decided to make people sign up with them in order to post anything on eHow. I dislike the new policy very much.
Dennis Edell@Direct Sales Marketing
January 23, 2011 (11:08 am)Reply
Wow that does suck. So what then, they pay salary or still per article or what?
January 23, 2011 (11:15 am)
Yes they pay per article. The pay ranges from $7-25.
January 21, 2011 (7:13 pm)Reply
Hi Ben
I have been wondering about the value of article marketing. Informative article without the hype. Thanks
As I am in a small niche, don't know how popular my articles would be there but can only give it a go and see where that takes me.
Patricia Perth Australia
January 21, 2011 (8:09 pm)Reply
Hi Patricia!
Since you have a small niche, your articles will actually thrive a lot better than the ones who are in a bigger niche such as SEO or sports.
Definitely give it a shot!
January 21, 2011 (11:36 pm)Reply
Thanks for the input Morgan. That works with some aspects of blogging. It depends how many people are looking for my topics. Not as many searches as the bigger niches but I may just give it a try….once I get all my other goals sorted 🙂
Patricia Perth Australia
January 22, 2011 (10:13 am)Reply
Everything is searchable. 🙂 I bet you'll be tapping into a market that is dying for information. You never know. Let me know how it goes for you or if you have any questions!
ipad apps uk
January 22, 2011 (2:06 am)Reply
Article marketing is useful only with legitimate and creative content. Thanks Morgan.
January 23, 2011 (9:49 am)Reply
Well, yes, that is the point. 🙂
Thanks for reading!
January 22, 2011 (5:39 am)Reply
Great post! I think article marketing is a really effective way of proving yourself as an entrepreneur. I'll definitely be utilizing some of the websites listed.
January 22, 2011 (10:17 am)Reply
It really is effective! You'll be known as the authority in no time. 🙂
January 22, 2011 (12:40 pm)Reply
Hey Samuel!
Thank you for reading! I'm so glad you're loving the idea more now! 🙂 I do hope it serves you well and if you have any questions, please feel free to let me know.
William Tha Great
January 22, 2011 (12:24 pm)Reply
Hey Morgan,
Thanks for the informational post!
These are some good tips for article marketing. I remember I used to do freelance writing for a little, but I never really got into it too deep. If i ever start arti le marketing again this would be a great post to refer back too!
Thanks again!
God bless,
William Veasley
January 22, 2011 (12:41 pm)Reply
Hey William!
I appreciate the sentiment about referring back to this post. 🙂
Article marketing can be a tiresome experience at first, but you'll definitely see results if done correctly.
William Tha Great
January 22, 2011 (2:47 pm)Reply
Hey Morgsn,
Thanks for the response!
It's great that you respond to your guest post. That is a great way to connect with people such as myself haha. No but, article marketing is very tiresome, but I'm sure the benefits are well worth it if you only promote your best quality.
Do you do article marketing?
Thanks again!
God bless,
William Veasley
January 22, 2011 (2:59 pm)Reply
Hey William,
I wouldn't have it any other way! I'd respond whether they let me or not. 😉 I think it's important to keep in touch with those that enjoyed your article or blog or what-have-you. Engaging is key!
I do a lot of article marketing for my clients and it's extremely useful to them. It shows them as the authority and brings back a lot of clients to them.
William Tha Great
January 23, 2011 (5:33 am)
Hey Morgan,
Thanks for the response!
I just wanted to say sorry for misspelling your name stupid iPad typo's haha.
I like your attitude respond either way and get your voice heard at all cost. A mindset like that is only bound to achieve great things! You are right on point, well at least I believe it's important to connect to those who connect with you.
It seems like people forget the original point of blogging, and for me what I enjoy most!
Thanks again!
God bless,
William Veasley
January 23, 2011 (9:53 am)
Hey William!
Exactly! Blogging is about building relationships! It's frustrating when I see blogs just to be blog hubs for their own selfish needs. But at least there are still plenty of blogs out there that understand the process. 🙂
William Tha Great
January 23, 2011 (8:47 pm)
Hey Morgan!
Thanks for the response!
I totally agree wi you that blogging is about building relationships. You need those people to guide, push, and help you when times are tough. It is very frustrating to see those types of blogs, and also ones that provide you with no form of communication.
You have to let your traffic have some way to communicate with you.
Just my 2 cents.
Thanks again!
God bless,
William Veasley
January 24, 2011 (1:56 pm)
Hey William!
Absolutely! You need that personal connection with your readers, otherwise your site is bound to fail. If there's no indication of human life on the website, people leave.
In the end, we all just want to be heard one way or another.
Plus, I love the blogging community! We're all awesome people who love to support each other. 🙂
Dennis Edell@Direct Sales Marketing
January 22, 2011 (2:31 pm)Reply
Definitely something I'm getting into this yer. I was always having trouble writing them, but I believe I have hit my secret.
January 22, 2011 (2:56 pm)Reply
Hey Dennis!
That's great to hear!! What is this secret you speak of? 🙂
Dennis Edell@Direct Sales Marketing
January 23, 2011 (11:15 am)Reply
Promise you won't laugh. I felt both relieved and kinda stupid when this revelation hit me.
Whenever I sat to write "an article", I always drew a blank…nuthin there…very overwhelmed.
However, I've never had issues of any kind blogging…
The biggest DUH moment of my life right here…..write as if I'm blogging, don't publish, send it off as an article. 🙂
January 23, 2011 (3:46 pm)Reply
It makes total sense! =) I bet that advice would be really helpful to a lot of people. 🙂
January 23, 2011 (1:28 am)Reply
Great post! It seems most people are too quick to cast aside the dirty elbow-grease marketing that works so well. Article marketing works–it isn't easy, but it works!
Of all those sites I think the best one is still Hubpages. The rest seem to have been taken over by low-quality spam. Ezine is the worst offender, in my opinion.
January 23, 2011 (9:48 am)Reply
You nailed it. And that's why it's usually a last resort these days.
Ezine definitely contains a lot of spam, however, it is also seen highly by the search engines.
Hector Avellaneda
January 24, 2011 (12:08 am)Reply
Awesome post! Ive always thought that it's really hard to connect with your readership and give them a sense of who you are and what your blog is about if you have someone else write for you instead of you writing your content yourself.
Personally, I am extremely passionate about promoting financial intelligence and therefore use my blog articles to educate and inspire people to increase their financial IQ. As someone who was brought up with no financial education, I made lots of mistakes during my college years but took the necessary step to correct my financial mistakes and am now an entrepreneur in the internet entrepreneurship and education space.
Thanks for all the tips!
January 24, 2011 (1:58 pm)Reply
Hi Hector!
I agree with you! I understand needing some help with editing, but if it doesn't sound like you're writing it, then you lose your audience.
I think what you're doing is fantastic! I bet you're helping a lot of people. Article marketing would work really well for you since I know a lot of people would be looking for that advice – especially now-a-days.
Thanks for stopping by!
January 24, 2011 (4:32 pm)Reply
Thanks for the post. I often submit to EzineArticles. Definitely have to try out the others though.
January 25, 2011 (10:51 am)Reply
Ezine is definitely a great place to submit.
Good luck! Thanks for reading!
Melbourne Graphic Design
January 24, 2011 (11:35 pm)Reply
Great post, Morgan! I actually focused on article marketing when I first started promoting my website. Although I am focusing on just three article directories now and submitting all original articles to them (basically, because I jumped into the social media bandwagon last year), I can say that they are steadily bringing in the backlinks and the traffic that I could expect from them.
– Wes –
January 25, 2011 (10:55 am)Reply
Focusing on just a few is a really good idea! I wouldn't overload people with your content being everywhere. This way, people will expect you to be certain places instead of everywhere.
Thanks for reading! Glad to hear it's working for you!
January 25, 2011 (7:30 am)Reply
Article marketing has been proven time over time that when done properly is an effective technique and good to create exposure, build credibility and "show off " as an authority within a specified niche.
What many people forget is to apply your guidelines here, such as taking the time to make it unique and after submitting the article, they think it will start generating loads of traffic. An article in eZine for example gets lost in the loads of other information available in there. How do you expect someone to find it? Well, promoting it equally as if it where in your own website. That gives the article more strength and prone to be found.
January 25, 2011 (4:52 pm)Reply
Hi DiTesco!
Yes, promoting it is just as important as writing it. 🙂
Thank you for reading!
Rasel Arnold
January 25, 2011 (12:00 pm)Reply
Article marketing is a sort of advertising in which businesses write short articles related to their worth industry.
January 25, 2011 (4:51 pm)Reply
Oh but there's so much more to it! I would suggest actually not mentioning your business until the very end or in your profile.
What I would suggest is writing about your industry and any help or information you can give, instead of marketing your business.
Thanks for reading!
Mindset Marketing
January 25, 2011 (2:48 pm)Reply
Really impressive results shown here – I think we will be seeing and hearing much more about real time email marketing in 2011 as it looks set to become a latest trend. Thanks for the post.
January 25, 2011 (4:52 pm)Reply
Sure! Thanks for reading.
January 26, 2011 (8:47 pm)Reply
I made a commitment in 2010 to blog for web traffic in hopes of obtaining more client leads. After about a year and 40 or so blog entries, I thought I was engaging in article marketing or simply a content marketing strategy. I didn't use any of the article directories to push traffic back to my site and I think that's where there was a BIG disconnect for me and where I made a biggest mistake. I somewhat relied on my on page SEO strategies, which DID generate traffic for me and lead to a few client leads. The next evolution for me will be video marketing and submitting articles to article directories since I see value in placing more emphasis on off page SEO.
January 27, 2011 (9:50 am)Reply
Video marketing is definitely a great next step! Glad to hear that article marketing has worked for you. Let me know how the video marketing goes!
January 27, 2011 (9:51 am)Reply
Hi Keith,
Outsourcing can be a great tool, though I would be careful that the articles sound like you and not a third part. Though I'm sure you're aware of that. 🙂
Glad it's working for you!
January 30, 2011 (11:30 am)Reply
Hi Kalen!
I agree about Ezine. It's a great place to get traffic from, but it's not such a good place to form a community. Hubpages has a very strong community.
Thanks for reading!
February 2, 2011 (4:28 pm)Reply
Hi Bit!
Yes guest posting is definitely a great way to attract attention. 🙂 Thanks for reading!
February 18, 2011 (3:03 am)Reply
Totally agree with you about article marketing. I think it's really one of the best ways to build backlinks.
Personally, I'm now heading into the direction of putting content on hubpages. I've noticed some referral traffic from my hubpages to some of my sites. It's also a better place to build a community and establish your presence more so than Squidoo and most article directories.
February 21, 2011 (2:08 pm)Reply
Hubpages is a GREAT community! Good luck with your article marketing efforts! 🙂
Female Voice Over Talent
January 17, 2012 (12:46 am)Reply
Really a superb post on Article Marketing. Would you like to share something on Off Page SEO Methods as well.
November 21, 2012 (1:35 am)Reply
Article marketing is considered as an important part for boosting your Google rank. It is advisable to use http://www.domadeed.com to start sharing your articles.


text:
I found this album pleasant to listen to and enjoyed the talent expressed within it. Oberhofer is named after Brad Oberhofer, the front man and founder whom is originally from Tacoma, Washington. Brad moved from Washington to New York City to attend New York University to study music composition. The band Obehofer formed in 2008 to perform Brad's compositions.
Horns and go, "Step Inside" insists immediately. The tune takes a nostalgic turn near the end, riffing on some old jazz, but it's largely a characteristic overture for Kings and Queens – a slight shift from the early days of John Brown's Body has taken form here, albeit grounded firmly in roots from the same tree.
After listening to the beginning of The 4onthefloor's new album, Spirit of Minneapolis, I had very mixed feelings. I put these feelings aside to listen to and write about this band on a non-biased opinion. I cannot find too much information about the bands' musicians as much as I can about their concerts, tours and origins.
Kurt Vile's fifth long player is a humble opus if there's such a thing. His lyrics could be hopeful, could be melancholy, it's tough to tell – the almost-eponymous opening track (substitute "Day" for Daze in the title) sums it simply, it's hard to explain, (my love) in this daze. Vile's vocals throughout Wakin On A Pretty Daze are soft and flatline, yet they tend to hint at a humble crispness around the edges.
The young D.C. trio going by Mission South is set to release their sophomore extended play, Migration Vol. 2. No doubt, these boys are rather proficient (and workers, too, as their extensive touring schedule can attest to). But aptitude doesn't necessarily dictate achievement, and that's where this particular band teeters.
Minneapolis-based band Night Moves is drawing fans in for a number of reasons, whether it's the addicting, breezy hooks that pepper their debut album Colored Emotions; singer John Pelant's distinct, gritty warble; or even their ode to Bob Seger.
With TAUK, it's best to ignore the foursome's wordy pandering (We really wanted this new album to be a statement... Homunculus has everything to do with how we as humans interpret ourselves and our surroundings. It acknowledges the inherent duality that exists between our mental and physical beings and to me that's where it relates to our creative process...
I was very impressed with the message and playing abilities found on this second studio album released by the band WhiteWater Ramble. The band's name specifies the style; beautiful, fluid, flowing music rambled with determination.
Can you seriously tell me that you can spin the mental Rolodex to a particular song or artist when I describe music as "Turkish influenced?" If so, stop reading now because you already know more about this style of music then I do and you damn sure know about 3 Trees.
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text:
A radical alternative is the Evo3 from iSkin. This is designed to be a total protection for your iPod covering it from head to foot. The iSkin is not subtle being available in a range of colours, all of which are garish. My one was red – or 'Blaze' as iSkin has dubbed it. Great.
Amazingly, the Evo3 is even less pleasant to touch than it is to look at. It's constructed of a high-grade silicon that is described as durable. Of that I've no argument but it feels quite unpleasant under the fingers. Wrapping my iPod in a tight fitting plastic skin I actually felt a bit dirty in a slightly sleazy kind of way. To top it off the plastic all smells a bit strange. In fact, one colleague asked me why I was putting a prophylactic over my iPod. Not pleasant.
Unlike the ProPorta on the iSkin, the touchwheel is full covered. This is a thin memnbrane which means you can still operate it. The screen is protected by a separate plastic cover but this model was to be designed for the 30GB iPod so as I had trouble getting it on my thicker 60GB unit.
A belt clip is integrated to the Evo3 so you don't get a choice of whether you want it on or not.
Certainly if you're in an environment that you really need to have your iPod fully protected, then the Evo3 will do the job. However, it absolutely robs the iPod of any sort of style and as such I'd look for alternatives.


text:
Stosur knocked out of home slam early, again
By Greg Stutchbury
MELBOURNE (Reuters) - Playing an unranked opponent, 5-2 up in the third set and serving for the match, world number nine Samantha Stosur agreed she had 'choked' to lose her Australian Open second round match against China's Zheng Jie on Wednesday.
The 6-4 1-6 7-5 'choke' ensured she crashed out of her home grand slam early, again. The Australian has not advanced beyond the fourth round at Melbourne Park in 10 main draws.
Her form in her home country is even more abysmal. Over the last two seasons, the 2011 U.S. Open champion has won just twice in eight matches.
"Obviously it's a pretty hard one to take when you get yourself well and truly into a winning position (and) playing really quite well," Stosur told reporters. "Then all of a sudden you get to 5‑2 and you lose five games straight.
"Whatever word you want to put on it. At 5-2 up in the third, double break probably is a bit of a choke, yeah."
Stosur had started terribly against Zheng, who had beaten her a week earlier in the first round in Sydney, with her kick serve drifting wide of the service line or beyond the "T".
However, battling to hold her serve in the first game of the second set kick-started a renaissance for the Australian and she won 11 of the next 14 games.
Serving for the match, and a third round clash with Germany's Julia Goerges, the Australian served a double fault and then blasted an easy forehand return straight into the net. Continued...


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Blessing for the Home Mezuzah
53 reviews
Bring a piece of the Holy Land to your home. This Mezuzah is made of stone from four different parts of Israel including the Holy cities of Jerusalem and Hebron. The Hebrew letters engraved on the front are the traditional blessing for the home. "Blessed are you as you enter. Blessed are you as you leave". Makes an ideal housewarming gift.
Please note that this product is made from natural stone and the color will vary. You can read more about Jerusalem stone in particular here .
Affixes with strong double-sided tape which is included.
Measures 4 inches
Takes 2.5 inch scroll (available in the product options)
See more: Mezuzah


text:
The old bull snorts angrily, but Hardbit is unmoved. He has stalked and killed scores of calves and cows on his own, and together with his mate, Tanjaw, large bulls like this one. And there is no moon in the sky overhead. A good night for a kill. Silently and methodically, the two hunters circle their prey. Unlike them, it cannot see in the pitch darkness and can only swing its horned head blindly from side to side. Suddenly, Tanjaw lunges and bites down on the bull's right thigh! The bull jerks its head to the right and in that instant, Hardbit comes in from the left, clamps his jaws down on the bull's frill, and forces the massive animal to the ground. Immediately, Tanjaw places her full weight on top of the bull and pins it down. Hardbit then plants a foot on the bull's shoulder, bites down on the frill even harder, and begins to tug with every ounce of his brute strength. The frantic bull struggles and screams as the skin around its neck stretches and tears. There is a sickening crunch of breaking bone, and then, with one last effort, Hardbit wrenches off the bull's head completely! He holds the dripping prize aloft for a few seconds, then lets it fall to the ground with a thump.
Tanjaw wastes no time in beginning to feed, but Hardbit pauses to catch his breath. He regards the severed head lying in a dark puddle at his feet. Although his mind does not allow much in the way of deep thought, he is still capable of feeling pride. This was indeed a good night for a kill . . .
Tyrannosaurus rex is THE dinosaur, plain and simple. Granted, some people have other favourites, and that's great, but the tyrant lizard king will always be iconic. It's the Batman, Spider-Man, Optimus Prime, Mickey Mouse, and Darth Vader of dinosaurs, the most famous and the most liked. And the most studied one too. The Dinosaur Toy Blog certainly attests to this popularity, as there are by far more T. rex reviews than any other animal. And now it's my pleasure and privilege to review one of the newest and most anticipated renditions, the 2017 Feathered T. rex from Wild Safari!
As is often the case when I acquire a new figure, I promptly presented this T. rex before my non-dinosaur loving wife. Her gut reaction this time? "It's really fat!" And yes, this certainly is one of the beefiest tyrant kings I've seen yet. The rib cage is nearly 6 cm wide and the torso is around 7 cm deep. The figure stands a majestic 14 cm tall and measures just over 31 cm long, positively dwarfing all the previous Wild Safari T. rex figures! It is also noticeably heavier than either of the Papo T. rexes or the CollectA Deluxe Feathered T. rex. So why is this figure so massive? Well, the truth is that we've all been duped for a long time into envisioning T. rex as slimmer than it really was. One reason is that a number of prominent museums such as the Royal Ontario Museum in Toronto, the American Museum of Natural History in New York City, and even the Field Museum in Chicago have their T. rex specimens mounted without gastralia, those belly ribs that would have made the animal's torso particularly bulgy. The Smithsonian's recently acquired specimen, however, will include its gastralia when it goes on display in 2019. Another reason is that even some of the best paleoartists have had a tendency to depict T. rex more along the lines of an NBA forward as opposed to the NFL linebacker it really was. Indeed, while it's true that Giganotosaurus was longer than T. rex, the latter was still heavier, with a thicker head, neck, and torso as well as far greater physical strength. And if the Ibrahim/Sereno reconstruction of Spinosaurus is indeed correct, then the spined lizard also lost out to the tyrant lizard in terms of mass if not length. Bottom line: I strongly advise against placing this figure up on a high shelf; you really wouldn't want it falling on your head!
This T. rex is posed with the head raised high and turned to the left, the jaws wide open, the left foot forward, and the powerful tail twitching slightly to the left and well off the ground. Unfortunately, I've heard a few people report balancing issues with their figures. Mine was stood well enough when I first got it, but after a couple of days, it became more prone to tipping forward. Fortunately, after softening the left foot in boiling water, bending it back slightly, and then running it under cold water, I have solved that issue. The figure can also be balanced on the tip of its tail if needs be.
One detail that I'm going to touch right now are the deep scars crisscrossing the muzzle, three one on side and three on the other. Looks like the result of a very nasty scrap with another T. rex. There does exist fossil evidence that tyrannosaurines at least occasionally engaged in intraspecific conflict. These could have arisen over food, territory, mating rights, or possibly even cannibalism. In any case, the scars give this individual a distinctly rugged, hard-bitten appearance, hence why I've named him Hardbit.
Hardbit's most prominent feature is, of course, his elaborate plumage. Nearly his entire body is covered in feathers. A thick mane, similar to that found on the wreathed hornbill (Rhyticeros undulatus) and various species of eagle, covers the back of his neck. A welcome bit of variety from the tall mohawks so frequently seen on feathered dinosaur depictions. There are even feathers extending onto his cranium. Feathered renditions of T. rex usually omit such a feature, so this is again a welcome change. The only bare parts are the muzzle, the mandible, the throat, the feet, and a large patch on the underbelly.
Like many of the 2017 Wild Safari figures, Hardbit here was crafted by artist (and fellow Canuck) Doug Watson, which alone should tell you about the quality of the sculpting. The featherless bits have a very fine pebbled texture, with thick folds of skin on the throat and large, overlapping scales on the fingers and feet. The feathers on the main body have a lush, shaggy feel, like on a rhea or a kiwi. The larger, spikier feathers comprising the mane have tiny grooves carved into them. The savage teeth are appropriately sharp and the roof of the mouth and the tongue are pitted. There are also rows of tiny osteoderms running along the brow ridges and along the top of the muzzle, as well as the aforementioned scars. Even the soles of the feet are textured. I can't imagine how much time, effort, patience, and heart must have gone into sculpting this figure, but I take my hat off to you, Doug.
Hardbit is easily one of the most colourful Wild Safari dinosaurs to date. The plumage on his main body varies between dark red and pale orange with black stripes and a slight dark grey wash. The plumage on his chest and the underside of his tail have a white wash and the mane and cranial feathers are a combination of crimson and black. The scaly parts are a mixture of grey and beige, the nostrils, claws, and the large scales on the feet are dark grey, the eyes are yellow surrounded by red, the osteoderms are red as well, the mouth, connective tissue, and scars are crimson, and the teeth are white. It really is a gorgeous scheme, one that utilises a wide variety of colours, yet at the same time is quite realistic and not overly bold. There are a few small nicks and blemishes here and there amidst the feathers, but keep in mind that finely textured figures carry a greater risk of paint rubbing when they are packed close together during shipping. This goes more so for large figures like this one. Safari may want to consider providing extra protective packaging for their products in future.
And now it's time to discuss the scientific accuracy of this figure. Doug Watson informed me that Hardbit was based on the famous Sue specimen, which is the largest T. rex known from near-complete remains (although Scotty might actually have exceeded Sue in mass). As such, Hardbit possesses all the correct anatomical features. The massive, T-shaped skull features stereoscopic vision and savage teeth, although the ones in the lower jaw could probably afford to be a bit larger. No question of shrink wrapping here; the orbits and fenestrae are completely hidden beneath the skin. There is a short, sturdy neck, a barrel-like rib cage, rightly tiny arms with non-pronated wrists, powerful-looking hind legs, and finally a stout tail whose enormous caudofemoralis muscles would have enhanced the animal's speed and allowed it to successfully pursue live prey. As for the plumage, we all know that's a divisive issue among dinosaur experts and enthusiasts. It's true that there's currently no direct fossil evidence of feathers on T. rex, but the same can be said for a good many theropods that we are still pretty certain had them. Fossilised feathers are the rare exception, not the norm. Phylogenetic bracketing certainly suggests the distinct possibility that the tyrant king had feathers. And shoot, no one has yet discovered a Megatherium with preserved fur, but you don't hear anyone claiming it had naked skin! Speculation will always be an essential and major component of paleontology, and I for one am all in favour of a tyrant king adorned with plumage. I will also note that for the life of me, I've never understood the notion that feathered theropods can't possibly be scary. I think the people who claim as such would be screaming at their top of their lungs if they were attacked by a Canada goose, let alone a great horned owl, a harpy eagle, or a cassowary (which actually has killed at least one person). And consider brown bears, which are covered in heavy fur that gives them a cuddly, friendly appearance. But have you ever seen The Revenant or the documentary film Grizzly Man? A feathered T. rex like Hardbit here would be no less lethal than a scaly one.
Wild Safari has long been one of the very best prehistoric lines, but 2017 will go down in history as the year they took it to a whole new level of awesomeness. This Tyrannosaurus rex is nothing short of a masterpiece, no two ways about it. Its gargantuan size and bulk alone make it stand out in any dinosaur collection, but add to that the expansive plumage, the handsome colour scheme, the exquisite sculpting, and those vital airs of majesty and ferociousness that accompany any proper depiction of the tyrant lizard king, and you have a truly phenomenal toy. One for the ages. Even people who don't agree with the feathered look ought to be impressed by it. Hardbit here is now my favourite T. rex figure, hands down. Yes, even more so than the CollectA version (which I still adore, mind you). Highest of recommendations!
A heartfelt thank you goes out to Dan's Dinosaurs for generously providing this figure for review.
Also available from Amazon.com here.


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Off-campus access to e-resources
ePrint Locations
- First floor across from the Perkins Service Desk (2 printers)
- First floor across from the Perk entrance (2 printers)
- Lower level in the Link (4 printers)
- Fourth floor near Bostock connector (1 printer)
- First floor in the Edge (2 printers)
- Lower level near Perkins connector (1 printer)
- Third floor near Perkins connector (1 printer)
Scanners are located near the Help Desks of both Perkins and Bostock Libraries on the first floors.
Bookeye Kicscanner
- Location: 1st floor Perkins
- Overhead scanner with foot peddle scan button
- Save to: USB, email, upload to cloud, etc.
- Scan format: PDF, Quick PDF, JPG, PNG, Rich Text, Audio
Book2net overhead
- Location: 1st floor Bostock / The Edge
- Requires USB drive
Epson flatbed
- Location: 1st floor Perkins
- Highest resolution
- Editing Software: Adobe Acrobat Pro, Adobe Photoshop Elements, and ABBYY Fine Reader
- Scan formats: PDF, JPG, TIFF, OCR
- Location: 1st floor Perkins
- Sheet feed; copies both sides
- Scan format: PDF only
- Location: 1st floor Bostock / The Edge
- Location: 1st floor Perkins
- Book edge and sheet feed; 2nd best resolution; requires no logon; no ePrint capability
- Tools: Crop, reposition, split-screen, etc.
- Save to: e-mail, phone, google drive, jump drive
- About Labs & Software
- Typing Non-Roman Languages on Library Computers
- About Wireless Connectivity
- About Ethernet Wired Connectivity
- OIT Help Desk
- Arts & Sciences Information Science & Technology (A&SIST)
- Ask a Librarian to learn how to search databases or to report a problem with a database
- Center for Instructional Technology


text:
Map of St. Thomas, Virgin Islands
Adding markers to the map ...
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text:
Huawei Pillar
320 x 240 pixels
US carrier availabilityDiscontinued: Cricket
- Hardware QWERTY keyboard
- Lacks Wi-Fi
- The camera lacks autofocus
- The camera lacks flash
- Does not support automatic screen brightness
- Low pixel density screen (200 ppi)
LinksHuawei Pillar manual
- Device type:
- Feature phone
- Weight:
the average is 5.4 oz (154 g) 3.50 oz (99 g)
- 3.50 oz (99 g)
- Features:
- Full keyboard, Soft keys
- Colors:
- Black
- Physical size:
- 2.0 inches
- Resolution:
- 320 x 240 pixels
- Pixel density:
Pixel density - The pixel density of a display represents the number of pixels over an area of one inch. It's measured in "pixels per inch", or ppi. The higher the number, the more detailed and good-looking the display is.
- 200 ppi
- Camera:
- 0.3 megapixels VGA
- Storage expansion:
- microSD, microSDHC up to 16 GB
- Capacity:
- 900 mAh
- Type:
- User replaceable
- Talk time:
the average is 26 h (1581 min) 4.50 hours
- 4.50 hours
- Stand-by time:
the average is 20 days (476 h) 10.0 days (240 hours)
- 10.0 days (240 hours)
CDMA - Code Division Multiple Access. A technique of multiplexing, also called spread spectrum, in which analog signals are converted into digital form for transmission. For each communication channel, the signals are encoded in a sequence known to the transmitter and the receiver for that channel. The foremost application is digital cellular phone technology from QUALCOMM that operates in the 800MHz band and 1.9GHz PCS band. CDMA phones are noted for their excellent call quality and long battery life.
- 800, 1700/2100, 1900 MHz
Phone features
- Notifications:
- Polyphonic ringtones, Vibration, Silent mode, Speakerphone
- Hearing aid compatibility:
- Other features:
- Bluetooth:
- Yes
- Yes
- Connector:
- microUSB
- Features:
- Mass storage device, USB charging
- Positioning:
Positioning - This field shows the positioning systems supported by the device. There are three main types: GPS, A-GPS and GLONASS. GPS - This is one of the most widespread global positioning technologies, developed and maintained by the U.S. government. It uses satellites in order to detect your location. Works best in clear weather. A-GPS - A-GPS stands for Assisted GPS and is the industry standard for positioning and navigation. "Assisted" means that it can use local wireless networks, in addition to satellites, for quicker and more precise localization. GLONASS - GLONASS is a global positioning system, developed by the Russian Federation. It's very similar to GPS, but isn't so popular in cell phones.
- Navigation:
- Turn-by-turn navigation
To see the full specifications with in-depth details click here.
This device is also known as Huawei M615
Despite our efforts to provide full and correct Huawei Pillar specifications, there is always a possibility of making a mistake. If you see any wrong or incomplete data, please LET US KNOW.
If you are interested in using our specs commercially, check out our Phone specs database licensing page.


text:
Cyanobacterial psbA gene family: optimization of oxygenic photosynthesis
- First Online:
- Received:
- Revised:
- Accepted:
DOI: 10.1007/s00018-009-0103-6
- Cite this article as:
- Mulo, P., Sicora, C. & Aro, EM. Cell. Mol. Life Sci. (2009) 66: 3697. doi:10.1007/s00018-009-0103-6
- Citations
- 1.2k Downloads
The D1 protein of Photosystem II (PSII), encoded by the psbA genes, is an indispensable component of oxygenic photosynthesis. Due to strongly oxidative chemistry of PSII water splitting, the D1 protein is prone to constant photodamage requiring its replacement, whereas most of the other PSII subunits remain ordinarily undamaged. In cyanobacteria, the D1 protein is encoded by a psbA gene family, whose members are differentially expressed according to environmental cues. Here, the regulation of the psbA gene expression is first discussed with emphasis on the model organisms Synechococcus sp. and Synechocystis sp. Then, a general classification of cyanobacterial D1 isoforms in various cyanobacterial species into D1m, D1:1, D1:2, and D1′ forms depending on their expression pattern under acclimated growth conditions and upon stress is discussed, taking into consideration the phototolerance of different D1 forms and the expression conditions of respective members of the psbA gene family.
KeywordsAdaptation Cyanobacteria Cyanophage D1 protein psbA gene Regulation of gene expression Stress responses
Cyanobacteria, algae, and higher plants have a unique capacity to use water as a source of electrons in reducing CO2 to various organic compounds. In organisms performing oxygenic photosynthesis, the linear electron transfer (light reactions) takes place in the thylakoid membrane-embedded protein complexes Photosystem II (PSII), Cytochrome b6f (Cytb6f), and Photosystem I (PSI). These multiprotein complexes harness solar energy and, together with ATP synthase, produce reducing power (NADPH) and chemical energy (ATP) for production of carbohydrates. These carbohydrates together with oxygen, the side product of photosynthetic electron transfer, enable all heterotrophic life on Earth.
The psbA genes and the D1 isoforms present in various cyanobacterial species, classified according to their functional features
D1 forms
Selected references
Synechocystis sp. PCC 6803
1 protein
1 protein
Anabaena sp. PCC 7120
1 protein
1 protein
psbA2 psbA3 psbA4
1 protein
elongatus BP-1
1 protein
1 protein
1 protein
Synechococcus sp. PCC 7942
1 protein
1 protein
Gloeobacter violaceus PCC 7421
1 protein
1, maybe 2 proteins
psbA4 psbA5
A vast number of studies focusing on regulation of cyanobacterial psbA gene expression have been published during the past two decades. Although more and more details are currently being disclosed, the ultimate mechanisms concerning especially the trans-acting factors regulating the psbA gene expression still remain to be revealed. It has also become clear that, although the increase in psbA transcripts is a general response of cyanobacteria upon shift of the cells to high light intensity, each strain seems to have its own characteristic regulation mechanisms which cannot be directly generalized to other strains. Here, we have summarized the current knowledge about regulation of psbA gene expression with major focus on studies performed with model organisms Synechococcuselongatus sp. PCC 7942 (hereafter, Synechococcus 7942) and Synechocystis sp. PCC 6803 and 6714 (hereafter, Synechocystis 6803 and 6714, respectively). Although at least in some species also valid to the psbA genes, we have here excluded the circadian regulation of gene expression, which was recently reviewed in [11]. We apologize for not being able to present results from all published experimental systems due to space limitation and for the sake of clarity.
Regulation of psbA gene expression in cyanobacteria
As typical to eubacteria, the psbA gene expression can be expected to be regulated at the levels of transcription initiation, elongation, and termination, mRNA stability as well as translation. In cyanobacteria, the initiation of transcription is considered to be the most crucial determinant of gene expression. RNA polymerase holoenzyme, composed of the catalytically active core and one of the several sigma factors, initiates transcription. The sigma factors are responsible for promoter recognition, and the chromatine structure along with various cis-acting elements up- and downstream from the transcription start site regulates the level of gene expression. It has been shown that the principal sigma factor (Group 1) specifically recognizes the hexameric −35 and −10 regions located in the promoter region of many cyanobacterial psbA genes [12, 13, 14]. The light-responsive expression of the psbA gene, however, seems to require the function of SigB, SigD, and SigE [15, 16, 17, 18].
The tertiary structure of DNA is known to have a marked effect on gene expression [19]. AT repeats on one face of the DNA, often found in the region −240 to −40 from the transcriptional start, influence the formation of DNA double helix, and these bends may modulate transcriptional activity. An intrinsic curvature composed of several AT tracts has been found in the upstream region of psbA genes in many organisms including Synechocystis 6803 and Microcystis aeruginosa K-81 [20, 21]. Modification of these tracts was observed to severely downregulate the transcription of the psbA gene, which, however, still remained light-responsive in nature.
Specific features of psbA gene regulation have mainly been addressed in studies with Synechococcus elongatus PCC 7942 and Synechocystis 6803 and 6714. Below, we focus on regulation of psbA gene expression in these species.
psbA gene expression in Synechococcus elongatus PCC 7942
In Synechococcus 7942, the three psbA genes encode two distinct D1 protein isoforms, D1:1 being encoded by psbAI and D1:2 by psbAII and psbAIII [22, 23, 24, 25, 26, 27]. Under low light conditions (125 μmol photons m−2 s−1), more than 80% of total psbA transcripts originate from psbAI, but a shift of Synechococcus 7942 cells to high light (750 μmol photons m−2 s−1) decreases the transcription of psbAI while the transcription of psbAII and psbAIII increases [28, 29, 30, 31]. Due to enhanced turnover and differences in transcription activity between the psbAI and the psbAII and psbAIII genes, a rapid interchange of the D1:1 form by D1:2 occurs upon shift of cells to high light, which in turn is important for adaptation of cells to changing environmental cues [23, 24, 27, 32, 33, 34]. Moreover, mutant strains of Synechococcus 7942 [22] in which the exchange of D1:1 to D1:2 is blocked suffer enhanced inhibition of PSII under UVB and high light illumination [35], showing that the two isoforms are functionally distinct.
All three psbA genes in Synechococcus 7942 give rise to transcripts of 1.2 kb with 5′ ends comprising 49–52 bases upstream from the coding region (Fig. 3a; [22]). The constitutive expression of the psbAII and psbAIII genes is driven by basal σ70 type promoter elements residing between −39 and +12 for psbAII and positions −38 and −1 for psbAIII [30], whereas in psbAI the σ70 promoter TATAAT is replaced by an atypical −10 element, TCTCCT [22]. In addition to the promoter of the psbA gene, there are regulatory elements within the transcribed region, which enhance gene expression and confer light-responsiveness.
psbAI promoter region encompasses nucleotides −54 to +1, and one or more proteins bind specifically to the psbAI upstream region (+1 to +43; [36, 37]). At least one of the regulatory factors is shared with psbAI and psbAII [37], whereas the gel mobility shift experiments have shown binding of a de novo synthesized protein factor specifically to the 5′ end (66 bp) of the psbAI coding region [38]. This so far uncharacterized protein factor is essential for transcriptional activation of the psbAI gene [38]. Moreover, PsfR protein has been identified as a regulator of psbAI gene expression [39]. Overexpression of psfR results in enhanced expression of psbAI without an effect on psbAII and III. However, knock-out of psfR did not prevent psbAI expression. Thomas and coauthors [39] have suggested that PsfR may rather regulate gene expression via protein–protein interactions than via direct binding to the psbAI promoter. Moreover, a recent study suggested that the D1 protein might regulate its own synthesis: gel mobility shift experiments provided evidence that the degradation products of the D1:1 protein bind to the upstream region (−106 to −10) of the psbAI gene, thereby possibly regulating the efficiency of transcription [40]. Additional upstream sequences enhance expression but are not needed for light-responsive regulation.
Both psbAII and psbAIII also bind regulatory proteins in the regions +1 to +41 in psbAII and −2 to +38 in psbAIII. One such protein is CmpR, which is required for expression of the best-characterized low-CO2 inducible operon cmpABCD [41]. The exact binding site of the CmpR protein has been shown to be the palindromic TTA-N7-TAA and TTA-N8-TAA sequences in the enhancer elements of psbAII and psbAIII, respectively [41]. Knock-out of CmpR, however, did not completely stall the expression of the psbA genes. Moreover, the region between the −10 basal promoter and the Shine–Dalgarno sequence of the psbAII gene in Synechococcus 7942 shares similarity with that in Microcystis aeruginosa K-81 [42]. This AT-rich region upstream from the Shine–Dalgarno sequence seems to function as a negative cis-element, which might bind regulatory factors and/or ribosomes affecting accumulation of the psbAII transcripts [42]. Additionally, upstream of the basal promoters are negative elements that depress the expression [30].
Besides the 1.2-kb psbAII transcripts, the psbAII gene also produces 1.6-kb transcripts which originate 419 bp upstream from the start site for the 1.2-kb psbAII mRNA. This dicistronic message contains a 342-bp ORF immediately upstream of the psbAII coding region [28]. Expression of the 1.6-kb transcript, however, is not light dependent [43]. Whether the gene product of this ORF is somehow involved in the regulation of psbA gene expression, for example as a regulatory factor, remains to be elucidated.
Induction of psbAII/III messages occurs not only at high light intensity, but also upon a shift of the cells to low temperature under constant low light intensity, to UVB irradiation as well as to anoxia and to light favoring excitation of PSI [35, 38, 44, 45, 46]. However, neither heat shock nor oxidative stress produced similar responses as generated by an increase in light intensity [32]. It is conceivable that the reducing power, produced by the two photosystems, cannot be used efficiently in carbon assimilation under stress conditions, thus leading to elevated levels of thiol reductants. Reducing conditions, in turn, are likely to be sensed as a signal to induce the expression of psbAII/III. Indeed, addition of reduced DTT to the cell suspension was shown to induce the psbAII/III gene expression and down-regulation of the psbAI gene expression. Inhibitors of photosynthetic electron transfer chain (DCMU and DBMIB) did not cause any changes in psbA gene expression when added under low light conditions, but both chemicals dramatically reduced the induction of psbAII/III when added upon a high light shift. These results indicate that the thiol redox state rather than the redox state of the plastoquinone pool regulates psbA gene expression in Synechococcus 7942 [46].
Besides redox regulation, the expression of the psbA genes has been suggested to be controlled via a blue light photoreceptor. Shift of cells to low-fluence blue light was shown to induce transcription of the psbAII/III genes, and this induction could be reversed by a subsequent pulse of red light [47]. Since blue light has been shown to regulate chloroplast gene expression in higher plants, it is possible that components of an ancient blue light photosensory pathway are evolutionarily conserved during the divergence of plant chloroplasts from cyanobacteria [48, 49]. One possible candidate for a blue light receptor is NblS, a putative histidine kinase, which is involved—among other things—in controlling psbA gene expression [50].
Concomitantly with upregulation of psbAII and psbAIII the amount of psbAI transcripts decreases as an immediate reaction when Synechococcus cells are shifted to bright light [29, 51]. This is mostly due to destabilization of the psbAI transcript (T1/2 = 10–12 min), which is dependent on the 52-nt untranslated leader sequence [29, 51]. Destabilization of the psbAI transcript is accompanied by a decrease in transcription activity of the psbAI gene [37]. Similarly, the degradation of the psbAIII mRNA has been shown to be accelerated at high light intensity, but the psbAII transcripts are long-lived and apparently not subject to post-transcriptional regulation [29]. Determinants of psbA mRNA turnover have been shown to reside within the untranslated leader regions of the psbA genes as well as within the coding region [51]. The region encoding the first membrane span of the D1 protein is essential for the stability of both the psbAI and psbAII transcripts, probably via pausing of ribosomes, which protects the mRNAs [51].
In contrast to an immediate response of cells to high light, prolonged exposure of Synechococcus 7942 cells to high light leads to an increased accumulation of all psbA transcripts, including psbAI. This was shown to result from restabilization of the psbAI transcript after several hours at high light intensity [32]. Nevertheless, despite vigorous reaccumulation of psbAI, no corresponding increase in the amount of D1:1 could be detected in the thylakoid membrane of high light-adapted Synechococcus 7942 cells [32] probably due to a high sensitivity of D1:1-containing PSII centres to photoinhibition.
psbA gene expression in Synechocystis sp. PCC 6803 and 6714
In Synechocystis sp., only one type of D1 (D1m, see below) protein, encoded by both the psbA2 and psbA3 genes, has been detected under normal growth conditions as well as under most stress conditions [52]. However, recent studies have revealed anaerobiosis-induced expression of the psbA1 gene [9, 10], which was previously thought to be a silent pseudogene. The transcript from psbA2 accounts for 90% and from psbA3 for 3–10% of the total psbA transcript pool under normal growth conditions (ca. 50 μmol photons m−2 s−1), whereas under these conditions the psbA1 gene remains silent [52, 53, 54, 55]. Intense illumination increases the transcription of the psbA2 and psbA3 genes, and this induction is not affected by addition of electron transfer inhibitors [56, 57, 58, 59, 60, 61]. Studies using S1 nuclease protection assay, micro-arrays [62] and northern blotting [63] have shown that UVB exposure of the cells also induces an increase in the total psbA transcript pool primarily through increased accumulation of psbA3 transcripts [57, 64]. However, inactivation of either psbA2 or psbA3 up-regulates the expression of the intact gene to the normal wild type level without marked effects on cell metabolism, indicating that either gene alone is sufficient to support autotrophic growth of the cells [54, 63]. Thus, Synechocystis pattern of supplemental expression of an identical protein isoform under excitation stress is distinct from the D1 isoform exchange found in Synechococcus 7942.
In Synechocystis 6803, both the −35 and −10 elements are present in the upstream regions of the psbA2 and psbA3 genes (Fig. 3b). The transcription start points for psbA2 and psbA3 have been mapped to positions −49 and −88, respectively, relative to the ATG site [52]. The promoter region of psbA1 differs significantly from those of psbA2 and psbA3, especially in the −35 element, which is identical in psbA2 and psbA3. psbA1 also lacks a Shine–Dalgarno sequence, which, however, is not absolutely required for successful translation [65]. The transcription start site of psbA1 has not yet been defined and hence the location of promoter is uncertain. In addition to the possible promoter next to the coding region of the psbA1 gene, there is a distal promoter-like region.
Synechocystis 6803 mutants with modified degradation rates of the D1 protein have provided evidence that not only light intensity but also the rate of D1 synthesis regulates transcription of the psbA2 and psbA3 genes [5, 66, 67, 68, 69], which additionally seems to require de novo synthesized protein factors [70, 71]. Transcription of the psbA genes during the recovery process after photoinhibitory treatment of Synechocystis 6714, however, was not prevented by inhibition of translation, and the photoinhibitory treatment longer than 40 min finally resulted in increased stability of the psbA messages [58]. Transcription of the psbA genes ceases rapidly upon shift of the cells to darkness, even in the presence of an external energy source, indicating that the energy status of the cells does not directly affect the transcription activity of the psbA genes in Synechocystis [60].
Numerous studies during the past 10 years have suggested an involvement of the intersystem redox status in the regulation of psbA gene expression in Synechocystis 6803. These experiments mainly followed the accumulation of psbA transcripts in cells exposed to different nutrient regimes, light quantities and qualities as well as to electron transfer inhibitors, DCMU and DBMIB. The presence of DCMU or DBMIB upon illumination of cells results in accumulation of PSII reaction centres with a reduced quinone at a QA site, which was suggested to act as a signal to transiently increase the amount of psbA transcripts in Synechocystis 6803 and 6714 [59]. A more recent study by the same authors suggested that the occupancy of the plastoquinone binding QO site in the Cyt b6f complex might be involved in regulation of the psbA gene expression. This conclusion was deduced from the fact that the reduction of the intersystem carriers activated the transcription of the psbA gene and destabilized the message, whereas oxidation of the electron transfer chain decreased transcription and stabilized the psbA message [72]. Li and Sherman have shown that long-term (6 h) treatment of Synechocystis 6803 cells with either DCMU or DBMIB has strong effects on accumulation of psbA transcripts and indicated that reduction of the plastoquinone pool (presence of DBMIB) decreases, and oxidation (presence of DCMU) increases the expression of the psbA genes [69]. RppA, a response regulator of a two-component system, was suggested to sense the changes in the redox poise and accordingly to adjust the stoichiometry between PSII and PSI via regulating the expression of photosynthetic genes, e.g., psbA [69]. The effects of blue, orange, and far-red light on the expression of psbA gene in Synechocystis 6803 made El Bissati and coworkers conclude that light quality regulates the expression of photosynthetic genes via a redox control occurring in the Cyt b6f complex [73]. However, data not supporting this interpretation also exist. Comparison of the action spectra of psbA transcription to that of PSII activity, photosynthesis and photoinhibition [63], as well as subjecting Synechocystis 6803 cells to over-saturating single turn-over flashes inducing photoinhibition but without affecting the oxidation state of the intersystem redox carriers [61], made the authors conclude that the redox state of the electron transfer chain is an unlikely candidate to carry information for regulation of psbA expression.
The half-life of psbA2 and psbA3 transcripts in Synechocystis 6803 under illumination is around 10–20 min, and independent of light intensity or the rate of PSII electron transfer [56, 58, 59, 66, 72, 74], whereas the stability of psbA transcripts increases remarkably in darkness [56, 60, 71, 75]. Stabilization of transcripts is dependent on cessation of photosynthetic electron transfer rather than on light per se [56, 60, 61, 72]. Other factors, such as polyamines, have also been suggested to affect the stability of psbA transcripts [71]. Transfer of dark-treated cells back to light induces rapid protein-synthesis-independent accumulation of psbA transcripts [60]. Indeed, the 170-bp upstream region of the psbA2 gene was shown to bind protein factors in the dark, suggesting that the transcription of the psbA2 gene is down-regulated in darkness via transcriptional repressor proteins [60]. Specifically, a hexanucleotide direct repeat, TTACAA-N4-TTACAA, found in the promoter region of Synechocystis 6803 psbA2 and psbA3 genes as well as in Anabaena 7120, has been shown to act as a binding site for a putative repressor in the dark (Fig. 3b; [76]).
Translational regulation of D1 synthesis in cyanobacteria
It is well known that translation is a key regulatory step in the chloroplasts of higher plants, and many trans-acting factors involved in translation of the D1 protein in chloroplasts have been characterized since the 1980s [77]. Evidence is accumulating indicating that the D1 protein synthesis in cyanobacteria is also not solely controlled at the transcriptional level, e.g., in Synechococcus 7942, practically no D1:1 protein accumulated upon stress in the thylakoid membrane, even though high amounts of psbAI transcripts were present [32, 35, 46, 78]. Moreover, upon anoxia and under thiol-reducing conditions in the presence of electron transfer inhibitors, a substantial amount of psbAII/III messages accumulates without synthesis of the D1:2 protein [45, 79]. It has been shown that, if there are psbA transcripts present in Synechococcus 7942, the messages are always efficiently associated with ribosomes, suggesting that initiation of translation does not play a significant role in regulation of psbA gene expression, whereas membrane targeting of nascent D1 protein ribosome complex might, at least under some conditions, be a rate-limiting step for D1 protein synthesis [5]. Elongation of translation is also under strict regulation in Synechocystis 6803: upon shift of the cells from light to darkness, the abundant psbA transcripts keep attached to ribosomes and D1 translation continues up to a distinct pausing site. However, the newly formed ribosome-nascent D1 chain complexes are not targeted to the thylakoid membrane, and therefore no complete D1 synthesis takes place in the dark [80]. Indeed, the ribosome complexes are targeted to the thylakoid membranes and D1 synthesis can be completed only in light to replace the damaged D1 protein, indicating that it is not the initiation of translation but rather the translational elongation that is an important regulatory step in expression of the psbA genes [80]. Translational elongation of proteins in general and in particular that of the D1 protein is also sensitive to singlet oxygen, which is generated especially during photosynthesis [81, 82]. An additional regulatory factor is the availability of chlorophyll: although the accumulation of psbA transcripts is independent of chlorophyll availability, the lack of chlorophyll seems to affect the initiation of psbA translation [83]. Further complexity to the regulation of psbA gene expression is provided by the membrane insertion and C-terminal processing of the D1 protein as well as the assembly of PSII (see [84] and references therein). The different steps of PSII assembly and function are assisted by a number of auxiliary proteins, including factors involved in degradation of the damaged D1 protein, in translation and membrane insertion of the new D1 copy, and in PSII (super) complex formation and activation [85, 86]. Putative antisense mRNAs against various psbA genes in Synechocystis 6803 might represent a novel regulatory network in psbA gene expression and are presently under intense research (Wolfgang Hess, unpublished results).
Functional classification of cyanobacterial D1 proteins
Nearly 40 fully sequenced cyanobacterial genomes together with functional characterization of several psbA gene families make it now possible to attempt to classify this important gene family and its product, the D1 protein. This, however, faces a serious difficulty since the psbA genes within a species tend to be more closely related to their family relatives than to functionally similar members of other species. Therefore, the classification of D1 isoforms based solely on the genomic data is unavoidably prone to errors. In order to be accurate, one has to take into consideration the specific response and behavior of the individual members of the gene family as a response to a variety of factors. From evidence accumulated so far, we attempt here a classification of the D1 isoforms based on the manner that their expression is regulated under typical growth conditions as well as under various environmental stresses (Fig. 2; Table 1).
- i)
D1m is a D1 form expressed and present in the PSII centres under normal growth conditions. D1m is also induced under most stress conditions (m denotes for "major");
- ii)
D1:1 is a D1 form expressed and assembled into PSII under normal growth conditions, but repressed under stress;
- iii)
D1:2 is a D1 form repressed under normal growth conditions, but induced and accumulated into PSII by stress;
- iv)
D1′ is a D1 form virtually silent under standard growth conditions, but induced under microaerobic/low oxygen conditions
In the following section, we will characterize the above-mentioned D1 isoforms one by one. The individual functional distinctiveness of each D1 form, and the justification of placing them in these distinct categories, is also discussed.
The D1m isoform
Charasteristic to the D1m isoform is its presence in PSII centres both under normal growth conditions and when the cells are exposed to stress. This isoform thus contributes to the maintenance of functional PSII, but the defining trait is its increased expression under environmental stress conditions that speed up D1 degradation. A typical example of D1m is the D1 isoform encoded by the psbA2 and psbA3 genes in Synechocystis 6803 (see "psbA gene expression in Synechocystis sp. PCC 6803 and 6714"). In this cyanobacterium, the psbA2 transcript is responsible for almost all the D1 protein produced under regular growth. When the rate of D1 degradation increases, e.g., upon exposure of the cells to high light or UVB stress, the transcription of the psbA3 gene is considerably enhanced thereby enabling an increased turnover rate of the D1 protein (Fig. 2b). When the stress condition is removed, the pattern of the psbA2 and psbA3 gene expression reverses. Both genes are alone sufficient to support normal autotrophic growth mode of the cells [54, 87].
Interestingly, Gloeobaacter violaceus PCC 7421, a cyanobacterium that shows deep molecular and ultrastructural divergence from other cyanobacteria and is considered very primitive, regulates its five psbA genes in a similar manner as Synechocystis 6803 [88]. Under standard growth conditions (10 μmol photons m−2 s−1, 25°C), the D1 protein is produced by psbAI (glr2322) and psbAII (glr0779) both showing considerably high levels of expression, whereas under stress, psbAIII (gll3144) is induced supplementing the available transcripts. All three genes are encoding the same D1 isoform. The other two psbA genes in Gloeobacter encode distinctly different D1 proteins of unclear function. One of them, psbAV (glr2656), encodes the most divergent psbA sequence known to date [88].
The D1:1 isoform
D1:1, also known as D1 form 1, is best described in Synechococcus 7942 and is, together with D1:2, part of a specific regulation mechanism in response to stress conditions. Recently, a similar stress-response mechanism was revealed in Anabaena PCC 7120, Thermosynechococcus elongatus BP-1, and the marine Synechococcus WH 7803 [89]. These are diverse cyanobacterial species with no easily traceable phylogenetic relationship. The defining characteristic of D1:1 is its high level of expression in well-acclimated cells, and a distinct down-regulation upon abrupt changes in the standard growth conditions (Fig. 2a). Once the cell acclimates to the new status quo or the stress is removed the expression of the genes encoding D1:1 increases and it becomes once again the dominant D1 form in PSII centres. For some time, D1:1 was also associated with the presence of a specific amino acid at position 130 proven to influence the potential of the redox active D1-Phe residue (see discussion in the next section). This is situated in a conserved portion of the D1 protein and it has been proven true, so far, that the D1:1 protein always contains a glutamine residue at this position. While this is true, the reverse is not: not every D1 with a glutamine at 130 is a D1:1. A clear example is the low-expressed, divergent D1′ encoding psbA genes from Anabaena 7120 (psbA0) and Synechocystis 6803 (psbA1) that have a glutamine at position 130 but functionally are clearly not D1:1 forms (see the following section on D1′).
The D1:2 isoform
The D1:2 isoform reflects the mirror image of its "sister" form D1:1 in its expression. D1:2 is expressed at low level in well-acclimated cells, but upon stress conditions the expression is markedly induced and D1:2 replaces D1:1 in PSII centres (Fig. 2a). Upon acclimation or removal of stress, the D1:2 will be repressed and replaced by D1:1. This D1 form exchange is one of the clearest mechanisms documented so far on gene regulation as adaptation to changing environmental conditions, and it was well documented in Synechococcus 7942 by numerous studies (see "psbA gene expression in Synechococcus elongatus PCC 7942"). The exchange is facilitated by the fast turnover rate of the D1 protein, establishing a correlation between the expression levels of distinct psbA genes, the form of D1 protein present in PSII centres, and the functional characteristics of PSII. The evolutionary development and maintenance of this regulation mechanism speaks for a direct requirement of the functional characteristics of both D1 forms, thus offering unique advantages under their specific expression conditions. A good question raised by the presence of D1:2 under stress conditions is why the cells do not always possess D1:2 in the PSII centres? The answer, while not clear yet, has to do with a discreet advantage of D1:1 over D1:2 under standard growth conditions. Maintenance of D1:1 in the PSII centres seems to give an evolutionary advantage over mutants containing only the D1:2 form, generated by random mutagenesis.
D1:2 is a functionally well-defined form and different from its counterpart D1:1, as shown by artificially made mutant strains of Synechococcus 7942 [22]. In these strains, the exchange of D1:1 to D1:2 is blocked, which results in enhanced PSII photoinhibition under UVB and high light illumination [35]. Furthermore, it has been shown that a transgenic Synechocystis 6803 strain expressing the Synechococcus 7942 D1:2 isoform possesses a faster decay of variable fluorescence in the presence of DCMU, reflecting faster recombination of reduced QA with positive charges on the donor side of PSII, compared to a Synechocystis 6803 mutant expressing only the D1:1 isoform from Synechococcus 7942 [25]. Thermoluminescence [27] and fluorescence life time data [26] from Synechococcus 7942 cells containing only the D1:1 or the D1:2 isoform also support enhanced charge recombination in PSII centers containing D1:2. It is important to note that this functional difference between the D1:1 and D1:2 isoforms is correlated with the presence of a glutamate residue in D1:2 instead of a glutamine residue at position 130 in the D1 protein sequence, which interacts with a key Phe co-factor [7, 90]. It was recently shown that Glu occupies D1-130 position in all high-light D1 isoforms identified so far, whereas Gln is found in the low light D1 isoforms, like D1:1 [91]. The correlation of the D1 Gln130Glu amino acid replacement with phototolerance is apparently related to accelerated charge recombination, i.e., to the existence of an Em (Phe/Phe−)-dependent photoprotection [92]. It is worth noting here that the single D1 protein in higher plants has strictly conserved Glu130 residue.
The D1′ isoform
The term D1′ was for a long time used only for the product of psbA1 gene in Synechocystis 6803. This gene was considered enigmatic as its open reading frame was intact but its transcript could not be detected by classic hybridization methods [54, 56, 75]. However, artificial activation of psbA1 in Synechocystis 6803 [53], via replacement of the 320-bp upstream fragment of the psbA1 gene with a 160-bp upstream fragment of psbA2, leads to production of a functional and light-responsive, albeit aberrant, D1 protein [93, 94]. Whole genome sequences and functional studies recently published from several cyanobacterial species (http://www.kazusa.or.jp/cyano/, http://genome.jgi-psf.org/) have revealed that, apart from Synechocystis 6803, a psbA1-type divergent, low-expressed and non-responsive psbA gene also exist in Anabaena sp. PCC 7120 (psbA0-alr3742) [95], Thermosynechococcus elongatus BP-1 (psbA2-tlr1844) [91, 96], Cyanothece sp. ATCC 51142 (psbA2) [10], and possibly Gloeobacter violaceus PCC 7421 [88]. So far, in cyanobacterial species where the expression of the psbA gene family has been characterized, Synechococcus 7942 is the only species that does not contain such a gene [32].
During the past year, two independent studies demonstrated induction of the psbA gene encoding the D1′ isoform under microaerobic [9] or low oxygen [10] conditions in Synechocystis 6803, Anabaena 7120, Thermosynechococcus elongatus BP-1, and Cyanothece sp. ATCC 51142 (Fig. 2c). Already earlier, the Anabaena psbA0 (alr3742) gene was found to be expressed at a very low constitutive level and was not responsive to photo-oxidative UVB stress, light stress, or nitrogen stress [95]. This behavior of psbA0 of Anabaena 7120 is very similar to that of psbA1 of Synechocystis 6803. Indeed, both these genes are in fact not silent even under normal growth conditions, but transcribed to a very low level insufficient to play a significant role as part of the cellular psbA transcript pool [95].
While still we do not have an answer regarding the direct functional role of the D1′ isoform, we can now with a high degree of confidence establish D1′ as a widely distributed D1 isoform important for adaptation of cyanobacterial cells to specific environmental conditions, particularly to a microaerobic pressure. The conditions requiring the presence of D1′ thus have a sufficient evolutionary importance to ensure that the gene has been selectively maintained in the genome. There are no major sequence characteristics that would individualize the D1′ from the other D1 forms. In the D1 forms functionally characterized [9], there are only three amino acid residues that are both conserved in all traditional D1 forms and mutated but conserved across all the D1′ proteins. These modifications are Gly80Ala, Phe158Leu, and Thr286Ala [9]. The low-oxygen-pressure-induced D1′ form encoded by psbA2 in Cyanothece [10] also has the same consistent modifications. Some initial molecular modelling studies did not show any major D1 conformational changes [9], and it is not yet clear what, if any, are the implications of these modifications on the general PSII function. It is worth mentioning, however, that at least two of the conserved modifications in D1′ (Phe158Leu and Thr286Ala) are located in the binding pockets of important Chl residues: ChlD1, the place of initial charge separation, and Chl PD1, the most probable location of the stable cation P680+. The fact that the genes encoding D1′ are present in such a diverse array of cyanobacterial species and that all these genes apparently respond to the same environmental cue makes it very likely that the encoded D1 protein belongs to a distinct category: D1′. The clear selective advantage of D1′ over the regular D1 is not obvious and remains to be identified. Also, while under standard laboratory growth conditions the cells do not usually experience periods of microaerobic growth, in the natural environment they do occur as a result from imbalanced cellular metabolism, specific environmental conditions or due to niche a given cyanobacteria inhabits, e.g., in microbial mats [97].
psbA genes in cyanophages
Cyanobacteria are responsible for a majority of primary production in oligotrophic regions of oceans. Cyanophages, viruses that infect cyanobacteria, are equally abundant in marine ecosystems, and probably exert major ecological effects on the marine environment [98, 99]. The first cyanophages were isolated in 1963 [100], but only recently it has been found that many of the cyanophages infecting Synechococcus and Prochlorococcus hosts possess psbA and psbD genes encoding the D1 and D2 proteins, respectively [101, 102, 103, 104, 105, 106, 107]. The photosynthesis genes in cyanophages originate from their host cyanobacteria, and thus show marked (up to 95%) identity to their host homologs at amino acid level [103, 105, 108, 109]. Accordingly, Prochlorococcus myoviruses and podoviruses encode the only D1 form found in Prochlorococcus, whereas in Synechococcus myoviruses, the stress-responsive D1:2 form of Synechococcus D1 protein has been selected over D1:1 [107].
It has been shown that the cyanophage psbA gene is indeed both transcribed and translated in cyanobacteria during infection [104, 110], and at the same time the expression of the host photosynthesis genes declines [104]. Since all bacteriophages rely on their hosts to provide energy and carbon sources for replication and assembly, it is conceivable that the expression of the cyanophage psbA gene allows continuous operation of the PSII repair cycle (Fig. 1), even if infection down-regulates the expression of the host psbA gene. Thus, the D1 protein encoded by the cyanophage is likely to replace the light-sensitive D1 protein of the host (despite possessing some unique features in structure [111]), thus allowing photosynthesis to continue efficiently even under bright light [104, 110, 112, 113], and thereby provide the energy needed by the virus for its replication. Obviously, the prevalence of photosynthesis genes in cyanophages serves as a valuable genetic reservoir for the host, and has probably played a role in driving host niche differentiation [105].
The D1 protein of PSII is a specialized protein component targeted to photodamage and rapid turnover in all organisms performing oxygenic photosynthesis. In cyanobacteria, which can inhabit various extreme habitats, the D1 protein is encoded by a small gene family. Obviously, the range of environmental factors cyanobacteria may face during their life cycle requires the possibility to modify the expression of the psbA genes, and more importantly the presence of a proper D1 isoform in PSII centres, in order to guarantee the best adaptation and fitness under given conditions. Optimal adaptation to varying environmental conditions seems to be obtained via complex regulation of not only one, but several, psbA genes. During the past two decades, it has become clear that there is not a single, universal pattern of regulation of psbA gene expression in cyanobacteria, but rather the ecological niche occupied by the individual strain has guided the evolution towards proper adaptation and satisfactory fitness. Ongoing release of genomic data from a vast number of cyanobacterial species will certainly provide additional tools to further study the regulatory factors of psbA gene expression in different cyanobacterial species. Understanding the adaptation processes of the photosynthetic machinery to changing environmental cues will be of utmost importance when biotechnological applications, for example the production of biofuels and improvement of crop yields, are to be developed.
Dr. Taina Tyystjärvi is thanked for critical reading of the manuscript. Research in the authors′ laboratory is financially supported by the Academy of Finland (110099, 118637), Nordic Energy Research Program (BioH2), European FP7 SOLAR-H2 Program (contract 212508), and Maj and Tor Nessling Foundation.
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When Cisco killed the Flip brand, it looked like the pocket Internet camcorder era had peaked. But most other companies have merrily continued to release models at this level regardless. Toshiba has been focusing on the budget end of the market for longer than most, although its camcorders haven't usually conformed entirely to the pocket Internet genre. The Camileo B10, however, fits quite squarely into this niche, although there are some notable differences.
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2013 Cadillac ATS GM Takes Aim at BMW. No, Seriously.
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I threw out the mailer I received from BMW a couple of days ago and also the email that I received. This is what I copied from the BMWUSA site:
Enjoy a Holiday Credit of up to $3,500 on select models at the BMW Happier New Year Event, now until January 2, 2013.1
Current and returning BMW owners can get $750 in lease/finance Loyalty Cash on any new 2012 or 2013 BMW.2
Existing BMW Financial Services customers can take up to $700 off a new 2012 or 2013 BMW. Just download the BMW Ultimate Drive App and take a test drive.
Offer ends December 31, 2012.3
I'll ask my dealer what cars BMW includes on this deal when I take my wife's car in for service for something they screwed up last week when it was there. I love wasting my time!
There's $1000 incentives on 2013 328s, $1500 on 328 coupes etc. $3500 is probably on 7 series or something.
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Bodie Hooks onto victory No. 1,000
Astros' Double-A skipper earns milestone in Corpus Christi
By Andrew Pentis / Special to MLB.com | May 11, 2012 7:55 PM ET
Then the Hooks started losing and Bodie started worrying.
"The way we've been playing," he said, "I was hoping we weren't going to lose 1,000 before we won 1,000."
Bodie can put that concern to bed.
Corpus Christi's skipper achieved the milestone Friday night in the Hooks' 4-2 victory over the visiting Frisco RoughRiders. He was doused with ice from the dugout water cooler after the final out was recorded.
"That woke me up to the moment," the 56-year-old Brooklyn, N.Y., native said.
Bodie became the third Minor League manager this season to reach 1,000. He was preceded by Double-A Trenton's Tony Franklin on April 11 and Double-A Erie's Chris Cron on April 21.
With 16 years of managerial experience, Bodie improved to 1,000-945, although he's managing for the first time since 2005.
"I was the hitting coach at Triple-A, in Oklahoma City, last year, and I felt like I was on the sidelines a little too much," he said. "I wanted that platform to have bigger impact on the players.
"It's like riding a bike. It feels very comfortable."
Bodie and the Hooks (14-20) received a strong start from Astros' No. 7 prospect Brett Oberholtzer, who allowed a run on five hits -- including Jurickson Profar's sixth-inning solo homer -- over seven innings.
"He threw outstanding," Bodie said. "He put the brakes on their whole offense, had three pitches working. I think he dominated. He threw one bad pitch, a 2-0 fastball, that was hit for a home run."
The guilty party there, Profar, extended his hitting streak to 21 games. The Rangers' No. 1 prospect also plated a run in the eighth with a sacrifice fly and has reached base safely in 29 consecutive games.
Frisco starter Jake Brigham gave up two runs on five hits, including Jair Fernandez's solo homer, over seven frames.
Andrew Pentis is a contributor to MLB.com. Follow him on Twitter at AndrewMiLB. This story was not subject to the approval of the National Association of Professional Baseball Leagues or its clubs.View More