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Zeta Petroleum To Benefit From Romanian Gas Market Deregulation 0 comments
Jul 26, 2012 1:27 AMZeta Petroleum (ASX: ZTA) will see an improvement to the economics of its Romanian projects after the Government put in place legislation to deregulate its domestic gas market.
Under the new Energy and Gas Law No. 123/2012, regulated prices for end clients will be gradually eliminated with the changes taking effect for non-household clients from 1 December 2012 and from 1 July 2013 for household clients.
"Zeta has long been attracted to Romania given its favourable and stable fiscal regime, proven hydrocarbon system and long established oil and gas industry," managing director Stephen West said.
"The announcement of the official conversion into Romanian legislation of the EC directive supports our strategy to invest in Romania as we expect domestic gas prices to commence converging with Western European prices from early 2013."
Gas prices in Romania currently average about US$4.25 per thousand cubic feet though West had previously said that deregulation could send prices up to about US$10.50 per thousand cubic feet, more than doubling its future revenue.
Bobocu gas field
Zeta has started its six well drilling aimed at bringing the Bobocu gas field back into production with the spudding of the Bobocu 310 appraisal and development well earlier this week.
The program targets identified mean contingent gas resources of 44.8 billion cubic feet (Bcf) of gas, or 7.71 million barrels of oil equivalent (MMboe), along with mean prospective gas resource of 68.73Bcf or 11.82MMboe.
This is based on the company's understanding of the geology and distribution of hydrocarbons across the field from the extensive technical studies, geological modelling and a 75 square kilometre 3D seismic survey that it carried out.
Additional potential that was identified in shallower and deeper horizons will be evaluated during this program.
Any gas discovered will find ready buyers in both Romania and the broader European market, where there is a shortage of gas supply.
Proactive Investors is a market leader in the investment news space, providing ASX "Small and Mid-cap" company news, research reports, StockTube videos and One2One Investor Forums.
Back To Proactive Investor's Instablog HomePage »Instablogs are blogs which are instantly set up and networked within the Seeking Alpha community. Instablog posts are not selected, edited or screened by Seeking Alpha editors, in contrast to contributors' articles.
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J.K. Rowling�is ushering in a new holiday celebration: the 12 days of Harry!The author announced on�Pottermore�that she would be releasing new�
Bones�star�Emily Deschanel�is pregnant,�
The Los Angeles Police Department has opened an investigation into allegations that�Bill Cosby�molested...
PEORIA — Manual came into its home opener and Big 12 Conference debut unbeaten and ranked No. 8 statewide in Class 3A.
PEORIA — Nine months after closing the book on the Mid-State 6 Conference, the Peoria High boys basketball team began writing Volume 1 of its Big 12 odyssey Friday night.And though the story begins...
High school girlsFriday's gamesMID-ILLINIPekin 83, Dunlap 58
CHICAGO — Most of the bounces — even crazy, hard-to-explain ones — seem to be going the Chicago Blackhawks' way. Brandon Saad scored with 27 seconds left in the third period and...
Willie Cauley-Stein scored a season-high 21 points and made several defensive plays to key host Kentucky's 18-2 blitz to open the second half, helping the top-ranked Wildcats pull out a 63-51 win over No. 6 Texas on...
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« Modes réels collectifs », for the first time remastered and presented in vynil and cd format, is the first work published in 1981 under the unique banner Vivenza, after he performed in the bands Glace (1976) and Mécanique Populaire (1979). It must be seen as a kind of genuine « Manifesto » that will then become emblematic of all his work.
By suggesting to us, on a philosophical as well as sonorous point of view, an idea of opening to the real directly inspired from the futuristic theories, Vivenza was then presenting to us what he brought from his relatioship with the machines and with the sound of the factories, to make us penetrate the brutal concrete of the heavy industry world, that's to say according to his « Modes réels collectifs » of existence.
This album, by virtue of its initial founder characteristic from 1981, shows this concern about participating to the interaction of the energy of the future, to the reciprocal mediation linked to the productive activity of the world. Like Vivenza will say: « By working and through the strengths of work, in the sonorous magma of the industrial society, at the heart of the forges and weirs, of the rolling mills and power stations, reactors and artificial intelligence, nature reveals its dynamic characteristic. The power of machines, and of industry in general, are the truth of the world. » Thus, industry and technique are what we could call the principle of any reality, because, like the futurists said accurately: « the machine is indeed the richest symbol of the mysterious human creator strength ». [1] 1. PRAMPOLINI, PANAGGI, PALADINI, Manifesto of Mechanical Art, 1923.


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Squirrels are Barking...Politicians, Too
It certainly feels like fall. Every time I walk outside I hear squirrels barking around my house. Eating pecans, chasing each other...just another week. I can't wait to get out there. I already scouted a good spot in the Homochitto. In years past, we have worked in an early trip or two, but kept it limited so as not to disturb bowhunters. Not this year, I figure as big as the Homochitto is, there's room for everybody.
With the election just around the corner, the politicians are at it, too....harder and dumber than ever. Our candidates, who both make claims about themselves that are just short of walking on water, are the exact people who got us in the mess we're in. (save Palin...she had nothing to do with it) Both promise unrealistic tax cuts, one says he will not raise taxes on households that take in less than $250,000 per year. (Wow, Gee thanks! Was an increase on those households ever even on the table?) Sad. Sad. Sad.
Two absolutely useless candidates.
I guess we should be happy that hunting season is here and we can forgot about these Bozo's for at least a few hours a week.


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Welcome, Sign In
Keloid Scar Tissue Treatment
Description:After suffering from chronic acne in his teenage years, Luke, 23, developed keloid scars on his chest, shoulders and back. Luke visits dermatologist Dr. Sandra Lee, who uses a laser treatment, in combination with intralesional steroids, to combat his chronic skin condition. (3:37)
Source: The Doctors
Tags: Keloid Scar Tissue Treatment , chest scars , Dr. Sandra Lee , gross anatomy


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Top Definition
An environmentally friendly resources conservation measure where the toilet is flushed only after the bowl contains the crap of multiple cable laying sessions. A specific sub-type of "going green".
Going brown could potentially save 150 million gallons of water annually.
wolfbait51가 작성
2011년 04월 04일 (월)
1 more definition
To fuck someone in the ass.
Jane is on her rag so I guess I'll be going brown tonight with her
2003년 01월 09일 (목)


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My bad.
Erhm.... That's a good point I didn't think about.
My bad.
Erhm.... That's a good point I didn't think about.
- Nate the
321 This place has gone quiet...
- Nate the
318 Quite honestly, before all of you came here (thanks, by the way) most interactions, especially late-night ones, were silent.I didn't want to talk because fatigue affects me a lot, I don't think (or post!) clearly and consequently avoid talking.
It's actually from Seihou, where Marisa was an EX-Boss with Reimu. They use a remix of her SOEW theme in the fighting games. (Love Colored Magic, Love Colored Master Spark, sometimes Oreintal Dark Flight and Magus night. Though Witches Ball is just from Seihou)
312 Oh yeah, forgot that the fighting games only make music for the characters they introduce.
Should I download IaMP or SWR?
Last edited by BritishLanguage; 13th June 2013 at 5:55 AM.
Yup, and for Yukari because the gap hag has that power. (Basically Broken Moon, Night Falls, Missing Power, Iku's Theme, Bhava-Agra seen through a child's mind, and Lost emotion.) HM's mix of Hartmann's Youkai Girl is really fitting for Koishi.
310 Oh, you didn't catch my edit, I made it too late.
Should I download IaMP or SWR?
If you're a story mode player, get SWR. If you don't care about it get UNL too and then tell your UNL where SWR is.
I suggest Sakuya, she's easy to learn and powerful. I main Reisen and Youmu though, and use the Gap Hag and Tenshi from time to time
308 Alright, probably not going to be on this thread for one of two reasons:
1. Enjoying Scarlet Weather Rhapsody
2. Punching my goddamn keyboard because it won't fucking work fuck fuck shit
Last edited by BritishLanguage; 13th June 2013 at 6:06 AM. Reason: never mind, takes a few minutes to download
Hmm, the only character you'll be missing in UNL is Okuu who is broken, and rebalanced characters. Also beware of Alice on Lunatic. Seriously, the AI if set up wil ldestroy you. Alice is argueable the best corner abuser in the game, anyways gl and enjoy SWR


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I have had my MX-600 Zoom for many years now. Bought it in Singapore, where consumer electronics are cheap. When I bought it it was state of the art and very feature rich. There are, compared to todays digital cameras, not a whole lot of buttons and features. It's easy to use, the image quality is great. Light, white ballance etc. can be adjusted manually - but it's very combersome. Back then it was a good camera, but it's slow and very noisy. Problems: The MX-600 Zoom uses SmartCard Flash ...
View review in forums Leave a comment on this reviewFujifilm MX-600 Zoom
Announced Feb 4, 1999
1 megapixels | 1.8″ screen | 38 – 115 mm (3×)


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This site is dedicated to providing the residents and Non Residents of Brevard County Florida, Cocoa, Rockledge, Merritt Island, Melbourne, Satellite Beach, Palm Bay, Cocoa Beach, Port St. John, Mims, Titusville, Scottsmore, Cape Canaveral, Suntree, Viera, Indialantic, Satellite Beach, and the State of Florida with viable information on Concealed Weapon Permit Training, Basic & Advanced Firearm Training, and information on the laws of firearm use and owner ship, along with Marksmanship Instruction in rifle, shotgun and handgun. Non-Residents see us about a non-resident Florida
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Register & Pay Deposit For Up Coming Course
When you pay the deposit it also serves as your
registration. You will later receive 2 e mail's
one from the credit card company which is
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the registration feel free to call.
Notary Service may be available at the course
from Mobile Notary Services for $10. per
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the usual fee is $20 per form. There is a place
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If you have a CCW and carry a firearm or other
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Other Links of Interest at our Market Place
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Thomas Jefferson quoting Cesare Beccaria: �Laws that forbid the
carrying of arms . . . disarm only those who are neither inclined nor
determined to commit crimes . . . Such laws make things worse for
the assaulted and better for the assailants; they serve rather to
encourage than to prevent homicides, for an unarmed man may be
attacked with greater confidence than an armed man.�


text:
Read the letter from 25 local officials opposing Chuck Reed's public pension proposal
In case you missed it, this week's State Worker column reports on a new twist in the ongoing tussle over San Jose Mayor Chuck Reed's public pension proposal: some local officials oppose the idea.
Reed has argued that local governments, whose budgets are particularly susceptible to pension-cost hikes, need the clear authority to prospectively cut benefits for current employees. His measure, aimed for the November 2014 ballot, would let them do that (while leaving accrued benefits unchanged). The letter, released by a union coalition fighting Reed's ballot proposal, is designed to chip away at that assertion. Here's the letter:
Local officials' letter to Mayor Reed
With just 400 to 450 words for our weekly State Worker column, much of what we learn each week never sees print. Column Extras give you some of the notes, the quotes, the documents and the observations that inform what's published.
About This Blog
Jon Ortiz launched The State Worker blog in 2008 to cover state government from the perspective of California government employees. Every day he filters the news through a single question: "What does this mean for state workers?" Join Ortiz for updates and debate on state pay, benefits, pensions, contracts and jobs. Contact him at firstname.lastname@example.com or 916-321-1043. Twitter: @TheStateWorker.The State Worker
Join the Discussion
The Sacramento Bee is pleased to provide this opportunity to share information, experiences and observations about what's in the news. Some of the comments may be reprinted elsewhere on the site or in the newspaper. We encourage lively, open debate on the issues of the day, and ask that you refrain from profanity, hate speech, personal comments and remarks that are off point. Thank you for taking the time to offer your thoughts.


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Save the Dates
Thursday, Jan 29, 2015: The 'worms are cold Join us in reading Smila's Sense of Snow by Peter Hoeg.
Thursday, Feb 26, 2015: The 'worms add and subtractThe February book is The Mathematician's Siva by Stuart Rojstaczer.
Thursday, Mar 26, 2015: Who said the Bookworm?March is for the Book of Unknown Americans by Christina Henriquez.
Thursday, Apr 30, 2015: 'Worms doing the romance thingSaltwater Kisses by Krista Lakes is the April choice (Do you dare to read the entire series?)
Thursday, May 28, 2015: The 'worms go AsianExplore a different culture as the Bookworms read Daughters of the Dragon by William Andrews
New Meeting Day and Time!
Beginning with the January 29th meeting, the McClellan Bookworms will be meeting on the last Thursday of each month. Meetings have moved to the library and will begin at 12:00pm. Oh, bring your lunch! We're going to eat and talk in the library!
Semester Reading
Smilla's Sense of Snow
Publication Date: 1995-10-01
She thinks more highly of snow and ice than she does of love. She lives in a world of numbers, science and memories--a dark, exotic stranger in a strange land. And now Smilla Jaspersen is convinced she has uncovered a shattering crime.
It happened in the Copenhagen snow. A six-year-old boy, a Greenlander like Smilla, fell to his death from the top of his apartment building. While the boy's body is still warm, the police pronounce his death an accident. But Smilla knows her young neighbor didn't fall from the roof on his own. Soon she is following a path of clues as clear to her as footsteps in the snow. For her dead neighbor, and for herself, she must embark on a harrowing journey of lies, revelation and violence that will take her back to the world of ice and snow from which she comes, where an explosive secret waits beneath the ice. (from Amazon)
It happened in the Copenhagen snow. A six-year-old boy, a Greenlander like Smilla, fell to his death from the top of his apartment building. While the boy's body is still warm, the police pronounce his death an accident. But Smilla knows her young neighbor didn't fall from the roof on his own. Soon she is following a path of clues as clear to her as footsteps in the snow. For her dead neighbor, and for herself, she must embark on a harrowing journey of lies, revelation and violence that will take her back to the world of ice and snow from which she comes, where an explosive secret waits beneath the ice. (from Amazon)
The Mathematician's Shiva
Publication Date: 2014-09-02
When the greatest female mathematician in history passes away, her son, Alexander "Sasha" Karnokovitch, just wants to mourn his mother in peace. But rumor has it the notoriously eccentric Polish émigré has solved one of the most difficult problems in all of mathematics, and has spitefully taken the solution to her grave. As a ragtag group of mathematicians from around the world descends upon Rachela's shiva, determined to find the proof or solve it for themselves—even if it means prying up the floorboards for notes or desperately scrutinizing the mutterings of her African Grey parrot—Sasha must come to terms with his mother's outsized influence on his life. (from Amazon)
The Book of Unknown Americans
Publication Date: 2014-06-03
*Starred Review* On a cold, bewildering night, the Riveras, who have just left their happy lives in Mexico, are dropped off at a dilapidated apartment building on the western edge of Delaware. Arturo has given up his thriving construction company to labor in a dark, grimy indoor mushroom farm, while his wife, Alma, lonely and afraid, with no English and little money, worries incessantly about their beautiful 15-year-old daughter, Maribel. She has suffered a traumatic brain injury, and her parents have sacrificed everything to send her to a special school. Their building turns out to be a sanctuary for Central and Latin American immigrants, and as the Riveras' dramatic tale unfolds, Henríquez brings their generous neighbors forward to tell the compelling stories of why and how they left Venezuela, Puerto Rico, Guatemala, Nicaragua, Colombia, Mexico, Panama, and Paraguay. As one man says, "We are the unknown Americans," those who are feared and hated. As Maribel opens up to Mayor, the infatuated boy next-door who is relentlessly bullied by his father and his classmates, terror of the unknown becomes a tragic force. Each scene, voice, misunderstanding, and alliance is beautifully realized and brimming with feeling in the acclaimed Henríquez's (The World in Half, 2009) compassionately imagined, gently comedic, and profoundly wrenching novel of big dreams and crushing reality, courageous love and unfathomable heartbreak. --Donna Seaman (Booklist)
Saltwater Kisses
Publication Date: 2013-06-22
When small-town girl Emma LaRue won a vacation to an exclusive tropical island, a last minute cancellation meant she would be going by herself. Shy and studious, she never had time to fall in love, and often wondered if she was just meant to be alone. However, that all changed when a handsome stranger literally walked into her life while on the beach and sparks began to fly. New York's most eligible billionaire bachelor Jack Saunders thought this vacation would be the perfect escape, one last hurrah, before taking full control of his father's company. When an innocent Emma didn't recognize him, he figured that he might get a chance to have a vacation from being rich. He didn't tell her about the cars, the yacht, or the penthouse. All he did was let her fall in love with him. Soon, Jack found that he was the one falling in love with Emma. When they enjoy a fantasy marriage ceremony on the beach, they thought it was a bit of harmless fun before returning to their normal lives. A bittersweet goodbye was supposed to be the end of their perfect vacation romance, but when photos of the ceremony were leaked to the press, everything changed. Feeling lied to and thrust into a world of wealth and privilege, Emma must choose between following her dreams or following her heart. Will she be content at being nothing more than the billionaire's wife, or will she return to her normal life with only memories of saltwater kisses? (from Amazon)
Daughters of the Dragon
Publication Date: 2014-01-01
2014 IPPY AWARD WINNER. DURING WORLD WAR II, the Japanese forced 200,000 young Korean women to be sex slaves or "comfort women" for their soldiers. This is one woman's riveting story of strength, courage and promises kept. In 1943, the Japanese tear young Ja-hee and her sister from their peaceful family farm to be comfort women for the Imperial Army. Before they leave home, their mother gives them a magnificent antique comb with an ivory inlay of a two-headed dragon, saying it will protect them. The sisters suffer terribly at the hands of the Japanese, and by the end of the war, Ja-hee must flee while her sister lies dying. Ja-hee keeps her time as a comfort woman a secret while she struggles to rebuild her life. She meets a man in North Korea who shows her what true love is. But the communists take him away in the middle of the night, and she escapes to the South. There, she finally finds success as the country rebuilds after the Korean War. However when her terrible secret is revealed, she's thrown into poverty. In the depths of despair, she's tempted to sell the comb with the two-headed dragon that she believes has no magic for her. Then one day she discovers its true meaning and her surprising heredity. And now she must find the only person who can carry on the legacy of the two-headed dragon… someone she abandoned years ago. Set within the tumultuous backdrop of 20th century Korea, Daughters of the Dragon by award-winning author William Andrews will make you cry and cheer for Ja-hee. And in the end, you'll have a better understanding of the Land of the Morning Calm. (from Amazon)
McClellan Bookworms
Who are the McClellan Bookworms?
- Well, way back in the year 2010 a group of intrepid folks decided that we were tired of competing to tell about our latest book find. We gathered together and drew book titles out of a pot in order to determine what to read. Since that time we have joyously read the same book each month. We meet, eat, laugh, and discuss our book choices.
Did it stop the problem of pushy book readers sharing their books?
- No, but it did add to the fun.
Who can be a bookworm?
- Anybody can be a McClellan Bookworm. Each month we share the web information about the book and discussion questions. We even have dedicated bookworms who fax in their responses! Yes, they are read to the group.
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12-03-2010, 07:36 PM | #11 (permalink) |
Join Date: Nov 2010
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That method was free! Not $1800! Anyway, there really isn't a ton of price gouging, although i do find it outrageous that, at least at my local hill, a 4 hour pass is $42 and a full day pass is $45. I mean come on... is $3 all that important to you!?
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12-03-2010, 08:53 PM | #13 (permalink) | |
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You surely exemplify the idiocracy i'm talking about. It's not a race. That's the mentality in Vancouver and city life talking. I'm not looking for cheap shite. I don't mind paying for good quality products at a reasonable price. A hand-crafted board is worth $750, I agree with that much. The problem is too much mass-produced, mass-marketed, overpriced junk that will fall apart faster than you can order your frappacrappa skinny milk mochaccinno at Starbucks. There was a time when Tom, Chuck, Jake and other pioneers did want to share the soul of snowboarding. A unique community was created. It was reasonable and still can be with good quality. You're attitude along with others can change and you have the power to demand quality without the hefty price tag. I think your problem, as many others is expensive 'race' winning gear, doesn't hide your inability to ride. And if you think you're race is worth it...well maybe try to define the word delusional. |
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To support, encourage, & promote interaction between Asian Men & Black Women.
I have decided to get a tat and would love to hear some creative ideas. I want to incorporate my name "NAM" into the design. Care to share? Cuz ya know, "sharing is caring." LOL.
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Started by tinker dotts. Last reply by Greg Sep 5, 2013. 3 Replies 1 Like
It all depends on how big u want it. I think they areAddictive lol I know 3 is nothin but I want more lolContinue
Started by Brittany Mc. Last reply by britt cole Aug 16, 2013. 13 Replies 0 Likes
My first one was the celtic symbol for trinty, the triquetra, an it means for me the Creator, Destroyer, Sustainer, i can't wait to get another one!!!!!Continue
Your tattoo is sick! I still haven't gotten my fourth one. For some reason, I can't find anything that speaks to me :(
i have a few more session im getting a full sleeve done
Maybe you could join the art group on here or post to Facebook for someone to modify it? Find someone who is willing and then tell them what you wanted added/taken away.
No, that's not me. I found that picture when I was browsing online. I'm not sure, but I think it is that girl's personal take on a dream catcher.
Anyway, it's very elegant and I like it. I want to change a few things to make more my style, but I suck at drawing lol
Is that you in the picture? I don't think it would look weird but I would probably change something about it, even a small detail, so it's not exactly the same.
Hi everyone! I'm so glad I found other people who share my love for tattoos lol. I have three so far, and I'm thinking about getting one mo
This is what I want! But I want to get it on my ribs on the left side of my body (from just below the armpit to about halfway down) What do you guys think? Would it look weird?
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Find a copy in the library
Finding libraries that hold this item...
Genre/Form: | Biographie Biography |
Named Person: | Irène Némirovsky; Irène Némirovsky; Irène Némirovsky; Irène Némirovsky |
Document Type: | Book |
All Authors / Contributors: |
Jonathan Weiss |
OCLC Number: | 636926555 |
Notes: | Originaltitel: Irène Némirovsky. |
Description: | XIII, 200 S : Ill ; 24 cm. |
Responsibility: | Jonathan Weiss. |
Publisher Synopsis
"In Nemirovsky's search to reconcile national, religious, and cultural identities, Weiss recognizes the struggle of many immigrants in France today. For this reason, more general readers may enjoy this biography as much for the fresh perspectives it bring to questions of national and cultural identity currently under debate in the Francophone world as for the insights it brings to Nemirovsky's life and literature. Scholars will value the complete bibliography of her published and unpublished works, Weiss's attention to the present controversy regarding Nemirovsky's anti-Semitism, and his sensitive and well informed readings of her work, including Suite francaise. This critical volume provides solid evidence that Nemirovsky should be included among the most important writers of twentieth-century French fiction." - Hollie Markland Harder, French Review "Immensely clarifying ... As Weiss's important and prodigiously researched biography makes clear, Nemirovsky was the very definition of a self-hating Jew." - The New Republic "[An] informative work, neatly structured to bring out the dramatic and tragic fate of a woman who, as [Weiss] puts it, 'died without ever having resolved the question of where she belonged.' It is a book well worth reading." - Australian Book Review "This short critical biography by Weiss, an expert on contemporary French literature, is a fine introduction to her work." - Publishers Weekly "[A] brief, but intensely thought-provoking biography by Jonathan Weiss." - The San Francisco Chronicle Read more...
Similar Items
Related Subjects:(16)
- Némirovsky, Irène.
- Némirovsky, Irène, -- 1903-1942.
- Authors, French -- 20th century -- Biography.
- Jewish authors -- France -- Biography.
- Authors, Russian -- 20th century -- Biography.
- Antisemitism -- France.
- œaAntisemitismœzFrance
- œaAuthors, Frenchœy20th centuryœvBiography
- œaAuthors, Russianœy20th centuryœvBiography
- œaJewish authorsœzFranceœvBiography
- œaNémirovsky, Irèneœd1903-1942
- Antisemitism.
- Authors, French.
- Authors, Russian.
- Jewish authors.
- France.


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Review posted for Sony Cyber-shot DSC-T33
Mike Tomkins
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The DSC-T33 continues a Sony tradition of high quality in the subcompact category, but with a new spin: Affordability!
Most impressive is how few tradeoffs Sony was forced to make to achieve this. The
shows good image quality, with good color, and high resolution. Its image sharpness and noise levels aren't quite up to the level of the best full-sized 5-megapixel cameras out there, but they're impressive for a subcompact model.
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Page 1
Triose phosphate isomerase, a novel enzyme-crystallin,
and s-crystallin in crocodile cornea
High accumulation of both proteins during late embryonic
Thandavarayan Kathiresan*, Kannan Krishnan*, Vaithilingam Krishnakumar, Raman Agrawal,
Amit Anand, Dasari Muralidhar, Anurag K. Mishra, Vishnu M. Dhople, Ramesh K. Aggrawal
and Yogendra Sharma
Centre for Cellular and Molecular Biology, Uppal Road, Hyderabad-500 007, India
Crystallins are defined as lens structural proteins, which
are classified as ubiquitous a-, b- and c-crystallins, and
taxon-specific crystallins [1]. The transparency and
refractive properties of lens depends on crystallins.
Taxon-specific crystallins are generally enzymes recrui-
ted as structural proteins in the lens to perform special-
ized functions, i.e., maintaining lens transparency [2].
Some examples of the species-specific crystallins are
d- and e-crystallins present in birds and reptiles, s-crys-
tallins in turtle and reptiles, q-crystallins in frog and
camel lenses [3]. Several of these crystallins are meta-
bolic enzymes: d-crystallin is argininosuccinate lyase [4];
s-crystallin is a-enolase [5]. They are believed to be
recruited by gene sharing or by gene duplication [2,6].
Some enzymes such as aldehyde dehydrogenase
class 3(ALDH3),aldehydedehydrogenaseclass 1
cornea; crocodile development; s-crystallin;
a-enolase; triose phosphate isomerase
Y. Sharma, Centre for Cellular and
Molecular Biology, Uppal Road, Hyderabad-
500 007, India
E-mail: firstname.lastname@example.com
R. K. Aggrawal, Centre for Cellular and
Molecular Biology, Uppal Road, Hyderabad-
500 007, India
Fax: +91 40 2716 0591
Tel: +91 40 2716 0222
E-mail: email@example.com
*These authors contributed equally to this
(Received 24 March 2006, revised 11 May
2006, accepted 26 May 2006)
Several enzymes are known to accumulate in the cornea in unusually high
concentrations. Based on the analogy with lens crystallins, these enzymes
are called corneal crystallins, which are diverse and species-specific. Exam-
ining crystallins in lens and cornea in multiple species provides great insight
into their evolution. We report data on major proteins present in the cro-
codile cornea, an evolutionarily distant taxon. We demonstrate that s-crys-
tallin⁄a-enolase and triose phosphate isomerase (TIM) are among the
major proteins expressed in the crocodile cornea as resolved by 2D gel elec-
trophoresis and identified by MALDI-TOF. These proteins might be classi-
fied as putative corneal crystallins. s-Crystallin, known to be present in
turtle and crocodile lens, has earlier been identified in chicken and bovine
cornea, whereas TIM has not been identified in the cornea of any species.
Immunostaining showed that s-crystallin and TIM are concentrated largely
in the corneal epithelium. Using western blot, immunofluorescence and
enzymatic activity, we demonstrate that high accumulation of s-crystallin
and TIM starts in the late embryonic development (after the 24th stage of
embryonic development) with maximum expression in a two-week post-
hatched animal. The crocodile corneal extract exhibits significant a-enolase
and TIM activities, which increases in the corneal extract with develop-
ment. Our results establishing the presence of s-crystallin in crocodile, in
conjunction with similar reports for other species, suggest that it is a widely
prevalent corneal crystallin. Identification of TIM in the crocodile cornea
reported here adds to the growing list of corneal crystallins.
ALDH, aldehyde dehydrogenase; IEF, isoelectric focusing; TIM, triose phosphate isomerase.
FEBS Journal 273 (2006) 3370–3380 ª 2006 The Authors Journal compilation ª 2006 FEBS
Page 2
(ALDH1) [7–10], transketolase [11] and isocitrate
dehydrogenase [12] in bovine and human cornea,
a-enolase⁄s-crystallin in chicken cornea [13,14] and
gelsolin in zebrafish cornea [15] are found in unusually
high concentrations. Because their expression in the
cornea, specifically in the corneal epithelium, is unusu-
ally high, they are thought to be functioning as struc-
tural proteins rather than enzymes. Based on the
analogy with lens crystallins, these enzymes are called
corneal crystallins [16]. The criteria for a protein to
qualify as a corneal crystallin are: (i) it must be
expressed at a high concentration, (ii) it should gener-
ally be an enzyme, though it might not perform an en-
zymatic function, and (iii) it should probably express
in corneal epithelium [9,16]. It is presumed that like
lens crystallins, the abundant corneal proteins might
contribute structurally to its transparency and optical
properties [9,17]. Because cornea is the outermost win-
dow or gate of the eye, some of the abundant corneal
proteins such as aldehyde dehydrogenase, also act as
an antioxidant filters and protect the cornea against
oxidative insults [7,18,19]. These tissue- and species-
specific crystallins represent recruitment of enzymes to
serve a structural role in the tissue, and this phenom-
enon has been termed as gene sharing and gene dupli-
cation [2,4,6,17].
Some of the corneal crystallins are also lens crystal-
lins. d-Crystallin, a major crystallin in birds and some
reptile lenses is present in bovine cornea [14]. ALDH
and glutathione S-transferase are both lens and corneal
crystallins in some species [20,21]. However, not all
corneal crystallins are found as lens proteins. s-Crys-
tallin is present in the mammalian cornea but absent
in mammalian lens. Just as there are taxon-specific lens
crystallins, there are taxon-specific corneal crystallins
e.g., gelsolin presents in zebrafish cornea, but not in
bovine cornea [15]. Similarly, aldehyde dehydrogenase,
which is present in bovine and human corneas, might
not accumulate in high concentrations in the chicken
cornea. It appears that corneas from various organisms
express different enzyme-crystallins, which could be
due to various divergent routes during evolution. It is
to be mentioned that corneas of only a few species
have been analyzed to date. Examining major proteins
in the cornea of multiple species provides great insights
into their evolution, as well as how crystallins adapt to
varying physiological demands. It is therefore interest-
ing to analyze corneas from diverse species and iden-
tify their corneal crystallins. In this regard, we have
selected crocodile cornea for the presence of corneal
crystallins. Crocodilians are evolutionarily interesting
vertebrates, supposed to be of the Triassic era. Cornea
(cyclophilin) and
of this taxon or its closely related species such as alli-
gator has not been analyzed previously. Any informa-
tion about such a species would add significantly to
our understanding of the origin and evolution of genes
and their functions.
In this work, we have performed the proteomics
analysis of the major proteins expressed in crocodile
cornea. We have found that a taxon-specific lens crys-
tallin, s-crystallin⁄a-enolase, is expressed in the cro-
codile corneal epithelium as a corneal crystallin.
s-Crystallin was first identified in turtle lens [22] and is
also expressed in the lenses of duck [23] and lamprey
[5]. We have reported earlier that s-crystallin is present
in crocodile lens, though as a minor fraction [24].
a-Enolase⁄s-crystallin has been identified previously in
the corneas of bovine and chicken [7,13,14]. In this
study, we show that s-crystallin is present in crocodile
cornea and have collected data on its expression and
abundance in corneal epithelium. We demonstrate that
the accumulation of s-crystallin in the corneal epithe-
lium increases with embryonic development, which
might coincide with the attainment of many optical
properties of the cornea, such as transparency and cur-
vature. In addition to s-crystallin, we have found triose
phosphate isomerase (TIM), a dimeric enzyme that
3-phosphate and dihydroxyacetone phosphate during
glycolysis [25], as another major enzyme-crystallin in
the soluble corneal fraction. Using immunofluores-
cence, we demonstrate that TIM accumulation in the
cornea increases during late embryonic development.
Our results suggest that s-crystallin is a widely preval-
ent corneal crystallin, and TIM is a novel enzyme-crys-
tallin expressed in crocodile cornea.
Results and Discussion
Though very diverse corneal crystallins have been iden-
tified in the corneas of bovine and chicken, no reptile
cornea has been studied to date [11,12,14,15]. Croco-
diles represent an evolutionarily important taxon
among various vertebrate species because it is believed
that they have survived and changed very little in
hundreds of millions of years, in spite of the global
catastrophe that caused the mass extinction of the
dinosaurs and many other animal species 65 million
years ago. Unlike the lens of many vertebrate species,
the crocodile lens has several different crystallins
(a-, b-, c-, d-, e-, s-crystallins and at least one crystal-
lin yet to be identified) [24], which makes this species
very interesting for studying various crystallins. We
were therefore interested in identifying and comparing
the major crystallins expressed in the crocodile cornea,
T. Kathiresan et al.
TIM and s-crystallin in crocodile cornea
FEBS Journal 273 (2006) 3370–3380 ª 2006 The Authors Journal compilation ª 2006 FEBS
Page 3
which would provide great insight into their evolution.
Some species of crocodiles, being endangered, are not
easily accessible for studies, thus further increasing the
significance of any experimental data, more so if these
are from the embryonic developmental stages.
SDS⁄PAGE yielded prominent protein bands in the
water-soluble fraction of crocodile cornea with molecu-
lar masses of approximately 20, 35, 47 and 50 kDa
(Fig. 1, lane 2). However, corresponding protein bands
of the same intensity are absent in the SDS⁄PAGE of
bovine corneal extract used for comparison (Fig. 1,
lane 3). The bovine corneal sample shows a prominent
54 kDa band corresponding to ALDH3 (earlier known
as BCP54) (Fig. 1, lane 3). Interestingly, a correspond-
ing band is not seen in the SDS⁄PAGE of crocodile
cornea. We further resolved the proteins expressed in
crocodile cornea by 2D gel electrophoresis and identi-
fied them by MALDI-TOF.
Identification of proteins expressed in crocodile
cornea by proteomics/MS methods
s-Crystallin/a-enolase is expressed as a major protein
in crocodile cornea
Two-dimensional electrophoresis of the soluble frac-
tion of corneal proteins revealed several spots scattered
all over the gel. Two of the major spots (spots 3 and
4) at pI ?6.5, adjacent to each other in the ?50 kDa
range (Fig. 2) were excised separately for MALDI-
TOF analysis. Both the spots matched to s-crystallin
with a score of 143 and 69 on mascot (Matrix Science,
www.matrixscience.com) (more than the accepted score
of 63) search for spot 3 and 4, respectively. The
molecular mass (48 kDa) of these proteins (spots 3
and 4) also matched with a-enolase⁄s-crystallin. In
total, 13 peptides generated after tryptic digestion
(Table 1). These MALDI-TOF results were further
confirmed by ESI-MS-QTOF MS⁄MS analysis (data
not shown). The difference in pI of two s-crystallin
spots could be because of partial post-translational
modification of the protein. Mobility difference of ver-
tebrate a-enolase isozyme has been observed earlier
on a native 2D SDS⁄PAGE due to phosphorylation
[26,27]. We suspect that there might be partial phos-
phorylation or some post-translational modification of
s-crystallin resulting in differential mobility on 2D gel
Triose phosphate isomerase (TIM) is a novel protein
in crocodile cornea
Another prominent spot (spot 1, Fig. 2) was analyzed
by MALDI-TOF. The scores on mascot (score 77)
and profound (score 2.43) programs (Rockefeller Uni-
versity, New York, NY, USA) used for the identifica-
tion were sufficiently high to establish the identity of
Fig. 1. SDS⁄PAGE [12% (w⁄v) gel] stained with Coomassie brilliant
blue. Lane 1, molecular mass markers; lane 2, crocodile total cor-
neal extract; lane 3, bovine corneal epithelium extract. Arrowhead
points to the prominent ALDH band.
Fig. 2. Protein profile of the total corneal extract of crocodile by 2D
gel electrophoresis on a 12% (w⁄v) gel. First dimension separation
by isoelectric focusing with pI range between 3 and 10; second
dimension separation by SDS⁄PAGE and stained with sensitive
Coomassie. Protein spots were excised and identified by MALDI-
TOF. Spot 1, triose phosphate isomerase; spots 3 and 4, s-crystal-
lin⁄a-enolase; spot 5, tropomyosin 2; spot 6, b-actin. Spots 2, 7, 8
and 9 could not be identified because of low confidence score.
TIM and s-crystallin in crocodile cornea
T. Kathiresan et al.
FEBS Journal 273 (2006) 3370–3380 ª 2006 The Authors Journal compilation ª 2006 FEBS
Page 4
this spot as TIM, an enzyme of the glycolytic cycle
that catalyzes the interconversion of d-glyceraldehyde
3-phosphate and dihydroxyacetone phosphate. Out of
the 17 peptides obtained from tryptic digestion of this
spot, 14 matched to TIM (molecular size 27 kDa) with
a score of 77 on mascot (Table 1). The accumulation
of TIM has not been identified earlier in the cornea of
any other species, and our data suggest that TIM qual-
ifies as a new enzyme-crystallin present in the crocodile
Other proteins in the crocodile cornea
We have attempted to identify the other major spots
obtained on 2D gel. We have identified tropomyosin 2
(spot 5), actin (spot 6), and myosin (spot 7) by
MALDI-TOF analysis (Fig. 2). Actin was reported as
a major intracellular water-soluble protein in zebrafish
along with gelsolin [15]. We could not establish the
identity of other spots (spots 2, 8 and 9 in Fig. 2)
because their scores on mascot and profound were
below the confidence level. We have not analyzed
the spots whose molecular masses were more than
70–80 kDa.
Localization and embryonic development studies
Different stages of embryonic development of crocodile
We followed the various stages of embryonic develop-
ment of crocodile as described by Ferguson [28],
however, with minor modifications based on our
experience with Crocodylus palustris development. The
embryonic development of crocodile is divided into 28
stages, spanning over 65 days. In the embryo of stage
9 (9 days after egg laying), the optic cup is large and
round but unpigmented. Up to stage 20 (28 days) the
embryo size is very small and fragile. In the embryo of
the 21st stage (31 days), a white ring in the iris sur-
rounds the outline of the lens of the eye and is over-
lapped by both upper and lower eyelids. Scales are
present on the dorsal and ventral aspects of the body
of the embryo of the 21st stage. In the embryo of the
22nd stage (35 days), the eye is almost fully developed.
Stages 23 and 24 represent 40 and 45 days after egg
laying. The embryo of the 25th stage (51 days) is like a
miniature version of the hatching. Hatching takes
place after 65 days (after the 27th stage of develop-
ment) of egg laying.
Expression of s-crystallin in crocodile cornea during
embryonic development
Although s-crystallin has been identified in the adult
bovine cornea, its expression during embryonic devel-
opment has not been studied. We have followed its
expression in crocodile cornea at various stages of
embryonic development by western blot analysis and
immunofluorescence using a polyclonal rat antiserum
raised against recombinant s-crystallin cloned from a
crocodile lens [29]. The western blot data showed that
Table 1. Protein identification for spots 1, 3 and 4 from 2D gel shown in Fig. 2 using MALDI-TOF peptide mass fingerprinting. Spots from
acrylamide gel were excised, protein was in-gel digested with trypsin, eluted and analyzed.
Spot on 2D gel
Measured mass
(error in p.p.m.)
score Residues
Matched peptide sequences
from database
Spots 3 and 4: s-crystallin⁄a-enolase1455.721()32)
1464.706 ()12)
1690.906 (11)
1804.041 (58)
1853.876 ()86)
1853.876 (18)
1918.038 (6)
1922.058 (36)
2003.205 (50)
2195.989 ()36)
2347.233 (21)
2533.182 ()37)
2669.344 (9)
143 and 181 2.43 and 2.2781–92
Spot 1: Triose phosphate isomerase1081.582 (11)
1457.804 (62)
1735.019 (69)
77 2.43
T. Kathiresan et al.
TIM and s-crystallin in crocodile cornea
FEBS Journal 273 (2006) 3370–3380 ª 2006 The Authors Journal compilation ª 2006 FEBS
Page 5
s-crystallin is present, albeit at a lower concentration,
in the corneal extract of the cornea from the stage 21
embryo and its expression increased during further sta-
ges of development as seen from the intensity of the
band (Fig. 3). These data suggest a gradual increase in
s-crystallin expression, assuming that the antibody spe-
cificity to s-crystallin does not change with develop-
We followed the expression of s-crystallin in croco-
dile cornea during embryonic development by immu-
nofluorescence using the above antibody. Corneas
obtained from different embryonic developmental sta-
ges (stages 21–25) and from a two-week old crocodile
were sectioned and stained for the presence of s-crys-
tallin. The cornea obtained from embryos of stages
earlier than 21st were fragile and were not studied.
Low or basal level of expression of s-crystallin was
observed in the corneas obtained from embryos in the
21st to 23rd stages of development (Fig. 4A,E,I).
Weak but detectable intensity of the antibody staining
was seen in the corneas, specifically in the corneal epi-
thelium from the 24th stage onwards (Fig. 4M,Q). The
intensity of s-crystallin staining in the cornea of a two-
week posthatched animal was the highest (Fig. 4U).
These results demonstrate that the accumulation of
s-crystallin begins towards the later stages of embry-
onic development and reaches its maximum as the
embryo attains maturity and hatches.
We also used a-enolase activity as a measure of its
level of expression in the cornea. The a-enolase assay
was performed with the water-soluble extract of croco-
dile cornea collected from various stages of embryonic
development and from a two-week old animal. As seen
from our analysis, the enzymatic activity increases in
the corneal extract with development. The corneal sol-
uble fraction from a two-week old animal exhibits
higher expression of s-crystallin (Fig. 5) provided there
is no change in its activity during development. These
results further support the above data suggesting that
the expression of s-crystallin in the cornea increases
during development.
Accumulation of TIM in crocodile cornea during
embryonic development
We also monitored the expression of TIM in the cor-
nea during embryonic development by western blot-
ting, immunofluorescence and enzymatic activity. The
antibody used was raised against recombinant TIM
cloned from Saccharomyces cerevisiae. As monitored
by western blot, the cornea obtained from embryos in
the 21st stage exhibited a comparatively low expression
of TIM, which increased in the cornea during further
development (Fig. 6).
Immunofluorescence data demonstrate a low level of
expression of TIM in the cornea obtained from the
21st and 22nd stages of embryos (Fig. 7A,E), which
increased with further development and reached its
maximum in the cornea obtained from a two-week
posthatched animal (Fig. 7U), confirming that TIM
observed that TIM is largely expressed in the corneal
epithelium with low expression in the stromal region
(Fig. 7). We also assayed the TIM activity in the total
corneal extract and found that it possesses significant
enzymatic activity even as early as the 21st stage of
embryonic development. TIM activity increased in the
corneal extract with the advancement of embryonic
development (Fig. 8). Maximum activity was seen in
the extract collected from a two-week-old crocodile
cornea (Fig. 8). These results suggest that similar to
development. We also
Fig. 3. Detection of s-crystallin by western blot using anti-s-crystal-
lin polyclonal IgG in the cornea collected from different stages of
crocodile embryos and from a two-week posthatched animal. Num-
bers above the lanes indicate the stages of embryonic development
(from 21st to 25th stages). PH indicates cornea from a two-week
posthatched crocodile. Solid arrowhead points to the prominent
band of s-crystallin. Lower panel: the blot after Ponceau staining,
demonstrating that equal amount of proteins were transferred from
gel to membrane.
TIM and s-crystallin in crocodile cornea
T. Kathiresan et al.
FEBS Journal 273 (2006) 3370–3380 ª 2006 The Authors Journal compilation ª 2006 FEBS
Page 6
s-crystallin, accumulation of TIM also increases in the
cornea with embryonic development.
s-Crystallin and TIM are putative corneal crystallins
in crocodile cornea
Our results demonstrate that (i) s-crystallin and TIM
are among the major proteins expressed in crocodile
cornea, (ii) they are expressed mainly in the corneal
epithelium, and (iii) they accumulate during late stages
of development. These are some of the criteria by
which a crystallin is defined [9,16]. These proteins
might be performing a structural role and are crystal-
lins in crocodile cornea. We have observed that the
cornea from an embryo of the 21st and 22nd stages is
Fig. 4. Immuno-localization of s-crystallin in
crocodile cornea during embryonic develop-
mental stages and in a two-week post-
hatched animal. (A–D) Stage 21 (31 days
after egg laying); (E–H) stage 22 (35 days
after egg laying); (I–L) stage 23 (40 days after
egg laying); (M–P) stage 24 (45 days after
egg laying); (Q–T) stage 25 (51 days after egg
laying); (U–X) cornea from a two-week post-
hatched crocodile. Column 1 (A, E, I, M, Q
and U) shows anti-s-crystallin IgG staining
(green); column 2 (B, F, J, N, R and V), propi-
dium iodide staining showing nuclei in red
colour; column 3 (C, G, K, O, S and W) shows
s-crystallin and propidium iodide staining
overlapped; column 4 (D, H, L, P, T and X)
represents transmission phase contrast.
Scale bar 20 lm.
Fig. 5. a-Enolase activity in corneal extract collected from the
embryos of different developmental stages and from a two-week
posthatched crocodile. h 21st stage; h 22nd stage; n 23rd stage;
s 25th stage; d two-week posthatched crocodile.
T. Kathiresan et al.
TIM and s-crystallin in crocodile cornea
FEBS Journal 273 (2006) 3370–3380 ª 2006 The Authors Journal compilation ª 2006 FEBS
Page 7
opaque. Therefore, our data on the accumulation of
s-crystallin and TIM during the late stages of develop-
ment suggest that they might have a role in providing
the necessary optical properties, such as transparency
and curvature to cornea during its development.
In conclusion, by proteomics and mass spectroscopic
analysis, we have identified s-crystallin, a prevalent
corneal crystallin, and TIM, a novel corneal crystallin,
as two crystallins in the cornea of crocodile. Our data
support the earlier observation that s-crystallin is a
widely prevalent corneal crystallin, present in the cor-
nea of bovine, chicken and crocodile. The expression
of TIM in the corneas of other species is yet to be
identified. Both crystallins accumulate during later sta-
ges of embryonic development. Our work further sug-
gests the diversity of corneal crystallins. Crystallins in
the corneas of diverse species need to be investigated
for comparative purpose, which would help in under-
standing the evolution of transparency of lens and
Experimental procedures
Crocodile cornea
Fertilized eggs of Indian mugger (Crocodylus palustris) were
collected from the Nehru Zoological Park, Hyderabad,
India on day zero of egg laying and were incubated in the
laboratory at 30–32 ?C. Proper guidelines of ARVO were
strictly followed in the handling of animals and were
approved by the Institutional Ethics Committee. Embryonic
developmental stages of crocodile have been classified previ-
ously [28]. Accordingly, there are 27 stages of embryonic
development, after which hatching takes place. Corneas
were dissected from the embryos between the developmen-
tal stages 20–25, and also from two-week old animals. Cor-
neas collected from crocodile embryos were homogenized in
50 mm Tris, pH 7.5, 100 mm NaCl, 0.02% (v⁄v) sodium
azide, 1 mm EDTA, centrifuged at 10 000 g for 15 min at
4 ?C and the supernatant collected.
2D gel electrophoresis
2D electrophoresis was performed using immobilized pH
gradient strips of 7 cm and pH range 3–10 (Bio-Rad, Her-
cules, CA, USA). The samples were resolved by isoelectric
focusing (IEF) in the first direction and by SDS⁄PAGE
[12% (w⁄v) acrylamide] in the second direction. In brief,
40–50 lL aliquot of the sample containing about 60 lg of
the protein was premixed with rehydration buffer [8 m urea,
2% (v⁄v) CHAPS, 50 mm dithiothreitol and 0.2% (v⁄v) of
3–10 ampholytes] and the rehydration of the immobilized
pH gradient strips was carried out for about 12–15 h. The
rehydrated strip was subjected to IEF at a current of
50 mAÆstrip)1with an end voltage of 8000 V. After the IEF
run, the gel strips were equilibrated with equilibration buf-
fer and placed on a precast SDS-polyacrylamide gel. The
gel was stained with Coomassie blue R-250 and destained.
The image was acquired using Fluor-S Multi-imager
(Bio-Rad) and the protein spots were identified based on
their discrete presence on the gel and processed further for
In-gel tryptic digestion
The selected spots were manually excised from the gel and
were subjected to in-gel digestion using sequence grade
bovine trypsin (Sigma, St Louis, MO, USA) reconstituted
in 25 mm ammonium bicarbonate at pH 8.0. The molar
ratio of protein to trypsin was maintained between 10 : 1
and 30 : 1, and the mixture was incubated at 37 ?C for
16–18 h. After the incubation, peptides from the gel were
extracted twice using 100 lL of 50% (v⁄v) acetonitrile con-
taining 5% (v⁄v) trifluoroacetic acid for about 30 mins by
vortex. The extracts were pooled, lyophilized and used for
MALDI-TOF analysis.
Fig. 6. Detection of TIM by western blot using anti-TIM polyclonal
IgG in different stages of embryos and from a two-week pos-
thatched crocodile. Numbers above the lanes indicate the stages of
embryonic development (from 21st to 25th stage). PH indicates
cornea from a two-week posthatched crocodile. Solid arrowhead
points to the prominent band of TIM. Lower panel: the blot after
Ponceau staining, demonstrating that equal amount of proteins
were transferred from gel to membrane.
TIM and s-crystallin in crocodile cornea
T. Kathiresan et al.
FEBS Journal 273 (2006) 3370–3380 ª 2006 The Authors Journal compilation ª 2006 FEBS
Page 8
Mass spectrometry
The peptides were analyzed for peptide mass fingerprinting
using the Voyager-DE-STR MALDI-TOF mass spectro-
meter (Applied Biosystems, Foster City, CA, USA) using
a-cyano-4-hydroxy cinnamic acid as matrix. The reconstitu-
ted peptides in 50% (v⁄v) acetonitrile containing 0.1% (v⁄v)
trifluoroacetic acid were spotted on a 96-well plate. The
MALDI-TOF spectra were recorded in the Reflectron mode
using the following parameters: accelerating voltage 20 kV,
grid voltage 72%, delay time 200 ns, low mass gate 750, scan
range 800–4000 and accumulation from 100 laser shots.
Determination of the internal sequence was done by
MS⁄MS fragmentation of the peptides using QSTAR
Pulsar ESI-QTOF (PE SCIEX, Applied Biosystems, Foster
City, CA, USA) and nano spray source. TOF-MS spectra
were obtained in the range m⁄z, 500–1700 at 1000 V spray-
ing voltage. The multiply charged species were subjected to
fragmentation by CID using collision energy ranging from
30 to 50 eV.
Protein identifications
A peak list was generated from the MALDI-TOF spectrum
screened for the presence of nonpeptide peaks. The proteins
were identified with peptide mass fingerprinting using NCBI
protein database and by mascot (Matrix Sciences) and
profound (Rockefeller University) search programmes.
The search criteria fixed were partial methionine oxidation,
and finally,
Fig. 7. Immuno-localization of TIM in croco-
dile cornea from early embryonic stages and
from a two-week posthatched crocodile. (A–
D) Cornea from embryo of 21st stage; (E–H)
cornea from 22nd stage; (I–L) cornea from
23rd stage; (M–P) cornea form 24th stage;
(Q–T) cornea from 25th stage; (U–X) cornea
from a two-week posthatched crocodile. A,
E, I, M, Q and U show anti-TIM IgG staining
(green). B, F, J, N, R and V propidium iodide
staining showing nuclei (red). C, G, K, O, S
and W show TIM and propidium iodide stai-
ning overlapped. D, H, L, P, T and X repre-
sent transmission phase contrast. Scale bar
20 lm.
T. Kathiresan et al.
TIM and s-crystallin in crocodile cornea
FEBS Journal 273 (2006) 3370–3380 ª 2006 The Authors Journal compilation ª 2006 FEBS
Page 9
carbamidomethylation of cysteines and up to maximum of
two miscleavages with a mass error of 100 p.p.m. For the
searches, molecular mass within ±5 kDa and PI within
±0.5 units of the molecular mass and pI observed in 2D
gels were used.
Protein ID was accepted if the scores on mascot and
profound were at least 63 and 1.5, respectively. Protein
identification was further validated by examining the num-
ber of peptides matched (at least 5) and the sequence cover-
age (15–20%). The ID of some proteins was further
confirmed by MS⁄MS fragmentation and searches were car-
ried out using sequences that were analyzed manually by
bio-analyst (PE Sciex, Applied Biosystems).
Purification of recombinant s-crystallin and
generation of polyclonal s-crystallin/a-enolase
and TIM antibodies
Recombinant s-crystallin was cloned and overexpressed
from crocodilian lens as described earlier [29]. Antibody
against recombinant a-enolase⁄s-crystallin was raised in a
six-month-old mouse using the standard procedure. Ani-
mals were bled three days after the final booster. Serum
was separated and antibody titer and specificity were
assessed by western blot using recombinant a-enolase. For
the preparation of polyclonal antibody against TIM, we
used recombinant TIM cloned from Saccharomyces cere-
visiae, a kind gift from P Guptasarma and S Ahmed
(IMTECH, Chandigarh, India).
SDS/PAGE and western blot
Corneas of Crocodylus palustris were removed surgically
and the epithelial cells were scraped and homogenized in
50 mm Tris, pH 7.5, 100 mm NaCl, 0.02% (v⁄v) sodium
azide, 1 mm EDTA. After sedimenting the insoluble debris
by spinning at 5000 g for 2 min total protein was estimated
using the Bio-Rad protein estimation kit. About 100 lg
protein was resolved using 12% (w⁄v) SDS-polyacrylamide
gel and transferred on a nitrocellulose membrane (Hybond-
C, Amersham Biosciences, Inc., Pittsburg, PA, USA).
Examination of the blot after Ponceau staining demonstra-
ted that equal amount of protein from the gel to membrane
was transferred. These membranes were probed either with
polyclonal mouse anti-a-enolase IgG (1 : 500) or with poly-
clonal anti-TIM IgG (1 : 300) followed by secondary anti-
body, anti-mouse-HRP conjugate (1 : 5000) and visualized
using an ECL kit (Amersham Biosciences, Inc.).
Immunofluorescence staining and confocal
Cornea was frozen-sectioned (10 lm) by a cryomicrotome
(Leica Microsystems, Bensheim, Germany) and the sections
were fixed in 100% methanol for one minute and washed
with 1· NaCl⁄PipH 7.2. The slides were air-dried and stored
till further use. The sections were rehydrated using 1·
NaCl⁄Piand the cells were permeabilized with 0.1% (v⁄v)
Triton X-100 in NaCl⁄Pifor 4 min. The cells were reacted
separately with anti-enolase IgG (mouse polyclonal anti-
body) at 1 : 100 dilutions or with anti-TIM IgG (mouse poly-
clonal antibody) at 1 : 50 dilutions. After three washes in
NaCl⁄Pi, anti-enolase or anti-TIM was visualized with
FITC-labeled sheep anti-mouse IgG (Jackson Laboratory,
Bar Haine, Maine, USA) at a 1 : 200 dilution. The nucleus
was stained with propidium iodide. Dual immunofluores-
cence confocal microscopy was carried out on a laser scan-
ning confocal microscope (model LSM 510 META) (Carl
Zeiss, Gottingen, Germany) using a 488 nm excitation and
540 nm long-pass emission filter.
a-Enolase and TIM activities in corneal
The a-enolase activity in the corneal homogenate of Crocody-
lus palustris was measured as described earlier [30]. Total
protein (40 lg) was taken with different concentrations of
glyceraldehydes-2-phosphate (0.1–1 m) prepared in imidazole
buffer pH 6.7. The substrate and enzyme were incubated at
37 ?C for 30 min and the conversion of glyceraldehydes-2-
phosphate to the phosphoenolpyruvate was measured by tak-
ing A240, which was then plotted as a function of substrate
concentration. TIM activity was measured using the method
described by Plaut and Knowles [31]. Glyceraldehyde phos-
phate was used as a substrate. Data were plotted as change in
absorption at 340 nm vs. substrate concentration.
We thank Dr Lalji Singh for his interest in the work
and initiating the program on crocodile development
Fig. 8. TIM activity in the soluble corneal extract collected from the
embryos of different developmental stages and from a two-week
posthatched crocodile. r 21st stage; h 22nd stage; m 23rd stage;
s 25th stage; d two-week posthatched crocodile.
TIM and s-crystallin in crocodile cornea
T. Kathiresan et al.
FEBS Journal 273 (2006) 3370–3380 ª 2006 The Authors Journal compilation ª 2006 FEBS
Page 10
and sex determination. We acknowledge Dr T. Rama-
krishna Murthy for his critical comments, G. Srinivas
for helping with frozen-sectioning, R. Nandini for con-
focal microscopy imaging and Dr Purnanad Guptasar-
ma and Shabbir Ahmed for providing yeast TIM. We
thank Hyderabad Zoological Park and Zoo Authority
of India for allowing us to collect crocodile eggs.
Raman Agrawal and Amit Anand are the recipients of
a senior research fellowship from the Council of Scien-
tific and Industrial Research (CSIR), Govt. of India.
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FEBS Journal 273 (2006) 3370–3380 ª 2006 The Authors Journal compilation ª 2006 FEBS


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Genre/Form: | Feature films Thrillers (Motion pictures) Legal films Video recordings for the hearing impaired Film adaptations Suspense films Drama |
Material Type: | Videorecording |
Document Type: | Visual material |
All Authors / Contributors: |
Brad Furman; Sidney Kimmel; John Romano; Michael Connelly; Matthew McConaughey; Marisa Tomei; Ryan Phillippe; Josh Lucas; John Leguizamo; Michael Peña; Bob Gunton; Frances Fisher; Bryan Cranston; William H Macy; Lions Gate Entertainment (Firm); Lakeshore Entertainment (Firm); Sidney Kimmel Entertainment (Firm); Stone Village Pictures (Firm); Lions Gate Home Entertainment.; et al |
OCLC Number: | 741292566 |
Language Note: | English dialogue; English or Spanish subtitles; closed-captioned. |
Notes: | Set of two videodiscs of the same film (119 min.): DVD and Blu-ray. Originally released as motion picture in 2011. Special features: Making the case: creating the Lincoln lawyer ; At home on the road ; One on one with Matthew McConaughey and Michael Connelly ; deleted scenes. |
Credits: | Director of photography, Lukas Ettlin ; editor, Jeff McEvoy ; music, Cliff Martinez. |
Cast: | Matthew McConaughey ; Marisa Tomei ; Ryan Phillippe ; Josh Lucas ; John Leguizamo ; Michael Peña ; Bob Gunton ; Frances Fisher ; Bryan Cranston ; William H. Macy. |
Target Audience: | Rating: R; for some violence, sexual content and language. |
Description: | 2 videodiscs : sd., col. ; 4 3/4 in. |
Details: | Blu-ray disc: region A; 1080p HD; aspect ratio: 16:9, widescreen; English 7.1 DTS-HD Master Audio; requires Blu-ray player.; DVD: region 1; 2.35:1 aspect ratio, widescreen; 5.1 Dolby Digital EX audio. |
Contents: | Disc 1. Blu-ray version of the film (119 min.) -- Special features -- Disc 2. DVD version of the film (119 min.) -- Special features. |
Other Titles: | Making the case. At home on the road. One on one. |
Responsibility: | Lionsgate and Lakeshore Entertainment present ; a Lakeshore Entertainment, Lionsgate production ; in association with Sidney Kimmel Entertainment, Stone Village Pictures ; producers, Sidney Kimmel ... [et al.] ; screenplay by John Romano ; based on the novel by Michael Connelly ; directed by Brad Furman. |
Michael 'Mick' Haller is a slick, charismatic Los Angeles criminal defense attorney who operates out of the back of his Lincoln Town Car. Having spent most of his career defending petty, gutter-variety criminals, Mick unexpectedly lands the case of a lifetime. However, what initially appears to be a straightforward case with a big money payoff swiftly develops into a deadly match between two masters of manipulation and a crisis of conscience for Haller.
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- Publisher: Watson-Guptill Publications Inc.,U.S.
- Format: Paperback | 144 pages
- Dimensions: 208mm x 276mm x 14mm | 581g
- Publication date: 13 October 1988
- Publication City/Country: New York
- Illustrations note: 275 b&w illustrations, index
- Sales rank: 137,743
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A guide to drawing cartoon characters - people of all ages, animals, birds and fish, with an emphasis on the personality, mood and character of the subject. The author deals with facial features and expressions, body language, drawing of hands and feet, head and body construction, conveying emotion and age, placing subjects in panels, making a layout, master reverse angles, medium shots, close ups, lights and darks, dialogue balloons and how to develop a comic strip. Christopher Hart is a former participant in the Disney Animation Program, has created numerous commercials and storyboards and was a contributor to "Mad" magazine.
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CHRISTOPHER HART is the world's bestselling author of drawing and cartooning books. His books have sold more than 6 million copies and have been translated into 20 languages. Renowned for up-to-the-minute content and easy-to-follow steps, all of Hart's books have become staples for a new generation of aspiring artists and professionals, and they have been selected by the American Library Association for special notice.


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Toppling Goliath Dorothys Old World LagerBrewed by Toppling Goliath Decorah, Iowa USA Dorothys older sister beer. Like a nurturing big sister, she will gently bring you into the big world of beers Toppling Goliath has to offer. Light amber in color with medium body and a smooth finish make this beer a true classic. Dorothy would be glad to have her name associated with such a great beer.
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SAC Capital accused of being "magnet for market cheaters"
Federal prosecutors allege SAC Capital, a hedge fund run by billionaire Steven A. Cohen, was a place where insider trading was "systemic and rampant." The hedge fund was indicted Thursday on charges of securities fraud. Anthony Mason reports.
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- C1: Appraisal and authorisation
- C2: Authorisation limits and process
- C3: Project proposal form and request for authority form
- C4: Revised request for authority
- C5: Post project appraisal and evaluation
- C6: Type of post project appraisal
- C7: Role of the committees
- C7.2: Infrastructure Coordination Group
- C7.3: Infrastructure Strategy Group
- C7.4: Council
- C7.5: Project Coordination Group
- C8: Procurement
- C9: University and College strategic goals
- C10: Self funded schemes
- C11: Key deliverables
- C12: Implementation costs and income
- C13: Operational costs and income
- C14: Inflation
- C15: Discount rates and expected rates of return
- C16: Reviewing alternatives
- C17: Appraisal narrative
- C18: Risk mitigation and avoidance
- C19: Project team competencies
C2: Authorisation Limits and Process
The process required to obtain authorisation for a major project under these procedures (Scope and Definition: Definition of a Project) is dependent upon the level of risk the University is taking. For simplicity, risk has been assumed to vary with gross expenditure. As such, if the estimated gross expenditure required by the project is known then the authorisation route and the process to be completed is noted below.
Where a project is considered to be high risk, novel, contentious or otherwise unusual the Infrastructure Coordination Group (ICG) may request a more onerous authorisation route. The tables below set out the key authority routes for projects:
All projects
The form of appraisal that projects will need to undertake is set out below.
Gross £ |
Investment Appraisal |
Post Project Appraisal |
£25-200k |
Yes |
Yes | Basic |
Basic |
£200k-£1m | Yes | Yes | Limited |
Limited |
£1m+ | Yes | Yes | Full |
Full |
See the CPP: Project Authorisation Workbook for full details of what must be included in each part of the appraisal. The workbook includes:
- Project Proposal Form (PPF)
- Request for Authority (RFA)
- EDS Strategic Risk
- Income and Expenditure
- Option Appraisal
- Revised Request for Authority (RRFA)
- Monitoring Statement
IT projects
All IT projects need to follow the governance approval process agreed for IT projects, with detailed consideration by IS Groups and approval/recommendation by the Project Coordination Group (PCG), prior to proceeding where required to ICG, ISG and C7.4: Council. Full details are available on the Academic Services website.
IT projects earmarked in the Infrastructure Fund, can be authorised by the Project Coordination Group (PCG) up to a value of £200k, but still need to be sent to ICG for coordination.
IT projects making a new call on the Infrastructure Fund can be recommended by PCG but can only be authorised by ICG/ISG/C7.4: Council.
A project is not authorised until it has been authorised by the appropriate authorising committee.
College/Service funded projects
Gross Expenditure |
Management Approval |
PCG (IT only) |
Dean/Head |
DVC/Registrar | ICG | ISG |
Council |
£25-100k |
Head of college/service |
Yes |
Yes |
For info |
£100-200k |
Yes |
Yes |
For info |
£200-500k | VCEG | Yes | |
Yes | |
£500k-£2m | VCEG | Yes | |
Yes | Yes | |
£2m+ | VCEG | Yes | |
Yes | Yes | Yes |
Infrastructure Fund projects
Gross Expenditure |
Management Approval |
PCG (IT only) |
Dean/Head |
DVC/Registrar | ICG | ISG |
Council |
£25-100k |
Head of college/service |
Yes |
Yes |
£100-200k |
Yes |
Yes |
£200-500k | VCEG | Yes | |
Yes | |
£500k-£2m | VCEG | Yes | |
Yes | Yes | |
£2m+ | VCEG | Yes | |
Yes | Yes | Yes |
All infrastructure funded projects need to be approved by ICG for co-ordinating purposes.
The rationale for the different levels of appraisal work required for different sized projects is one of cost / benefit.
In terms of the above levels of Investment Appraisal refer to Appendix D2 for definition and content.


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ABC Video on Demand
Watch the latest video programs:
Inside ABC TV
Program Summary
Pacific Stories
Land Of The Morning Star
1:00pm Saturday, May 20, 2006
Join some amazing people on this journey through the Pacific and share in an extraordinary meeting of cultures and contemporary history. This episode looks at an extraordinary land with a rich and turbulent history.
Education Subtitles CC G