text:
Making Star Wars better
Posted in Film by Ben
Digital warlock: George Lucas (clothes added in post)
What really bugged me about the original release of Return of the Jedi were the doors to Jabba's palace. I was only 5 when I saw this version and I remember someone on my row shouting 'BULLSHIT!" very loudly when they saw the reverse of the doors after R2D2 and C3PO walk in. He then started crying and vomiting loudly. The whole of Cannon Cinemas smelt like Parma Violets and stomach acid. From that moment on, the audience was lost; if they couldn't believe the exterior doors to Jabba's lavish palace were that size then they'd hardly believe in a Rancor beast, would they? I felt in some ways that I lost my innocence that day, my childhood cruelly molested by Uncle George and his silly doors.
Lucas attempted to repair my childhood when he made the doors slightly larger in the third iteration of the special edition. Sure, I'd believe you could fit a smaller skiff through these doors but the onset of adulthood had left me with a cold and cynical nature. I didn't really want to believe you could fit a smaller skiff through – let alone the full-girth of the Master's sail barge with those pig-men stood on top. Yeah, I was dead inside. But thank the maker for the fifth pass on Blu-Ray. George has made the doors massive and I now look back on the film with a warm fuzzy glow. You could fit anything through those bastards. ANYTHING. It also emphasises the impressiveness of Jabba before you even meet him for the second time, because nothing says don't fuck with the owner like 'my doors is bigger than yours.' Holla!
Original release: The search for doors
Third special edition: door-ish
Blu-Ray: BAM! Doors.
More subtle alterations are done later on in the new release of the film: When Luke is electrocuted by the Emperor, after revealing to Darth Vader that he is his son (OMG SPOILZ!), I always thought it daft of Darth Vader to just stand there. People probably have their own ideas about what was going on; I like to think that maybe Vader didn't see anything, because he was wearing a giant bucket on his head and because it's later revealed that he's about 170, so his eyes are probably fucked. Anyway, like Yoda says, "doubt leads to arguments, arguments lead to fights, fights lead to nuff probz", so it's only right for George Lucas to come back and stick his dick in the metaphorical mashed-potato, by adding a subtle audio enhancement that you probably won't even notice, unless you're really paying attention. Check-it:
In case you missed it…
Your ears aren't lying, he says 'NOOOOOOOOOO'. It's a neat addition and also ties the old films in with the original trilogy of Episodes I-III. There are so many people wearing stuff on their heads in the entire saga of seven films* that it can be confusing knowing who's who. When Darth Vader shouts "NOOOOOOOOOO" in Episode VI it's a neat tie-in to when that guy with the long hair in part 3 wakes up and can't get the black bucket off his head and is all like "NOOOOOOOOOOOO" before walking into things and knocking them over. Ha ha. This is just great synergy from George Lucas, or as I like to call it: 'cinemargy'.
* 1-6 + Episode 7: The Caravan of Courage, or 'Campervan of Crap' as it was sometimes known – despite being the second best entry to feature Ewoks. Maybe Uncle George should take a pass at that using his new maxim: "If it don't blink, it don't think."
Casualties of cinematic war
Posted in Film by Ben
Everyone hates Ryan.
The following contains spoilers for Saving Private Ryan and World War II.
It's funny how a film takes on a new dimension over the passage of time. Take Saving Private Ryan, for example. Ostensibly, 'Ryan' was a chance for Steven Spielberg to explore the nature of conflict and themes of guilt and regret on a broad canvas, while the viewer reflects on what a bunch of cock war is. But in the intervening years, the viewer learns not only to mourn the senseless loss of life during the most widespread war in history, but also begins to wonder just what happened to the young men who didn't return from Europe to be welcomed by a hit trilogy of spy thrillers.
Saving Private Ryan is like a yearbook of late 90s acting talent. There's Vin Diesel and the guy from Battlefield Earth – not to mention those blokes who were sometimes on Friends. And one by one, through the course of the film, they perish on their quest to rescue Matt Damon, who, it doesn't need to be said, has had a pretty stellar career since 1998. Sadly this wasn't the case for Spielberg's expendables, who've been kind of sidelined in the intervening years.
Perhaps if the actors had realised that real-life would mirror the events of Saving Private Ryan they would have improvised a scene where they escape France on snowboards, but not before Vin Diesel smashes all the light bulbs and leaves Private Ryan to be eaten by aliens in the muddy streets of Ramelle. After all, 'The Deez' is supposed to be a filmmaker of some note – at least according to his website – while Edward Burns originally caused a stir with his directing and Tom Hanks has won one or two awards for acting. But alas, as each cast member meets his fate in Private Ryan, his career goes tits skyward*.
Anyway, enough of my yapping, I'm now going to examine the wilderness years of the Private Ryan crew.
Tom Sizemore
If publicity were the career apex of an actor, then Tom Sizemore would straddle Hollywood like an inebriated colossus. In the 90s, if you were looking for someone to play a heavy with more than two-dimensions, you'd look for Sizemore. Following a string of solid supporting turns in Heat and Natural Born Killers – not to mention his admirably unconventional turn as a lead in The Relic, which is notable, considering the film concerned a statue that came to life and ate people, it looked like 'The Size' (as he was probably never known – least of all after his sex-tape) had carved out a successful niche playing goons with substance on-screen, and substance abuse issues off.
The afore-mentioned sex-tape was probably the low-point of Sgt. Horvath's career, or it could be 'Slumber Party Slaughter', I don't know; I haven't seen it. But I'm going to take a guess and suggest that if it doesn't feature a wasted Tom dressed in cycling shorts attempting to throw a football with more success than he finds summoning his mucky 'man-pride', it's probably a couple of rungs up on the ladder of career respectability. The good news is that Mr. Sizemore is now holding down a steady job on the remake of Hawaii Five-O, so perhaps a filmic resurgence isn't out of the question.
Vin Diesel
'The Deez' seemed to come out of nowhere with a string of good roles, including Private Ryan, The Iron Giant, Pitch Black and that funny video where he break-dances. Diesel seized the mantle of Generation Y's go-to action star with roles in extended nu-metal music videos like Fast and the Furious and XXX. Unfortunately, Diesel's Achilles' heel appears to be a reach that exceeds his grasp – in this case the desire to be treated as a serious thesp, and not just a deep voice with muscles.
While he could have happily ploughed a furrow of sequels and anaemic action roles, 'The Dieselator' decided to stretch his talents and re-brand himself as a proper actor. This is either admirable or completely misguided, I don't know – but he wouldn't be the first, as evidenced by the 'Oscar' Stallone's probably not very happy to talk about. Anyway, the dark years of The Pacifier appear to be behind him now and he's gradually accepting the fact that people only really want to see him denying Paul Walker's amorous advances while driving cars and scowling…or break-dancing.
Barry Pepper
Tom Sizemore may have lived through his real-life Abel Ferrera film with booze, drugs and hookers, but Barry Pepper's the cast-member I feel most sorry for. Not only did he appear in the series of painfully elongated war clichés that was We Were Soldiers Once, but after surviving that nightmare he was then door-stepped by John Travolta, offering a free copy of Dianetics by L Ron Hubbard and a starring role in a film that was going to be better than Star Wars, because it was based on true events. Or something.
There are no suggestions for the existence of Battlefield Earth. Clearly, Travolta must have some epic powers of mind-control to convince people that investing millions of dollars in a sci-fi film that featured him matching his giant face with the body of a 9-foot tall alien rasta-goth was a good idea. Maybe there's something in that Scientology stuff after all? Might be worth a try. Anyway, Pepper also survived this episode and has made something of a comeback, winning an award for his role in The Kennedys and appearing in True Grit. The only caveat appears to be a contractual requirement to have really bad teeth.
Giovanni Ribisi
I once thought Giovanni Ribisi a good actor. Then I re-watched the film and discovered completely new things about it the second time round. And then I realized it was a completely different film, just the same performance by Giovanni Ribisi. And then I started to worry about Giovanni Ribisi. Did he even know he was in a film? Maybe he's actually like that in real-life.
The best thing I can say for the wilderness years of Giovanni Ribisi is that I no longer felt I had to worry about Giovanni Ribisi's well-being. And then I watched Avatar, and saw visual spectacles I'd never seen before, like aliens shoving their hair up dragons' orifices. And then Giovanni Ribisi showed up being all Giovanni Ribisi-like and I started to think that maybe I had seen this before, which is either an indictment of the well-trodden path James Cameron's story walks, or Giovanni Ribisi and his Giovanni Ribisi-ness.
The others
There's not much to say about the others; I don't know much about Edward Burns, but I suspect that's because he appears to play Edward Burns in every Kate Hudson film that doesn't feature Matthew McConaughey or Owen Wilson. Anyway, his character doesn't die, so escaped 'the Curse of Ryan'.
Adam Goldberg appears to be the go-to guy when producers are looking for a hotheaded motor mouth, which veers from the mighty Zodiac to TV's Medium.
I was genuinely surprised to find out that the guy who played the bookish Upham recently appeared as redneck sociopath Dickie Bennet in the second series of Justified. Anyway, as good as he was in that role (and he really was), Upham survived the film, so is also excluded.
In conclusion
The main star of Spielberg's films is usually the man himself. His films are littered with actors who've been massive by proxy one minute and then vanished into the cinematic ether the next. So with that in mind, Matt Damon's success is probably down to talent, luck and shrewd decisions – not the fact that he survived Saving Private Ryan (I know, right?). But, regardless, I bet at least one of the above examples still think he's something of a spawny bastard.
* Tom Hanks is exempt from this rule – he's survived much worse (as Joe Versus the Volcano and Bonfire of the Vanities will attest).
Mortal Kombat
Posted in Games by Ben
The acceptable face of violent decapitation.
The existence and commercial acceptance of Portal 2, Heavy Rain and LA Noire suggests that we're entering a brave new world of electronic storytelling, where journeys amuse, surprise and delight in equal measure, and our decisions are informed by morality and characterised by more shades of grey than the lost episode of 24 that featured Kilroy airing his scrotum during an alien invasion in the centre of Manchester.
LA Noire's acceptance to the Tribeca Film Festival may have had the faint whiff of PR tokenism, but it also shows that the lines between both art forms are likely to converge over the next few years, with cinema embracing the young upstart to prove its future offers far more than plastic glasses and blue cat-aliens, and games returning the favour to gain more artistic legitimacy. And then there's Mortal Kombat, happy to sit at the back of the class and throw rubbers at the smart kids' heads while they're busy earning respect for our hobby.
Though Mortal Kombat is no stranger to experimentation; it's nearly 20 years old and has spent a large part of its teens trying new things that both failed and were ultimately fairly embarrassing. But hey, we've all been there. And if your teenage failures were witnessed by millions of adolescents – the demographic most likely to not let you forget – you'd probably be a little bit angry too. So Mortal Kombat 2011 aims to rewrite the rules, not by refining a system of combat that was never the most sophisticated in the first place, but by embracing its coarse charms, smashing your face in with 'kontent' and spelling things inkorrectly.
By pitting the kind of generic martial artists you'd normally find generously distributed on the bottom shelves of a video shop against the kind of creatures you'd find on Napoleon Dynamite's schoolbook, Mortal Kombat was obviously a product of its time. But the new iteration's greatest strength is resisting the urge to forego its fantasy elements in pursuit of realism. So you won't find Goro working as a bus driver, or Kano played by Danny Dyer. Instead it offers a compendium of violent brutes, robot ninjas, scantily clad ladies and ugly people with four-arms beating the crap out of each other in two-dimensions. And it does so wonderfully.
As unlikely as it may sound for a fighting game, the story mode is Mortal Kombat's most compelling element, condensing the narrative of the first 3 games into an enjoyably ridiculous soap opera that intersects every violent bout with wilfully cheesy dialogue and ridiculous plot twists. In fact the levels of treachery and backstabbing surrounding the tournament are so ridiculous that Mortal Kombat's story mode is arguably a more accurate FIFA game than FIFA ever could be, with Shao Khan and Shang Tsung filling the roles of Sepp Blatter and Jack Warner, respectively. And of course there's Goro, who's like a more socially advanced Wayne Rooney.
The Mortal Kombat story mode is generously long. In fact by the time you've finished, you'd be forgiven for not wanting to spend another second ripping people's heads off and vomiting down their necks. Not that I found the fatalaties all that compelling – my fingers aren't as nimble as they used to be and the graphical fidelity on display actually made me feel quite bad about performing them. But anyway, that's Mortal Kombat in a nutshell: a wealth of riches wrapped up in a deceptively simple outer-casing – a bit like an aggressive Karl Pilkington. And even if you don't move beyond the basic arcade mode, you'll still find a suitably authentic Mortal Kombat experience that's been lovingly updated to please those who remember 'ABACABB' and the yoof of today, who probably recognise the aforementioned collection of letters as the correct way to recite the alphabet.
Alongside arcade and story, there's a Challenge Tower, which consists of 300 tasks, including needlework and pet grooming. Just kidding, of course it involves ripping people's arms off before the timer runs out and shooting zombies with missiles, that kind of stuff. Each challenge unlocks 'Koins', (natch), which can be redeemed in the Krypt (also natch) for fatalaties, violent pictures and new togs for your Kombatants (um, yeah).
Despite the detail involved you still get the impression that game play was at the forefront of this reinvention – and they've actually gone for a 'less is more' approach. So there are no swords, side-steps or superheroes. Though if you're really desperate for a multimedia crossover then one of the world's most famous deceased child molesters is available for download (don't get your hopes up, Ready 2 Rumble fans). Combat is meaty and immensely satisfying and the moves of yore are updated and replicated perfectly – alongside some new additions like Super Meters, which seem to be de-rigueur for the modern fighter.
There's not a lot to complain about with the new iteration of Mortal Kombat. Sure, it's cheesy, and stupid, but after years spent in denial it has finally realised that these are the things that made it great in the first place – and it's now embracing them with all four arms. It's never going to be a brain surgeon, or a nobel-prize winner, but it would make a fine plumber. And there's no shame in that.
Chest bumps and severed love pumps
Posted in Film by Ben
(I originally wrote this review when I saw Piranha 3D at the cinema. After waking from my coma I totally forgot about it, much like the film itself. Anyway, in the absence of a new article this week here are my thoughts on Alex Aja's Piranha 3D from the heady days of 2010. In short: it wasn't very good, but don't worry, at least Takers is out next week!)
Piranha 3D: a film about killer fish and tits that's more gory than a heroic bloodshed retelling of The Passion of The Christ, and contains more severed man members than you might expect find in Michael Barrymore's pool filter – if you believe everything you read in the News of the World, natch.
If you were such a purveyor of quality journalism then you may be wondering what more there could be, beyond gore, tits and fish – not just in the film, perhaps even in real life. And while you could be accused of having criminally low standards, you wouldn't exactly be wrong.
But is it wrong to expect internal logic and character development from something that prides itself on showing little more than fish and tits in 3D? The part of my brain that tries to get me to acknowledge the existence of Chris Moyles would say 'probably' (if it understood the question), but I disagree.
The original Piranha was from the Roger Corman school of filmmaking. While the quality of some of Rog's directorial output could be questioned, at least from a traditional critical viewpoint (though I remain convinced time will reveal the true genius of She Gods of Shark Reef) his ability to spot talent remains without equal. Corman's movies were where the aspiring A-list could cut their teeth working on B-level productions. So while you could expect little of these productions beyond a snappy title and often hilariously misleading poster, you'd often be pleasantly surprised with some of the artistry on display. And these directors would be justly rewarded with careers making better films with bigger budgets.
Unfortunately those times seem to be over; the best today's director can expect is to get hired by a studio to remake the B movies of yesteryear or turn a boardgame/cartoon/cracker joke into a filmic franchise. So while people bemoan the absence of another Coppola or Scorsese they fail to realise that he might be out there right now, making Yogi Bear in 3D, hoping to earn enough juice to remake Slugs for Michael Bay's Platinum Dunes, before inevitably descending into a creative abyss fuelled by self-loathing and friendship with Brett Ratner.
Alexandre Aja's career seems to revolve around making well-directed horror films with a nod to whatever's popular at the time. So Switchblade Romance, a taut efficient slasher for the most part, ends with a ridiculous trope from the 90s cinema that, thankfully, seems to have been banished to the M Night Shymalan landfill of bad ideas. The Hills Have Eyes was one of the better horror remakes, but a remake all the same. I didn't see Mirrors but as it was made in 2008 I'm going to assume it featured Kiefer Sutherland going mental after his iPhone 3G wouldn't stop playing Leona Lewis. Or something.
Anyway, what I'm trying to say that perhaps the only way to survive when you have a degree of intelligence but keep getting fed scripts like Yahtzee 3D, or whatever's supposedly hot at the moment, is to embrace the ridiculousness of it. In which case a lack of the key components that normally gauge a film's quality: character development, plot and narrative logic, become the film's strengths. That would explain the appearance of Oscar winners Elisabeth Shue, and Richard Dreyfuss and, to a lesser-extent, Oscar watcher, Jerry O'Connell. I suppose it's like the Emperor's New Clothes, but the emperor knows he doesn't have any clothes, and everyone else knows it too, but it's OK because they don't have any clothes either, and nobody cares because there are tits and cock-eating fish everywhere…in 3D!!!
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subset: commoncrawl


text:
Stop the Madness: Don't Overschedule Your Child
These days, many children have schedules that could challenge most CEOs. Their days start early, end late, and are full of organized activities. Sports practices, music lessons, playdates, and homework for school-age children can result in stressed-out, unhappy children—and parents.
While parents want to prepare their children for the future, and believe they're doing what's in their child's best interests, involving your child in too many extracurricular activities is not always a good idea and can actually be detrimental to their health.
Many children who are over scheduled will begin to show signs of being stressed out.
- They may say they don't want to go to their scheduled activities or cling to you when you try to leave.
- They may complaint, or be tired all the time.
- They may be angry, whine, or cry when it's time to go to a scheduled activity.
- They may have frequent stomachaches.
- They may not eat or sleep well or have nightmares.
- They may show low self-confidence.
Here are a few things to keep in mind if you want to avoid overscheduling in your household:
Balance is Key: While child experts acknowledge that organized activities have many positive effects, as with most things, finding a balance is key. Young children are not often able to function well with so many responsibilities and may start to feel anxious about all the added pressure. Research shows that children need relationships more than they need activities. Pediatricians continue to stress the importance of family bonding.
Free Time: Children also need to have times without planned activities. Downtime allows for relaxation, free play, and family time. Adding this downtime to their schedule can help a child see that these unstructured times are just as important as their organized activities.
Boredom is Sometimes OK: Parents sometimes worry their children will be bored if every minute of their lives aren't planned out. However, boredom can actually be beneficial to children! If your child complains of boredom, your first instinct may be to organize something for him to do. But really, you should be letting them figure it out for themselves. The ability to think critically, gather information, make decisions, and put all that into action is an important skill and we need to give children practice in that skill. Children who are able to do these things are generally more successful in school, and later in life.
So many parents lament that childhood goes by too quickly. Limiting the extracurricular activities is one way of slowing it down. You want your child to think fondly of their childhood, remembering all the time you spent together playing board games and talking walks outside. Children only have one chance to be a kid, they should enjoy every minute of it!


text:
begin Rich Bodo quotation of Fri, Aug 29, 2008 at 01:51:29PM -0700:
> The fact that sword manufacturers are paying people not to manufacture
> pens is interesting, though. It's an unnecessary validation for me,
> perhaps not for others.
It's not that they're being paid not to _manufacture_
pens -- just not to sell or promote them. As the
usefulness of the available pens goes up, the payment
not to work with them has to go up as well.
A "cynic" funding free software as a bargaining chip
might actually be more motivated to fund "plumbing"
work than an FSB that had to make a sustainable living
out of somehow charging for one piece of the product.
Don Marti +1 415-734-7913 mobile
email@example.com Linux device driver unconference: http://freedomhec.org/


text:
The Pre-master's programme is designed to meet the needs of international students. You will be taught English language, academic subjects and study skills to help you prepare for studying a postgraduate degree at Keele University.
The Pre-master's is a two-term programme. You will study a combination of core and subject-specific modules that focus on academic areas most useful to your chosen degree.
To ensure you make good progress and achieve the results you need, we'll give you regular assessments. These include coursework, unseen written exams, presentations, extended essays and case studies.
When you receive confirmation of your place to study at the International Study Centre, you will receive a conditional offer of a place on one of the specified master's degrees at the university. Once you successfully complete the programme, and achieve the required grades, you will join your chosen master's degree.
Please note that not all intakes may be available for all subject routes.


text:
|Reviews for Ruatha's Grammar Review
| storiesbyarat chapter 5 . 9/29
Wow! I never knew that semi colons could be used to seperate phrases in a list, that clears up so much. Thanks a bunch!
| Marjulie chapter 6 . 5/1/2016
I found this chapter to be most helpful since I don't remember running across it in any of my English classes ever. All the examples were great and understandable except for the last example of number five, "Sylvia, Ben's, his and my movie was a real bore." I think it's avoidable if it's rephrased, but if a writer insists on a sentence similar as that, can you expand on it? I feel like I'm half way understanding it.
A suggestion from someone who's still a little confused with writing and English, a section explaining 'Was vs. Were', when to end a paragraph, and run on sentences would be amazing. It's something that I've always struggled with.
| Bob Story Builder chapter 3 . 3/16/2016
Really enjoyed this so far! I love dialogue and still figuring out my style in terms of writing dialogue. Reviewing rules for dialogue is helpful because sometimes I forget. I like the idea of writing down "said" and "ask" words and think I am going to do that. You got an entertaining writing style. I read this in the voice of a sarcastic young grandmother funny enough.
| AikaRikaru chapter 5 . 1/10/2016
I am so happy I stumbled upon your page.
This was a much-needed refresher on semicolons, so thank you!
| FictionFox'95 chapter 3 . 12/15/2014
Finally! Found something that gives sense to writing. It's like a grammar for dummies. Thanks!
| Aleatoric chapter 5 . 10/17/2014
Ch 5:
"English" is a proper noun and should be capitalized. (ie. [E]nglish teacher).
"In fact, she admitted to me that even SHE, the teacher, wasn't exactly comfortable with semicolons and thus tends to avoid them." I think "tends" might need to become "tended," due to parallelism.
| Aleatoric chapter 4 . 10/17/2014
Ch 4:
"You're just lucky the Bad Pun Police don't read your site!" I think it should be "doesn't" instead of "don't." Unless that was a stylistic choice.
The "you're" vs. "your" is at the top of my pet peeve list, haha.
| Aleatoric chapter 3 . 10/17/2014
Ch 3: "I see that tend to err on the side of caution..." Missing an "I" before "tend." (This is in the last few paragraphs).
| VeronaViridian chapter 3 . 7/3/2013
This chapter helped me a lot!
Thank you for posting this . . .
I'm sure you have mentioned this in your writing, but I feel the need to state it again . . . It's so hard to find a good grammar book that doesn't bore people. Keep on writing! :)
| yo buddy chapter 6 . 10/25/2012
so lame!
| MrGagaSlashLover chapter 1 . 9/17/2012
I praise you. I do like half the stuff in here so thank you!
| Abbytjie chapter 4 . 8/13/2012
When I was still struggling with to and too, I found a palindrome that really helped out:
"Too hot to hoot."
If it can be read the same backwards and forwards, you have the right to/too in the right place.
| Ayana Parker chapter 6 . 5/9/2012
This is an awesome Grammar Guide but Sheeps isn't a word just thought I'd tell you. But anyway so far so good.
| darkwrider chapter 2 . 5/3/2012
thank this help a lot.
| Wendy Thompson135th chapter 2 . 3/25/2012
I added this url to my bio page; I rant often and loudly on some of the topics your raise.
Good work. ~~Lynn K Hollander.


text:
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text:
Roger '56 and Janice Johnson
Spring 2003
"I just didn't know where to begin."
That statement was quite a surprise, coming from Roger Johnson '56. Roger W. Johnson is one of a very small group whose career includes leadership positions in four institutional segments, i.e., CEO of a Fortune 500 company, head of a major Federal Agency as well as significant leadership roles in non-profit art organizations and two major universities. But as a high school student in 1952, the first in his family to go to college, the process of finding a school was daunting. "We didn't know where to begin. It was through the help and guidance of Coach Hodge and others at Clarkson that I believed it could be possible. My Clarkson years were some of the best of my life, both on the baseball field and in the classroom," said Roger. The nurturing social and academic atmosphere, along with the personal attention from faculty and coaches like Hank Hodge were critical to Roger's development and propelled him to graduate first in his class. It also allowed him to meet his love and lifelong partner, Janice (Potsdam '56).
"We knew exactly what we wanted to accomplish."
This sounded much more like Roger, as he spoke of his and Janice's plans for charity: "We were determined to support both Potsdam State and Clarkson. We created a family trust to help manage our assets, and we included bequests to both Potsdam and Clarkson. Our gift to Potsdam will create an endowment to bring world-class artists to Potsdam to provide master classes and performances for the entire Potsdam community. Our bequest to Clarkson will create an endowment to provide scholarship support to students in the School of Business. Scholarships made it possible for me to attend Clarkson, and now I can help future generations benefit from the same Clarkson experience that I had." Roger contacted Clarkson to create guidelines for how the scholarship will be managed when the gift ultimately goes to the University. "The University staff was very helpful in documenting my desires. I feel confident that my gift will someday help those most in need of financial assistance. An added bonus is that our gift will benefit the local economy by keeping assets in the North Country."
"Find the courage to take the first step."
"Everything worthwhile takes courage," said Roger. "It took courage for all of us to go away to college, to manage careers and families, to make a difference in the world. Now it takes courage for us all to begin or update our estate plans and determine what we can do to make a difference for the future of the world through a place like Clarkson. It's a wonderful feeling of accomplishment."
Footnote: Roger Johnson passed away in 2005. This article is published in thanks and admiration to a loyal Clarkson alumnus.
Roger '56 & Janice Johnson
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Taste buds are gustatory endorgans which use an uncommon purinergic signalling system to transmit information to afferent gustatory nerve fibres. In mammals, ATP is a crucial neurotransmitter released by the taste cells to activate the afferent nerve fibres. Taste buds in mammals display a characteristic, highly specific ecto-ATPase (NTPDase2) activity, suggesting a role in inactivation of the neurotransmitter. The purpose of this study was to test whether the presence of markers of purinergic signalling characterize taste buds in anamniote vertebrates and to test whether similar purinergic systems are employed by other exteroceptive chemosensory systems. The species examined include several teleosts, elasmobranchs, lampreys and hagfish, the last of which lacks vertebrate-type taste buds. For comparison, Schreiner organs of hagfish and solitary chemosensory cells (SCCs) of teleosts, both of which are epidermal chemosensory end organs, were also examined because they might be evolutionarily related to taste buds. Ecto-ATPase activity was evident in elongate cells in all fish taste buds, including teleosts, elasmobranchs and lampreys. Neither SCCs nor Schreiner organs show specific ecto-ATPase activity, suggesting that purinergic signalling is not crucial in those systems as it is for taste buds. These findings suggest that the taste system did not originate from SCCs but arose independently in early vertebrates.
2. Introduction
Taste buds are the gustatory end organs in vertebrates ranging from lamprey to mammals [1]. These end organs respond to a variety of sapid chemicals, and transmit signals to afferent nerves fibres arising from three cranial ganglia: facial, glossopharyngeal and vagus. While other nerves, e.g. trigeminal, may heavily invest the epithelium surrounding the taste buds (the so-called perigemmal innervation), the trigeminal fibres do not enter the taste bud itself. Taste buds can be recognized by three key features. (i) Taste buds are an aggregate of elongate taste cells of multiple morphological and functional types. (ii) Taste cells extend from the basal lamina to an apical pore or other opening in the epithelium. (iii) Finally, taste buds are innervated by sensory fibres of the cranial nerves containing cells derived from epibranchial placodes: i.e. the facial, glossopharyngeal or vagus nerves [2].
Throughout the vertebrate lineage, numerous epithelial chemoreceptors can be identified, but only taste buds meet the three criteria listed above. For example, most fishes possess solitary chemosensory cells (SCCs) scattered across the body surface [3]. SCCs share some features with taste buds, i.e. they are elongate, span the height of the epithelium, form synapses with afferent nerves and even may express common receptors [4,5]; but SCCs are not merely dispersed taste cells. The SCCs mostly appear singly, not in clusters, and, unlike taste buds, can be innervated by non-gustatory (e.g. spinal or trigeminal) nerves [3,6,7]. Previously, some have speculated that SCCs may even be phylogenetic forerunners of taste buds [8]. Hagfish possess another distinct chemosensory endorgan, the Schreiner organ, which shares many features with taste buds, but again is not identical to taste buds [4]. Schreiner organs are an assembly of several cell types and form functional contacts with nerve fibres entering from their basal aspect. But Schreiner organs do not extend to the basal lamina and can be innervated by non-gustatory nerves, e.g. the trigeminal nerve [4]. Thus, Schreiner organs are not considered to be taste buds, but may be functionally or phylogenetically related to taste buds.
An unusual feature of taste buds, at least in mammals, is their dependence on purinergic neurotransmission to convey the signal from taste buds to the nervous system [9]. Taste cells release ATP upon stimulation by tastants [9–12] and the gustatory nerve fibres express two ionotropic purinergic receptors, P2X2 and P2X3, which are required for activation [9]. Many taste cells express a highly specific ecto-ATPase (NTPDase2) [13], necessary for inactivation of the purinergic neurotransmitter. This ectonucleotidase isoform is highly selective for ATP over ADP [14] and so can be identified by classical histochemical methods employing these two different substrates. As nucleotidase activity has been reported in teleost taste buds [15], it is likely that taste buds in teleosts also employ purinergic signalling. The present study was undertaken to test whether taste buds in all vertebrates use purinergic neurotransmission, and whether this unusual mechanism is also associated with other epithelial chemoreceptor systems.
Ultrastructural and immunochemical studies show that taste buds consist of a heterogeneous population of cells. Mammalian taste buds cells are classified into three principal types of mature, elongate cells, i.e. type I, II and III, along with proliferative basal cells lying just outside the taste bud proper [8,16–18]. The elongate cells extend an apical process into an opening in the epithelium, and a basal process, which may reach the basal lamina of the epithelium. Although the cell types were originally described based purely on ultrastructural criteria [19], each of the cell types appears to have a unique function. In mammals, the type II cells express metabotropic taste receptor proteins and elements of the G-protein transduction cascade required for sweet, bitter and umami tastes [20–22]. Similarly, type III cells express the channels used in sour and possibly salt transduction [23,24]. The type III and type II cells apparently transmit this information to the gustatory nerves through the agency of ATP acting on P2X-type ATP receptors situated on the nerve fibres [9]. Type I cells are considered to be supporting cells like glia because type I cells often envelop other elongated cells and express proteins such as GLAST, which are expressed by glia in the central nervous system. More importantly, the type I cells also express the ecto-ATPase (NTPDase2) highly specific for ATP [13,14]. Presumably, this ecto-ATPase is crucial in inactivating the ATP signal release by the taste cells. Thus, the presence of this specific ATPase can serve as a marker for the utilization of ATP as a neurotransmitter in the taste bud system.
In fish, the cellular organization of taste buds is less well understood, but consists of at least three types of cells [15,25,26]. According to their morphological features, they are termed tubular or light cells, filamentous or dark cells and basal cells. Multiple classes of light cells have been reported [27]. Both light and dark cells are elongate cells and are generally believed to be receptor cells and supporting cells, respectively. One prominent type of basal cell in fish taste buds are Merkel-like in terms of morphology and neurotransmitter content [8,15]. In addition, proliferative marginal cells lie along the basolateral margin of the taste bud similar to basal cells of mammalian systems [27]. Light cells, dark cells and Merkel-like basal cells all have synaptic connection to afferent fibres [15]. Synaptic neurotransmitters in taste buds of fish are unknown although the Merkel-like basal cells accumulate and presumably release serotonin, similar to type III cells of mammalian taste buds [28] and Merkel-like basal cells in amphibia [29].
The purpose of this study was to determine when during the evolutionary history of taste buds and epithelial chemoreceptor cells a highly specific ecto-ATPase appeared. Is the presence of ecto-ATPase coincident with the appearance of vertebrate taste buds, or is ecto-ATPase present in or around other epithelial chemosensory endorgans suggestive of a more ancient origin for ATP neurotransmission by chemosensory systems? The present results show that ecto-ATPase is present in taste buds of fish including lamprey but not at all in Schreiner organs in hagfish, nor is it associated with SCCs of any vertebrate. These findings suggest that utilization of ATP as a transmitter co-evolved with the taste system in early vertebrates.
3. Material and methods
3.1. Animals
In this study, we examined six species of Teleostei (channel catfish, Ictalurus punctatus; sea catfish, Plotosus japonicus (n = 10); Japanese sea robin, Chelidonichthys spinosus (n = 6); common carp, Cyprinus carpio (n = 3); goldfish, Carassius auratus (n = 4); and zebrafish line P2X3.2 : gfp [30] (n = 2)), as well as one elasmobranch species (cat shark, Scyliorhinus torazame (n = 2)), two species of lamprey, Lethenteron japonicum (n = 4) and Petromyzon marinus (n = 10), and one species of hagfish, Eptaretus burgeri (n = 10). Apart from the transgenic zebrafish, all species were obtained commercially or caught with fisheries nets except for P. marinus: tissue was kindly supplied by Sorensen (University of Minnesota) and amnocetes by Nicholas Johnson, Hammond Bay Biological Station (Millersburg, MI, USA). The P2X3.2 : gfp line of zebrafish was generously supplied by Mark Voigt (St Louis University) and maintained at the University of Colorado Anschutz campus aquatics facility. The experiments for zebrafish, channel catfish, goldfish and lamprey were performed at the University of Colorado, School of Medicine and for others at Kagoshima University. All experiments were carried out with the approval of the local animal care and use committees (University of Colorado IACUC or guidelines of Kagoshima University).
3.2. Enzyme histochemistry for light microscopy
Ecto-ATPase activity was examined by lead precipitation as described previously [31,32]. Specific ATPase activity was determined by comparison of tissues reacted with 1 mM ATP substrate compared with those reacted with 1 mM ADP substrate. Purinergic signalling and ecto-ATPase enzymes are phyletically ancient, present even in singe-celled organisms [33]. Among vertebrates, the different ectonucleotidase isoforms are characterized in part by their substrate specificity. The ecto-ATPase associated with taste buds in mammals, NTPDase2, is highly specific for ATP over ADP or AMP and this property distinguishes this isoform from other, related ectonucleotidases [14]. Hence, histochemical results comparing hydrolysis of ATP with ADP is strongly indicative of NTPDase isoform. In this paper, we refer to enzyme activity particular to the ATP substrate but absent with the ADP substrate as ecto-ATPase.
Each species of fish was deeply anaesthetized with dilute tricane methansulfonate (MS222), and either fixed by transcardial perfusion or with immediate removal of chemosensory tissues from the specimen. Tissues were immersed for up to 1 h in 2 per cent paraformaldehyde, 0.2 per cent glutaraldehyde, in 0.1 M Tris–maleate buffer (pH 7.4) with 2 mM CaCl2 and then placed overnight in the same buffer at 4°C with 20 per cent sucrose for cryoprotection. The tissues were cut on a freezing microtome or cryostat at 12–40 μm. Free-floating sections or slides were rinsed three times for 10 min each with 0.07 M Tris–maleate buffer (pH 7.4).
In the beginning of the experiments, histochemical tests of ATPase were performed in the barbels of Plotosus and Carrassius. The free-floating sections of barbels, lips or palatal organ were incubated first with the following medium for 30 min at room temperature: 2 mM Pb(NO3)2, 5 mM KCl, 2 mM CaCl2 and 1 mM of substrate, either ATP or ADP. The incubation was followed by three 10-min washes in 0.07 M Tris–maleate buffer. The lead precipitate was visualized by treating the sections for 1 min with 1 per cent ammonium sulfide. After several rinses in distilled water, the sections were collected on slides, then coverslipped with fluoromount (Fisher Biotec). Some sections were counterstained with Giemsa dye. The sections were viewed under a Nikon or Olympus light microscope. After observation of significant activities in the presence of ATP, the incubation medium was modified; three kinds of inhibitors were added to the above mentioned medium: 1 mM levamisole (inhibits alkaline phosphatases), 1 mM ouabain (inhibits Na+,K+-ATPase) or 50 μM α,β-methylene ADP (inhibits 5′-nucleotidase). Incubation medium without CaCl2 was also tested.
Optimal conditions for the specific detection of ecto-ATPase involved fixation times of 30–60 min followed by prolonged washes in buffer. These optimal reaction conditions were used in processing tissues from the diverse species. Specific ecto-ATPase activity described below is defined as reaction product observed with ATP in the presence of inhibitors of other phosphatases and nucleotidase, but not the ADP substrate in the same reaction medium (figure 1a–d).
3.3. Enzyme histochemistry for electron microscopy
On one specimen each of goldfish and catfish (I. punctatus), we carried out the ATPase reaction for electron microscopy slightly modified from the protocol of Barry [31]. As the lead precipitate is electron dense, little modification is necessary. Fixation was as for light microscopy, but after fixation the tissue was cryoprotected in an ascending series of glycerin/sucrose in Tris–maleate buffer ending with 15 per cent glycerin, 20 per cent sucrose [34]. Free-floating 40 µm sections were cut on a cryostat into 70 mM Tris buffer and were then reacted as for light microscopy using either ATP or ADP as substrates. Following the reaction, sections were not placed into ammonium sulfide but rather were postfixed in 4 per cent EM grade glutaraldehyde in cacodylate buffer. After overnight postfixation, the tissue was rinsed in cacodylate buffer and placed into 1 per cent osmium tetroxide for 30 min. After rinsing in cacodylate buffer, the specimens were dehydrated in a graded series of ethanol and propylene oxide and embedded in Epon-Araldite (Electron Microscopy Sciences, Hatfield, PA, USA). Ultrathin sections (silver to gold) were stained with uranyl acetate and lead citrate and examined with a FEI Tecnai G2 electron microscope (Philips, Eindhoven, The Netherlands).
3.4. Immunohistochemistry
In order to delineate the different types of cells and innervation of taste buds, we carried out a series of immunohistochemical experiments in tissues from some teleosts and the lamprey. In all cases, omission of the primary antiserum yielded no specific reactivity reported below.
3.4.1. Single-label staining after ATPase reaction
Following distilled water rinses, the sections were washed for 10 min in 0.1 M PB (phosphate buffer, pH 7.2–7.4) and then 2 × 10 min changes of 0.1 M PBS (phosphate-buffered saline, pH 7.2–7.4). The tissue was then incubated for 1 h in blocking solution (3% normal donkey serum, 1% bovine serum albumin, 0.3% Triton in PBS) before an overnight incubation in rabbit anti-serotonin (1 : 5000, lot: 924005, Immunostar, Hudson, WI, USA) or (for zebrafish) chicken anti-GFP (1 : 2000, lot: 0609FP10, Avēs labs, Tigard, OR, USA) diluted in blocking solution at 40°C. After 3 × 10 min washes in 0.1 M PBS, the sections were incubated in secondary antibodies for 2 h at room temperature: DyLight 550 anti-rabbit (1 : 500, lot: GR32373–2, Abcam) or Alexa 488 anti-chicken (1 : 500, lot: 102758, Life Technologies, Gand Island, NY, USA). A far red draq5 counterstain was applied during the secondary antibody application (1 : 1000, lot: 402DR50050, Abcam). Slides were then coverslipped in Fluoromount-G.
3.4.2. Double-label staining
Alternate sets from the ATPase tissue were used for immunohistochemistry. Slides were washed 3 × 10 min in 0.1 M PBS and incubated for 1 h in blocking solution. Overnight incubation of the primary antibodies, mouse anti-acetylated tubulin (1 : 5000, lot: 118K4821, Sigma, St Louis, MO, USA), rabbit anti-serotonin (1 : 5000, lot: 924005, Immunostar) or chicken anti-GFP (1 : 2000, lot: 0609FP10, Avēs labs) was carried out at 4°C. Three 10 min washes in 0.1 M PBS preceded secondary incubation for 2 h at room temperature: DyLight 550 anti-rabbit (1 : 500, lot: GR32373–2, Abcam) or Alexa 488 anti-chicken (1 : 500, lot: 102758, Life Technologies) or Alexa 488 anti-mouse (1 : 500, lot 811493, Life Technologies) was diluted in the blocking solution. A far red draq5 counterstain was applied during the secondary antibody application (1 : 1000, lot: 402DR50050, Abcam). Slides were then coverslipped in Fluoromount-G after three more rinses in 0.1 M PBS.
4. Results
4.1. Ecto-ATPase in teleosts
Taste buds in teleosts reacted strongly for ecto-ATPase activity, but the labelling was slightly different for taste buds innervated by the facial nerve (lips and barbels) compared with taste buds innervated by the vagus nerve (palatal organ; figure 1). Facially innervated taste buds tended to show stronger labelling of elongate cells within the bud, but similar reactivity of elongate cells could be seen in intraoral taste buds of goldfish when using slightly shorter fixation conditions (30 instead of 60 min).
Taste buds in teleosts are organized into two distinct compartments: apically situated elongate taste cells, occupying the upper two-thirds of the taste bud, and a basal nerve plexus surrounding serotonergic Merkel-like basal cells (figure 1e–g; [35]). Heavy ecto-ATPase activity is evident within the basal nerve plexus above and below the Merkel-like basal cell (figure 1e–g). Distinct ecto-ATPase staining was evident in both compartments of the taste epithelium: elongate taste cells, within the basal neural plexus, and along the shaft of the nerve bundle below the base of the taste buds. In P2X3.2 : gfp zebrafish, gfp label is driven by the promotor for the puringergic receptor P2X3.2. In these fish, gfp-labelled nerve fibres entered the base of the taste bud to form a plexus just above the level of the serotonergic basal cell (figure 1g), suggesting purinergic neurotransmission in this region.
In more heavily reacted or less fixed specimens of different species, numerous elongate taste cells display reaction product along their membranes. This is particularly evident in sections from the barbels of the two species of catfish (figure 2). Beneath the taste buds, heavy reaction product also is apparent along the incoming nerve bundles (figures 1 and 2). When ADP is substituted for ATP as the substrate, some reaction product remains, but this is confined to the area below the Merkel-like basal cell and includes the incoming nerve bundles. The presence of reaction product with ADP substrate indicates that at least some of this reactivity is attributable to a less specific ectonucleotidase.
Ecto-ATPase staining of elongate taste cells was most apparent in tissue from lips or barbels, especially when fixation time was limited to 30 min. The apparent membrane association of reaction product seen at the light microscopic level was confirmed by electron microscopy (figure 3). Because the reaction product forms on the external face of the membrane, it is impossible to determine unequivocally at the electron microscopic level whether the enzyme activity is present on one or both of the facing cell membranes. Based on light microscopy, some elongate cells appear more reactive than others because the taste bud is not completely filled with reaction product (figure 2a). Similarly, in the electron microscope, some light cells are surrounded by reaction product (figure 3a,b), whereas dark cells and other light cells are not. The reaction product is more evident in the basal half of the taste bud, fading out as one proceeds towards the apical pore (figure 3a). In both goldfish and catfish, some elongate cells within each bud are surrounded by reaction product, probably indicative that these are the cells producing the ectoenzyme (figure 3a–d). In addition, nerve processes within the taste bud (figure 3c) as well as in the basal plexus (BP in figure 3e) are surrounded by reaction product.
In Plotosus, staining of the nerve bundles was especially prominent. In many taste buds, two fascicles of nerve fibres approached the bottom of the taste bud (figure 1d). Darker strands are visible within these fascicles, as if some elements are more reactive than others. It is not, however, apparent whether it is the nerve fibres or the ensheathing cells that display this ecto-ATPase activity.
In the searobin, oral and buccal taste buds were similar in size to those in the barbel of catfishes and lip of goldfish, but more widely spaced. Numerous elongated cells show ecto-ATPase activity (figure 4a) but nothing of ADPase activity (figure 4b). Similar to other teleosts, substantial nucleotidase activity was present in the basal nerve plexus but some of this product was likely due to non-specific nucleotidase activity. ADPase activity could be detected in nerve bundles and strands below the taste buds, but was fainter than those in the other teleosts.
4.2. ATPase in taste buds of sharks
In the cat shark, taste buds are present on the lip as well as in oral, buccal, branchial and pharyngeal epithelia. Ecto-ATPase activity was present in the basal region of the buds with fainter labelling of the overlying elongate cells (figure 4e). Reaction product in the elongate cells was weaker than that in the Teleostei studied. ADPase activity occurred only in the nerve components (figure 4f).
4.3. ATPase in taste buds of lamprey
Taste buds in lampreys lie along the branchial arches [36,37]. The organization of taste buds in lampreys is somewhat different than in teleosts and elasmobranchs in that lamprey taste buds lack serotonergic Merkel-like basal cells but do have elongate serotonergic cells similar to those in taste buds of mammals (figure 5c,e).
Reactions of whole mounts of the lamprey oropharynx showed ecto-ATPase activity in taste buds which appear as circular patches 50–100 µm in diameter along the branchial arches (figure 5a). Reactions of the whole mounts using ADP as a substrate showed little reactivity, even when examined in sections (figure 5b,f).
Sections through the taste buds showed reactivity within the basal nerve plexus (figure 5d,e,g,h), which in lampreys lies below the basal processes of the serotonergic cells (figure 5g,h). The heavy ecto-ATPase activity then lies in the region where the elongate serotonergic cells contact, and presumably synapse with, the afferent nerve fibres.
4.4. Hagfish Schreiner organs lack ecto-ATPase activity
Schreiner organs are widely dispersed throughout the epidermis of hagfish [4], with especially high density on the nasal and oral barbels. Neither Schreiner organs nor nerve fibre bundles nearby showed substantial ATPase or ADPase activity (figure 6). Some scattered cells deep in the epithelium showed reaction product with ATP as substrate but not with ADP. The exact nature of these cells is unclear, but they were not associated with Schreiner organs or any other obvious epithelial endorgan.
4.5. Solitary chemosensory cells lack ecto-ATPase activity
Solitary chemosensory cells are scattered across virtually the entire external epithelium of most teleosts [3], including on the barbels of catfishes, where SCCs are scattered between the taste buds [5]. In our preparations of catfish barbels, as shown in figure 1, ecto-ATPase activity is not apparent in the epithelium outside of the taste buds and therefore is not associated with SCCs in these locations.
The pectoral fin of sea robins has a unique specialization in which SCCs are closely packed together along the anterior three fin rays, which lack fin webbing and which thus form specialized non-taste chemoreceptor organs [38,39]. These modified free fin rays possess numerous SCCs in the epidermis, but no taste buds. Despite the high density of SCCs, the fin rays showed no detectable ATPase or ADPase activities (figure 4c,d). Similarly, the nerve fibre bundles that innervate SCCs showed no specific reaction product.
5. Discussion
In all species examined, including lamprey, an elasmobranch and all teleosts, taste buds exhibit pronounced ecto-ATPase activity. This is consistent with previous findings showing ecto-ATPase in mammalian taste buds [13,31,40,41], in an amphibian [42] and in another teleost [26]. By contrast, neither Schreiner organs in hagfish nor areas containing densely packed SCCs exhibited this trait in any species examined. Thus, the presence of ecto-ATPase appears coincident with the appearance of taste buds in the vertebrate lineage.
Mammalian taste buds are known to use ATP as a key transmitter between taste cells and nerve fibres [9], which express two ionotropic purinergic receptors, P2X2 and P2X3 [43]. The presence of ecto-ATPase is probably necessary to inactivate the ATP neurotransmitter once it is released into the extracellular space because P2X receptors will desensitize rapidly if exposed to high levels of extracellular ATP [44]. The presence of ecto-ATPase in association with taste buds of non-mammalian species suggests that all vertebrate taste buds similarly use purinergic signalling to transmit information from taste cells to nerve fibres. Indeed, zebrafish, like rodents, express P2X2 receptors on the nerve fibres innervating taste buds (figure 1g) [30].
SCCs are single sensory epithelial cells, present in all vertebrates from hagfish to mammals [4,8,45,46]. Like taste cells, SCCs are chemosensory endorgans consisting of secondary sensory cells, i.e. they lack an axon. Despite the similarity in function and neural relationships between SCCs and taste buds, no ecto-ATPase activity occurs in association with SCCs even in epithelia with densely packed SCCs such as the fin rays of searobins (figure 3c). SCCs often occur in epithelium near taste buds, as well as in respiratory passageways [4,47,48]. Yet, despite proximity to taste buds showing ecto-ATPase reactivity, the SCCs and nerve fibres innervating them lack such reactivity.
Like taste cells, SCCs have synaptic connections onto afferent nerve fibres. Whereas SCCs are innervated by either spinal or cranial nerves appropriate to the epithelium in which they reside, taste buds are only innervated by facial, glossopharyngeal or vagus nerves. Thus, SCCs are innervated by ganglion cells arising from neural crest (e.g. dorsal root ganglia), while taste buds are innervated by ganglion cells derived from epibranchial placodes [2].
Schreiner organs are sensory organs, scattered throughout the epidermis of hagfish. Schreiner organs are superficially similar in appearance to taste buds, being multicelluar aggregates composed of several types of elongate epithelial cells [4,49]. Schreiner organs are similar to taste buds as being specialized, multicellular epithelial chemosensory endorgans, but are not homologous to taste buds. They can be distinguished by several morphologic features. First, Schreiner organs lack dermal papilla, i.e. they do not sit adjacent to the basement membrane of the epithelium as do all taste buds. Secondly, Schreiner organs are innervated by spinal or cranial nerves. On the basis of these differences, Braun [4] suggested Schreiner organs are not the forerunners of taste buds. Rather, they may be a specialization of accumulated SCCs. Why hagfish lack taste buds and the associated purinergic signalling system is enigmatic. Recent molecular data indicate that hagfish are monophyletic with lampreys [50–52]. If so, the absence of taste buds probably indicates that this is not a primitive trait, but merely another of the collection of vertebrate traits that hagfish have lost during evolutionary time [53].
More than a decade ago, Finger [8] had suggested that taste buds may have evolved as an aggregation of Merkel-like sensory cells becoming associated with SCCs. This hypothesis was based in part on the similarity of morphology of the sensory cells of these two systems. But a stronger similarity was seen in the commonality of receptor mechanisms between a subset of taste bud cells and SCCs in catfish; both SCCs and a subset of taste cells express similar lectin binding indicative of an arginine receptor [5]. In mammals, the SCCs of the airways use the bitter taste (T2R) receptor cascade to detect toxins where the SCCs release acetylcholine as a neurotransmitter [47,48]. As many chemoreceptor cells in diverse organ systems rely on taste transduction cascades [48,54–59], we no longer believe that the common expression of taste receptors by SCCs and taste buds necessarily suggests a phylogenetic linkage. Furthermore, in fish, SCCs do not appear to rely on the G-protein-coupled receptor cascade characteristic of taste buds [60]. Thus, the evolutionary relationship of SCCs and taste buds is unclear.
The gustatory nerve fibres in teleosts form a plexus in the basal part of the bud surrounding the Merkel-like basal cells [15,35]. In the teleosts studied, the region within the taste buds containing the nerve fibres exhibits heavy ecto-ATPase activity. In mammalian taste buds, it is the glial-like type I cells that express the ecto-ATPase, NTPDase2. In the teleost taste buds examined at ultrastructural levels in the present study, we see specific ecto-ATPase activity associated with the plasma membranes of both elongate taste cells and nerve fibres (figure 3). Thus, in teleosts, unlike in mammals, the nerve fibres themselves appear to express a specific ecto-ATPase enzyme. In both mammals and the fishes studied herein, non-specific ectonuleotidase is associated with the nerve bundles, including glia cells, below the taste buds.
In mammals as in all species examined in the present study, the ectoenzyme associated with taste buds is highly specific for extracellular 5′-triphosphates, i.e. strongly preferring ATP over ADP [13]. The high substrate specificity is unique to this isoenzyme and accounts for the high levels of staining seen with the ATP substrate over the ADP substrate [61]. The presence of a highly specific ecto-ATPase associated with taste buds in all classes of vertebrates suggests that purinergic transmission may be one of the defining features of the gustatory periphery. The necessity for purinergic transmission is unique for the taste system although other neural systems use ATP as a co-transmitter or cofactor which modulates the effectiveness of a co-released primary neurotransmitter [62]. The use of a purinergic signal may relate to the epithelial origins of the taste system, i.e. release of ATP is a common response of epithelial cells to external stimuli [63–65]. Furthermore, in taste buds, release of ATP from type II cells is via an unusual non-vesicular mechanism involving gated hemichannels [11,12,66], with gating largely dependent on action potential-mediated depolarization of the taste cells [10]. By contrast, typical neural systems use a variety of other neurotransmitters, including acetylcholine, amino acids and amines, all of which are released via snare-protein-mediated vesicular mechanisms. In mammalian taste buds, type III taste cells make obvious morphologically typical synapses onto nerve endings [67] and use a vesicular mechanism [68] to release serotonin and GABA which act on receptors expressed by the type II cells [66,69]. Thus, taste buds appear to use both neuronal-type (vesicular) and epithelial-type (hemichannels) mechanisms to release neurotransmitter.
In other epithelial secondary receptor cells such as hair cells of the ear and lateral line organs, the sensory cells are more neuronal in terms of mechanism of transmitter release. They use vesicular release of glutamate as the primary means of transmission from sensory cell to nerve fibre [70,71]. Unlike taste buds, hair cells in these systems originate from neurogenic placodes [72], hence a vesicular, neuronal type of transmitter release is not surprising. By contrast, the more epithelial-like release of ATP via hemichannels is consistent with the origin of taste bud receptor cells from local epithelium, rather than from neurogenic placodes or neural crest [73,74].
In summary, we find that ecto-ATPase activity, indicative of purinergic signalling, is common to taste buds in all vertebrates examined to date, including lampreys, elasmobranchs, teleosts and amniotes. The commonality of taste buds and purinergic signalling mechanisms throughout the vertebrate lineage places the origin of taste buds alongside of the evolutionary origin of the earliest vertebrates. The lack of evidence for purinergic transmission from SCCs does not support the previously hypothesized relationship between taste buds and SCCs [8]. Rather, SCCs may be more primitive than taste buds, being related to the secondary sensory cells described in the epidermis of amphioxus [75,76].
6. Acknowledgements
The authors thank Dr Mark Voigt of St Louis University for providing the P2X3.2 : gfp zebrafish and Kristin Artinger of University of Colorado Anschutz Medical campus for helping to maintain these animals. We also express sincere thanks to Dr Takayuki Shoji of Tokai University for his help in collecting shark and hagfish. The authors thank Drs Linda Barlow and Kristin Artinger, University of Colorado Anschutz Medical Campus, for critical reading and comments on earlier versions of this manuscript. This work was supported by grants from the following sources: National Institutes of Health (RO1 DC007495 and RO1 DC009820 to T.E.F., and P30 DC04657 to D. Restrepo); and by grant-in-aids (no. 22580205) to S.K. from the Ministry of Education, Science, Sports, and Culture of Japan.
- Received January 24, 2013.
- Accepted February 14, 2013.
© 2013 The Authors. Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0/, which permits unrestricted use, provided the original author and source are credited.


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(Here Are Three Ways to Make Sure You're Not Anymore)
Are You Guilty Of Being A Distracted Dater?
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March 16, 2018
| Product
This kit features a nail file, nail clipper, 4-in-1 Nail White Pencil, and shiner and buffer.
November 14, 2016
| Photo Gallery
Browse our gallery for stocking stuffer ideas that will surely make you say holi-yay!
June 1, 2015
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Flowery's 4-in-1 Catty Nail File & Case Color Pop Collection includes 180/240 grit nail file with three new eye popping designs.
April 8, 2015
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Flowery's new retail design will spruce up your salon.
December 17, 2014
| Article
Flowery announces new File Catty Candy Lush Collection for winter.
August 31, 2014
| News
Professional beauty products distributor Spilo Worldwide has rebranded five of its signature lines — including Flowery and ToolWorx.
April 22, 2013
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The 5" nail files come in a patented protective case and feature six new fun and fanciful exclusive designs.
December 1, 2011
| Photo Gallery
NAILS announces 2011 Readers' Choice Award winners and finalists.
January 3, 2008
| News
Several well-known nail product manufacturers have been purchased by larger beauty industry companies. Realys, Flowery Beauty Products, and Chéci have all been bought.
December 1, 2005
| Photo Gallery
The votes are in! Your favorite products from this year are all here!
December 1, 1985
| Article
Anniversaries are a time to reflect on past accomplishments and to evaluate plans for the future. Flowery Beauty Products takes the occasion of its 75th Anniversary to do just that.
Foot Forward Summit 201832 photos
Cosmoprof North America 201837 photos
System established under state law that provides payments, without regard to fault, to employees injured in the course and scope of their employment.
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We've officially launched
It's a great day for elmah.io! The beta sign is gone and a lot of exiting stuff have happened since the first alpha. We really think that we have built a product, which will save you hours and make your everyday fun and focused.
Why this post then? Along the official launch, we're also ready to introduce paid plans on elmah.io. Luckily for all of you beta testers, we still offer a free plan and all of your logs are automatically moved to the free plan. So why introduce pricing at all? Well as you probably know, running a website isn't free. To keep the releases flowing and to give us a change to innovate on this already awesome platform, we really hope that you will consider supporting us, by purchasing one of our paid plans. We have a lot of ideas on how to make elmah.io even cooler and we need your help to do that. Check out the Plans and Pricing page for details. To purchase a plan, log in to elmah.io and visit the profile page.
If you haven't visited us for a while, let us use a few lines to introduce some of the great features we have been building recently. In November we introduced Lucene search, making it possible to write advanced queries against your errors. In December we gave you the much sought HTTPS support. In January Ignore Filters which later on matured into an advanced rule engine, making it possible for you to react through mail and/or web hooks, when new errors occur. Recently we introduced the new Overview tab, which in a simple way visualizes your recent error activity.
As usual please reach out if you have ideas for elmah.io.
Thank you for your support!
This post was brought to you by the elmah.io team. elmah.io is the best error management system for .NET web applications. We monitor your website, alert you when errors start happening and help you fix errors fast.
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PAUL MITCHELL CLARIFYING SHAMPOO TWO 500ml
Color-safe formula deeply cleanses oily hair and scalp, leaving hair fresh, shiny and full of body. Helps minimize oil production, so hair becomes less oily over time. Paraben free and Colour safe.
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Please allow 10 working days from dispatch of your order before notifying us of any late deliveries.
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Army football coach Rich Ellerson mentioned Jared McFarlin in the same conversation as Giants receiver Ramses Barden last year.
Army football coach Rich Ellerson mentioned Jared McFarlin in the same conversation as Giants receiver Ramses Barden last year.
McFarlin, a 2010 Monroe-Woodbury graduate, was a sophomore wide receiver with a lot of raw talent. At 6-foot-5, he had the height similar to Barden, who is 6-7.
Army's staff thought McFarlin had the potential to be as good as Barden.
It doesn't appear McFarlin, who started three games in 2011, will have a chance to further impress Army's coaches.
The Army sports information department confirmed Tuesday that McFarlin is no longer with the team. McFarlin's name had been removed from the team's roster on its website.
McFarlin couldn't be reached for comment. The reason for his departure wasn't known at press time.
McFarlin was a feel-good story early in the 2011 season. He earned an opening-day start and made four catches, including his first college touchdown against Northern Illinois.
McFarlin started two of the next three games. But McFarlin suffered a concussion before Army's fifth game against Tulane on Oct. 1, and made just one catch the remainder of the season.
McFarlin finished the season with seven catches for 74 yards.
Competition at the two wide receiver positions was open this spring due to graduation of starters Davyd Brooks and Austin Barr.


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And what makes you think your Mom or Grandma wouldn't love some nameplate jewelry as much as you would? Why not? What women doesn't love some gold (or silver) and something shiny?? You could go with something traditional that say's "I Love You" or "Mom", maybe throw in some colored (or real, you baller you!) stones in her birthstone or even in yours! Or why not get her her name or a short and sweet phrase or a nickname she calls you. Maybe you and your siblings go in on it together and get all your names on a necklace or bracelet. The sky is the limit! Make it something sweet, thoughtful and meaningful…hey, maybe mom would appreciate some bamboos, but at least two pair 😉
Top Row: Jewelry By Net Bottom Row: Elim Jewelry
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What are you doing Thursday evening? No plans?? Why not come out, have some fun and get some cool stuff!?! [Um, some FREE stuff!! Keep reading….]
This Thursday happens to be Last Thursday, that one special day of the month the Alberta Arts District turns their charming little neighborhood into a spectacle of art and entertainment for all ages to enjoy. Even better, DRY Soda's Savor the Flavor Tour is making Last Thursday their official Portland stop that night!
So what better place for me to hang out with my pals from DRY Soda and meet up with some of my other favorite pals: YOU GUYS!! I'm officially inviting all of you to come hangout and enjoy the festivities with us, rain or shine—but let's hope for some sunshine!
Not only that, but if you stop by the DRY Soda Truck and tell us the "secret" password (see below), you'll get a special treat from Stumptown Lunch and DRY Soda!
Could anything be cooler than that?
Well…the DRY Soda Truck is going to be parked right outside Salt & Straw (they make delicious small-batch, homemade ice cream that even Oprah loves), so you can pop in and buy yourself a couple of creamy scoops for dessert! That's pretty darn cool, too! (Again, let's hope for sunshine….)
Here's the information:
When: Thursday, March 29, 2012
Where: DRY Soda's Tour Truck will be parked just outside of Salt & Straw
Map: 2035 NE Alberta Street
Time: 5 to 8 p.m.
Secret Password: Put A Bird On It!
Feel free to shoot me an email or tweet me to let me know if you're going to stop by! Hope to see you Thursday!!
The Lunch Lady


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Phu Quoc, also known as Pearl Island, is the largest island in Vietnam. Blessed with cool ocean breezes and sunny weather, this is a great destination for relaxing on the beach or by the pool.
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The national development and Reform Commission suspended production licenses for electric vehicles earlier this year to control capacity expansion. At present, about 2000000 new energy vehicles are expected to be added every year, and the relevant institutions are re evaluating the production of electric vehicles.
The national development and Reform Commission to re open license news declined to comment, said it was to amend the existing license program, and hope to improve the threshold of new entrants.
Since March 2016, China has issued 15 manufacturing licenses for electric vehicles. Local manufacturers, Wanxiang Group, and the joint venture between Volkswagen and domestic manufacturers are among them, which has also promoted the competition in the automotive industry. Then, smart TV and air conditioning manufacturers are planning to enter the new energy automotive industry.
The most important is that foreign auto makers Ford and Tesla are ready to enter the China electric car market, in order to meet the increasingly stringent local carbon emissions and fuel economy requirements. Ford and the Anhui auto company's new joint venture plans to apply for the establishment of a factory, the annual production 100 thousand units of pure electric vehicles. According to the report said Tesla has reached a preliminary agreement in the local production and the government of Shanghai. Chief executive Ilon Mask (Elon Musk) this month told investors that the company from production and three years in China.
The license plan is also a key measure for the Chinese government to manage the world's largest new energy vehicle market. The Chinese government announced last week that it will launch a pilot program by June next year, allowing foreign automakers to set up wholly-owned electric vehicle factories in the free trade area. Starting in 2019, China will require most automakers to score new energy vehicles, or if they do not meet the requirements, or will face rectification or punishment.
With the support of the state, China surpassed the United States in 2015 as the world's largest new energy vehicle market, including pure electric vehicles, plug-in hybrid vehicles and fuel cell vehicles. According to the China Automobile Industry Association, 507 thousand new energy vehicles were sold nationwide last year. China suspends the expansion of traditional gasoline vehicle capacity, aiming to increase the annual sales of new energy vehicles to 2 million vehicles by 2020.
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Learn how to protect private wells at workshop
The University of Rhode Island Cooperative Extension and the Rhode Island Department of Health will conduct an educational workshop on private well water protection on Thursday, Jan. 29, from 6 to 8 p.m. The workshop will be held at the North Kingstown Free Library, 100 Boone St. in North Kingstown.
"Approximately 10 percent of Rhode Island residents depend on private wells for their drinking water. While the state does test public drinking water sources, it is the private well owner's responsibility to make sure that the water is safe to drink," explains Alyson McCann, with the University of Rhode Island Home*A*Syst Program.
The Rhode Island Department of Health recommends that private well owners test their wells every year for contaminants such as total coliform bacteria, nitrate, and sodium. Tests are also recommended every three to five years, and every five to 10 years for a series of other contaminants.
"Ultimately, we hope that workshop participants will be able to use the information they receive in order to ensure that they and their families have healthy drinking water," McCann said.
Funded by the RI Department of Health Capacity Development Program, this free two-hour long workshop will include a presentation, informational hand-outs, and the chance to speak with well water experts about individual concerns and questions.
Pre-registration is required, as space is limited. Call the URI Cooperative, 874-4918 for more information or to register.


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Although the physical "Space" is closed... We are no longer located in a specific location... This online space will remain! Yes, you'll still find Frank's recipes :) and we'll be posting some other things as well. We'll take some time to re-group, then look for more information :) Thank you so much for all your support over the last 5 years!! Ryan, Wanda, and Julie … [Continue reading...]
The "Space" is Now Officially Closed
Final Yard Sale Today! 9-3
Sadly... Today's the day... 9-3 Final Closing Yard Sale... Come check out our great deals, or just come and visit a little bit... If you haven't been in, it's indoors, 12 Bay Street, Wilmington... Great Deals that you won't want to miss! … [Continue reading...]
Final Yard Sale – July 11 – 9am-3pm
Our FINAL "We're Closing" Yard Sale is coming up on July 11, that evening we are officially moving out of the space… Please come and give us less to move!! You are not going to want to miss this final sale!! Mark your Calendars!!!
[Continue reading...]
Yard Sale – Day 2!
Day 2 of our Massive "We're Closing" Yard Sale series here. Please remember that we will open promptly at Noon today (Sunday). We will not open early for previews. The doors will remain locked and will not be opened for anyone prior to the full opening at Noon. The same rules apply for for the final sale day […]
[Continue reading...]
Yard Sale Reminder!!
Day 1 of our Massive "We're Closing" Yard Sale series is almost upon us. Please remember that we will open promptly at 10am on Saturday morning. We will not open early for previews as we have a lot to prepare. The doors will remain locked and will not be opened for anyone prior to the full opening […]
[Continue reading...]


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Side Chair DSW, 1950
Diseñada originalmente por Charles y Ray Eames en 1950. Charles Ormond Eames, Jr (1907–1978) and Bernice Alexandra "Ray" (néeKaiser)Eames (1912–1988) /ˈiːmz/ were American designers who worked in and made major contributions to modern architecture and furniture. They also worked in the fields of industrial and graphic design, fine art and film Charles Eames, Jr (June 17, 1907 – August 21, 1978) was born in St. Louis, Missouri. Charles was the nephew of St. Louis architectWilliam S. Eames. By the age of 14, while attending Yeatman high school, Charles worked at the Laclede Steel Company as a part-time laborer, where he learned about engineering, drawing, and architecture (and also first entertained the idea of one day becoming an architect). Ray-Bernice Alexandra Kaiser Eames (December 15, 1912 – August 21, 1988) was an American artist, designer, and filmmakerwho, together with her husband Charles, is responsible for many classic, iconic designs of the 20th century. She was born inSacramento, California to Alexander and Edna Burr Kaiser, and had a brother named Maurice. After having lived in a number of cities during her youth, in 1933 she graduated from Bennett Women's College in Millbrook, New York, and moved to New York City, where she studied abstract expressionist painting with Hans Hofmann. She was a founder of the American Abstract Artists group in 1936 and displayed paintings in their first show a year later at Riverside Museum in Manhattan. One of her paintings is in the permanent collection of The Whitney Museum of American Art Fuente: Wikipedia


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"Grim experience.."
About: Manchester Royal Eye Hospital Manchester Royal Eye Hospital Manchester M13 9WL
Posted via NHS Choices
What I liked
What could be improved
Better communication from reception/medical staff...
Anything else?
I attended the emergency eye dept after my GP advised me I needed to be seen urgently..I live an hour away from MRI.& am physically Disabled..Live on my own & need someone to drive me..I managed to get help from a family friend to drop me off..arrived at 4pm.then waited 4hours in the overfilled waiting area.My daughter came to pick me up at 8pm.I still had not been seen by the doctor.& when I enquired from reception staff member how much longer before I would be examined I was promptly told that their had been some emergincies so she said it would be a long wait.also the nurse said I may have to return next morning if they where running over..seems that the unit closes at 9pm..I couldn't expect my daughter to wait around for that amount of time she as to work the next day..so I left without being seen..so this service is abysmal..my eye condition goes untreated..I may take this further..see what the Trust thinks of this..
Response from Manchester Royal Eye Hospital 5 years ago
Thank you for your comments posted on the NHS Choices website. We were very sorry to learn of your concerns regarding your attendance at the Emergency Eye Centre in December 2012. This is not the level of service we want our patients to experience. It is, however, very difficult to respond to the specific concerns you have raised without being able to investigate in detail. We take all issues surrounding patient care very seriously and would very much like to hear from you directly about this. If you contact our Patient Advice and Liaison Service on 0161 276 8686 or by e-mailing firstname.lastname@example.com they will be happy to take the details of your attendance which will enable an investigation to take place.
Eve Koutidou
Patient Experience & Quality Project Officer
Central Manchester University Hospitals, NHS Foundation Trust
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Livestock . . .
Cross Creek Rodeo Company, owned by Ed and Vana Doolittle, can provide a number of bovine and equine superstars that have been voted by the top cowboys to go to the International Finals Rodeo held in Oklahoma City, Oklahoma or the Southern Rodeo Finals held in Asheville, North Carolina.
Some of the four legged superstars are #67 Mud Flap, International Finals Rodeo bucking bull and son of World Champion bucking bull Skat Kat; #611 Smarty Pants, PBR Bud Cup Finalist bull featured in Las Vegas; #08 Copenhagen Backwash, son of Wrangler National Finals bucking bull Copenhagen Crazy and #140 Lil Erkel who also has made his way to the International Finals Rodeo.
Nationally televised bucking horses #80 Goose Creek, a four time Wrangler National Finals Rodeo bucking horse; #X6 Yukon; #L1 Lucky Lady; #839 Teddy Bear; #005 Cotton Fields; #734 Ariat; #L199 Shanook; and #831 Mustang Sally have all respectively made their way either to the Wrangler National Finals Rodeo, the International Finals Rodeo or the Southern Finals Rodeo.
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During my tenure as editor of this magazine I wasted many hours studying a big, multi-colored map of the world pinned above my bookshelf, imagining where the next J-Bay or Nias might be. When well-traveled surfers or photographers would pass through the office, I'd steer them to the map and ask what they knew about various uncharted surf zones, hinting that if a spot had potential I might arrange to send them there. I hungered for info about places like the Kuril Islands, between Japan and Kamchatka. Or Isla Socorro, below Cabo. Or a tiny dot named Europa in the Mozambique Channel.
For some reason, my internal compass kept pointing to Alaska. I was convinced that somewhere along the 2,500-mile sweep of inlets and islands on that cold Pacific coast there had to be a Klondike of good surf. But Alaska's expensive, so we didn't dare risk such a mission without reliable intel, and no one I talked to knew dick about the place.
Then in 1991 the mag ran a short piece about Dominic Rice, who'd been stationed there by the U.S. Coast Guard. "You wouldn't believe the surf in Alaska," Rice told writer Gary Lynch. "There can be 30-foot seas outside pushing swell 20 miles up an inlet into clean, mirror-smooth waters. Lefts, rights, points, and reefs."
That did it. We eventually recruited one of the sport's best wordsmiths, Dave Parmenter, one of its most reliable photographers, Bob Barbour, and two of its most versatile pros, Brock Little and Josh Mulcoy, and sent them north.
They scored.
"All of my most memorable surf trips have included a fear of frostbite."
Barbour captured both the magnificence of the landscape and the quality of the surf in a single image from Yakutat, which ended up on the cover (January 1993): Mulcoy leaning into an easy green left with 18,000-foot Mount Saint Elias as backdrop. And Parmenter crafted a witty, informative, impassioned story that I still regard as one of the best articles ever published in any surf magazine anywhere.
"It resembled a small day at Third Point, Malibu, may it rest in peace," Dave wrote about a pointbreak they found near Sitka. "Except Gidget would have some pretty stiff nipples here, and the bears have even less patience with the Vals than Dora did. This was as far away from the teeming mouse hole of surf culture as you could get in America."
It probably would have happened anyway, but that article (coupled with ongoing advances in wetsuit technology) seemed to trigger a coldwater gold rush. In the 15 years since we've seen surf shots and video from pretty much every icy, gray outpost on the planet: Iceland, Norway, Greenland, Kamchatka.
Parmenter's piece was one of the few in my tenure at SURFER that affected me personally. I have never been to Indo, but I have followed that first crew's footsteps to Yakutat and beyond. I've surfed within view of white-tipped volcanoes in the Aleutians, thawed my hands in a smoke-scented sauna on Vancouver Island, heard the sound of ice bits crackle against my fins on a roping right-hander in Antarctica. I've had longer and better sessions in the tropics, but all of my most memorable surf trips have included a fear of frostbite.
So when I glance at a globe these days, the extreme latitudes north and south have lost much of their mystery. But then I look close and realize how much unsurveyed coast remains worldwide. Take the Kurils. I've been asking about the place for almost two decades now, Googling for maybe half that, and I still haven't heard dick about the surf there. Part of me hopes I never will.


text: [Federal Register: February 1, 2000 (Volume 65, Number 21)] [Notices] [Page 4812-4813] From the Federal Register Online via GPO Access [wais.access.gpo.gov] [DOCID:fr01fe00-42] ======================================================================= ----------------------------------------------------------------------- DEPARTMENT OF EDUCATION Federal Family Education Loan Program AGENCY: Department of Education. ACTION: Announcement of guaranty agencies selected to negotiate participation in a Voluntary Flexible Agreement. ----------------------------------------------------------------------- SUMMARY: The Secretary announces the guaranty agencies that have been selected as possible participants in Voluntary Flexible Agreements under Section 428A of the Higher Education Act of 1965, as amended (the ``HEA''). FOR FURTHER INFORMATION CONTACT: Mr. Cameron R. Ishaq, Department of Education, Student Financial Assistance, 7th and D Streets SW, Room 4616 Washington, DC 20202, (202) 260-5076. Individuals with disabilities may obtain this document in an alternate format (e.g., Braille, large print, audiotape, or computer diskette) on request to the contact person listed in the preceding paragraph. SUPPLEMENTARY INFORMATION: On July 28, 1999, the Secretary published a Notice in the Federal Register (64 FR 40859) inviting guaranty agencies participating in the Federal Family Education Loan (FFEL) Program to apply to participate in the FFEL Program under a Voluntary Flexible Agreement (VFA) as authorized by Section 428A of the HEA. The Notice informed the guaranty agencies of the deadline for submission of the application, the information that needed to be included with the application and an outline of the evaluation and selection process that the Secretary planned to use. Nine guaranty agencies submitted timely VFA applications. Eight of the agencies made oral presentations to the Secretary's representatives during September and October. The Secretary has evaluated the information provided by the nine agencies in their applications and oral presentations and has selected six agencies with which to try to negotiate a VFA. The selected agencies are identified in this notice. The Secretary believes that each of the selected VFA applications includes specific proposals that could improve the FFEL Program. However, a number of the applications also include proposals that the Secretary believes are outside the appropriate scope of a VFA or are contrary to the Secretary's policies and goals. In light of this situation, the Secretary has decided not to publish the selected VFA proposals at this time. The Secretary understands, however, that each of the selected guaranty agencies has made their proposal public. In the list of selected agencies below, the Secretary is also providing the guaranty agency's contact to obtain a copy of the proposal. The agencies selected by the Secretary are: 1. American Student Assistance (ASA) Proposal available from: www.amsa.com 2. California Student Aid Commission/EdFund (CSAC) Proposal available from: www.csac.ca.gov 3. Colorado Student Loan Program (CSLP) Proposal available from: www.cslp.org 4. Great Lakes Higher Education Assistance Corporation (GLHEAC) Proposal available from: www.glhec.org 5. Pennsylvania Higher Education Assistance Authority (PHEAA) Proposal available from: Peggy Shedden (717) 720-2660 or email@example.com 6. Texas Guaranteed Student Loan Corporation (TGSLC) Proposal available from: www.tgslc.org Electronic Access to this Document You may view this document, as well as all other Department of Education documents published in the Federal Register, in text or Adobe Portable Document Format (PDF) on the Internet at either of the following sites: http://ocfo.ed.gov/fedreg.htm http://www.ed.gov/news.html To use the PDF you must have the Adobe Acrobat Reader Program with Search, which is available free at either of the previous sites. If you have questions about using the PDF, call the U.S. Government Printing Office (GPO), toll free, at 1-888-293-6498; or in Washington, DC, area at (202) 512-1530. Note: The official version of this document is the document published in the Federal Register. Free Internet access to the official edition of the Federal Register and the Code of Federal Regulations is available on GPO Access at: http://www.access.gpo.gov/nara/index.html. Program Authority: 20 U.S.C. 1078-1. [[Page 4813]] Dated: January 20, 2000. Greg Woods, Chief Operating Officer, Office of Student Financial Assistance. [FR Doc. 00-2195 Filed 1-28-00; 11:11 am] BILLING CODE 4000-01-U