text:
Friends and all characters within are copyrighted to NBC and are not my
creations, although I'd be rich if they were. Damn!
This work is copyrighted to the author (c) 1997. Please do not remove the
author information or make any changes to this story. You may post freely to
non-commercial "free" sites, or in the "free" area of commercial sites. Thank
you for your consideration.
Friends: The Contest Part 3 (m/f)
by Scarlet Pimp(ernel)
For the next two days nothing more happened. Monica, Rachel and Phoebe went
shopping, each bringing home at least two new bikinis and a tiny suit for the
men, while the guys went about their lives normally.
On Friday, down at Central Perk, Joey and Rachel sat on the couch. Chandler
walked in the door, carrying his briefcase.
"You will not believe the day I've had!" he said then plopped down on the
couch next to Rachel.
"Why, what happened?" asked Joey.
"Um, well... okay, fine. Nothing. Are you happy now? You've found me out!
Nothing particularly strange happened today."
Chandler got up and went to get himself a cup of coffee. As he left, Joey
got up as well.
"I'm going to the bathroom," he told everyone and no one in particular.
Some people looked at him, then went about their business. Joey left.
Sensing an opportunity, Rachel, too, got up and went to the bathroom. She
waited outside the door until Joey emerged.
"Joey, did you wash your hands?"
"Huh? Uhm, no."
He hung his head and turned to go back into the room. Rachel looked around
and saw no one looking, and pushed him into the tiny bathroom.
"Rachel, what are you doing?"
"I'm gonna show you something about personal hygiene!"
"Okay, I'm sorry, I forgot! Jeez, you're worse than my mom."
She turned on the tap and he put his hands under the water, washing them.
When he was done, he shook them dry and turned to go.
"There, done. Happy?"
"No, aren't you forgetting something?"
"What now?"
"Didn't you just pee?"
"Did you wash off your big cock?"
Joey jumped back from Rachel, confused by this question.
"It's all right, I'll show you," she said, and reached for his zipper.
Shocked, Joey just stood there while she pulled his dick out of his pants,
splashing it with warm water and rubbing it. His mouth hung wide open.
"There, much better, no? Oh, look, now it's all wet. I'll dry it off. I'll
bet your mom never did this!"
She took it in her mouth and started blowing him. Joey moaned.
"What're you doing, Rachel?"
"I'm just ensuring that I have your support on Saturday, if you know what I
Sadly, he didn't.
"What?" he stammered.
"Vote. For. Me," she said, punctuating each word with a tug on Joey's
hardening cock. "I want to win, and I want it bad."
Okay, thought Joey.
"I can do that."
"That's good, now sit back and enjoy."
With that, she returned to her business, sliding her lips up and down Joey's
shaft, her red lips a perfect 'o'. She reached into Joey's pants and pulled
out his balls, massaging them with her hands, while her mouth teased his
Rachel released his cock from her vacuum mouth and took a minute to suck on
each of his testicles, while she jerked him with her hand. There wasn't much
room in the small bathroom, so he had to stand while she knelt before him and
sucked his hard cock.
Her free hand found its way down her panties and she began to wank herself
while she pleasured Joey. He saw her fondling her twat and placed his hands
on her head, bringing her mouth back to his rod. He controlled her motion
with his hands, fucking her mouth while she felt herself up and tweaked her
own nipple.
He smiled down at Rachel, kneeling in front of him. Her lips and tongue
worked on the head of his penis, summoning his orgasm. Joey let go of her
head and forced her off of his cock. She pouted at him, then stood.
"What's wrong, Joey?"
"This isn't fair," he said, "you're not getting enough."
She smiled at him and hiked up her skirt.
"Okay, you want some of this?"
He pulled her sopping wet underwear aside and stuck a finger into her.
Rachel closed her eyes, and grabbed his cock once more.
Joey grabbed her by the ass and lifted her onto the sink. He wedged himself
between her legs and she guided his cock into her steaming cunt. Slowly, but
with increasing speed, he began to pump her, her legs locked behind his back.
He lifted her from the sink and continued to thrust in and out of her. She
slid her hand down and toyed with her clit, while he reached up under her
blouse and tore it open. Her lacy bra soon fell to the floor, torn and
unwearable ever again.
Rachel put her arms around Joey's shoulders, while he bent to her breasts,
kissing both on the nipple, then sucking on them. He put both hands on her
ass and held her aloft, rutting into her. She began to pant, her orgasm
He turned and started pumping her against the wall. He bit down on her
nipple and she nearly screamed in pleasure, but held it back. She had no
desire to let everyone know what was going on in the bathroom.
She began to writhe in his grip as the throes of her orgasm crashed through
her. After a minute, the waves subsided and she calmed down, her breath
once more coming in even gasps. She smiled seductively at Joey.
He pulled out of her, and she once more got to her knees, returning his cock
to her hungry lips. Rachel could taste her own womanly juices on his prick
as she took it onto her tongue, using her tastebuds to scrape roughly across
oey's sensitive cockhead.
"Oh. That's it," he moaned, then began to unload into her mouth.
Rachel immediately removed his cock and directed his sperm into the sink,
spitting what little she had already taken in her mouth. Joey was a little
disappointed that she didn't swallow, but didn't really mind. This was an
unexpected BJ, so how could he complain?
"Now, who are you going to vote for?"
"You, Rachel."
She carefully washed Joey's cock clean once more, then returned it to his
pants and zipped him up. She then picked up her bra and stuffed it into her
inside pocket and buttoned her blouse.
"Remeber, Joey, this is our little secret, right? Don't tell anyone."
"I promise."
Rachel peeked out the bathroom door, then snuck into the woman's toilet.
Joey went and sat down on the couch.
"What the hell were you doing in there?" Chandler asked.
"You know, guy stuff."
"Oh, okay," Chandler said sarcastically, but left it alone.
Minutes later, Rachel emerged from the woman's bathroom, looking as
immaculately clean as she had before, bearing no trace of the last few
minutes' actions. (Unless one was observent enough to notice that she no
longer wore a bra under her blouse.
At work, Phoebe was busy giving Ross a massage. He'd started coming to the
parlour a few weeks ago, trying to relieve some of the tension that had been
building up while he and Rachel fought and then after they'd broken up. Now
he'd started getting used to it, and looked forward to his weekly session.
"Uh, Ross! You are sooo tense! You've got to relax!"
Phoebe kneaded the muscles in the back of Ross' neck, pounding on the little
knots of flesh that she found there.
"I can't help it, Phoebs. Ever since I lost Rachel, every little thing that
goes wrong at work has just been driving me over the edge. It's like the
world has it in for me."
"You see? It's negative remarks like that that make bad stuff happen to you!
You've got to be positive! I can't take your bad vibes! Here, try this
herbal tea," she said, holding a box out to him. Ross reached for it, and
she pulled it away from him, saying, "Hey! I was just showing you, get your
own! Now lie back down, and RELAX!"
Ross put his head back onto the table and closed his eyes, thinking of Rachel
naked in his bed. He pretended that Phoebe's hands were hers, massaging his
troubles away.
"There, much better. Now you're really relaxing," Phoebe said, pleased with
her progress. "Keep breathing, in and out. Slowly."
He ignored her, fantasizing about Rachel. Rachel knelt in front of Ross, her
admriable, tanned boobs squeezed together between her arms in the classic
Playboy 'fuck me' pose, her lips pouty and full, whispering "I love you" to
him. He reached out and took her left breast in his mouth, sucking on her
nipple delicately. She cooed in pleasure, putting her hands on his huge cock
(it was a fantasy) and pulling on it gently.
Ross felt his real penis swelling under his towel, to its full length (which,
although not huge, was not small). Rachel pushed him onto his back and
slurped up the drop of pre-cum on his tool, then deepthroated the monster,
gagging on its magnificent girth and length.
Phoebe worked her way down Ross' back, then stopped.
"Okay, Ross, turn over."
He snapped back to reality and began to turn over, not thinking of his
excited state. As he shifted position, his towel fell, exposing his erect
manhood to his friend. He quickly retrieved the towel and held it in front
of his, blushing.
"Sorry about that."
"Don't worry, Ross, this happens to a lot of my customers. It's nothing to
be ashamed of. In fact, I take it as compliment to my skills. Now lie down
and relax."
Ross lay down on his back and placed the towel over his crotch, although it
still tented embarrassingly on his still-hard cock. Phoebe went to her shelf
and retrieved some massage oils, and poured them onto his chest, continuing
the massage. He was more tense than before.
"Ug, Ross, you're worse than before! Well, I can see that you're not gonna
relax as long as you're like this, so I guess I'll have to do something to
help you. Now close your eyes and concentrate on relaxing."
Phoebe put her hand over his eyes and he shut them, trying once more to
relax. She continued the massage, and he continued his fantasy.
She grabbed his cock in her hand and continued to work on the rest of him
with the other hand. She expertly began to jerk him off, using her fingers
for full effect on his shaft and head.
Ross opened his eyes in surprise, but shut them again when she told him to
relax. He pictured Rachel again, her mouth once more around his throbbing
cock. She gobbled his tool up hungrily, sucking and licking him as only she
knew how.
Phoebe switched hands on Ross' cock and began to fondle his balls with her
other hand. Efficiently, she began to massage Ross' cockhead, using the
massage oil as a lubricant. She felt him stiffen and grabbed some kleenex,
anticipating his load.
Rachel slid her tongue up Ross' sack, taking his balls one at a time into her
sweet lips, while stroking his member in her hands. He moaned. She returned
her lips to his penis, licking its eye before she swallowed him. With his
cock buried in her throat, he began to cum, spraying his load into her
He came, and Phoebe held the kleenex over his cockhead, collecting the warm
fluid, then wiped his cock clean when he was done. She tossed the ball of
paper into the trash, then resumed her massage of his body. Ross was
smiling, his eyes closed.
"Now you're really relaxed," she said.
Later that night, Monica crept into the guys' apartment. She opened the door
to Chandler's bedroom and began stripping out of her clothes. She hid them
in his closet, then got into his bed, pulling the covers up to her chin.
A little while later, Chandler walked into his room, tossing his coat on the
bed and dropping his briefcase on the floor. He opened the closet and
started to remove his tie, looking in the mirror hanging on the inside of the
closet door.
He stopped, his mind assessing what he'd seen as he walked in the door.
Monica was in his bed? Chandler spun and looked at her.
"Oh, hi, Mon. What're you doing here?" he asked, confused.
In answer, she threw back his sheets and covers, her naked body exposed to
"I saw how you were looking at me the other day, Bing. I know that you came
back here and beat off in this bed, thinking about me," she said. She cupped
her boobs, squeezing the nipples. "Thinking about my big, firm tits, and how
you'd like to touch them, to fondle them, to make my nipples hard with
pleasure. Wouldn't you?"
"If you're kidding me, Monica, I'm not gonna be able to be your friend
anymore. 'cos that's just cruel."
"Shut up and get over here, Bing."
"Okay, you asked for it," he laughed, jumping into the bed.
Monica ripped his shirt open, sending buttons flying everywhere. He just
shrugged out of it and took her breasts in his hands, squeezing them, his
thumbs rubbing against her erect nipples.
"Ooh, you're good at this," she cooed, her head back and eyes closed in
Chandler responded by enveloping her nipple with his lips and sucking on it
hard. Monica gasped at the force of the suction. He removed his lips from
her nipples and licked it. She felt the saliva evapourate from her nipple as
Chandler blew across it.
"I guess you want me to vote for you, too," he said. "Right, Monica?"
"Forget about that silly contest, boy, just keep doing what you're doing. We
can discuss that all later."
Obliging her, Chandler returned his concentration to her truly gorgeous
globes, licking and sucking Monica to her first orgasm, her pussy juicing
He then lowered his focus, kissing his way down her tight stomach, all the
way down to her wet hole. She smiled at him, then pushed his head into her
snatch, licking her lips in anticipation.
Running his tongue along her lips, Chandler massaged Monica's clit
delicately, teasing the sensitive flesh. She moaned her appreciation, and
put her hands on his head, forcing him closer. Not one to argue, he inserted
his tongue into her honey pot, tasting her sweetness.
"Keep going, honey," she sighed, close to orgasm.
"Your wish..." he trailed off, licking her clit, before once more sticking
his tongue into her. She gasped with pleasure, grinding her hips against his
wonderful tongue, wanting it deeper.
He slowly inserted a finger, then another, testing her readiness. Slowly, he
worked in a third finger, twisting them in unison.
"Oh god, I'm... yeeee!" she screamed, cumming.
When her orgasm had subsided, she ran her hands down Chandler's body,
signalling him to stop by grabbing at his belt. He gratefully removed his
pants, standing before her, his underwear straining to hold his cock inside.
She stood with him, running her hands over his body, down to his ass, where
she squeezed his buttocks, surprising him. He jumped a bit, thrusting his
pelvis into hers, his cotton chained cock pressing against her. Wanting to
get a look at his tool, she reached down and tugged at the waistband, peering
down his underpants.
Chandler's cock was at full hardness, held in place only by the cloth,
pointing straight up at her. She gladly knelt before him, pulling his
underpants down as she went, freeing his member, which she immediately took
in her mouth.
"Mmm, I see you've done this before," he joked, her expert lips and tongue
caressing his cockhead.
"Once or twice," she laughed, then suddenly deepthroated his entire length,
easily supressing her gag reflex, allowing his tool to enter her throat.
She kept it there a while, using her mouth and tongue while sucking him like
a pro. Chandler grunted, enjoying the experience.
Soon, Monica pulled him out, then began giving him a tremedous blow, sliding
her lips and fingers tightly, up and down the length of his shaft. Chandler
forced himself not to cum, feeling the familiar urge as it built up in his
But Monica was not to be denied. She wanted to taste his seed, and doubled
her efforts, massaging his balls and butt, bringing him to the brink.
Chandler cried out, then shot his load into Monica's hungry mouth. She
swallowed most of it, letting some escape, once more displaying her skill.
The excess she licked from her lips, winking at Chandler.
"Not bad, big boy."
"I try."
He got onto his knees, bringing himself to her level, and took her face in
his hands, kissing her deeply, probing her mouth with his tongue. His hands
began to stray, one tracing a line down to her pussy, where it immediately
found a home, the other cupping her left breast, lifting it, as though
testing its weight.
Monica scraped her nails across his back as he touched her clit, inciting
another orgasm. He brought his tongue down to her bosom, and wrapped his
lips tightly around her nipple, suckling. With her eyes shut in ecstasy, she
kissed his ear, whispering dirty words to him.
"Ooh, yeah. You're hand feels so good in my pussy. Why haven't we done this
before? You wanna tit fuck me, baby? Come on, cum on me. Shoot your load
all over my face. I want to see it."
Her suggestion sounded good to him, so Chandler let her lie down on the
floor. She mashed her bountiful globes together, and blew an air kiss at
him. He stroked his rod, then positioned himself over her chest his cock
once more rock hard.
"Come on, Bing. I'm waiting," she whispered in a husky voice, meant for
maximum sexiness.
Complying, Chandler rubbed his pre-cum between her boobs, lubricating them.
He then began to thrust between her tits, closing his eyes, enjoying the
When he opened them, he watched Monica as she watched him thrusting between
her breasts, jiggling with each stroke. He, too, watched his penis thrusting
his friends' firm titflesh.
Quickly, he built to an orgasm, shooting his load on her chest and neck, then
aiming a few drops onto her face as she squealed her delight. When his load
was spent, he kissed her once, then she grabbed at his cock and licked him
clean, remarking how good he tasted.
Soon as she was done, she went to the closet and grabbed her clothes. Putting
on her jeans and t-shirt (no panties), she turned back to Chandler and kissed
him. She hugged him, then grabbed his limp cock and kissed it, too.
"Thank you," she said. "Now, about your vote..."
Continued in part 4...
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text:
Ye gods, but the NPCs are MEAN when teasing poor Alistair. Funny, but MEAN.
Now, of course, I have to see what gets said with other romances. But then, the others aren't as likely to blush all pretty, either. LOL
Dragon Age - Melodramatic, corsetted mistress of the obscure
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text: ASOS have free, yes FREE next day delivery today (when ordered between 9am and 5pm) so what are you waiting for? Cheer up your mid-week blues and ordering something glam for the weekend, or maybe a new piece for your Autumn collection? Their clearance section is still brimming with bargains so head to www.asos.com before they're all gone!


text: Our Team We founded ManufactOn to help the construction industry leverage the power of prefabrication, with tools that integrate supply chain, prefab production, and planning. We are AEC and technology veterans who are working with construction industry leaders, designing for efficiency, usability and best practices. Let us help you bring the best of manufacturing to construction. Raghi Iyengar Founder & CEO Murali Iyer CTO Dirk Ruiz Dir. Product Design Brad Hartnagle VP Sales & Marketing Steve Burri VP Customer Success Jordan Easterling Dir. Customer Success Danny Small Sr Account Executive Advisors Paul Teicholz Sanjay Sarma Josh Kanner Andy Deschenes OUR INVESTORS


text:
A renewal of spirit seems to take place
While following a path along a meandering brook.
Walking through a forest of beauty and serenity
One's mind is filled with the sights and soft sounds
Of nature's peaceful surroundings.
Time stands still and you are enveloped
Within the outstretched arms of trees as if
Welcoming a friend,
The brook we follow has a sound of its own –
Like a beautiful melody of joy.
For a few moments, one is mesmerized
By the feeling of being one with the
Natural rhythm of a forest so alive.
As the brilliant sun sends its golden rays
Upon the canopy of green above us,
One hears a quiet whisper of this forest –
It is the gentle wind flowing through the trees
And sheltering us as we end our Peaceful Walk.


text:
The Owl:
Name: Heartbeak
Age: Adolescent
Gender: Female
Coloring: Brown and white feathers
Physical size: Small
Current rank: Eli
Heartbeak loves to sing, or call, which is where she got her name. Even though she is not very good at it, that does not stop her. She is a social owl who is very friendly and kind. She can lie, however, so you shouldn't trust her completely.
The Human:
Human information private
E-mail: XXXXX
Back to the member page
Back to the main page.


text:
I graduated from Clarksville High School in Clarksville, TX in 2002. I then attended Texas A&M - Commerce in Commerce, TX, were I received a Bachelor of Arts degree in 2006.
Bachelor of Arts
Certified in Spanish and Music
This is my 5th year teaching in Roxton, and I am currently teaching Music, Band, Spanish I, and Spanish II.
I am the music minister for Paris New Generation church, where I direct the choir and play organ and piano. I also enjoy playing golf, riding motorcycles, and working on old cars.


text:
It has been a year of change in how we police Doncaster – but we now have plenty of reason to be optimistic for the future.
During 2018, we have changed the structures to make sure we are more proactive than we may have been historically.
We have seen the return of a CID department based in Doncaster, rather than operating as a centralised South Yorkshire team.
It means we have local ownership and therefore crime investigation in Doncaster can be more flexibly managed. We've also now created four tasking teams who proactively intervene to manage and reduce the activities of organised crime groups.
It has been successful. We've seen a considerable decrease in the activity of crime groups in Doncaster particularly in Mexborough. The tasking teams completed over 100 warrants against organised crime group members and seized hundreds of thousands of pounds in drugs and cash in 2018 but there is still much to be done and this work will continue in 2019.
The work with our partners has also developed including agencies such as Doncaster Council, St Leger Homes, the NHS and the Probation Service, and they have contributed significantly.
With the reintroduction of the Police neighbourhood teams, the partnership work has been about tackling community problems from different angles. For example when we recently worked in an area of Doncaster, the council teams undertook street cleaning work, and removed graffiti. Trading standards officers completed test purchases at licensed premises for alcohol, whilst the NHS did health screening. Together with the police enforcement activity, this partnership approach ensured we addressed communities issues from a variety of perspectives.
The work we are doing also includes going into school to talk to children about antisocial behaviour, so that they understand some of the issues. It helps them understand that rowdy groups of young people on the street can be intimidated for older people, which they may not realise. We have completed presentations to over 600 school children about the signs of child criminal exploitation, and the dangers of getting involved in gangs
It is about improving an area with a rounded response and as a result we have seen a fall in reports of crime and antisocial behaviour. There has been a 12 per cent fall in the number of incidents reported in 2018 compared to the previous year.
The effects of this partnership approach are most evident in the town centre where there are now only 14 known people begging and sleeping rough in the town centre – it was 67 three months ago.
Taking a more preventive / problem solving approach means demand for the police is starting to reduce.
But we are also set to receive extra resources.
South Yorkshire Police is going to take on an extra 40 officers, and 20 of those are due to come to Doncaster.
Taking this together with the work we have done to reduce demand, I'm confident that we will be able to provide an improving service. We are starting 2019 in a healthier position that we have been for several years.
We have doubled the size of our neighbourhood teams, and 17 of the 20 new posts will be put into the neighbourhood teams. I'm hoping this will mean a more visible police presence, so we can tackle the problems that matter to people and allow us to develop a trusting and engaging relationship with our communities.
The other three new officers will work with the prisons investigations team. We have started a new prison partnership board to work together with other organisations like the Prisons, Probation Service and the Crown Prosecution Service. We will be training prison service personnel in techniques such as crime scheme preservation and taking statements. This will provide great assistance to our staff and significantly increase the capacity we have to deal with the investigations and demand generated by the prisons.
Nine of the top 10 locations for crime have seen a substantial reduction in calls for the police. The only one that has not done is the Doncaster Royal Infirmary, and we are looking to create a bespoke way to address issues there. Discussions are ongoing.
Overall the structures we are developing will mean that the police take a more problem solving and preventative approach to tackling crime and this has started to reduce demand. As our resources are also to increase for the first time in several years, I am optimistic about the future and hope that we can increasingly deliver a good service for the people of Doncaster during the next twelve months..


text:
The DNA (National Anticorruption Directorate) prosecutors have put Victor Ponta, the Romanian PM, under criminal investigation. He is accused of forgery, money-laundering, conflict of interest and tax evasion.
The president of Romania and all political parties (except for his own) asked for his resignation but he refused. People from all over the country are preparing massive protests because of this "unprecedented situation" in the EU; No other EU leader has ever kept such a high political position after being officially accused and prosecuted for serious corruption acts.
The DNA statement also said that an investigation would continue into alleged conflict of interest during Ponta's term as prime minister, for making his closest business associate a minister.
Are these kind of democratic deficiencies a problem for the EU as a whole?
How should the European Commission and the rest of the EU leaders respond when the judicial system and democratic values in a member state are at stake?
Sources: Reuters, The Guardian


text:
Looking to bring down your scores in 2019?
Fed up overshooting greens, or leaving yourself short?
Then, you need a way to accurately measure your distance from the flag for each shot.
In this 2 minute guide we've picked out the best Golf GPSs for 2019. We've got a mix of handheld GPS and watches on our list.
Let's get started!
At A Glance: 2019's Best Golf GPS
Pushed for time? Here's a quick rundown of our picks for the best handheld golf GPS and watches. Scroll down for the full reviews.
Best Handheld Golf GPS Units
1. Golf Buddy Voice 2
Why is it called voice? Because it talks to you that's why. Yes, the Golf Buddy Voice 2 will actually tell you your yardage in either a male or female voice (your choice, we won't judge).
But, if you find that a bit annoying (or your playing partners do), then the good news is that the Voice 2 also has a digital display, giving you the option to turn the feature off. The previous model shipped without a display, so an electronic voice was all you got.
Function wise, it's incredibly easy to use. Just turn it on and it will figure out which course (and hole) you are on and tell you how far you are from the front, centre, and back of the green. You can also pull up the 'dynamic green view' to see the shape of the green on the display.
Anything else?
Yep, you'll get shot distance measurement, and an impressive 14 hour battery life. So one charge should get you through 3-4 rounds.
And if you decide you want to wear it as a watch, you can just attach it to the included wrist band. You can even clip it onto your bag.
The Golf Buddy Voice 2 is a small, lightweight golf GPS that will give you an accurate measurement with no fuss setup.
Our pick for: best cheap golf GPS in 2019.Check Current Price
2. Garmin Approach G10
The Garmin Approach G10 is another handheld golf GPS.
This time it won't speak to you, but it will do everything else the Golf Buddy does (and more).
You'll get distances to the front, middle, and back of the green, a green view mode, plus you'll also be able to see yardages to hazards on each hole. Which if you're like me and have an uncanny nack of finding bunkers is very useful.
The Approach G10 also takes into account dog legs, so you view yardages for layups if you don't fancy trying to shape the ball round the corner.
And you can track your stats (fairways, GIR, putts per round etc) to help improve your game.
Again, it's very portable and can be clipped onto your belt or bag with ease.
Another excellent choice for a handheld golf GPS.
Our pick for: the best golf GPS under $200 in 2019.Check Current Price
Best Golf GPS Watches
3. Garmin Approach S20
The S20 is considered to be Garmin's entry level golf GPS watch. But that doesn't mean to say it doesn't pack in a ton of great features.
This is the latest version of my personal golf watch, the Garmin Approach S2.
You'll get the usual yardage information, plus layup/hazard details. The green elevation display also gives you the option to move the flag around, so you can get a more precise yardage to the hole.
Of course you'll need to know where the flag is to be able to do that. It can be a little dificult to pinpoint if you are over 200 yards out (easy 5 iron right…?).
Any other cool features?
Yep, with the S20 you can actually get specific feedback on your swing. The TruSwing sensor will tell you your swing speed and your tempo, which is useful information to have – particularly if your tempo drifts a little during your round.
This watch doubles as a fitbit replacement, so you can track you steps, activity etc. And you can also keep track of all your scores and stats by connecting to the free Garmin Connect app (wireless sync via bluetooth).
The navigation can be a little tricky to get your head round at first, so read the manual. But once you've got it down, you'll find the S20 pretty straight forward to use.
Overall, an excellent Golf GPS watch.
Our pick for: the best affordable golf watch in 2019.
Check Current Price
4. Callaway GPSy
Callaway is one of golf's biggest brands, so it's no surprise they have a finger in the golf GPS pie.
And the Callaway GPSy is certainly a great choice for your GPS watch.
The first thing you'll notice is it looks the part. In fact, I would say it's probably the most stylish smartwatch at the affordable end of the market.
It's reasonably no frills in terms of functionality. It focuses on accurate stats for yardages (usual front, middle, back of green), and recording information on your round. And it does that very well.
The GPSy comes loaded with 30,000 courses, which is less than some golf GPS wathches on the market, but still likely to include any course you are likely to play.
Battery life in GPS mode is 10-12 hours, so likely to get you through 2 rounds comfortably, and 3 at a push.
Overall, it's good, but we still prefer the Garmin.
Buy it if: you want to rock Callaway on your wrist.
Check Current Price
5. Bushnell NEO Ion
Bushnell make some pretty decent golf accessories at the affordable end of the market. And the NEO Ion has most of the features you'll be looking for in a golf watch.
You'll get stats on yardages, hazard and layup distances, and it also has a built in step counter.
Most impressive is the battery life. You'll get up to 16 hours on GPS mode, which should get you through about 4 rounds between recharges.
If you are on a budget, the Bushnell is about $50 cheaper than the other watches on our list, so is a pretty good option.
Buy it if: you want a great golf GPS watch on a budget.
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6. Garmin Approach S60
At the other end of the price scale we have the Garmin Approach S60. You'll be looking at around $400 for this watch, so what do you get for the extra cash?
Well, first you'll get a full colour display, on which you can view full course maps. The display works particularly well in direct sunlight, where other golf watches can be difficult to view. The maps include hazards, doglegs, layup positions etc.
You'll also get advanced stats on your swing and your round, which you can analyse afterwards on the Garmin Connect app.
The Approach S60 has a slope mode, which means you can get a "plays like" distance, taking into account hills etc. But remember, this functionality is not legal for tournament play, so only use it for practice. Still, a very useful feature to have.
You will of course get the standard yardage stats, with distances to front, middle, and back of green.
Finally, you can sync your S60 with your smartphone to get notifications of incoming calls, texts, emails etc.
All in all, if you have the cash, it's our pick for the best golf smartwatch on the market in 2019. Otherwise, we'd opt for the Garmin Approach S20 which is less than half the price.
Our pick for: the best premium golf watch in 2019.
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Our Pick: 2019's Best Golf GPS
- Accuracy
- Features
- Connectivity
- Value for money
All in all, if you have the cash, it's our pick for the best golf smartwatch on the market in 2019. Otherwise, we'd opt for the Garmin Approach S20 which is less than half the price.
What I Use: The Garmin Approach S2
Let me start by quickly covering the golf GPS I currently use, the Garmin Approach S2.
It's a couple of years old now (and the model has been replaced), but it's still working as well as ever.
Basically all I need to do is click to "Start Round" and the watch will figure out which course I'm on, which hole I'm on, and give me an accurate yardage to the green (front, middle, and back).
Very simple to use.
My house overlooks my local course, and I can actually get a yardage to the green from my back garden (shown in the photo). A 4 iron over my neighbour's shed should do it!
I haven't had any problems locating satelites, and as I'm played on some remote courses in the North of Scotland, it's safe to say that the watch will work pretty much anywhere.
So from personal experience, I can recommend Garmin Approach watches. I've included the latest models in my run down above.
Why Should You Buy A Golf GPS?
When you first start playing golf, your main focus will be getting a good strike on the ball, and advancing it towards the hole. But as you improve, you'll want to start hitting more greens in regulation. And that means selecting the right club for the yardage, and of course, knowing what the actual yardage is.
Most courses will have markers at 200, 150, and 100 yards. But, notwithstanding the fact they can be a little inaccurate, when you're in between (or within 100 yards) you're going to be guessing how far you are from the green. And relying on guessing in golf is never a good idea. When it comes to your swing, aiming, or distance, you want to remove as many variables as possible. An accurate yardage takes care of distance.
Most modern golf GPSs (a horrible plural) will give you a distance to the front, middle, and back of the green. Generally I would recommend going with the club that would take you to the back of the green on a pure strike. Because let's be honest, we don't always hit our max distances. So if you club for the back of the green and hit it a little short, you should still be on.
Something I should point out:
When working out your yardages for each club, I would recommend doing it on the course with your own balls. Many driving ranges use limited flight balls, so there could be a 10-15 yard distance between your range distance and course distance. That's a lot.
And on that note, most golf GPSs and watches will give you an accurate yardage for your shot. Just set the device to the correct mode, hit your shot, then walk to your ball. Very handy.
Should You Buy A Handheld GPS, Or A Golf Watch?
Handheld GPS devices generally offer more information (for example hazards) and perhaps an overhead map of the hole, while golf watches have the advantage of convenience. It's easier to glance at your wrist for a yardage than dig into your bag and pull out your GPS.
Some golf GPS systems also allow you to record (and track) your scores and important stats (like GIR, putts etc).
Personally I use a watch (The Garmin Approach S2, which I've reviewed below), but either is going to provide the main function you need: accurate yardage. It's your call whether you want the extras of a GPS, or the convenience of a golf watch.
So now let's look at the best golf GPSs and watches around this year. We've picked out two GPS systems and 4 watches, and we'll start with the GPSs.
Golf GPS FAQs
Which golf gps watch is best?
In terms of value for money and features, the Garmin Approach s60 is the best golf GPS available in 2019.
How accurate are golf gps watches?
Most golf GPS watches are accurate to between 3 and 4 yards under optimal conditions.
Which golf gps is most accurate?
We found the Garmin Approach s60 to be the most accurate golf GPS available in 2019.
Are golf gps devices legal?
Golf GPS systems are legal for tournament play, however, if the GPS has a built in slope mode (which takes into account hills etc) this must be disabled.
golf gps vs rangefinder
Rangefinders are normally slightly more accurate than golf GPS systems. They also have the benefit of being able to target obstacles on the course (such as bunkers or streams) and get an accurate yardage to the hazard.
What's Next?
Thanks for reading our guide to the best golf GPSs and watches in 2019. Check out our guide to the best golf shoes for the 2019 season here.
And for more golf tips and gear guides, check out our golf section.


text: After seeing the standing room only crowd at the Secret Society meeting at Turner Ignite in New York this past week, one can see that the secret is finally out. Mitch Oscar, USIM Advanced Television Strategist, created the Secret Society "to talk about issues that affect the Advanced TV space," and this particular session was filled with speakers on the subject. The primary takeaways follow below.


text:
Che bella giornataSome days are ordinary, some are bad, but today is a special day, a red letter day, a crazily fantastic day, un giorno magnifico! Let us count the ways.
It's Fabio's birthday! Though he would never have let on if we hadn't used some of our sneaky NY spy action to find out...
Last night I had a glass of wine with an ex-pat German who asked me, "But how do you speak Italian so well?" First I laughed and laughed because of how far there is still to go, but then I just was grateful to Fabio, because his friendship includes, as a very small part, fifteen or so minutes of endlessly patient conversation every day at pickup time. I (F) wanted to get him a copy of Mangia, Prega, Ama, but instead we found him a rice paper photo album and a candle with some dancing Indian gods on the sides.
Count number two--K got another gym job! This time at one of those fancy palestre where the reception staff all smoke and guzzle shots of coffee from the vending machine, while the widescreen TVs cycle through the various cameras tucked in every nook and cranny of the place.
After calling the risponsabile several times, we decided to just show up, which worked in one of those inexplainable ways that things sometimes require in Italy. Other than a little illustrative "stretching" casually outside the office window, all it took was dropping the words "New York," "hip-hop," and "Shakira," and a quick viewing of a Tracy Anderson on her iPhone and K was hired on the spot and her class will have a launch party in a few weeks--ideally with a velvet rope. Or at least using the NYC-style turnstyle that keeps out the riff-raff.
Then of course we had a terremoto / earthquake that shook the region and more pertinently canceled school tomorrow when T was going to have two verifica / tests. So T celebrated by making perhaps the best French onion soup I've ever had complete with slices of baguette and a melted mound of Pecarino on top.
Plus also finally I have passed my teoria exam for the driver's license. After ten months of study and two prior frustrating failures, it came down to the last 22 seconds where at the last moment I correctly went with the gut feeling that a sign was consigliata rather than obbligatoria. Now I get to pick up my foglio rosa / pink paper (learner's permit) next Tuesday and then after a month I can give the practical exam a try, but it's probably a given that I shouldn't count on thirty-one years of driving experience alone to pass!


text:
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Review ARTICLE
Localization and function of GABA transporters GAT-1 and GAT-3 in the basal ganglia
- Division of Neuroscience, Yerkes National Primate Research Center and Department of Neurology, Emory University, Atlanta, GA, USA
GABA transporter type 1 and 3 (GAT-1 and GAT-3, respectively) are the two main subtypes of GATs responsible for the regulation of extracellular GABA levels in the central nervous system. These transporters are widely expressed in neuronal (mainly GAT-1) and glial (mainly GAT-3) elements throughout the brain, but most data obtained so far relate to their role in the regulation of GABAA receptor-mediated postsynaptic tonic and phasic inhibition in the hippocampus, cerebral cortex and cerebellum. Taking into consideration the key role of GABAergic transmission within basal ganglia networks, and the importance for these systems to be properly balanced to mediate normal basal ganglia function, we analyzed in detail the localization and function of GAT-1 and GAT-3 in the globus pallidus of normal and Parkinsonian animals, in order to further understand the substrate and possible mechanisms by which GABA transporters may regulate basal ganglia outflow, and may become relevant targets for new therapeutic approaches for the treatment of basal ganglia-related disorders. In this review, we describe the general features of GATs in the basal ganglia, and give a detailed account of recent evidence that GAT-1 and GAT-3 regulation can have a major impact on the firing rate and pattern of basal ganglia neurons through pre- and post-synaptic GABAA- and GABAB-receptor-mediated effects.
GABA is the main neurotransmitter used in the basal ganglia network, and abnormal transmission at specific GABAergic synapses underlies some of the pathophysiological features of various basal ganglia diseases. A tight regulation of GABA homeostasis is essential to mediate normal basal ganglia functions. In this manuscript, we will provide a brief overview of the main characteristics of the different subtypes of GABA transporters in the mammalian CNS, and then discuss some of our recent findings and those from other laboratories about the localization and functions of GABA transporters (GATs) in the basal ganglia. This review does not intend to cover the extensive literature on GATs, but will specifically focus on the distribution and regulatory mechanisms by which these transporters modulate neuronal activity and synaptic transmission in the basal ganglia. Because of the limited amount of data available, this review does not aim at generating integrative concepts about GATs function in the basal ganglia. It is rather focused on the presentation of recent findings that have been gathered about these transporters in specific basal ganglia nuclei, and their potential importance for basal ganglia function and dysfunction. For a more comprehensive account of our current knowledge of GAT function in other brain regions, readers are referred to previous reviews (Borden, 1996; Gadea and Lopez-Colome, 2001; Dalby, 2003; Conti et al., 2004).
General Features of GABA Transporters
GABA is the main inhibitory neurotransmitter in the mammalian brain. After release from presynaptic terminals, GABA is rapidly removed from the extracellular space by GATs, a regulatory mechanism that terminates inhibitory synaptic transmission (Borden, 1996; Richerson and Wu, 2003), regulates GABA spillover to neighboring synapses (Borden, 1996; Overstreet and Westbrook, 2003), and maintains GABA homeostasis to prevent excessive tonic activation of synaptic and extrasynaptic GABA receptors (Borden, 1996; Semyanov et al., 2004). In addition, the reversal of the GATs function may result in additional GABA release under certain pathological and physiological conditions (Allen et al., 2004; Wu et al., 2007).
GATs Cloning and Pharmacology
To date, four different GATs have been described, GAT-1, GAT-2, GAT-3, and the Betain/GABA transporter type 1 (BGT-1). These transporters are members of a large family of 12-transmembrane spanning Na+/Cl− coupled transporters (for review, see Borden, 1996). GAT-1 was the first GAT to be cloned (Guastella et al., 1990). The GAT-1 protein sequence in rat (Guastella et al., 1990), mouse (Liu et al., 1993), and human (Nelson et al., 1990) displays a high degree of homology and nearly identical pharmacological properties (Borden, 1996). The GAT-2 and GAT-3, cloned by Borden et al. (1992), display a higher degree of amino acid identity between each other (67% identity), and with the fourth GABA transporter, BGT-1 (68 and 65% identity for GAT-2 and GAT-3, respectively) than with GAT-1 (∼52% amino acid identity). The amino acid sequence of GAT-3 in human, rat, and mouse is virtually identical with only a few substitutions (for review, see Borden, 1996). In contrast to other transporters, BGT-1, cloned by Yamauchi et al. (1992), utilizes both GABA and betaine as substrates.
GATs exchange GABA for Na+ and Cl−. The GABA-transporting function of GATs is particularly dependent on the Na+ gradient across the membrane. Although Cl− can significantly enhance the rate of transport, Cl− alone does not drive GABA uptake in the absence of Na+. The proposed stoichiometry for GAT-1, GAT-2, and GAT-3 is 2 Na+:1 Cl−:1 GABA (Loo et al., 2000; Sacher et al., 2002; Karakossian et al., 2005).
GAT-1 can be pharmacologically isolated from GAT-2, GAT-3, and BGT-1. Various drugs have been identified as highly specific GAT-1 inhibitors (for instance, Cl966, SKF 89976A, NO-711, and Tiagabine), while SNAP 5114 is a semiselective blocker of GAT-2 and GAT-3, with a higher affinity for GAT-3 than GAT-2 (IC50 ∼5 and 20 μM, respectively). However, because GAT-3 is far more abundant in neurons and glia than GAT-2, SNAP 5114 is commonly used as a GAT-3 blocker in studies of GATs regulation of synaptic transmission in the central nervous system. Microdialysis experiments in rodent hippocampus and thalamus have shown that either local or systemic application of GAT-1 antagonists can increase extracellular GABA concentrations by up to 1.5- to 4-folds the basal levels (Richards and Bowery, 1996; Dalby, 2000). Similarly, application of the GAT-2/GAT-3 blocker, SNAP 5114 (100 μM), increases GABA levels in the thalamus by almost 250%, but has no significant effect on hippocampal GABA concentration (Dalby, 2000).
GATs Localization in the CNS
The cellular localization of GABA transporters has been studied in the rat brain using both in situ hybridization for mRNA (Rattray and Priestley, 1993; Brecha and Weigmann, 1994; Augood et al., 1995; Durkin et al., 1995; Jursky and Nelson, 1996; Nishimura et al., 1997; Yasumi et al., 1997; Ficková et al., 1999) and immunocytochemistry for transporters protein (Ikegaki et al., 1994; Augood et al., 1995; Minelli et al., 1995; Itouji et al., 1996; Ribak et al., 1996; Conti et al., 1998). The GAT-1 mRNA is expressed throughout the brain, but particularly enriched in the olfactory bulb, basal ganglia, interpeduncular nucleus, cerebellum, and retina (Augood et al., 1995; Durkin et al., 1995; Yasumi et al., 1997). Immunohistochemical studies using antibodies raised against recombinant proteins have shown that GAT-1 is not only expressed in GABAergic neurons, but also in non-GABAergic cells and glia in certain brain regions (for review, see Eulenburg and Gomeza, 2010), although their function in these neurons remains poorly understood.
GAT-2 mRNA is weakly expressed throughout the brain, primarily in arachnoid and ependymal cells, and to a much lesser extent, in neurons and astrocytes (Durkin et al., 1995; Conti et al., 1999). GAT-3 mRNA and protein are found predominantly in glial cells (Radian et al., 1990; Ikegaki et al., 1994; Durkin et al., 1995). The strongest GAT-3 expression is found in the glomerular layer of the olfactory bulb, the inner nucleus of the retina, the thalamic paraventricular nucleus, and the globus pallidus (GP; Clark et al., 1992; Ikegaki et al., 1994; Durkin et al., 1995; Minelli et al., 1996). Some of these studies showed that GAT-3 is nearly absent from the neocortex and cerebellar cortex, and very weakly expressed in the hippocampus (Clark et al., 1992; Brecha and Weigmann, 1994; Ikegaki et al., 1994; Durkin et al., 1995), while others provided evidence for significant neocortical expression in rodents (Minelli et al., 1996, 2003; Pow et al., 2005). Finally, low to moderate levels of BGT-1 are expressed in most brain regions (Durkin et al., 1995; Zhou and Ong, 2004).
GATs Regulation of Synaptic Transmission and Plasticity
The effects of GAT-1 modulation on synaptic transmission have been most studied in the CNS. A summary of the main effects of GAT blockade on GABA release and postsynaptic currents in various CNS regions is shown in Table 1. GAT-1 inhibitors increase the decay of evoked IPSCs, while not having significant effects on IPSC amplitude in many brain regions (Roepstorff and Lambert, 1992; Thompson and Gähwiler, 1992; Engel et al., 1998; Overstreet and Westbrook, 2003). GAT-1 inhibitors also increase GABAA receptor-mediated tonic conductances in cerebellar granule cells (Rossi et al., 2003) as well as in granule cells and pyramidal neurons of the hippocampal dentate gyrus (Nusser and Mody, 2002; Semyanov et al., 2003; Sipilä et al., 2007). A recent study also demonstrated that GAT-1 blockade or genetic deletion of GAT-1 specifically impairs long-term potentiation (LTP) induced by theta burst stimulation (Gong et al., 2009) in the CA1 region of mouse hippocampus. While there is compelling evidence that GAT-1 regulates GABAergic transmission in the hippocampus (Thompson and Gähwiler, 1992; Isaacson et al., 1993; Draguhn and Heinemann, 1996; Engel et al., 1998; Nusser and Mody, 2002; Overstreet and Westbrook, 2003; Semyanov et al., 2003), cerebral cortex (Keros and Hablitz, 2005; Bragina et al., 2008; Gonzalez-Burgos et al., 2009), and cerebellum (Rossi et al., 2003), much less is known about the functional role of GAT-1 in the basal ganglia (Rossi et al., 2003; Galvan et al., 2005; Kinney, 2005; Kirmse et al., 2009). Despite its widespread and abundant expression in many brain regions (see Borden, 1996; Eulenburg and Gomeza, 2010 for reviews), the role of GAT-3-mediated regulation of GABAergic transmission remains poorly understood compared with GAT-1 functions in most CNS regions, except for the cerebral cortex and some basal ganglia nuclei (Table 1).
Table 1. Summary the effects of GABA transporters blockade on extracellular GABA levels and postsynaptic currents in the CNS.
GAT-1 and GAT-3 in the Basal Ganglia
The relative importance of GAT-1 and GAT-3 in the normal and pathological functioning of the basal ganglia, and the possibility that their regulation could be used to achieve beneficial therapeutic responses in basal ganglia disorders remain largely unexplored. In the following sections, we describe the current knowledge of the localization and function of GAT-1 and GAT-3 in various basal ganglia nuclei, and critically discuss their potential relevance as targets for drug therapies of basal ganglia disorders, such as Parkinson's disease.
GAT-1 and GAT-3 Localization
Most neurons in the striatum, including medium spiny projection neurons and several interneuron subtypes are GABAergic (Kawaguchi et al., 1990). The maintenance of homeostasis in extracellular levels of GABA and GABAergic transmission is, therefore, critical for normal striatal functions. Although the chemical phenotype of most striatal GAT-1-positive cells remains to be determined, it is clear that a significant proportion of the GABAergic neurons in the striatum (including medium spiny neurons and a large proportion of parvalbumin-positive interneurons), express mRNA and immunoreactivity for GAT-1 (Augood et al., 1995; Durkin et al., 1995; Yasumi et al., 1997; Wang and Ong, 1999).
In contrast, the evidence for striatal GAT-3 expression remains controversial. Some of the existing in situ hybridization studies have reported negative data (Clark et al., 1992; Durkin et al., 1995), while others using similar methods or polymerase chain reaction (rt-PCR) techniques have demonstrated a significant level of striatal GAT-3 mRNA expression in the rat caudate–putamen (Yasumi et al., 1997; Ficková et al., 1999). GAT-3 immunoreactivity was also demonstrated in the monkey striatum (Ng et al., 2000). The cellular and chemical phenotypes of GAT-3-positive striatal elements remain poorly characterized in both primates and non-primates.
Functional Role of Striatal GAT-1
Systemic or local application of the selective GAT-1 inhibitor, SKF 89976A, doubles the extracellular concentration of GABA in the rat striatum (Waldmeier et al., 1992). Consistent with this finding, GAT-1 blockade induces GABAA receptor-mediated tonic inhibition of striatal neurons (Kirmse et al., 2008), similar to previous reports in the hippocampus (Jensen et al., 2003; Semyanov et al., 2003; Scimemi et al., 2005) and cerebellar Purkinje cells (Chiu et al., 2005). Electrophysiologic brain slice recording studies have demonstrated that bath application of the GAT-1 inhibitors NO-711 prolongs the decay time of IPSCs evoked locally in striatum (Kirmse et al., 2008) and decreases the amplitude of eIPSCs produced by intrastriatal stimulation. The latter effect is most likely caused by a presynaptic mechanism because it was associated with a significant increase of the paired-pulse facilitation ratio (PPR; Figures 1A,B). Interestingly, coapplication of NO-711 and the GABAB-receptor antagonist CGP55845 only partly restored the GAT-1 blockade-mediated effects on the amplitude of eIPSCs but reduced the PPR to control levels (Figures 1C–E), suggesting that the effects of GAT-1 blockade upon eIPSCs are partially, but not fully mediated by GABAB receptor-dependent mechanisms (Kirmse et al., 2008). This observation was recently extended to the hippocampus (Safiulina and Cherubini, 2009; Lindsly and Fraxier, 2010). Other mechanisms, including postsynaptic shunting and GABAA-receptor desensitization due to a persistent activation of GABAA receptors by high ambient GABA concentration in the presence of NO-711 could also contribute to these effects (Overstreet et al., 2000; Keros and Hablitz, 2005; Kirmse et al., 2008).
Figure 1. Tonic activation of presynaptic GABABRs induced by GAT-1 blockade in rat brain slices. (A) eIPSCs in response to paired-pulse stimulation (ISI 250 ms) in control solution and in the presence of the GAT-1 blocker, NO-711 (10 μm), in slices of P12-14 rats. Traces represent averages of 20 responses. (B) Quantification of results for NO-711 effects on the mean eIPSC amplitude, PPR (PPR50: ISI 50 ms, PPR250: ISI 250 ms), rise time (20–80%) and decay (T0.10) of striatal eIPSC. Data from 17 striatal output neurons (SONs), except PPR50. (C) Evoked IPSCs in response to paired-pulse stimulation (ISI 250 ms) in control solution and in the presence of either NO-711 or NO-711 plus the GABAB-receptor antagonist CGP55845 (1 μm). Traces represent averages of 20 responses. (D,E) CGP55845 partially restored the NO-711-induced reduction in eIPSC amplitude and completely reversed the NO-711-induced increase in PPR (ISI 250 ms). ns – not significant, *P < 0.05, **P < 0.01, ***P < 0.001. Reprinted from Kirmse et al. (2008) with permission from John Wiley and Sons.
GATs, acting in reverse direction, also contribute to the action potential-independent release of GABA in the rat striatum (Bernath and Zigmond, 1989; Del Arco et al., 1998; Schoffelmeer et al., 2000). For instance, microdialysis studies have shown that nipecotic acid, a non-selective GAT blocker, attenuates the amphetamine-induced increase in extracellular concentration of GABA in the striatum of freely moving rats, using a calcium-free microdialysis medium (Del Arco et al., 1998). Nipecotic acid also attenuates glutamate NMDA and dopamine D1-like receptor-mediated [3H]-GABA release from striatal slice and cultured striatal neurons in the presence of the sodium channel blocker tetrodotoxin (Schoffelmeer et al., 2000). These results demonstrate that the release of GABA induced by amphetamine or activation of D1-like or NMDA receptors involves a GAT mechanism. This reversal of GAT function may play a role in the behaviorally activating effects of psychostimulant drugs (Schoffelmeer et al., 2000).
Functional Role of Striatal GAT-3
Although the anatomical evidence for striatal GAT-3 expression is controversial compared with GAT-1 (Clark et al., 1992; Ficková et al., 1999; Ng et al., 2000), there is functional evidence for an active role of GAT-3 in the rat striatum (Kirmse et al., 2009). Despite the fact that the application of SNAP 5114 alone, a semiselective GAT-3 inhibitor, has no effect on striatal GABAergic transmission, and does not affect GABAA receptor-mediated tonic currents in striatal projection neurons (Kirmse et al., 2009), the coapplication of SNAP 5114 with the GAT-1 inhibitor NO-711 (10 μM) was shown to reduce the frequency of miniature IPSCs (mIPSCs) without affecting their amplitude or kinetic in striatal neurons (Figures 2A–C). As shown for GAT-1, GAT-3 blockade reduces the mean amplitude of eIPSCs through GABAB receptor-mediated presynaptic effects in the rat striatum (Kirmse et al., 2009). As the functional effects of GAT-3 blockade on GABAergic transmission may be masked by concomitant GAT-1-mediated effects, it is possible that GAT-3-mediated GABA uptake comes into play only under specific physiological or pathological conditions that result in increases in neuronal activity and GABA release beyond normal levels in the striatum (Kirmse et al., 2009). In other brain regions, it has been shown that that GAT-1 and GAT-3 are largely segregated in pre-terminal neuronal elements and glia, respectively, thus, it is possible that these transporters might play complementary and synergistic roles towards the regulation of GABAergic transmission in the striatum.
Figure 2. Functional expression of GAT-2/3 is unmasked by suppression of GAT-1 activity in the rat striatum. (A) Sample traces showing mIPSCs in the presence of NO-711 (10 μM) and NO-711 plus SNAP 5114 (40 μM) in striatal slices of P12–14 rats. (B) Cumulative mIPSCs amplitude distribution was not significantly affected by SNAP 5114 (P > 0.9, Kolmogorov–Smirnov test). Plot represents data from the cell shown in (A). (C) SNAP 5114 – selectively decreased the frequency of mIPSCs. *P < 0.05. Reprinted from Kirmse et al. (2009) with permission from John Wiley and Sons.
Globus Pallidus
GAT-1 and GAT-3 Localization
The rat GP expresses strong mRNA for both GAT-1 and GAT-3 (Durkin et al., 1995; Yasumi et al., 1997). Consistent with these findings, our studies have demonstrated strong GAT-1 and GAT-3 immunoreactivity in the rat and monkey GP (Galvan et al., 2005, 2010; Jin et al., 2009, 2011; Figure 3). At the electron microscopic level, GAT-1 is largely expressed in small unmyelinated axons in the rat GP (Figure 3), and in both unmyelinated axons and glial processes in the external and internal segments of the GP (GPe and GPi, respectively) in the monkey (Figure 3). The pattern of GAT-3 immunoreactivity in the rat and monkey pallidum is strikingly different from that of GAT-1, being almost exclusively expressed in glial cell processes which, in some cases (Figure 3) are closely apposed to putative GABAergic terminals forming symmetric synapses or wrapped around axo-dendritic complexes consisting of numerous unlabeled terminals and dendrites of pallidal neurons (Figure 3). Despite significant alterations in GAT function, there is no significant change in the general localization pattern of GAT-1 and GAT-3 in the GPe and GPi of MPTP-treated Parkinsonian monkeys (Figure 3 and below; Galvan et al., 2010).
Figure 3. Light and electron micrographs of GAT-1 and GAT-3-immunoreactive elements in the rat and monkey globus pallidus. (A,B) Light micrographs of GAT-1 and GAT-3 labeling in the rat striatum (STR) and globus pallidus (GP). (C) GAT-1-immunoreactive axon terminal (Te) and astrocytic process (AS) in the rat GP. Note that the labeled terminal forms a symmetric synapse (arrowhead) with an unlabeled dendrite. (D) GAT-3-positive astrocytic processes in close contact with unlabeled terminals and dendrites in the rat GP. (E) Quantification of the percentage of GAT-1- and GAT-3-labeled elements in the rat GP. Note that GAT-1 is predominantly found in unmyelinated axons, whereas GAT-3 is mainly expressed in glial processes. A total of three rats were used in these studies (see Jin et al., 2009, 2011 for details). (F,G) GAT-1 and GAT-3 immunoreactivity in the monkey putamen (Put) and globus pallidus (external and internal segments, GPe, GPi). (H) GAT-1-immunoreactive unmyelinated axons (AX) and astrocytic processes (AS) in the monkey GPe. Note that the labeled AS is in close contact with an unlabeled terminal (u.TE) that forms an axo-dendritic symmetric synapse [arrowhead; (I,J)] GAT-3-positive astrocytes in the monkey GPe (I) and GPi (J). Note the close association between the immunoreactive AS processes and unlabeled axon terminals forming symmetric synapses [arrowheads in (I)]. (K) Quantitative distribution of GAT-1 and GAT-3 across different neuronal and glial elements in the monkey GPe and GPi (see Galvan et al., 2005, 2010 for more details). Scale bars: (A,B,F,G): 1 mm; (C,D,H,I,J): 1 μm.
Functional Role of GAT-1 in the Globus Pallidus
In normal monkeys, local intrapallidal administration of the GAT-1 antagonist (SKF 89976A) significantly increases the ambient GABA level in GPe, as measured by microdialysis (Figure 4A), and reduces the firing rate of GPe and GPi neurons (Figures 4B1,B2; Galvan et al., 2005). We found that the inhibitory effects of GATs blockade on GPi firing are strongly decreased in monkeys rendered Parkinsonians by systemic treatment with the dopaminergic neurotoxin 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine (MPTP), while the effects on GPe discharge rates remain unaffected (Galvan et al., 2010). In line with our monkey data, systemic administration of tiagabine, a GAT-1 antagonist, increases extracellular levels of GABA by up to threefold in the rat GP (Fink-Jensen et al., 1992). Furthermore, bath application of tiagabine has several significant effects when used in rat brain slice recording experiments: (1) GAT-1 inhibition prolongs the decay time of IPSCs evoked by striatal stimulation, without affecting their amplitude (Figures 5A,B). Similar observations have been made in many brain regions, suggesting that it represents a general GAT-1 function in the CNS (Roepstorff and Lambert, 1992; Thompson and Gähwiler, 1992; Engel et al., 1998; Overstreet and Westbrook, 2003; for reviews, see Borden, 1996; Richerson and Wu, 2003). (2) GAT-1 blockade induces GABAA receptor-mediated tonic currents in rat GP neurons (Jin et al., 2011), another general role reminiscent of GAT-1-mediated effects in other brain regions (for reviews, see Borden, 1996; Richerson and Wu, 2003; Eulenburg and Gomeza, 2010). (3) The effects of GAT-1 blockade on spontaneous and miniature IPSCs in rat GP are controversial. On one hand, application of tiagabine prolongs the decay kinetics and reduces the frequency of spontaneous and miniature IPSCs, in part through activation of presynaptic GABAB autoreceptors (Chen and Yung, 2003). However, we found that the frequency and amplitude of spontaneous, but not miniature, IPSCs is increased following GAT-1 blockade (Jin et al., 2011). The sources of the discrepancy between these different sets of data remain to be established. (4) GAT-1 blockade reduces the frequency, but not the amplitude of mEPSCs (Figures 5E,G), most likely through GABAB receptor-mediated presynaptic inhibitory effects upon glutamatergic transmission (Jin and Smith, 2009). Thus, together with data from the striatum, these findings indicate that GAT-1 regulates striatal and pallidal activity through modulation of persistent synaptic GABAA receptor-mediated currents (IPSCs), extrasynaptic GABAA receptor-mediated tonic currents, and presynaptic GABAB receptors in both GABAergic and glutamatergic terminals (Galvan et al., 2006).
Figure 4. Effects of SKF 89976A and SNAP 5114 on GABA levels in GPe and pallidal discharge rate in monkeys. (A) Dialyzate samples were collected every 10 min. SKF 89976A or SNAP 5114 were administered during sample 4 (horizontal bar). Data area means ± SD from three experiments for each treatment in a single monkey. Difference from ACSF experiments (Mann–Whitney UU test): *P < 0.05. (B1) Example of effect of SKF 89976A on discharge rate of a GPe cell. (B2) Discharge rate of GPe and GPi cells during baseline period and at the point of maximal effect of SKF 89976A. (C1) The discharge rate of this GPe cell is inhibited after administration of the GAT-3 blocker, SNAP 5114. (C2) The discharge rate of GPe and GPi cells during baseline period and at the point of maximal effect of SNAP 5114 injection. In (B1,C1), horizontal bars indicate duration of drug infusions. Dashed lines represent mean discharge rate ± 2 SD. For more details see Galvan et al. (2005).
Figure 5. Effects of GAT-1 and GAT-3 blockade on GABAergic and glutamatergic synaptic transmission in the rat GP. (A) Application of SKF 89976A increases the decay time, but not the amplitude of IPSCs evoked in GP neurons after striatal stimulation. (B) Bar graph showing that SKF 89976A increases the decay time, but has no effect on the amplitude and base line holding currents of eIPSCs expressed as percent of control ± SEM (*P < 0.005). (C) Application of SNAP 5114 increases the amplitude and decay time of IPSCs evoked in GP neuron by striatal stimulation. (D) Bar graph summarizing the effects of SNAP 5114 on eIPSCs amplitude, decay time, and holding current expressed as percent of control ± SEM (*P < 0.005). (E,F). Sample traces showing mEPSCs recorded in control condition and during SKF 89976A or SNAP 5114 application. (G,H) The summary bar graphs show that SKF 89976A or SNAP 5114 significantly reduce the frequency, but not amplitude of mEPSCs. * P < 0.01. For more details see Jin et al. (2009, 2011).
The behavioral effects of GAT-1 blockade in the GP are poorly characterized. Apart from evidence that the unilateral administration of tiagabine in the rat GP induces ipsilateral rotations in rats (Chen and Yung, 2003), very little is known about the influence of GATs on behavior. Given the prominent role of altered GABAergic transmission in parkinsonism (Galvan and Wichmann, 2007), it would be particularly interesting to examine the potential antiparkinsonian effects of GATs. Taking into consideration the physiological effects of GAT-1 blockade on pallidal activity and the proposed pathophysiology of basal ganglia networks in parkinsonism (DeLong, 1990), one could predict that GAT-1 blockade in GPe could induce or exacerbate parkinsonism, due to the increased GABAergic transmission along the indirect pathway, while GAT-1 blockade in GPi could alleviate parkinsonism through increased inhibition of the overactive basal ganglia pallidal outflow to the thalamus and brainstem (DeLong, 1990). However, because we found that the effects of GAT-1 blockers in GPi of Parkinsonian monkeys are altered from normal (Galvan et al., 2010), these speculations are, at best, incomplete with the available data.
Functional Role of GAT-3 in the Globus Pallidus
The strong expression of GAT-3 in the pallidum of monkeys and rats suggests that this transporter may play an important role in the clearance of extracellular GABA. Supporting this contention, local in vivo blockade of GAT-3 increases pallidal GABA levels (Figure 4A), and inhibits the firing rate of pallidal neurons in monkeys (Figures 4C1,C2). In rat brain slice recording studies, GAT-3 blockade increases the decay and amplitude of evoked IPSCs after striatal stimulation (Figures 5C,D), increases the frequency and amplitude of spontaneous IPSCs, and induces GABAA receptor-mediated tonic currents in GP neurons (Jin et al., 2011). The mechanisms by which GAT-3, but not GAT-1, blockade increases the amplitude of evoked IPSCs are unknown. One hypothesis put forward in our recent study (Jin et al., 2011) relates to the fact that GAT-3 blockade may result in the activation of a large pool of striatal GABAergic projections neurons and interneurons which, under specific conditions, can depolarize other projection neurons, thereby increase GABA release in the GP (Bracci and Panzeri, 2006; Ade et al., 2008).
GAT-3 blockade also reduces glutamatergic transmission, most likely from the subthalamic nucleus, through presynaptic GABAB-receptor activation (Figures 5F,H). In contrast with the striatum where the effects of GAT-3 blockade alone are fully masked by GAT-1 (Kirmse et al., 2009), the blockade of GAT-3, alone or in the presence of a GAT-1 blocker, produces significant effects on both GABAergic and glutamatergic transmission in the rat GP (Jin et al., 2009, 2011). These differential effects are consistent with localization studies showing that GAT-3 expression in the striatum is comparatively light, compared with the very strong glial expression in the GP (Figure 3; Clark et al., 1992; Durkin et al., 1995; Yasumi et al., 1997; Ng et al., 2000; Galvan et al., 2005; Jin et al., 2009, 2011). GAT-1 and GAT-3, thus, represent differential target sites through which GABA reuptake may subserve a complementary regulation of GABAergic and glutamatergic transmission in the pallidum.
Subthalamic Nucleus and Substantia Nigra
Despite their distinct glutamatergic phenotype, rat and human STN neurons, display intense GAT-1 mRNA expression (Yasumi et al., 1997; Augood et al., 1999). There is also evidence from other brain regions such as cerebral cortex, inferior colliculus, and the deep cerebellar nuclei, that the number of GAT-1 mRNA-containing cells is much larger than that of GABAergic (GAD67 mRNA-) positive cells (Swan et al., 1994; Yasumi et al., 1997), suggesting that GAT-1 expression extends beyond GABAergic neurons in these brain regions (Yasumi et al., 1997). The functional significance of GATs in non-GABAergic neurons remains unknown (Augood et al., 1999). However, it is interesting to note that an ongoing gene transfection clinical trial in PD aims at changing the phenotype of subthalamic neurons into GABAergic cells in order to reduce the overactive glutamatergic outflow from the subthalamic nucleus (Lewitt et al., 2011). The expression of GAT-1 into subthalamic terminals may increase the likelihood of success of this approach. As in other basal ganglia nuclei, GAT-3 immunoreactivity is expressed in astrocytic processes throughout the monkey STN (Ng et al., 2000).
The effects of GAT activity on the neuronal activity in the STN are unknown. Because of the preponderance of GABAergic pallidal terminals in the STN, and because of the fact that extrasynaptic GABA-B receptors appear to play important roles in the modulation of burst firing and the pallidosubthalamic "pacemaker" system (Plenz and Kitai, 1999; Bevan et al., 2002), a better understanding of the mechanisms by which GATs modulate GABAergic STN activity is clearly warranted (for review, see Bevan et al., 2007).
Both GAT-1 and GAT-3 mRNAs are moderately expressed in the rat substantia nigra as a whole. Surprisingly, GAT-1 expression is stronger in dopaminergic pars compacta (SNc) neurons than in the GABAergic pars reticulata (SNr) cells (Durkin et al., 1995; Yasumi et al., 1997), serving as another possible example of GAT-1-mediated function in non-GABAergic neurons. GAT-3 immunoreactivity is also strongly expressed in the monkey SN, where it appears to be preferentially associated with astrocytes (Ng et al., 2000). GAT-1 inhibition significantly reduces [3H] GABA uptake in synaptosomes prepared from the rat SNr (Bahena-Trujillo and Arias-Montano, 1999), and significantly increases extracellular GABA levels in the rat substantia nigra (Fink-Jensen et al., 1992). However, the overall regulatory functions of GATs on synaptic transmission in the SN remain unexplored.
GATs and Parkinson's Disease
The data discussed in this review highlight the fact that GAT-1 and GAT-3 represent different target sites through which GABA reuptake may regulate GABAergic and glutamatergic transmission in the basal ganglia. Although still poorly understood, recent in vitro and in vivo data from our laboratory and others have demonstrated that GAT-1 and GAT-3 are strongly expressed in the GP, and that their blockade significantly impacts the activity of pallidal neurons under normal and Parkinsonian conditions, by increasing GABA levels and subsequent overactivation of GABAA- and GABAB-receptors (Galvan et al., 2005, 2010; Jin et al., 2009, 2011).
Our recent data from primate experiments demonstrate that the impact of GAT-1 and GAT-3 blockade upon neuronal activity is reduced in the GPi of Parkinsonian animals compared with the normal state, despite the fact that the overall pattern of GAT distribution does not change (Galvan et al., 2010). The possible use of drugs that regulate GAT function to treat patients with Parkinson's disease remains speculative. The essential need of tightly regulated GABA homeostasis for normal brain functions reduces the likelihood that systemic administration of GAT blockers could be done without the risk of significant side effects, although such an approach has shown some therapeutic benefits in patients with epilepsy and anxiety (Gadea and Lopez-Colome, 2001; Dalby, 2003; Sarup et al., 2003; Conti et al., 2004; Schwartz and Nihalani, 2006).
Recent evidence suggests that the interaction of GATs with other receptor systems may provide an alternative strategy for the development of Parkinson's disease therapy. For instance, a recent study has demonstrated that adenosine inhibits GAT-1-mediated GABA uptake in the rat GP (Gonzalez et al., 2006). This finding, combined with the fact that A2A-receptor antagonists have significant antiparkinsonian effects (Kanda et al., 2000; Chase et al., 2003), raise the possibility that the antiparkinsonian effects of A2A-receptor antagonists are partly due to presynaptic modulation of GABA release at striatopallidal synapses through disinhibition of GAT-1 function. GABA uptake is also modulated by activation of cannabinoid CB1 receptors in the rat GP (Venderova et al., 2005), providing another mechanism that could be used to regulate the overactive GABAergic striatopallidal transmission in Parkinson's disease (Romero et al., 2002; Brotchie, 2003). These mechanisms of indirect modulation of GAT activity may represent a more promising therapeutic strategy in the treatment of Parkinson's disease than use of primary GAT blockers (for review, see Conti et al., 2004).
This research was supported by NIH grant RO1 NS 0432937 and the Yerkes Primate Center NIH base Grant (RR-00165). Thanks to Jean-Francois Pare and Xing Hu for technical assistance.
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Keywords: GABA transporter, striatum, globus pallidus, substantia nigra, patch clamp recording
Citation: Jin X-T, Galvan A, Wichmann T and Smith Y (2011) Localization and function of GABA transporters GAT-1 and GAT-3 in the basal ganglia. Front. Syst. Neurosci. 5:63. doi: 10.3389/fnsys.2011.00063
Received: 26 April 2011;
Paper pending published: 10 May 2011;
Accepted: 13 July 2011; Published online: 28 July 2011.
Edited by:James M. Tepper, Rutgers – The State University of New Jersey, USA
Reviewed by:Atsushi Nambu, Graduate University for Advanced Studies, Japan
Hitoshi Kita, The University of Tennessee Health Science Center, USA
Copyright: © 2011 Jin, Galvan, Wichmann and Smith. This is an open-access article subject to a non-exclusive license between the authors and Frontiers Media SA, which permits use, distribution and reproduction in other forums, provided the original authors and source are credited and other Frontiers conditions are complied with.
*Correspondence: Xiao-Tao Jin, Division of Neuroscience, Yerkes National Primate Research Center and Department of Neurology, Emory University, 954 Gatewood Road NE, Atlanta, GA 30322, USA. e-mail: firstname.lastname@example.com


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Approximately 55-80% of customers are considering opening an account in a digital-only bank.
More than a third (40%) of customers in developed Asia are willing to shift to branchless or digital-only banks, according to a report from management consultancy firm McKinsey.
The openness of customers to digital alternatives without the need for a physical distribution network has been correspondingly met by the local market in developed Asia.
"Over the last three years, several new banking propositions have been launched across the Asia-Pacific region," McKinsey noted.
In Korea, for instance, Kakao launched Kakaobank to become the fastest-growing mobile bank in teh world with over 5.5 million users and $6b in deposits.
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Title: No Sleep (9KB)
Author: faroutdragon1
Date archived: August 25, 2004
K+ - Angst/drama - Daniel/Janet
Spoilers: None
Summary: Janet has accidentally given Daniel a highly lethal virus. What will she do? (Chall. #32)
Title: Torture (10KB)
Author: Faroutdragon1
Date archived: June 24, 2004
T - Action/adventure; romance - Daniel/Janet, hinted Sam/Jack
Spoilers: Everything up until Season 4, really
Summary: On a mission to P3X-473, Daniel is captured, and when they find him, Janet is called to the planet to help him.