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6. Chapter 6: Mechanics
"Game Mechanics are constructs of rules and feedback loops intended to produce enjoyable gameplay. They are the building blocks that can be applied and combined to gamify any non-game context. "
— The Gamification WIKI
Mechanics are the most visible part of gamification and tend to be the primary focus of most gamification projects. We like to think of game mechanics as paints in an artist's palette. We cannot create great art just by adding many colors to our picture, unless we also have an artistic vision, talent, and training to begin with. Similarly, successful application of game mechanics depends on a well-designed gamification strategy built on a good understanding of the player, the mission, and human motivation.
In this chapter, we present a curated list of game mechanics that may be used as building blocks and combined in strategic ways to achieve the positive engagement loop in your application.
6.1 Curated list of Game Mechanics
There are a few collections of game mechanics available online (footnote 1). We find these lists are at different levels of granularity and not all of them are pertinent to enterprise gamification. Therefore, we have curated a list of game mechanics that can help business software designers embark on their gamification journey.
When used appropriately, game mechanics can leverage a natural motivational driver in the player. For example, the motivation driver for collection may be addressed with badges, and the achievement motivation may be addressed by leaderboard. However, in a holistic gamification design, many motivational drivers may be at play. For example, FourSquare players may be motivated by collecting points and badges, achieving the status of mayor in their favorite establishment and connecting with their network while doing so. We will point out the primary motivational driver addressed by each of the game mechanics discussed below.
6.1.1 Points
Points are the granular units of measurement in gamification. They are single count metrics. This is the way the system keeps count of the player's actions pertaining to the targeted behaviors in the overall gamification strategy. For example, FourSquare counts each check in, and LinkedIn counts each connection.
Points provide instant feedback to the player, and thus address the feedback motivational driver. Players may also motivated by collection, to see their points count go up.
6.1.2 Badges
Once the players have accumulated a certain number of points, they may be awarded badges. Badges are a form of virtual achievement by the player. They provide positive reinforcement for the targeted behavior.
Foursquare awards badges when the player has accumulated enough check-ins. Another example of a badge is eBay's top seller virtual "ribbon".
Badges address the motivational driver of collection and achievement.
6.1.3 Leaderboards
Leaderboards bring in the social aspect of points and badges, by displaying the players on a list, typically ranked in descending order with the greatest number of points at the top. The possible disadvantage of a leaderboard is that it could be demotivating to a new player. For example, if player A has 10,000 points, and is on top of the leaderboard, and a new player B has 10 points and is at the bottom, it is likely that player B may become demotivated and give up playing the game. She, or he, may believe that he/she is never going to compete with player A, and therefore why should he/she even try?
Foursquare has a leaderboard modified into a cross-situational leader board. This variant places the logged-in player (Matt Healy in Figure 6.7) in the center and shows similar scoring fellow players above (Tristan Walker) and below (Tim Vetter) for context. The ranking (points) is limited to a set of players who are close to the logged-in player. The goal with this variant of leaderboard is to motivate the player to compete with the players closest to him. Note that a cross-situational leaderboard may be different for each player since it is limited to his or her context. It does not convey an overall ranking of all players.
The achievement motivational driver is addressed via a leaderboard.
6.1.4 Relationships
Relationships are game mechanics based on the motivational driver of connection. We are social beings, and relationships have a powerful effect on how we feel and what we do.
Peer pressure is not restricted to school-age children. Adults succumb to it too. In 2010, one of the authors (Janaki Kumar) did research on personal sustainability and one of the findings was that a trusted person in a participant's network had more impact on his or her day-to-day choices than the media. For example, participants were more likely to recycle if a trusted member of their community did so, than if they were told to do so by the media.
Relationships reduce stress in people and are positive motivators. People who are trying to quit bad habits such as alcoholism, or deal with a loss of a loved one have found that support groups offer emotional support and encouragement during a time of need. In the technology world, developer communities are a good example of a support group for developers where they offer and receive technical help.
Relationship addresses the motivational driver of connection.
6.1.5 Challenge (with epic meaning)
Challenge is a powerful game mechanic to motivate people to action, especially if they believe they are working to achieve something great, something awe-inspiring, and something bigger than themselves.
As we mentioned earlier, scientist at the University of Washington challenged the public to play Foldit (footnote 2), a game about protein folding. This game has a scientific purpose behind it. Knowing the structure of the protein in a cell was the key to understanding how it works and how to target it with drugs. Since proteins are part of so many diseases, they can also be part of the cure. Folding proteins provides important clues to the scientists on how to prevent or treat diseases such as HIV/AIDS, cancer and Alzheimer's. A team of experts had worked on this problem for over 10 years and had not solved it. Once the scientific challenge was launched in the form of a game, 46,000 volunteer players solved the puzzle in 10 days.
The challenge game mechanic addresses the achievement motivational driver. However, in the case of the Foldit challenge, the feeling of connection and perhaps reciprocity, (if the player had known someone dear to him/her suffering with the illness the challenge was seeking to cure) may have played a part in its overwhelming success.
6.1.6 Constraints (with urgent optimism)
Interestingly, constraints such as deadlines, when combined with urgent optimism, motivate people to action. Urgent Optimism (footnote 3) refers to extreme self-motivation. It is the desire to act immediately to tackle an obstacle combined with the belief that we have a reasonable hope of success.
Some registration sites use gamification to reduce the drop off rate by limiting the amount of time the user can take to complete the registration process.
Gilt, a fashion ecommerce site, constrains the time allowed for their customers to bid on items to motivate them to action.
Players are motivated by achievement when they are faced with these constraints and are driven to overcome them.
6.1.7 Journey
The journey game mechanic recognizes that the player is on a personal journey and incorporates this element into the experience. Here are three examples of implementations of this game mechanic:
220.127.116.11 Onboarding
A new player needs to be on boarded since they are just starting the journey. Offering help and a brief introduction to the features and functions motivate the player to embark on the journey.
220.127.116.11 Scaffolding
Scaffolding is a way to help the on boarded, yet inexperienced, user avoid errors and feel a sense of positive accomplishment. A product could progressively disclose more features as the player gains more experience using the product.
18.104.22.168 Progress
Progress refers to providing feedback to the user on where he or she is in the journey, and encouraging him/her to take the next step.
Journey addresses the player's need for blissful productivity, by presenting the right set of features appropriate to the player's level in the game.
6.1.8 Narrative
The narrative game mechanic draws the players into a story within the game. Zombie Run, a fitness game, uses narrative to make the players believe that zombies are after them, and they need to run as fast as they can to get away. The object of the game is to motivate the players to get fit without making it explicit.
Narrative offers the players a chance to express themselves via role-play. In the case of Zombie run, players are motivated by achievement by out-running the zombies.
6.1.9 Emotion
As Don Norman eloquently argues in his book Emotional Design (footnote 4), our emotions do play a role in how we experience a product.
In many ways, emotional design is a large category in and of itself. In the context of gamification, we are not attempting to cover the topic as a whole. Rather, we want to draw inspiration from it, to enrich our gamification designs.
Game designers have led the way in investing in high quality artwork in their products that appeal to our emotions. Consumer products (iPhones, iPads) and websites (Pinterest) are following this trend. Employees experience emotional delight in the consumer software they use, and have similar expectations with enterprise software.
Humor is another emotion pertinent to game mechanics. The tone of the product can be conveyed in the micro-copy, or in the informational text and messages on the user interfaces. Humor has the power to deflect a negative experience into a (somewhat) positive one.
Humorous micro-copy addresses the motivational driver of feedback, while people may choose to use esthetically pleasing designed products as an avenue of self-expression.
6.2 The Game Plan
With a solid understanding of the player, mission, and motivation, the gamification system may be designed using game mechanics as building blocks. The three main considerations in pulling together the game plan are the game economy, game rules, engagement loops.
6.2.1 Game Economy
Garner describes game economy as follows:
There are four basic currencies that players accumulate in game economies — fun, things, social capital and self-esteem — that are implemented through game mechanics, such as points, badges and leaderboards. These game mechanics are simply tokens of different currencies of motivation that are being applied to reward players.
Fun refers to rewards such as discovery, excitement, fantasy, awe, delight, and surprise. Things refer to cash, resources, and prizes or gifts. Social capital refers to likes, friends, and status. Self-esteem refers to praise, access, mastery, and conquest.
As part of the game plan, you can decide the mechanics you want to use as currencies in your game economy.
6.2.2 Game Rules
Once you have decided what mechanics to use, the next thing is to come up with a set of rules of the game. If you are designing a system to motivate call center employees to undergo training, and you have decided to use points in your game economy, you will need to decide how many points you award for the action. If the employee only took 50% of the training, do they receive all the points, none of the points, or 50% of the points?
If you are designing a community where you are trying to increase participation, do you award points for all comments, or only for thoughtful comments, and who decides if a comment is thoughtful?
The rules of the game pull together the mechanics into a flow to motivate the player to achieve the mission.
6.2.3 Engagement Loop
The core engagement loop refers to game mechanics combined with positive reinforcement and feedback loops that keep the player engaged in the game. This concept has been discussed by Amy Jo Kim (footnote 5), a renowned game designer.
The four main stages in the loop are:
- Motivate emotion,
- Call to action,
- Re-engage,
- Feedback and reward.
For example, a new player to a community site may:
- visit out of curiosity,
- participate via a challenge,
- be reminded by the system to check-in periodically to explore and discover new features,
- receive feedback and rewards for participation.
This frequent invitation to interact with the system creates positive reinforcement and the player will be motivated to stay engaged with the gamified system.
6.3 Summary
Game mechanics are the most visible aspect of gamification. To be successful, mechanics need to be selected based on a thorough understanding of the player, the mission and human motivation.
In this chapter, we have presented a curated list of game mechanics that can help you start your journey into gamification. They are points, badges, leaderboards, relationships, challenges, constraints, journey, narrative, and emotion.
We guide you in creating a game plan that uses these mechanics as building blocks. The main elements to consider in the game plan are game economy, game rules, and the engagement loop.
The gamification system needs to engage the player continuously as part of a sustainable strategy.
6.4 Insights from SAP Community Network
In 2005, it was not common for communities to award points for participation. SCN ventured carefully into applying game mechanics by designing the system as follows:
- A regular blog post earned 40 points. Exceptional posts were awarded additional points while points were deducted for posts with short-comings
- Responses to questions were awarded between 2 and 10 points. Additional points were awarded for answers that were thoughtful or exceptional in solving the problem
- Points were awarded for wiki-edits depending on the amount of contribution.
- Badges were introduced to distinguish SAP employees. Community members could reach out to them directly to get more information from an SAP perspective.
- Members with administrative authority on forums and blogs were given Moderator badges.
- SCN offers leaderboards to showcase expertise in the community. The system also tracks life-time points for each player.
This simple combination of points, badges and leaderboards created a positive dynamic. However, it also led to some unintended consequences, which we will discuss in the next chapter.
To be continued at the end of next chapter...
6.5 References
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Table of contentschapter TOC
- 6 Chapter 6: Mechanics
- 6.1 Curated list of Game Mechanics
- 6.1.1 Points
- 6.1.2 Badges
- 6.1.3 Leaderboards
- 6.1.4 Relationships
- 6.1.5 Challenge (with epic meaning)
- 6.1.6 Constraints (with urgent optimism)
- 6.1.7 Journey
- 126.96.36.199 Onboarding
- 220.127.116.11 Scaffolding
- 126.96.36.199 Progress
- 6.1.8 Narrative
- 6.1.9 Emotion
- 6.2 The Game Plan
- 6.2.1 Game Economy
- 6.2.2 Game Rules
- 6.2.3 Engagement Loop
- 6.3 Summary
- 6.4 Insights from SAP Community Network
- 6.5 References


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A couple who battered the woman's four-year-old son and left him to die in a locked, unheated box room have been convicted of murder.
The woman and her partner starved Daniel Pelka for months and force-fed him salt during a campaign of physical abuse. A serious case review will now try to establish why a number of opportunities to save the boy's life were missed by authorities, who were convinced by his killers that he was suffering from a rare eating disorder.
Fourteen months before he died in March last year, Daniel became the focus of a police inquiry after his arm was broken. But the file was closed when his killers convinced officers that he hurt himself when he jumped from a sofa at their home in Coventry.
Just weeks before his death, the boy was sent to school with black eyes and was described as a "bag of bones". When he was at home, he was punished with arduous exercises and locked in a tiny room with no food or sanitation.
Magdelena Luczak, 27, and former soldier Mariusz Krezolek, 34, who were from Poland, showed no emotion after they were found guilty of murder at Birmingham Crown Court.
The couple, who were heavy drinkers and abused drugs, had blamed each other for the boy's death but text messages revealed they worked as a team to inflict the injuries. The messages showed they "took relish" in the abuse.
The pair carried out searches on their computer which suggested Daniel was beaten, almost drowned and poisoned with salt.
The court heard that he was a healthy boy when he started school in September 2011 but was subjected to suffering that left him resembling a concentration camp victim by the time he died seven months later.
Daniel, who weighed less than 11kgs and was in a state of "skeletal emaciation" when he died, had been seen by a paediatrician three weeks before he was found dead who reported that he was underweight but not "wasted".
In evidence, the couple admitted they had gone to sleep on both nights that Daniel lay dying in the box room, and even had sex after his death.
In a statement released by police, Eryk Pelka, Daniel's biological father, said: "It's a great tragedy that such a little angel had to leave this world. I hope that those responsible will be punished severely."
The couple will be sentenced on Friday.
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Organizations Directory
Academic & Pre-Professional (Academic)
Clark students are involved in many different activities on campus - from sports, student government, the arts, political activism, and journalism, to community service or cultural exchange. These clubs and organizations provide all types of experiences
Active Minds
Active Minds aims to remove the stigma that surrounds mental health issues, and create a comfortable environment for an open conversation about mental health issues on Clark's campus.
Activists United (formerly Peaceworks)
Activist United members are dedicated to advocacy, education and action for a just and equitable society. Members work on local and international issues with a focus on creating strong communities and community organizing.
African Diaspora Dance Association (ADDA)
ADDA is a student run dance performance group that draws from music and dance styles from the African Diaspora which includes afro beats, coupe decale, hip hop, r&b, dancehall and much more. Our shows are put on in both the Fall and Spring semester.
All Kinds of Girls (AKOG)
The All Kinds of Girls program aims to cultivate pride, leadership, self-esteem and self-expression in girls ages 9-15. Run by undergraduate women at Clark University, this program provides girls a safe space in which to use their voices,
Amnesty International
Amnesty International works to shine a light on local and global abuses of human rights and injustices to achieve dignity for all people.
Arboretum Advocates of Clark University (Arboretum Advocates)
We the Arboretum Advocates of Clark University speak for the trees of the Hadwen Arboretum, Clark Campus and the Worcester Community. Our weekly meetings are Wednesdays at 7pm in Jefferson 202
Arts and Performance
Clark students are involved in many different activities on campus - from sports, student government, the arts, political activism, and journalism, to community service or cultural exchange. These clubs and organizations provide all types of experiences
Asian Culture Society (ACS)
Founded in 1986, the Asian Culture Society enhances the Clark community by providing opportunities to learn about and experience Asian cultures. It has also worked in conjunction with other organizations to sponsor international cultural celebrations.
Association of Martial Artists of Clark (AMAC)
The Association of Martial Artists at Clark (AMAC) is dedicated to providing a space for martial artists to continue their training, regardless of style or past experience. See full description for training times.


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Arena Apartments
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Arena Apartments
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Property Location
With a stay at Arena Apartments in South Brisbane, you'll be minutes from The Markets and Brisbane Convention and Exhibition Centre. This 4-star apartment is within close proximity of Queensland Performing Arts Centre and Brisbane City Hall.
Make yourself at home in one of the 50 air-conditioned rooms featuring kitchens with full-sized refrigerators/freezers and microwaves. Rooms have private balconies or patios. Complimentary wireless Internet access keeps you connected, and cable programming is available for your entertainment. Conveniences include washers/dryers and ceiling fans, and housekeeping is provided weekly.
Be sure to enjoy recreational amenities including an outdoor pool and a fitness center. This apartment also features complimentary wireless Internet access, a picnic area, and barbecue grills.
Business, Other Amenities
Featured amenities include luggage storage, laundry facilities, and an elevator (lift). Free self parking is available onsite.
Hotel Facilities
- Barbecue grill(s)
- Garden
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- Year Built 2015
- Elevator/lift
- Total number of rooms - 50
- In-room accessibility
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- Number of floors - 11
- Smoke-free property
- Number of buildings/towers - 2
- Fitness facilities
- Free self parking
- Luggage storage
- Picnic area
Room Facilities
- Free toiletries
- Washer/dryer
- Private bathroom
- Ceiling fan
- Air conditioning
- Television
- Iron/ironing board
- Rollaway/extra beds (surcharge)
- Separate living room
- Balcony or patio
- Free WiFi
- Separate dining area
- Shower only
- Microwave
- Cribs/infant beds (surcharge)
- Cable TV service
- Premium TV channels
- Full-sized refrigerator/freezer
- Weekly housekeeping provided
- Kitchen
- Hair dryer
Hotel Policy
Know Before You Go
- This property's policy is to refuse certain bookings for the purpose of group events or parties, including pre-wedding stag/bachelor and hen/bachelorette parties.
- Only registered guests are allowed in the guestrooms.
The following fees and deposits are charged by the property at time of service, check-in, or check-out.
- Crib (infant bed) fee: AUD 20 per night
- Rollaway bed fee: AUD 30 per night
The above list may not be comprehensive. Fees and deposits may not include tax and are subject to change.Show all hotel policy Show less hotel policy


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The arrival of the BEAST
In the immortal words of Hunter S. Thompson: 'He who makes a beast of himself, gets rid of the pain of being a man'. The mountain biking world has been a buzz about the arrival of the limited edition, almost mythical 'Beastnut' from Calibre, and we're delighted to say we've got the first video footage of it out in the wild.
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Return to Never Land (2002)
Should of been a direct to video sequel!OK animation and songs
In this sequel to Disney's Peter Pan we see Wendy all grown up with children of her own. Wendy's daughter Jane is having family problems as London is sending all the children away during World War 2.
Wendy is kidnaped by Captain Hook and his pirate crew. Hook kidnaps her thinking he will finally get Peter Pan. Peter Pan discovers that Hook accidently brought Wendy's daughter instead. Peter tell Jane the only way is to fly. This sounds like something from the "The Wizard of Oz" type story. Jane gets herself into a deal with Hook which she eventually regrets.
Hook manages to capture Peter Pan and the Lost Boys. It's up to Jane and Tinker Bell to save them. There is also a sub-plot with Tinker Bell dying . Jane saves everyone and goes home. The ending of the movie is a happy one as Disney always seems to do. One question though . Was the trip to Neverland all a dream?
The animation was like watching one of the many direct to video sequels Disney has done over the years. The songs were OK. If you really want to see this wait until it gets to the cheap movie theaters.
Dragonfly (2002)
A cross of The Sixth Sense, Ghost and What Lies Beneath! Semi-predictable! Felt like a made for TV movie
In Kevin Costner's latest movie, Costner plays a doctor who recently lost his wife while she was away doing work for the Red Cross. There is accident and his wifes dies. She is also pregant.We see the funeral etc.
Then Costner is told by his co-workers, boss etc to stop working so much. Costner starts to see things with hospital patients who have near death trips. He believes that they are talking to his dead wife. Eventually Costner finds out all along what his wife was trying to tell him about. I will not give away the ending as it was kinda sappy and sugar coated.
This movie is basically a cross of "The Sixth Sense", "Ghost" and "What Lies Beneath". It's been done before so we did not get anything new from Hollywood again. If you've seen any of these movies, it will be semi-predicatable for you.
The movie at times felt like a made for TV movie thriller. Sure you get to see Costner going crazy over his wife from beyond the grave. But the supporting players as just window dressing, Kathy Bates has done a lot better work than this. So has Joe Morton and also Ron Rifkin. Overall , watch this movie if your are Kevin Costner fan ,otherwise wait for it to hit the cheap movie theaters or DVD.
Impostor (2001)
One of 2002 worst movies of the year! Should of gone straight to DVD & video! Lame! Stupid!
This movie was shelfed for over a year until Disney could find a dumping date to bring this to movie theaters. It should gone straight to DVD and video! This movie was very much a "paint by numbers" film for bad science fiction and movies in general.Not to mention it ripped off several other science fiction movies like "Total Recall","The Thing","Invasion of the Body Snatchers". and "Blade Runner'.
I saw this movie today at the last matinee after I find it out it did not get into the Box Office top ten.Perhaps a bad marketing killed it at the movie theaters. The cast does what little they have in terms of a script. There even a "cat and mouse" game in the movie. I suspect that this movie will be on DVD very fast. Needless to say the audience was laughing out loud at this flop!
I find hard to belive that with 2002 being only 6 days old,I'm already writing a review for one the worst films of the year. The cast all has done better work than this pathetic movie.
This movie also a very close boardline "R" movie. The amount of swearing ,violence , and nudity almost make it a "R" movie.Maybe the MPAA was asleep when they rated this box office turkey?This film is just as bad as the movie "Supernova".
Wait for this turkey on DVD or better yet wait for it hit the cheap theaters.
A Beautiful Mind (2001)
A wonderful biopic about John Forbes Nash,Jr and his life's problems to the Nobel Prize
I guess I'm one of the lucky people who have seen this movie before it gets a wide release.
This movie is a biopic of math prodigy John Forbes Nash,Jr and many of his problems in life.When we first meet John(played by the awesome Russell Crowe) he is a student a Princeton. John makes a few friends at the school. But John's problems are about to increase in his life big time. John graduates from Princeton and moves on to bigger things.
A few years later( the mid-1950's) John is working for the Dept. of Defense at MIT. John is recruited by a strange man in black to decode stuff for the government. John meets a beatuful woman who understands him.
After John and Alicia(jennifer Connelly) get married, John's life seems pretty normal. But their married problems do not get any better.Eventually John ends up in a mental hospital for his schizophrenia. John gets the medicine he needs to get better. But John is still seeing people and talking to people who do not exist. Eventaully John Nash wins his Nobel Prize in 1994.
There's lots more but I've already said enough.
This movie has Oscar written all over it. The cast should gets Oscar nominations for thier work.Ron Howard finally made a movie far better than his last movie(How the Grinch Stole Christmas). I hope this movie gets awards as it a very well done film about a real life person,
If you are sick and tired of all the typical mainstream trash out there, check out this wonderful biopic. It's about 2 1/2 hours but it goes by quite fast. This movie was better than the movie trailers before it! If not check it out when it comes out on DVD!
Spy Game (2001)
In a word: BORING!
This movie was not like the trailers/ TV ads for it.It is a long boring movie with Robert Redford and a bunch of men sitting around a table for most of the movie. There are too many flashbacks back to the past about Redford's and Pitt's relationship.Eventually I got up and refilled my soda becasue I was bored. Since this a mainstream movie, expect a typical mainstream ending. I predicted the ending of this movie at the halfway point. If this movie did not have so many flashbacks, maybe I would'nt of looked at my watch often during the movie.
This movie is more of a heavy drama, rather than a thriller. The only action in the movie is in the beginning and end.Other than that, it's mostly men talking around a table interviewing Redford. If I fell asleep, I doubt I would of missed much in this movie!The trailers and ads are misleading! Not a thriller but a long BORING drama!
So if you like BORING long movies, see this movie, Otherwise wait for this movie to hit the cheap movie theaters or DVD/videotape.
Harry Potter and the Sorcerer's Stone (2001)
Special effects take over and it has a very small plot!Wait for it on DVD/Videotape!
Well today, I finally caved in and saw this movie. For starters I'm sick and tired about "Harry Potter". What I saw was just as I was expecting for a very big hyped movie. The movie itself had a very small plot(mainly Harry's first year at special school for wizards) and a small subplot about a special stone. This acting at times seems very wooden from both the actors, the extras etc. There is special effects overkill in this movie.
The running time is a little over 2 1/2 hours. This movies moves very slowly a lot of the time. Even when they have special effects it seems like it's a too long of a movie. I liked the adult actors, but I hope the child actors playing Harry Potter etc do not get typecast as they could if they play their roles until they are grown up. I also wondered if parts of this movie were left out on the cutting room floor.
I say if you are die hard fan of the series, see this movie, But if you are causal fan or just sick and tired of the whole "Harry Potter" thing wait for it on DVD/videotape.
Monsters, Inc. (2001)
A cute story about life inside our closets with Monsters!
This movie tells the story of two monsters who accidently let a little human girl into their world inside our closets. The two monsters must find a way to get her home. Along the way, the human girl becomes friends with these two monsters. Eventually word spreads though out the monster world to find the human girl at all costs. Since it's a Disney movie it has a happy ending. The bad guys get what they had coming to them. The monster world is saved and a new power source is found to power their world.
Behind Enemy Lines (2001)
Behind Enemy Lines = 1991's Flight of the Inturder recycled just new location!
If you've seen the 1991 movie "Flight of the Intruder" which starred Brad Johnson and Danny Glover, you've seen this movie before. All Hollywood did was change the location to Bosnia! Big deal! The movie itself is OK but at times I'd swear this movie was misrated! It should of been rated R for all the war violence! I felt like I was watching Pearl Harbor's battle scenes over a 90 minute period!
Gene Hackman gives us his usual miltary performance in this movie. Although he acts like he is just doing it for the paycheck. Owen Wilson was OK in the role of Bennet who is lost in Bosnia over the holiday season. Both of these actors have done better work. I think Owen Wilson should stick to comedy or else try a new genre in his next flick. Hackman can do well in almost everything . I just find it hard to believe that i like him better in his past movies.
If you must see this, wait for it to reach your cheap movie theaters. Or better yet wait for on DVD! I belive there are several times in the movie which has bad editing.
Or better yet, save yourself your hard earned cash and watch 1991's Flight of the Intruder! This movie is a rip-off of it!
Black Knight (2001)
Direct rip-off of "A Connecticut Yankee in King Arthur's court" One of 2001's worst movies!
Needless to say I went to go see this movie before I went back to my job working in retail which is very busy right now for a laugh. I like Martin Lawernce but this movie was not at all funny . The best parts of the movie were in the trailer and TV ads. This movie is so bad I felt like I was watching a direct to video movie on the big screen. If was Martin Lawernce, I'd be embrassed to star in it. Maybe he had becasue he has a contract with Fox? Big Momma's House anyone? The movie itself had nothing great about it. The way the ending is will a sequel be made out this trash. Please don't make a sequel to this junk!
Save your $$ and wait for this movie on DVD and video! Also why was this movie not direct to video?
This movie is one of 2001's worst movies for me. I wonder if it will get any "Razzies" next year for bad movies?
The Substitute: Failure Is Not an Option (2001)
An OK sequel! Rent it first!
In the fourth "The Substitute" movie we find Treat Williams going undercover at miltary school for a old friend worried about his son. The story was OK, the action scene totally cheesy and the dialogue pathetic. This movie also stars Angie Everhart as love interest for Treat Williams. Too bad she could'nt say this movie for her few scenes.Also there are really bad signs saying stuff like "Another minority owned installation" and others. Like they would do that in reality? The movie is rated R for swearing, violence and some partial nudity.
I hope if they make a fifth movie, it is better than this installment of the series. I often wonder who plays role of the Substitute better: Treat Williams or Tom Berenger? I wish Berenger keep the role as he was real at least.
If you really want to see this movie , rent it first before you buy it!


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Phototransduction in ganglion-cell photoreceptors
- First Online:
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- Accepted:
DOI: 10.1007/s00424-007-0242-2
- Cite this article as:
- Berson, D.M. Pflugers Arch - Eur J Physiol (2007) 454: 849. doi:10.1007/s00424-007-0242-2
A third class of photoreceptors has recently been identified in the mammalian retina. They are a rare cell type within the class of ganglion cells, which are the output cells of the retina. These intrinsically photosensitive retinal ganglion cells support a variety of physiological responses to daylight, including synchronization of circadian rhythms, modulation of melatonin release, and regulation of pupil size. The goal of this review is to summarize what is currently known concerning the cellular and biochemical basis of phototransduction in these cells. I summarize the overwhelming evidence that melanopsin serves as the photopigment in these cells and review the emerging evidence that the downstream signaling cascade, including the light-gated channel, might resemble those found in rhabdomeric invertebrate photoreceptors.
KeywordsMelanopsin ipRGC Photoreceptor Rhabdomeric Retina Ganglion cell Suprachiasmatic nucleus TRPC
Within the past decade, the existence of a previously unknown class of photoreceptor in the mammalian retina has come to light. These directly photosensitive neurons differ radically in form and function from the well-known rod and cone photoreceptors. They are a rare variety of retinal ganglion cells (RGCs), the class of retinal neurons sending axons through the optic nerve to synapse in the brain. For at least the prior century, it had gone without question that ganglion cells were entirely dependent on polysynaptic influences from rods and cones for their responsiveness to light (indeed, even today there is no reason to doubt this for most ganglion cells). However, as the new millennium dawned, this dogma was being challenged by behavioral studies showing the persistence of certain reflex responses to light in photoreceptor degenerate animals, e.g., [14, 15, 35, 36], and the presence of a possible photopigment, melanopsin, in a rare population of ganglion cells [18, 19, 21, 49, 50]. In short order, it became clear that these cells possessed a capacity for autonomous phototransduction [5]. These intrinsically photosensitive RGCs (ipRGCs) are now recognized as playing key roles in synchronizing circadian rhythms to the day–night cycle, mediating the pupillary response to light, and modulation of melatonin release by the pineal gland. Many aspects of this rapidly developing story have been thoroughly summarized elsewhere [4, 6, 14, 16, 31, 44, 54, 61]. The reader is referred to these reviews for background on the intellectual origins of the discovery of melanopsin and the ipRGCs, on their structure and physiology, and on their contribution to circadian photoentrainment and other nonimage-forming visual functions. The goal of the present review is to summarize the current state of knowledge concerning the nature of the transduction process in ipRGCs.
Structural and functional properties of ipRGCs relevant to the phototransduction process
The ipRGCs represent a tiny minority of the RGCs of mammalian retinas (<1–2%) [9, 21, 51]. Like other ganglion cells, ipRGCs extend dendritic processes into a synaptic lamina of the retina, the inner plexiform layer, and send a single axon through the optic nerve to terminate in the brain. The ipRGCs are distinguishable from other RGCs, however, by their selective expression of a novel opsin called melanopsin and of a peptide neuromodulator, pituitary adenylate cyclase activating polypeptide [19]. These cells also exhibit a distinctive pattern of central axonal projection, with major targets including nuclei coordinating circadian rhythms and mediating pupillary responses [21]. Their most notable functional idiosyncrasy, of course, is their ability to respond to light when totally isolated from any synaptic influence [5, 9, 60, 62]. It should be noted, however, that in addition to this direct sensitivity to light, ipRGCs receive excitatory and inhibitory synaptic inputs that permit rods and cones to modulate their electrical behavior [2, 46; Wong et al. unpublished observations].
The direct photoresponse of ipRGCs to light can be evoked not only by diffuse illumination but also by stimuli restricted either to the soma or to the dendrites [5], indicating that the phototransduction apparatus is widely dispersed through the cell. Illumination induces a conductance increase and membrane depolarization, which, if sufficiently large, can trigger repetitive regenerative action potentials. The responses are generally very sluggish, with a latency of several seconds from light onset to response peak when probed with stimulus intensities in the middle of the dynamic range. However, the intensity–latency relation is steeply inverse, with latencies ranging from several hundred milliseconds for saturating stimuli to >1 min for near-threshold stimulation. Responses to prolonged light steps generally decay significantly from their initial peak, in part because of light adaptation [63], but the steady-state response is generally remarkably stable. This stability seems likely to be a key functional difference from rods and cones, although a systematic exploration of this difference has not yet been attempted. The ability of ipRGCs to signal the presence of steady illumination over many minutes is especially relevant in the context of the reflexive responses to ambient light to which they contribute. Like the ipRGCs themselves, these visual responses are remarkably tonic and/or exhibit unusually long integration times. Response recovery in ipRGCs, like response onset, is strikingly slow by comparison with rods and cones. Typically, several minutes in darkness are required for a photoactivated ipRGC to return to its prestimulus voltage [5, 60].
Identity and properties of the ipRGC photopigment
There is overwhelming evidence that melanopsin is the functional photopigment of ipRGCs. This opsin was first discovered in a screen for opsin-like genes in cDNA library from Xenopus dermal melanophores, a nonneural cell type that redistributes its pigment in direct response to light [49]. Orthologs of the gene were then identified in mammalian genomes, including those of mice, monkeys, and humans [50]. A striking feature of melanopsin is that its predicted amino acid sequence exhibits greater homology with the opsins of invertebrate rhabdomeric photoreceptors ("r-opsins") than with the opsins of vertebrate ciliary photoreceptors ("c-opsins") [1, 30, 49, 50, 57].
In mammals, melanopsin is apparently expressed almost exclusively within a rare population of RGCs ([9, 19, 22, 50, 51] but see [11] and [45]). These RGCs are the main source of retinal input to the suprachiasmatic nucleus of the hypothalamus, the brain's circadian pacemaker [18, 19, 21, 22]. Experiments combining recording, intracellular dye filling, and immunofluorescence established that melanopsin-expressing cells are directly photosensitive [5, 22]. The spectral tuning of the ipRGC photoresponse adheres closely to the form expected for an opsin-based response [5]. Immunohistochemistry identified the presence of melanopsin protein throughout the dendritic arbor of ipRGCs as well as in the soma [9, 19, 22, 50, 51]. This distribution bolsters melanopsin's hypothesized status as the ipRGC photopigment, because, as already noted, both the dendrites and somata of ipRGCs have an independent capacity for phototransduction. In fact, melanopsin has been observed even in the proximal, intraretinal portion of ipRGC axons. This suggests that the axons, too, could be at least weakly photosensitive, although this has not been tested. Melanopsin appears localized almost entirely to the plasma membrane of ipRGCs [2]. There is no evidence for any membranous specializations in ipRGCs analogous to the disks of rod or cone outer segments or the microvilli of rhabdomeric photoreceptors. Those specializations serve to dramatically increase membrane surface area and thus the density of the photopigment. Thus, an incident photon is presumably much less likely to be captured by the melanopsin in an ipRGC than by the photopigments of rod or cone outer segments or ommatidial rhabdomeres. This is one likely contributor to the relatively low sensitivity of ipRGCs by comparison with rods and cones [5, 9].
Key support for the hypothesis that melanopsin is the ipRGC photopigment came from the observation that the direct photoresponse of these cells was abolished by targeted deletion of the melanopsin gene [37; see also 23, 42]. By itself, however, this finding did nothing to exclude an alternative model in which melanopsin served not as a photopigment but rather as a photoisomerase. According to this model, melanopsin might be essential for providing cis retinaldehyde chromophore to the actual ipRGC photopigment, which remained to be identified [3]. However, parallel studies provided evidence that melanopsin could form a functional photopigment, inasmuch as it mediated light-triggered G-protein activation in a biochemical assay [40]. Further, evidence soon emerged that heterologous expression of melanopsin conferred intrinsic photosensitivity on each of three otherwise light-insensitive cell types [38, 41, 52]. Moreover, the action spectrum of the induced photoresponses closely matched that of the ipRGCs, peaking in the blue portion of the visible spectrum (∼480 nm; [41, 52]; see also [30]). This close spectral congruence has not been seen universally. Spectrophotometric observations on purified melanopsin initially suggested a peak absorption closer to 420 nm [40], and a similar value was obtained from electrophysiological measurements in one heterologous expression system [38]. More recent evidence suggests that if the composition of the extraction buffer is altered, purified melanopsin does indeed absorb maximally near 480 nm (Walker et al., personal communication; see also [30]). Thus, with the exception of the report of Melyan et al. [38], there is good agreement that melanopsin can account for the spectral behavior of ipRGC photoresponses. The question of whether the shorter wavelength absorbance maximum seen in some experimental settings occurs under physiological conditions in any native cellular environment is unresolved.
The identity of the melanopsin's retinaldehyde chromophore is unknown, although the opsin can form a functional photopigment when reconstituted with all-trans, 9-cis, 11-cis, or 13-cis retinaldehyde ([17, 38, 40, 41, 52], and Qiu and Berson, unpublished observations). The various states through which the pigment passes during photoexcitation and pigment regeneration are also largely mysterious. However, clues may come from the invertebrate-like structure of melanopsin generally and the invertebrate-like identity of the residue thought to serve as the Schiff-base counterion in particular [49]. These suggest that melanopsin may form a bistable, photoreversing pigment. In such pigments, photon absorption and isomerization of cis to trans retinal generates a metarhodopsin species that is thermally stable. Rhabdomeric metarhodopsin can function as a photoisomerase, absorbing light to convert its retinal chromophore from trans to cis retinal and reverting to photoexcitable rhodopsin [24]. This is very different from the arrangement in rod and cone photopigments ("c-opsins"), in which the metarhodopsin is thermally unstable and quickly dissociates into free retinal and naked opsin apoprotein. Pigment regeneration requires multiple enzymatic steps, most of which are localized to the retinal pigment epithelium (RPE). Were melanopsin truly bistable, it could help to explain why ipRGCs retain photoresponsiveness for many hours in the absence of the RPE [5]. Indeed, photoreversal has been observed spectrophotometrically in melanopsin from amphioxus, a primitive chordate [30]. Several studies have shown that functional photopigment can be obtained by loading melanopsin with all-trans retinal [17, 38, 41, 52], presumably because the opsin can catalyze formation of cis retinal. Melyan et al. [38] also obtained evidence that exposure to relatively long wavelengths sensitized subsequent melanopsin-based photoresponses, presumably by driving a trans-to-cis isomerization of retinal chromophore and thus increasing the concentration of excitable melanopsin. We have recently observed electrophysiological evidence for photoreversal in ipRGCs themselves [Qiu and Berson, unpublished observations]. When a saturating light stimulus is extinguished, the light-evoked depolarization decays very gradually (over >2 min) toward prestimulus levels. This poststimulus persistence resembles the persistent depolarizing afterpotential (PDA) of rhabdomeric photoreceptors, a phenomenon attributable to a persistent pool of thermally stable metarhodopsin [24]. During the PDA in ipRGCs, exposure to yellow light (560 nm) drives the ipRGC membrane potential back toward baseline, just as it does in rhabdomeric photoreceptors, strongly suggesting that melanopsin is bistable in mammalian ipRGCs. There is at least one observation, however, in apparent conflict with the view that intrinsic photoisomerization of melanopsin's chromophore is primarily responsible for pigment regeneration in ipRGCs. Fu et al. [17] reported that knocking out RPE65, an essential retinal isomerohydrolase of the RPE, reduced ipRGC sensitivity as assessed indirectly from pupillary responses. This result appeared to suggest that melanopsin depended at least in part on an extrinsic enzymatic retinoid cascade for its normal function. However, more recent evidence indicates that this effect is secondary to the devastating effects of the knockout on rod and cone function ([59]; see also [13, 34]). In mice with degenerative loss of rods and cones, neither RPE65 deletion nor acute pharmacological disruption of the retinoid cycle has any significant impact on ipRGC sensitivity [59]. These results do not entirely exclude a retinoid cycle for ipRGCs localized within the inner retina and involving proteins other than melanopsin itself, but at present, there seems no strong evidence favoring such a mechanism. Nonetheless, it is worth noting that the poststimulus depolarization of ipRGCs decays spontaneously (if sluggishly) in darkness, after which ipRGC sensitivity returns to prestimulus levels ([5, 64]; Qiu and Berson, unpublished observations). These results suggest that photoexcited melanopsin is not indefinitely thermostable and that there may be light-independent pathways capable of regenerating photoexcitable melanopsin.
Phototransduction cascade: the role of G proteins
By comparison with the widespread consensus about the identity of the photopigment in ipRGCs, the current state of understanding of the signaling cascade linking melanopsin to the light-gated conductance can only be described as sketchy. Opsins are heptihelical G-protein coupled receptors (GPCRs), and photoisomerization triggers heterotrimeric G-protein activation in all well-characterized bilaterian phototransduction cascades. There is some evidence that this is true in ipRGCs [63], although the evidence is not decisive. Nonetheless, melanopsin can trigger G-protein activation in a number of different biochemical assays and cell systems [38, 40, 41, 52], and it seems very likely that it does so in ipRGCs.
If a G protein is in fact the first element downstream of melanopsin in the ipRGC phototransduction cascade, the identity of that G protein is likely to be very informative not only about other probable signaling components but also of the evolutionary origins of ipRGCs. Well-studied phototransduction cascades in the photoreceptors of bilaterian animals fall into one of two broad frameworks [1, 30, 48]. The first, shared by all ciliary photoreceptors (encompassing all vertebrate photoreceptors but ipRGCs), involves G proteins of the Gi/o class (including transducin) and cyclic nucleotides as second messengers. The second, characteristic of the rhabdomeric photoreceptors of invertebrate eyes, employs G proteins of the Gq class and phospholipase C (PLC) as the effector enzyme. As vertebrate photoreceptors that use a rhabdomeric opsin, ipRGCs might conform to either of these canonical cascades, or to neither.
Evidence against a ciliary, cyclic-nucleotide cascade
The use of Gi/o class G-proteins and cyclic nucleotides as second messengers appears highly conserved among ciliary photoreceptors. Such cascades are most familiar from the vertebrate rods and cones, in which photoexcited rhodopsin activates transducin, a G protein belonging to the Gi/o class. The liberated alpha subunit of transducin then activates cyclic guanosine monophosphate (cGMP)-specific phosphodiesterase (PDE). The resulting hydrolysis of cGMP reduces the cytosolic concentration of this cyclic nucleotide, closing cGMP-gated channels and attenuating the current that otherwise persists in darkness ("dark current"). The net result of cascade activation is thus membrane hyperpolarization. Other ciliary photoreceptors, such as those of the chicken pineal [28] and lizard parietal eye [56] and even those in invertebrate eyes [10], improvise on this basic theme, but the involvement of Gi/o class G proteins and cGMP appears to be highly conserved.
The only evidence that such a cascade might operate in ipRGCs is that melanopsin can mediate light-dependent activation of rod transducin alpha in a biochemical assay [40]. However, promiscuous interactions of GPCRs with a wide range of G proteins in biochemical assays are well established, so this capacity sheds little light on the native G protein in the ipRGC cascade. Indeed, ipRGC photosensitivity appears intact in knockout mice lacking rod transducin alpha [23]. Furthermore, robust ipRGC photoresponses have been recorded in the presence of antagonists of PDE ([63]; Carlson and Berson, unpublished observations). The persistence of ipRGC photosensitivity and melanopsin-mediated visual behaviors in rd mutant mice, e.g., [43, 55, 60], are also relevant here, as the rd phenotype results from a nonsense mutation in the gene coding for the PDEβ subunit [47]. With respect to the ion channel, it is clear from studies in knockout mice that at least the cone cyclic-nucleotide-gated (CNG) channel (CNG3) is not required for melanopsin-dependent light responses in ipRGCs [23]. Pharmacological evidence weighs against CNG channels being the light-gated channel in ipRGCs ([63]; Carlson and Berson, unpublished observations). Taken together, there is no compelling evidence that a cyclic-nucleotide-based signaling cascade like that in ciliary photoreceptors mediates ipRGC phototransduction.
Evidence for a rhabdomeric phosphoinositide cascade
In invertebrate rhabdomeric photoreceptors, phototransduction invariably involves opsin activation of a G protein of the Gq/11 family, resulting in the stimulation of PLCβ and hydrolysis of phosphatidylinositol (4,5)-bisphosphate (PIP2) [1, 12, 20, 29, 33, 58]. Less complete information is available about the downstream components of the cascade. Hydrolysis of PIP2 generates the membrane-associated second messenger diacylglycerol (DAG) and the cytosolic inositol 1,4,5 trisphosphate (IP3). Both have been implicated to some degree in rhabdomeric phototransduction. In Drosophila ommatidial photoreceptors, IP3 is not essential, and a membrane-delimited cascade involving DAG, polyunsaturated fatty acids, and/or PIP2 itself appear to be playing an essential role [20]. The light-gated channels have been well characterized and represent the original members of the transient receptor potential (TRP) channel proteins, although their gating mechanism remains uncertain. In Limulus ventral photoreceptors, IP3 acting at IP3 receptors to mobilize intracellular Ca2+ may play a significant role in some photoresponse components [12].
In ipRGC phototransduction, although a G protein is almost certainly an essential component, there is no direct evidence that this belongs to the Gq class nor that PLC serves as the effector enzyme. Indeed, there is no information available at all concerning which G proteins and PLC isoforms might be expressed in these cells. There is, however, indirect evidence for the plausibility of a Gq–PLC cascade in ipRGCs. Gq has been colocalized with melanopsin in amphioxus rhabdomeric photoreceptors [30]. There is compelling evidence that melanopsin can signal through Gq-class G proteins and PLC from other cellular systems. In amphibian dermal melanophores, in which melanopsin is the presumptive photopigment, light elevates intracellular IP3 levels, and PLC appears to be the effector enzyme for the light-driven pigment dispersion response [27]. Another indirect line of evidence comes from studies of chick RGCs, a small percentage of which express melanopsin [7]. In relatively pure cultures of embryonic chick ganglion cells, transcripts for Gq but not transducin have been identified [8]. In the same cultures, light suppresses the release of melatonin and this effect, which may be mediated by melanopsin, requires PLC. In two separate heterologous expression systems, mammalian melanopsin signals through a Gq-class G protein and PLC to generate the transmembrane photocurrent [41, 52]. Data from a third expression system was at least partly congruent with this picture, implicating a G protein that did not belong to the Gi/o class [38]. However, PLC appeared not to be the effector enzyme in this system, making it unlikely that the G protein in question belongs to the Gq class. Taken together, the available evidence builds a circumstantial case that melanopsin may be coupled to the light-gated channel in mammalian ipRGCs through a 'rhabdomeric' G-protein cascade. Key evidence on this point is lacking, however. For example, it remains to be shown that ipRGCs express any Gq-class G proteins or PLCβ. The most glaring evidentiary gap, however, is the absence of data from pharmacological, knockout, or knockdown studies implicating Gq-class G proteins, PLC, or associated downstream second messengers in ipRGC photoresponses. Our own efforts to block the cascade in intact retinas with antagonists of Gq-class G proteins and PLC have failed (Carlson and Berson, unpublished). Although this could spell trouble for the hypothesis that a rhabdomeric phosphoinositide cascade is mediating ipRGC light responses, it is possible that such negative results are attributable merely to technical limitations, such as poor access of pharmacological agents to the critical sites of transduction. There is a precedent for such failures in Drosophila photoreceptors, although there is no obvious equivalent in the ipRGCs for the sort of diffusion barriers that are presumably responsible for the negative pharmacological findings in flies.
Identity of the light-gated channel
The identity of the plasma membrane channels that mediate the ipRGC light response is not known, although recent evidence suggests TRP canonical (TRPC) channels as the most likely candidates ([41, 52, 63], Carlson and Berson, unpublished observations). Light induces a conductance increase in ipRGCs, and the photocurrent reverses near 0 mV, suggesting mediation by a nonspecific cationic channel. Ion substitution experiments have suggested that sodium contributes relatively little to the overall current [62], although in our hands, removal of extracellular Na+ substantially alters current–voltage relations [Carlson and Berson, unpublished observations]. In any case, Ca2+ appears to be a significant charge carrier through these channels. This is consistent with the findings of Sekaran et al. [55], who imaged intracellular Ca2+ in retinas largely lacking functional rods and cones. Light triggered a substantial increase in intracellular Ca2+ in a small population of ganglion cells (presumably melanopsin-expressing ipRGCs), and this increase was abolished by the replacement of extracellular Ca2+ with Cd2+ [55]. Although this would seem to support the hypothesis that the light-gated channels have substantial Ca2+ permeability, alternative interpretations are possible. For example, Ca2+ influx through voltage-gated Ca2+ channels (VGCCs) has not been excluded as a source for the recorded Ca2+ signal. VGCCs, which are widely expressed in RGCs, would presumably be activated by the light-evoked depolarization of ipRGCs and would have been blocked by extracellular Cd2+. Further, although transmembrane Ca2+ flux clearly plays an important role, it would be premature to exclude a role for intracellular Ca2+ mobilization because influx of Ca2+ across the plasma membrane can trigger a secondary release of Ca2+ from intracellular stores ('calcium-induced calcium release') in many cells. This would be in addition to any Ca2+ mobilization that might occur as a result of PLC activation and consequent liberation of Ca2+ from IP3-sensitive stores [32, 52].
The similarity of both the opsin and the presumptive signaling cascade in ipRGCs to those in rhabdomeric photoreceptors has encouraged speculation that the light-gated channel might belong to the TRP family. Originally identified in Drosophila rhabdomeric photoreceptors [39], these channel proteins are now recognized as forming a large family of tetrameric, calcium-permeable channels involved in a variety of functions ranging from sensory transduction to intracellular calcium homeostasis [53]. Of particular relevance to ipRGCs are the vertebrate TRP proteins of the TRPC family, as these bear the greatest resemblance to Drosophila TRP including in their gating by activation of GPCRs coupled to phosphoinositide cascades. Indeed, the current–voltage relations of the ipRGC photocurrent ([62]; Carlson and Berson, unpublished observations) resemble those of TRPC-mediated currents in some heterologous expression systems [25, 26, 53], with reversal near 0 mV and both inward and outward rectification. TRPC proteins form nonspecific cationic channels with substantial Ca2+ permeability, matching known features of the ipRGC light-activated channel. Pharmacological agents known to effectively block TRPC channels attenuate or abolish the ipRGC photocurrent ([63]; Carlson and Berson, unpublished observations), but none of these agents is specific for TRPC channels. In heterologous expression systems, melanopsin has been shown to be able to mediate light-triggered gating of TRPC channels [41, 52]. At least one TRPC protein (TRPC6) has been detected in ipRGCs by immunohistochemistry [63]. Thus, there is ample evidence to view TRPC channels as good candidates for the light-gated channel in ipRGCs. A stronger case awaits better pharmacological tools or evidence from genetic manipulation experiments that the expression of these channels is necessary for the ipRGC photoresponse. It will also be important to catalog the full range of TRPC proteins expressed in ipRGCs in view of the potential for formation of heteromultimeric channels [53].
The road ahead
Although substantial progress has been made toward identifying the phototransduction cascade in mammalian ganglion-cell photoreceptors, there are many glaring gaps in our understanding. The evidence that melanopsin forms the photopigment in these cells is overwhelming, although much remains to be understood about the specifics of its behavior under physiological conditions in ipRGCs. Of particular importance will be to establish the identity of its chromophore and to understand the relative contributions of intrinsic photoreversal and extrinsic enzymatic retinoid cycles in the regeneration of photoexcitable pigment. Also largely unexplored are mechanisms for response termination and adaptation, melanopsin's selectivity of interaction with a range of G-protein partners, and the identity of its intracellular residues contributing to those interactions.
As for downstream signaling, there is clearly mounting evidence for the involvement of a phosphoinositide signaling cascade resembling that in rhabdomeric photoreceptors. There may be a particularly close resemblance to the cascade worked out in Drosophila photoreceptors, inasmuch as the TRPC channels are emerging as the best candidates for the light-activated channels. However, there remain troubling gaps in the empirical foundation for this view, especially the inability so far to demonstrate that Gq-class G proteins or PLC are essential for ipRGC photoresponses. One likely barrier to obtaining such evidence is the difficulty of exerting sufficient pharmacological control over cells with transduction machinery localized in part to very fine dendritic processes deeply embedded in the retinal tissue. An important challenge for future work, then, is to develop methods for recording photoresponses from dissociated ipRGCs. Ultimately, obtaining definitive evidence on the nature of the cascade will probably require experimental manipulation of critical genes, either in mouse models or transfection studies.
One of the most exciting aspects of the emerging picture of the ipRGC phototransduction cascade is that it provides a critical test of the emerging hypothesis that ipRGCs and invertebrate rhabdomeric photoreceptors share a common evolutionary origin [1, 48]. If this proves to be true, it will be of interest to pursue comparative studies to learn which aspects of the ipRGC response to light are inherent in the highly conserved signaling cascade on which their photosensitivity is based and which are adaptations to their particular functional roles in mammalian vision.


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Media Arts and Science Bachelor of Science
The focus of Media Arts and Science is on creativity and digital storytelling while enhancing contemporary career-building skills. Our program offers education and experience on various integrated media venues in today's convergent society. We strive to produce dynamic websites, smartphone apps, interactive education, serious and entertaining games and simulations, 3-D motion graphics, digital illustrations and animations, and audio and video.
Our program is designed to be flexible, with ample opportunity to concentrate or broaden your studies in one or more specialty areas. Some students develop a passion for a particular specialty, while others aim to become mixed-media virtuosos. Our specialty areas include:
- 3-D Graphics and Animation
- Digital Storytelling
- Game Design and Development
- Video Production and Sound Design
- Web Design and Development
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Media Arts Careers
Our graduates design and develop projects that focus on entertainment, serious game design, mobile applications, video and audio production, animation, and innovative web design and technologies.
Media Arts and Science graduates work for many big name companies such as Pixar, Cantina Creative, ChaCha, Blizzard Entertainment, ExactTarget, Pacers Sports & Media, Wiley and Sons (Publishers of Dummies books), Tour Design (Livenation), Emmis Publications, USA Gymnastics, Capital Records, BET – Black Entertainment Television, and Girls, Inc. Positions at these companies range from UX designers, to 3-D modelers to project managers and design leads.
Why study at SOIC?
- You'll have access to excellent faculty and state-of-the-art facilities, including: the Media Arts Research and Learning Arcade, the Advanced Visualization Lab and a virtual reality theater.
- You may customize your degree with specialized courses helping you achieve your career goals.
- You can finish your education with a portfolio of work showcasing your skills to employers.
- SOIC offers a myriad of opportunities to utilize the skills you develop in real-world applications, not only through research but through collaborative projects done outside of the classroom and in various student organizations.
- The School of Informatics and Computing is regionally accredited as a school of Indiana University by the North Central Association of Colleges and Schools.
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To pursue a degree in Media Arts and Science, you must first apply and be accepted to IUPUI. You'll then want to sit down with one of our undergraduate advisors to review the program's plan of study and find out what courses to take.
You'll likely start with N100 Foundations of New Media, where you'll be introduced to the evolution of digital media and begin exploring your particular interest areas with hands-on projects.
Specialty Areas
From there, you'll take additional core courses and electives in your chosen specialty area(s), becoming fluent in the use of contemporary media tools and project management principles. A specialty area is your opportunity to customize your education in those aspects of media and production best-suited for your career goals.
Capstone Project
With your skills sets firmly in place, you'll design and complete a faculty-mentored capstone project during your final semester. Your capstone project signals your readiness to graduate and represents the culmination of your knowledge and skill within your chosen specialty area(s).
And as a graduating senior, you'll showcase that education and passion to your fellow classmates, family, faculty, alumni, and visiting employers during the School of Informatics and Computing Capstone Event held at the close of each semester.
Learn more about upcoming Capstone Events you can attend, as well as examples of past student capstones.
Career Development
Before you graduate, you'll work with our Career Services Office to learn how to job search, assemble a portfolio of your work, pursue internship opportunities, and connect with employers.


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Tim hawkins
Udforsk beslægtede emner
i went to see him live with my aunt pam in Dothan . he is my fav comedian.
Tim hawkins
trying to cut down on the old cuss words.... this is for you!!!
lol everytime I go for an oil change they try to tell me 5+ things that they found wrong
funny tim hawkins quotes - Google Search
Tim Hawkins. Oh yes, belive everything the cereal box tells you. Who else will give you the important guidance you need about which cereals are 'magical'??
One does not simple watch tim Hawkins without laughing.. So hard that you can't breathe at points up to five minutes at a time
Seriously, if you have not seen this guy go do it!! His facial expressions are THE best.
"Christian cuss words" a fun list of things you don't have to bleep/delete.
tim hawkins funny quotes - Google Search


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List all words ending with alienly
Only the word alienly was found, nothing longer.
Some random words: he amin wrick ufological eohippus oka ebullience
This is not a dictionary, it's a word game wordfinder.
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-- Students --
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Group A – Vulnerable Bursary of *£1,200 is for those of you who are in receipt of income support/universal credit (in your own name); children looked after (in care); care leavers and disabled young people in receipt of employment support allowance and those of you who are also in receipt of disability living allowance/personal independence payments.
Additional travel subsidies will be made available when it is assessed that travel costs could be a barrier to your attendance. This could be in the form of a free bus pass.
Group B - Oaklands Discretionary Learner Support Fund (DLSF) Bursary is for those of you whose family is in receipt of any or all of the following benefits: Income support; Universal credit; Employment Support Allowance; Job Seekers Allowance; Pension Credit Guarantee; Working Tax Credit and have an annual gross income of no more than £26,600; Housing Benefit; Council Tax Benefit (excluding single person allowance) or on a family income which is below £26,600.
Group C – Oaklands Emergency Funding is for those of you that are in a temporary situation which is of a personal urgency or are experiencing a change of circumstances. This award is discretionary and will be based on an individual basis.
Residential Award – Is for those of you 16-18 year olds studying on one of our Sports or Land Based specialist courses, and moving/studying away from home.
At Oaklands we do our best to make learning affordable for you, therefore if you are struggling financially please discuss any of your queries or concerns about coming to college with our Student Advice Team.
For further information about all fees and to find out what financial support may be available for you, please contact our student advisers via an email to firstname.lastname@example.com or call us on 01727 737000.
*Fees subject to change.


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A Skull with Flowing Hair
Forever goes by. Ryan paces back and forth at the edge of the roof, looking to the sky. He yells every once in a while in the hope that Sparky will hear. But the only sounds I catch are my heartbeat and the sounds of the Jacks.
Over and over, there are screams echoing from the building. It sends shivers down my spine. It's definitely Janice. I can't tell if she's screaming out of fear or pain or anything, but when it starts to keep going on forever, I'm sure it's out of terror.
The sun starts to set, and still Spark hasn't showed. Ryan just stands with his back to the sun, creating a lovely outline if it weren't for the sick fear gripping me.
He looks at me, then whispers, "It's quiet."
He's right. The Jacks are barely making any sound at all. Janice isn't screaming. Neither are the kids. I wonder if they're gone. If the Jacks gave up.
The silence is broken again by another scream, and the sound of metal being torn. I know immediately what's going on and cry out, "They're tearing down the door! Ryan! We have to help them!"
He just slumps to the ground, head in his hands. So this is it. He's giving up. Letting the Jacks win.
I crawl to his side, wrap my arms around him, and sob into his shirt. There's nothing else for me to do. The freezer is being ripped to shreds, Janice is pleading for help, and in a matter of minutes, the screaming is silenced. I shudder as I imagine the Jacks down there, fighting over their bodies. Blood on the floor, pooling up.
Ryan and I are frozen. He whispers quietly, "It's my fault... it's my fault... oh good grace this is my fault..." I can't comfort him. I'm cold down to my bones. I bring my knees to my chest and rest my forehead on them. I can't take anymore.
A few minutes pass.
I don't respond.
"Sparky. ******* bird."
No words come from my mouth.
"YOU'RE THE BIGGEST PILE OF CHICKEN CRAP I HAVE EVER LAID MY EYES ON SPARKY!" he screams. I jump, then look up.
There she is. Perched carefully on the side of the building. She looks at us sadly, knowing she's done wrong. Her large violet eyes gaze at Ryan, apologizing.
He stands up and strides to her, furious. He punches the giant bird in the side of her face. She must be too large for that to hurt, but she chirps in soft pain anyways.
"You're a stupid bird. I can't believe you left. Did you not hear? You must have known that the Jacks were coming, you stupid beast," he punches her again, and she doesn't fight him. "You're a piece of crap. You better hope they're alive down there, or I'll kill you."
He punches the creature again, and she whimpers patheticly. Then she jumps down and crawls into the building through the front doors. It's quiet, followed by several minutes of fighting, roaring, hissing.
Silence again.
Sparky's head sticks through the ceiling door behind us, purring. There's blood on her beak, but she seems to be unharmed.
"Son of a gun," Ryan says, walking over to stroke the place where he had struck her, "Did you kill them?"
She purrs a little louder, then glanced back down.
"Oh, Ryan! What if they're alive? We need to go down there."
Ryan nods, wrapps his arms around Sparky's neck, and she lowers her head down, and comes back up. I follow Ryan's lead, and when I'm down there, I go numb.
The first thing I see is blood. And the Jacks laying in it. One of the dead Jacks has a skull in its mouth. The face is gone, but the hair flowing from it... Janice.
Ryan is completely silent. He walks among the bodies. I know he's seen Janice's head. Tears flow from his eyes.
I walk behind him. There's a tiny torso on the ground. It's stained with blood, so I can't tell whose it is. Pieces of Jacks and Janice and the child are everywhere. I go to the freezing, praying with all my might.
There's whimpering in the back of the freezer. I numbly pull a box away. Joseph sits there, untouched but in shock.
He's the only surviver.


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Shopping Options
Ken Wilber
Ken Wilber is the author of over twenty books. He is the founder of Integral Institute, a think-tank for studying integral theory and practice, with outreach through local and online communities such as Integral Education Network, Integral Training, and Integral Spiritual Center.
Please also see the Integral Theory Primer for a great guide to the world of Ken Wilber and Integral Theory
Photographer: Julie Harris, email@example.com, 720.938.7775
Books & Audio
A Sociable God
Toward a New Understanding of Religion
In one of the first attempts to bring an integral dimension to sociology, Ken Wilber introduces a system of reliable methods by which to make testable judgments of the authenticity of any religious movement. A Sociable God is… Read More
A Sociable GodPaperback
The Simple Feeling of Being
Embracing Your True Nature
"As you look deeply into your own awareness, and relax the self-contraction, and dissolve into the empty ground of your own primordial experience, the simple feeling of Being—right now, right here—is it not obvious all at once? Were you not… Read More
The Simple Feeling of BeingPaperback
A Novel That Will Set You Free
Ken Wilber's latest book is a daring departure from his previous writings—a highly original work of fiction that combines brilliant scholarship with tongue-in-cheek storytelling to present the integral approach to human development that he expounded in more conventional terms in… Read More
The Eye of Spirit
An Integral Vision for a World Gone Slightly Mad
In this groundbreaking book, Ken Wilber uses his widely acknowledged "spectrum of consciousness" model to completely rewrite our approach to such important fields as psychology, spirituality, anthropology, cultural studies, art and literary theory, ecology, feminism, and planetary transformation. What would… Read More
The Eye of SpiritPaperback
A Theory of Everything
An Integral Vision for Business, Politics, Science, and Spirituality
Here is a concise, comprehensive overview of Wilber's revolutionary thought and its application in today's world. In A Theory of Everything, Wilber uses clear, nontechnical language to present complex, cutting-edge theories that integrate the realms of body, mind,… Read More
A Theory of EverythingPaperback
Quantum Questions
Mystical Writings of the World's Greatest Physicists
- edited by
- Ken Wilber
Here is a collection of writings that bridges the gap between science and religion. Quantum Questions collects the mystical writings of each of the major physicists involved in the discovery of quantum physics and relativity, including Albert Einstein, Werner… Read More
Quantum QuestionsPaperback
A Brief History of Everything
Revised Edition
A Brief History of Everything is an altogether friendly and accessible account of men and women's place in a universe of sex, soul, and spirit, written by an author of whom New York Times reporter Tony Schwartz… Read More
A Brief History of EverythingPaperback
Eye to Eye
The Quest for the New Paradigm
In this book Wilber presents a model of consciousness that encompasses empirical, psychological, and spiritual modes of understanding. Wilber examines three realms of knowledge: the empirical realm of the senses, the rational realm of the mind, and the contemplative realm… Read More
Eye to EyePaperback
No Boundary
Eastern and Western Approaches to Personal Growth
A simple yet comprehensive guide to the types of psychologies and therapies available from Eastern and Western sources. Each chapter includes a specific exercise designed to help the reader understand the nature and practice of the specific therapies. Wilber presents… Read More
No BoundaryPaperback
Sex, Ecology, Spirituality
The Spirit of Evolution
In this tour de force of scholarship and vision, Ken Wilber traces the course of evolution from matter to life to mind and describes the common patterns that evolution takes in all three of these domains. From the emergence of… Read More
Sex, Ecology, SpiritualityPaperback
One Taste
Daily Reflections on Integral Spirituality
As one who has written extensively about the interior life, meditation, and psychotherapy, Ken Wilber—the leading theorist in the field of integral psychology—naturally arouses the curiosity of his numerous readers. In response to this curiosity, this one-year diary not only… Read More
One TastePaperback
The Essential Ken Wilber
An Introductory Reader
Ever since the publication of his first book, The Spectrum of Consciousness, written when he was twenty-three, Ken Wilber has been identified as the most comprehensive philosophical thinker of our times. This introductory sampler, designed to acquaint newcomers… Read More
The Essential Ken WilberPaperback
The Collected Works of Ken Wilber: Volume Eight
The Marriage of Sense and Soul, One Taste
Volume Eight of The Collected Works of Ken Wilber includes: In The Marriage of Sense and Soul: Integrating Science and Religion (1998), Wilber takes on the centuries-old problem of the relationship between science and religion.… Read More
The Collected Works of Ken Wilber: Volume EightPaperback
The Collected Works of Ken Wilber: Volume Five
Grace and Grit
Grace and Grit is the compelling story of the five-year journey of Ken Wilber and his wife, Treya Killam Wilber, through Treya's diagnosis of breast cancer, treatments, and finally, death. During this period, Ken put his own work on… Read More
The Collected Works of Ken Wilber: Volume FivePaperback
The Collected Works of Ken Wilber: Volume Four
Integral Psychology, Transformations of Consciousness, Selected Essays
Volume Four of The Collected Works of Ken Wilber includes: Integral Psychology, a concise version of Wilber's long-awaited textbook of transpersonal psychology, presenting one of the first truly integrative models of consciousness, psychology, and therapy.… Read More
The Collected Works of Ken Wilber: Volume FourPaperback
About the Author
Ken Wilber is the author of over twenty books. He is the founder of Integral Institute, a think-tank for studying integral theory and practice, with outreach through local and online communities such as Integral Education Network, Integral Training, and Integral Spiritual Center.
Please also see the Integral Theory Primer for a guide to the world of Ken Wilber and Integral Theory.
Videos & More
It's Just Love (Ken Wilber: ISE 2 Keynote) from Integral Life on Vimeo.
Ken Wilber delivers his Keynote Address at the second annual Integral Spiritual Experience practice event.
Want to see more? Check out the Integral Spiritual Experience Media Collection here: IntegralLife.com/ISE2-Love


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Caretta Beach Hotel
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So this is what comes from Watching too much Mama
... OK! Enough of the mama Mia wedding now.
So the long and the short of it is that Dyana & I only had one day to ourselves. Free from all the pomp and everything. So, as Frederick Forsythe the famous author ended one of his books with the phrase, 'Let loose, the dogs of war,' at last……. we were free to do what we normally do…… PARTY! And have a good time exploring. Yes!
Now, let me ...
French Castle in Greece
North of Olympia we find Clermont (Chelmoutsi) Castle, built in 1223 by a French Prince Geoffrey, after the Crusades when he claimed the region for himself. The castle was built to the highest standard of its time and still most impressive. Only taking 3 years to build, it is taking a lot more time and probably money (EU money) to restore. However it is one of ...
Olympic champions
... olympia
went to Archaeological site. Saw temple of Hera. Zeus. Athlete training grounds and athlete hangout. Visited stadium where first Olympic Games took place. Stood on marble start line and took off to race. Soaked up the spirit of the Olympic champions.
Dis is some shopping in Olympia and had lunch. Spanokapita ouzo. Gyros and Greek pizza
on return train was offered wine tasting from local winery
Sailing from Santorini and a day in Katakolo
... with the restaurant manager of Corriander, would you believe he is called Damian!!!). I will ask her for details for the wine lovers! Truly a good match. Dinner was Papadums with a selection of yoghurts, raita, hot spice and pickles. Next shared appetizers - lamb kebabs, onion pakora, tandoori chicken, spiced vegetables. Mains (from a long selection) chicken muggu, (Mary Ann), lamb jalfrese (David), shrimp and fish curry (chefs choice), spinach, curried lentils, spiced corn, ...
A Monday, a Monday is very, very good
... The swim was preceded by a short walk along the beach, a foray into some of the boutique shops and a glass of cold lemonade at a beachside caf�. Philip and I were enjoying cooling down under the shade, and idly chatting with a Welsh couple doing the same thing, when an old Holden came burning up and out jumped a couple of young Greeks.
They each had a musical instrument, an oud (I think) and a drum, and they came straight to our table and ...


text: State Police are investigating the deaths of a man and woman whose bodies were found inside a house in Webster Plantation, about 20 miles east of the town of Lincoln. The bodies were discovered Saturday afternoon in a mobile home on the Tucker Ridge Road, which is believed to be the residence of the couple.The bodies were transported late Saturday night to the State Medical Examiner's Office in Augusta for autopsies, which began late Sunday morning. The identities of the couple, believed to be in their late 40s, are being withheld pending the completion of the autopsies.Â


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1970s automotive zombies - CAFE comes again?
A recent appeals court decision, first reported in Detroit News, upholding the new Corporate Average Fuel Economy (CAFE) standard, means a stricter set of fuel economy rules are now in place. The first time this happened, it was the 1970s and it resulted in models whose names were legendary squandering their infamy on choked down excuses for scrap metal or, worse, became re-badged foreign imports.
The 1970s were an odd time in American history. It was a time that saw the hippies emerge from counter-culture into disco dweebs and the muscular pony give way to the limp noodle. When the newly-minted Environmental Protection Agency's CAFE standards began in 1975, American automakers rushed to comply with the strict rules meant to get us off foreign oil imports and force fuel efficiency into a culture that, up to that point, had only rarely considered it.
To meet requirements, the Detroit 3 all downsized in a big way, compromising everything in the name of compliance. These are the automotive zombies created during that time.
1973 Datsun B210
We start the list with an import car that set the stage for the atrocities to come. The B210 had two things going for it: the OPEC gasoline shortage and a cheap price tag. Everything about this car was horrible. Its chief attribute was its ability to rust in even the driest of environments thanks to the paper-thin metal that made up its bodywork. Spilling coffee on the floor virtually guaranteed a hole in the pan by midday.
1975 AMC Pacer
Nearly forty years later, the name "Pacer" still means "loser" in automotive parlance. Possibly the ugliest car ever made, the Flying Fishbowl still appears to be a model for "smart" vehicles today, re-emerging in today's lame duck "efficiency" models just as it did then. Party on, Wayne.
1975 Chevrolet Corvette
Growing into a real sports car with refinement, the Corvette entered a hiccup in its 1975 emissions-regulated edition. Its engine was strangled down to 165 horsepower for its base option and its best offering (the L-82) had only 205 horsepower. Despite this, though, the '75 'Vette was still respectably powerful and exceptionally fast. The fact that it looked cool didn't hurt.
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1975 Chevrolet Monza
This car was hit-and-miss with owners. The stock engine was a 78 horsepower 2.3-liter, 4-cylinder sputtering box. The original Wankel Rotary Engine to be showcased in this car was (rightfully) scrapped at the last minute.
Please SHARE with friends and include TorqueNews in Google Alerts for tomorrow's interesting stories.


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