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Posted by ESPN.com's Mike Sando
Paul from Phoenix writes: Sando, I think you need to give the AZ fans another 5 years to build a loyal ticket buying fan base and I want to emphasis that there is a difference between a loyal ticket buying fan base and a plain old fan base.
I am 25 years old and probably at the old edge of Arizona natives who grew up with the Cardinals in the Phoenix area. Anyone much older than me probably had time to develop a fondness for another team outside of the state, and the Cardinals haven't been good enough over the last 20 years to steal any fans from other teams.
Anyone younger is probably feeling the economic crunch just as bad as I am. (25 year olds as a rule do not have a lot of money to spend on playoff tickets.) I am a loyal Cardinal fan and I hope to make it to the game but I can understand how anyone in my position or in even a slightly tougher economic spot would be smart to cut a luxury item (I don't care how much of a bargain everyone says Cardinal tickets are) out of the budget.
Give me 5-7 years to start earning a salary worthy of season and playoff tickets and Ill be there every week. Right now Im going to focus on buying Christmas presents and meeting essential needs for my family while I watch the games on tv with my jersey on.
When the Arizona-native Cardinal fans like me hit their 30s and 40s, you can start looking for a serious increase in ticket buying diehard fans. Ill probably even bring my son and start a whole new generation of Redbirds.
Mike Sando: That makes a lot of sense. The Cardinals moved to Arizona for the 1988 season. You were roughly 5 years old at that time. Anyone even 5 years older than you would have had time to develop an allegiance to another team. And with the Cardinals struggling so profoundly, Arizona natives had little reason to latch on. The area also features quite a few transplanted residents who developed allegiances elsewhere.
I was 25 years old when the Raiders moved back to Oakland. I had grown up in Northern California. I naively bought into the initial hype predicting long waiting lists if fans didn't act quickly to purchase tickets. I could not afford two tickets. I could not really afford one ticket, but my former neighbor and I each purchased one ticket costing about $50 per game to sit near the front of the third deck. I watched a lot of bad football from that seat.
Mark from Olympia writes: Since the NFC West is basically just watching for the draft now...what positions are each of the teams looking for in the upcoming draft and what players could you see fulfilling those places?
Mike Sando: I'm heading to Phoenix on Friday for some football, so we're not quite in draft mode all the way, but the positional needs seem pretty well established. I wouldn't guess on specific players yet given that we don't even have a coaching staff in St. Louis, we're still expecting staff changes in Seattle and San Francisco, and the combine is still weeks away.
If draft needs match available talent, I would give the Rams a franchise offensive lineman second overall. The Seahawks would get a pass-rusher fourth overall. The 49ers would get a right tackle or defensive player at just about any position. The Cardinals would find a starter for their offensive line or possibly a linebacker to give them flexibility should Karlos Dansby leave in free agency.
Jared from Bend, Ore., writes: Sando, wouldn't Al Saunders be a logical new OC in SF? Of course I'm assuming he wouldn't be retained by St. Louis, but he's a Coryell disciple (like Turner) and plays a similar system. He also has Bay Area ties with San Jose State and Stanford. Do you think he will become available and if so, do you think he would be a fit in SF?
Mike Sando: Good idea. Also, Saunders would know the NFC West quite well. I'm not sure what will happen to Saunders in St. Louis, but that might be a pretty good fit if he became available. He is an established coach, too.
Jeremy from Portales, N.M., writes: Sando, Has there been any word on Staff changes for the Seahawks? I figured Mora jr would bring in his own people as soon as possible. Thanks
Mike Sando: Nothing yet. Mike Holmgren is still in the building this week. I'm thinking we'll see more movement once the dust settles a little bit.
Doug from Garden State writes: Hey Sando great job this season with the blog. Quick question I was wondering if you know who is on the list of potential OC for my niners and if Singletary plans to hire a WCO guy?
Mike Sando: Thanks, Doug. My understanding is that the 49ers have a list, and that Scott Linehan's name is on that list, but that doesn't necessarily make him a favorite. I know of no preference for a West Coast offense guy.
Chad from Oshkosh writes: Sando, Do you think there is any regret that the Niners pulled the trigger so quick on Singletary now that Shanahan is available? Let me go on the record as saying I have no problem with hiring Singletary after the way the team rallied around him and I thought those that poo-pooed the decision as having been made too quickly were just complaining to hear themselves talk.
However, now that Shanahan is on the market I can't help but wonder if the team wouldn't have been better served to wait and at least interview Shanahan. Perhaps he wouldn't have been interested, but he has ties to the bay area and has a history of success. Plus it may have brought some consistancy to the offense which has been lacking.
Mike Sando: I think moving so quickly on Singletary left the 49ers vulnerable to criticism on the matter, and that the criticism is founded even if Shanahan wouldn't have taken the job. The organization could have scheduled a meeting with Singletary for Monday or Tuesday with full intentions of naming him the head coach at that time. I see no tactical advantage in naming Singletary the coach minutes after the final regular-season game. He wasn't going to take a job elsewhere in the first 48 hours after the final game.
On the other hand, if Singletary was going to be their choice no matter what, then nothing was lost by hiring him at that time.
Mpier from Tucson writes: At the Cardinals and Seahawks game on the last face-masking penalty of the game, I believe it was the Seahawks last scoring drive, it appeared that the runner grabbed his own faskmask on the replay. There was a facemask penality called a couple of plays prior and it appeared legitimate, but on this play it looked like a good open field tackle. When they showed it on the jumbotron they stadium booed so I thought the saw it too. I have not seen any replays of this so I am wondering if I really saw what I thought I saw.
Mike Sando: You saw it the way others saw it. A league official in the press box wondered if the runner had been suggesting a facemask occurred earlier in the play, but the referee specifically said the facemask occurred at the end of the run.
Matt from Scottsdale, Ariz., writes: Sando, You seem like a great numbers/breakdown kind of guy. Would you be a good fit for a advance scout for an NFL team? Assume your family was cool with everything and had no impact on your decision, would you take a scouting job with an NFL team? Would the team interested in hiring you sway your decision? If not a scout, do you think there is a position with a team that would better suit your skillset?
Mike Sando: I don't know football well enough to do that sort of job well. I would have loved that sort of job in my 20s but, as noted, I would not have been qualified, then or now. I do enjoy the analysis stuff, and so
me NFL people I know humor me by looking at it, but that's as far as it goes.


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Doctor insights on:
How Many Days After Ovulation Does Period Start
What is the cause breast soreness after 4 days post ovulation period?
hormones: After you ovulate your body produces progesterone. This is the hormone that causes increased breast tenderness before your period. ...Read more
Ovulation (Definition)
Ovulation is the act of the ovary releasing the egg after it has been hormonally prepared, to be ...Read more
I am 16 days post ovulation. I took a pregnancy test this morning and it was negative. I have no pms sysmptoms like I normally would before a period. the only symptoms I have had this month were for 5 days after ovulation. severe bloating, cramping, dizzi
After missed period: Conception, as defined by the generation of a fertilized ovum, occurs within day or so of the sex. However, it takes about two weeks for the embryo to make enough HCG to produce a positive pregnancy test. Blood test may be done a day after a missed period. Home pregnancy tests should be done 4-5 days after the missed period. If the result is negative repeat the test in 4-5 days. Use first morning urine and follow instructions for the test carefully. If you do not wish to be pregnant, use contraception all the time, every time. You may consider implanted contraceptive, or IUD. Practice safe sex. ...Read more
Cycle usually 21-28 days. I am on cycle day 30(20 days post ovulation), had spotting on cycle day 20, have had 2 negative tests but still no period ?
Wait 10-14 days and: repeat the test. You might get your period by then, or if not maybe the test is then positive. If still no period and no positive tests, either wait another 2 weeks or see a doctor for more detailed testing to be done, usually a blood test. ...Read more
I had a number of positive pregnancy tests very early - 7 days before my missed period, 8 days post ovulation. Can this be right?
Unlikely: If you miss your period by 4-5 days, do a home pregnancy test. Use first morning urine and follow the instructions for the test carefully. Given the aberrant earlier results, it may be prudent to see your doctor for a blood test for HCG level. ...Read more
How many days after ovulation do you get your period?
12-14 days: After ovulation the follicle (egg sac of the ovary) is called a corpus luteum. It produces Progesterone to change the uterine lining to support implantation. If you are not pregnant the corpus luteum will die off in 12-14 days. If you are pregnant the HCG produced by the placenta keeps the Progesterone going to support the pregnancy. ...Read more
How many days after ovulation should I get my period?
Usually two weeks: Menses will usually be about 14 days after ovulation--but this is variable between individuals. ...Read more
How many days after ovulation should your period come?Tested positive for ovulation on 11th & tried on 11th-13th.Last period was on 24/12.Ihave cramps
Ovulation normally: Occurs 14 days prior to your menstrual period. Test for pregnancy if you miss your period. Take care. ...Read more
How many days after ovulation should your period come?I tested positive on 11/1 and tried on 11th-14th. Last period was 24/7. I have a lot of cramps
Please Clarify: Your last period was in july? And you have not been to see your gynecologist or ob? Your question is incomprehensible with these dates. ...Read more
How many days after ovulation do most early pregnancy symptoms begin to occur besides missed period?
Varies : This varies from person to person I have reports of breast tenderness occurring within a day or 2 of conception. It is hard to gauge as most of it is so subjective. ...Read more
Started spotting light pink day after ovulation, it hasn't stopped and its been 2.5 weeks started brown now pink, and now period is 3 days late?
Are you pregnant?: You certainly need a pregnancy test if you haven't had one yet. If you're not pregnant, the spotting is not normal and needs to be investigated by a qualified doctor. ...Read more
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movies, A lot of em, i tend to stay away from the dumbed down for the masses stuff, but im not above it.
music... any with soul, Punk , texas country, metal, rockabilly, electronic stuff, blues, classical ( yead i do go to the symphony too), grassroots hip hop. wtf ever as long as i can feel it.
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text: m_k Play via YouTube OPEN IN APP NSFW Safe Is this playlist safe for work? Pump It April 20, 2012 | 16 tracks | 2,043 78 | Stats EminemLil WayneDephonic workoutgymfitnessrap & hip hophiphop Liked Collect Share Edit celebrate Liked! Share this mix with friends: Pump It Wrong video Collect Share Liked! Share this mix with friends:


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Writing Committee drafts for the 15th National Congress on School Transportation (NCST) have been posted on the NCST Website, www.ncstonline.org. To view the drafts, click the "Writing Committees" link on the home page.
All comments, requests for additional changes or modifications, or consideration of new proposals are submitted to the respective Steering Committee coordinator shown on the Website. Coordinators are responsible for forwarding comments to the appropriate writing committee chairperson.
Feb. 19, 2010, is the deadline to submit comments to coordinators.
To view information on the activities scheduled for the 15th NCST, to be held May 16-19 in Warrensburg, Mo., click the "Congress Activities" link on the Website. The page includes the congress agenda and information on lodging in Warrensburg and surrounding communities.
Registration information is scheduled to become available in early January.
Get the latest news and most popular articles from SBF delivered straight to your inbox. Stay on top of the school bus industry and don't miss a thing!


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Adolf Hitler
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Of massacres and mistresses: Himmler diaries reveal chilling details of life under Nazi rule
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Uneven Structure stream album
Love this band. Can't wait for the album.
This is pretty cool. Really dig the heavy ambient influence. Definitely feels like an album that you'll appreciate more the more you listen to it as well.
not bad
I hear great things about this new album
Digging: Edan - Beauty and the Beat
I hear great things about this new album too, but I don't hear great things when I actually listen to it.
This is relevant to my interests, I think I'll listen now.
"I hear great things about this new album too, but I don't hear great things when I actually listen to it."
I've only heard it twice so far but it seems like a pretty subtle album. The focus isn't really on the riffs but more on the atmosphere and progression of the whole thing. Some people will like that more than others for sure.
yea the ambiance basically takes the front stage along side the grooves so their pretty much balanced. Def. a unique mix of post-metal and groove metal-makes for a pretty intense listening experience.
Digging: Artifex Pereo - Passengers
It's simpler than that really. Out of the massive djent (ughthatname.) scene there is, this band does nothing to carve out it's own identity or build something that would make you want to keep coming back and spin the record.
I haven't personally heard a djent album that sounds like this. The closest thing is Tesseract but the ambiance on this is dialed way, way up. Also, it has a much more carnal, powerful sound whereas Tesseract is more ethereal and melody-centric.
"I haven't personally heard a djent album that sounds like this. The closest thing is Tesseract but the ambiance on this is dialed way, way up. Also, it has a much more carnal, powerful sound whereas Tesseract is more ethereal and melody-centric."
Wow. This is the same band? holy shit lol this sounds awesome
Sounds pretty damn good. I also agree fully with Kubrick and his last post.
this might be my aoty. so fucking good.
At last!!
Called it.
holy shit this rules. dat atmosphere
"Wow. This is the same band?"
Was thinking the same thing first time I heard it. I listened to their EP a while back and I don't even remember it. This on the other hand grabbed me almost immediately. I really feel this has the potential to grow into one of my favorite albums in this style. I already really enjoy it and it doesn't feel like a flimsy, instant gratification kind of enjoyment.
Great album so far. Needs a few more listens, but it might make my end of year list.
uneven structure vs amia venera landscape vs comity.... i love my life
Digging: Norma Jean - Polar Similar
btw doesnt anyone else think that tesseract is like mudvayne?
toss out the cabbage.... im ready... but at least ill have integrity when ill speak to my friends about it in the real world...
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Lemon Chicken Fettuccine With Snow Peas
Total Time
Prep 5 mins
Cook 20 mins
Clipped out of a magazine, this is a fresh chicken recipe that is ready in a flash!
Ingredients Nutrition
- 12 ounces fettuccine
- 1 tablespoon vegetable oil
- 1 lb boneless skinless chicken breast
- 1 garlic clove
- 1 1⁄4 cups chicken broth
- 2 tablespoons lemon juice
- 1 tablespoon cornstarch
- 1⁄4 teaspoon salt
- 1⁄4 teaspoon pepper
- 2 cups snow peas (fresh is best)
- 2 tablespoons chopped fresh parsley
- Cut chicken into 1" pieces. Mince the garlic.
- Cook pasta according to package directions. Drain and keep warm.
- Heat oil in a large skillet over medium-high heat. Add chicken and garlic and cook, stirring frequently, for about 5 minutes until browned. Reduce heat to medium.
- In a bowl, whisk together the broth, lemon juice, cornstarch, salt and pepper. Add to chicken in skillet. Heat to boiling; reduce heat and simmer about 2 minutes, until thickened, stirring frequently.
- Add snow peas and cook 3 minutes longer.
- Toss hot pasta with sauce and garnish with parsley.
Most Helpful
Photo may be seen here (as Fettuccine With Snow Peas) in the Savor gallery: www.annacia.com. What a delicious recipe. I did skip the chicken but made all else by the recipe and the pasta was so yummy, I used the fresh snow peas (tagged this recipe to use some of them :D) and you wouldn't believe how good this simple sauce is until you taste it. For the protein I used stuffed crab in it's shell (had a lot of chicken lately, lol). The pairing worked beautifully and dinner was very well enjoyed. :D
Unfortunately, I didn't have fresh snow peas. I love them but had to use frozen. This was a yummy recipe! We enjoyed it alot. I doubled the lemon juice and the garlic. YUM! I agree with Deantini- this recipe tastes even better than you'd expect it to from looking at the simple list of ingredients. Thanks loof.
Delicious dinner! I used snow peas from the garden and they tasted fabulous with the light lemon flavor. The only changes I made were to double the garlic and add red pepper for color. Quick, easy, and tasty - what's not to love?


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A Community of Catholic Youth Ministry Leaders.
Support. Training. Collaboration.
Support. Training. Collaboration.
WCACYM is not just an association of those in Youth Ministry in Western Canada. We are a really large, quirky, can't-live-without-them extended family. Whether you are a paid youth ministry coordinator, educator, volunteer, financial supporter, or a prayer warrior for youth, we all have a desire to see our youth know Christ. That makes us a family!
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Come, Sabbath.
WCACYM Fall Retreat on Beautiful Vancouver Island, BC.
October 7-9th, 2016


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When: Fri., Oct. 29, 8 p.m. 2010
Let Malapropos turn your wine into jazz juice when the jazz fusion group (featuring members of Kung Fu Kongress and Reverend Slanky) plays the Missoula Winery, 5646 W. Harrier, at 8 PM. $5. Call 830-3296.
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Sharp XVZ-30000 home theatre projector
This home cinema projector handles 2D and 3D with aplomb
- Excellent 2D and 3D video quality
- Powered zoom for flexible setup
- 2D setting on glasses to remove 3D effect
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- No motion smoothing, fast-moving 3D can flicker
Bottom Line
Sharp's XVZ-30000 is a worthy competitor for any other premium home theatre projector. Its 2D and 3D video quality is excellent, and the projector is only slightly let down by no 2D-3D conversion and no frame smoothing.
Price$ 4,999.00 (AUD)
The Sharp XVZ-30000 is the company's top home theatre projector, with the other prospect being the cheaper XVZ-17000. The XVZ-30000 stands up to other high-end home theatre projectors like the BenQ W7000 and Panasonic PT-AE7000, with great picture quality and some convenient features included.
Sharp XVZ-30000: Design and setup
The Sharp XVZ-30000 is an almost-cylindrical projector, with a shape that vaguely reminds us of an old Sony Discman. The lens is set at the centre of the projector's front fascia, surrounded by a rectangular bezel. The rest of the XVZ-30000's front half is taken up by fan exhaust ports. Three touch-sensitive buttons for power and control are on top.
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The Sharp XVZ-30000 is easy to set up whether you're ceiling- or rack-mounting it, thanks to a powered 2x zoom lens with powered focus and powered vertical and horizontal lens shift. These features are beginning to become more popular on high-end projectors, and we welcome them — it's much easier to stand by the screen and make fine adjustments via the projector's remote control.
Sharp XVZ-30000: Specifications and performance
The XVZ-30000 has a Full HD 1080p DLP chip in its innards, and is able to produce a contrast ratio of 50000:1 (in High Contrast mode) while maintaining a reasonably high brightness rating of 1600 lumens. This isn't as extreme as the 2000-lumen BenQ W7000 and Panasonic PT-AE7000 but we found the XVZ-30000 equally capable in everything but the brightest possible conditions.
We had the Sharp XVZ-30000 set up to a screen size of 100in at its minimum zoom distance of approximately 3.1m, running 3D Blu-rays and Full HD videos from a Sony BDP-S580 Blu-ray player and Apple MacBook Pro notebook.
The 2D video performance of the Sharp XVZ-30000 is excellent. It's the equal of the BenQ W7000 and Panasonic AE7000 in almost all areas. We thought the default picture presets of the XVZ-30000 were well set up, with a variety of settings for movies, gaming or daylight viewing. If you do want to get the projector professionally set up or calibrated, the custom setting has plenty of range for adjustment of individual colour channels and plenty of other minor tweaks as well.
Sharpness is good in default settings — not overly sharp like most projectors — and colours are vibrant without being oversaturated. We dropped brightness significantly for our dim viewing room, but there's plenty of headroom for any room that might have a lamp or in-ceiling lighting.
Black performance from the projector was good. The XVZ-30000 isn't better than the Panasonic AE7000, but we'd put it on par with the BenQ W7000 and it's a close race between all three overall. Contrast levels in each projector's movie-optimised modes were impressive, with bright whites displayed alongside deep yet detailed black areas. A dynamic iris in the XVZ-30000 works well to adjust brightness in darker or lighter scenes and works almost imperceptibly fast.
3D performance is also very good. We didn't notice any crosstalk on the high quality scenes of 3D video that we watched — some Cloudy With A Chance Of Meatballs, some Ocean Wonderland 3D and some Avatar 3D — and the depth effect from such a large screen size is impressive. The Sharp XVZ-30000's 3D glasses also have a nifty anti-3D mode that lets users view any currently-shown 3D video in 2D mode (by setting both glasses lenses to the left lens' refresh rate) — great if you've got someone in the audience who's not a 3D fan. There's no inbuilt conversion to watch 2D content in 3D, though. Extra glasses are $199.
We did notice that the lack of a motion smoothing option in the Sharp XVZ-30000's menus did mean a small amount of motion judder and flicker, especially during scenes of fast motion in 3D Blu-ray video. It's not a large problem and it's likely some users won't even notice it, but if you're looking for super-smooth motion this projector won't be the best choice — we'd recommend a good LED TV or plasma for this purpose.
Sharp XVZ-30000: Conclusion
The Sharp XVZ-30000 is a high quality 3D-capable home theatre projector. If you've got the money to spend on setting up a home cinema properly, we'd be happy with the XVZ-30000 as its centrepiece.
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text:
Gundam MS08 (DVD 4 of 4)
Title: Gundam MS08
Volume: DVD 4 / 4
Running time: 75
Distributor: Bandai Entertainment
Release date: 2002-01-22
Suggested retail price: $19.98
Age rating: PG-13
Universal Century 0079. The last battle in Southeast Asia is about to take place. As the Zeon's being to evacuate their base, the Federation moves in for the attack! Zeon ace, Packard, rushes to engage the enemy in order to buy his troops enough time to escape, but Gineas has other plans...
Driven mad by his quest to finish the Apsalus, he turns on his sister and threatens to destroy the assembled Federation Forces! With the battle almost over, the 08th MS Team are left without a leader – Will Michael and Kiki find Shiro and Aina and put an end to the war once and for all?
Previously identified as UPC 669198171390 (SKU 1713) with $24.98 MSRP
(added on 2001-09-30, modified on 2001-09-30)
- Reviewed by Paul Fargo (March 3, 2004)
- Encyclopedia information about Mobile Suit Gundam: The 08th MS Team (OAV)
The Fall 2016 Anime Preview Guide
Available Now: DREAM FESTIVAL ― Welcome to Anime News Network's Fall 2016 Preview Guide! Here's how it goes: our team of critics write up their impressions instantly and they get posted as they go along. Each critic will cover as many shows as they can handle, resulting in alternate takes ...


text:
Recipe: Strawberry Frappe
This Test Kitchen Approved recipe has been tested for accuracy.
- 4 cups Homemade Cool-Ade*
- 1 lb. frozen strawberries (about 4 cups)
- 2 x limes, juiced
*Click here for the Homemade Cool-Ade recipe.
Place all ingredients in a blender and blend until smooth.
Serving Suggestions
Serve in chilled glasses and garnish with slices of fresh strawberry.
Yields4 servings
Thanks ToSavory Spice Shop Test Kitchen
Make This Recipe!
"You are all so very awesome. Thanks for carrying amazing products!"


text:
U.S. DOT mulling new ban on in-flight cell calls
The U.S. Department of Transportation may seek a formal ban on in-flight cellphone calls, reports The Wall Street Journal. Following a Federal Communications Commission proposal to allow in-flight cellular calling, airlines have suggested that they can self-determine whether cellphone service should be allowed, and have accused the DOT of overstepping its authority on this matter; some carriers hope to offer "passenger-friendly" ways to allow calls. According to the report, the DOT is reportedly planning to propose an in-flight calling rule nonetheless, though the department has said that "there is no final determination" on what that rule will be. Notably, a June report said that the DOT also seeks to give the National Highway Traffic Safety Administration the ability to regulate navigation in cars, as part of the Grow America Act.
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Inns of Court
Inns of Court information
All London barristers work from within one of the four atmospheric Inns of Court, positioned between the walls of the old City and Westminster. It would take a lifetime working here to grasp all the intricacies of their arcane protocols, originating in the 13th-century. It's best just to soak up the dreamy ambience of the alleys and open spaces and thank your lucky stars you're not one of the bewigged barristers scurrying about.
A roll call of former members includes the likes of Oliver Cromwell, Charles Dickens, Mahatma Gandhi and Margaret Thatcher.
Lincoln's Inn still has some original 15th-century buildings. It's the oldest and most attractive of the bunch, with a 17th-century chapel and pretty landscaped gardens.
Gray's Inn was largely rebuilt after the Luftwaffe levelled it.
Middle Temple and Inner Temple both sit between Fleet St and Victoria Embankment. The former is the best preserved, while the latter is home to the intriguing 12th-century Temple Church , built by the Knights Templar and featuring nine stone effigies of knights in its round chapel. Check the church's website or call ahead for opening hours.


text:
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Transport of GluA1-containing AMPA glutamate receptors to synapses in the nucleus accumbens, a process that involves phosphorylation of key serine residues by CaMKII, is associated with the reinstatement of cocaine-seeking behavior. A growing body of evidence indicates that the dorsal striatum contributes to aspects of cocaine addiction. However, the potential role of CaMKII-mediated phosphorylation of GluA1 subunits in the dorsolateral (DL) striatum during cocaine reinstatement has not been examined. In this study, rats were trained to self-administer cocaine and were partnered with saline-yoked rats that received injections of saline. Following extinction, each pair of rats received either a systemic priming injection of cocaine (10 mg/kg, i.p.) or saline. As expected, cocaine-experienced rats displayed robust reinstatement of cocaine seeking in response to a challenge injection, whereas yoked saline controls did not. The DL striatum was dissected immediately following the reinstatement test session. Results from Western blotting experiments showed increased pGluA1-ser831 and pCaMKII-thr286 in the DL striatum of saline-yoked rats given an acute injection of cocaine. This effect was absent in cocaine-experienced rats that received a saline injection, and no changes were observed following a priming injection of cocaine in cocaine-experienced rats. These results indicate that acute exposure to cocaine in drug naïve rats increased CaMKII-mediated phosphorylation of GluA1-containing AMPA receptors in the DL striatum, an effect that was not observed during cocaine priming-induced reinstatement of drug seeking. It is possible; therefore, that increased phosphorylation of CaMKII and GluA1 following acute cocaine is a compensatory mechanism in the DL striatum.
It is well documented that changes in glutamatergic transmission in subregions of the striatum play a role in the reinstatement of cocaine seeking, an animal model of relapse [25]. Although the majority of this work has focused on the nucleus accumbens [1, 11, 27], recent evidence indicates that pharmacological inhibition of the dorsolateral (DL) striatum attenuates cocaine seeking in rats following short-access (1- or 2-hour daily) self-administration [23, 29]. In terms of glutamatergic transmission, infusion ofα -amino-3-hydroxy-5-methyl-4-isoxazolepropionic (AMPA) glutamate receptor antagonists into the DL striatum blocked cue-controlled reinstatement and attenuated cocaine self-administration when injected into the dorsomedial (DM) striatum [11, 22, 28, 31]. Though there is an established role for the DL striatum in cue-induced reinstatement, there is little to no data on DL striatum involvement in cocaine priming-induced reinstatement of drug seeking.
AMPA glutamate receptors are ionotropic, excitatory ion channels comprised of four subunits (GluA1-GluA4) that are involved in several forms of neuronal plasticity [6]. The GluA1 subunit in particular has been associated with synaptic plasticity in animals exposed to cocaine [2, 5]. Activity-mediated autophosphorylation of calcium/calmodulin-dependent protein kinase II (CaMKII) at threonine 286 promotes phosphorylation of GluA1 at serine 831, resulting in enhanced GluA1 subunit trafficking to the plasma membrane and associated increases in synaptic strength [6, 18]. Indeed, there is evidence that reinstatement of cocaine seeking in rats involves CaMKII phosphorylation of the GluA1 subunit in the nucleus accumbens, and pharmacological inhibition of CaMKII in the accumbens prevents cocaine seeking [1, 19].
A single cocaine injection promotes glutamate release in the DL striatum of cocaine-naïve rats [3, 21]. Acute or repeated experimenter-delivered cocaine increases levels of phosphorylated GluA1 AMPA receptor subunits and GluA1 surface expression in the DL striatum [12, 16]. Here, we extend this work by examining expression of phospho-CaMKII and –GluA1 in the DL striatum during cocaine priming-induced reinstatement of drug seeking.
Male Sprague-Dawley rats (Rattus novergicus) weighing 250–300 g were obtained from Taconic Laboratories (Germantown, NY, USA). Animals were individually housed with food and water available ad libitum. A 12h light/dark cycle was used and all experiments were performed during the light cycle. All experiments used Med-Associates (East Fairfield, VT, USA) operant chambers equipped with response levers, house light, and pumps for injecting drugs intravenously. Operant chambers were enclosed within ventilated, sound attenuating chambers. All experimental procedures were consistent with the ethical guidelines of the U.S. National Institutes of Health and were approved by the University of Pennsylvania School of Medicine Institutional Animal Care and Use Committee.
Cocaine hydrochloride was obtained from the National Institute on Drug Abuse (Rockville, MD, USA) and dissolved in bacteriostatic 0.9% saline.
Prior to surgery, the rats were anesthetized with 80 mg/kg ketamine (i.p.; Sigma-Aldrich, St. Louis, MO, USA) and 12 mg/kg xylazine (i.p.; Sigma-Aldrich, St. Louis, MO, USA). An indwelling silastic catheter (CamCaths, UK) was placed into the right jugular vein and sutured in place. The catheter was routed to a mesh backmount platform and implanted subcutaneously dorsal to the shoulder blades. Catheters were flushed daily with 0.3 ml of an antibiotic (Timentin, 0.93 mg/ml, Henry Schein, Melville, NY, USA) dissolved in heparinized saline. The catheters were sealed with plastic obturators when not in use.
Following a 7-day recovery period from surgery, 50% of the rats were randomly selected to self-administer cocaine. The remaining 50% were paired to a cocaine self-administering animal as yoked-saline controls that only receive infusions of saline. Cocaine self-administering rats were placed in operant chambers and allowed to lever press for intravenous cocaine infusions (0.25 mg/59 µl saline over 5 s) over a 2-hour time period daily for 16 days. Yoked-saline controls received the same number and temporal pattern of infusions as the paired cocaine self-administering rat. Rats were initially trained using a fixed ratio (FR) 1 schedule of reinforcement, with each daily self-administration session initiated by a priming injection of cocaine (0.25 mg, i.v.) to fill the catheter (limited to 30 injections per 120 min). Once animals achieved stable responding, as defined by at least 20 infusions over the 2 h session for 2 consecutive days, they were transitioned to an FR5 schedule of reinforcement (limited to 30 injections per 120 min). For both schedules of reinforcement, a 20s time-out period followed each cocaine infusion, during which time active, drug-paired lever responses were tabulated but had no scheduled consequence. Each operant chamber was also equipped with an inactive lever. Responses made on the inactive lever, which had no scheduled consequence, were also recorded during all training sessions.
Following cocaine self-administration, drug-taking behavior in cocaine self-administering rats was extinguished by replacing cocaine with 0.9% bacteriostatic saline. Extinction continued until responding on the drug-paired lever was <15% of the response rate on the last day of cocaine self-administration using the FR5 schedule of reinforcement. Typically, it took rats 5 days to meet this criterion.
Following extinction, animals entered the reinstatement phase of the experiment. During reinstatement, satisfaction of the response requirements for FR5 resulted in saline rather than cocaine infusions. The FR5 schedule was used for the reinstatement test session. Pairs of cocaine self-administering/yoked-saline rats were randomly assigned to receive saline or cocaine (10 mg/kg, i.p.) injections immediately prior to the start of the reinstatement test session. Rats were placed into the operant chambers immediately following injection of saline or cocaine. Responding was recorded for 30 minutes, after which the pairs of rats were removed from the operant chambers and immediately decapitated. Whole brains were extracted and flash-frozen in isopentane on dry ice, then stored at −80° C. Brains were sliced on a cryostat (300 µm sections) and striatal subregions (+1.7 mm to −0.08 mm A/P; −3.8 mm to −6.0mm D/V; DM: ±1.2 mm – 2.8 mm M/L and DL: ±2.8 mm – 4.4 mm M/L) dissected by tissue punch (2.0 mm Harris Uni-core stainless steel punchers, Ted Pella, Inc.). Tissue samples were stored at −80° C until processing for Western blot as described below.
Whole-cell tissue was processed for Western blot as described previously (Anderson et al., 2008). For all samples, protein concentration was quantified using a Pierce BCA Protein Assay Kit (Thermo Scientific). Equal amounts of protein (10 µg for whole-cell) were loaded and separated in 10% Tris-Glycine gels (Invitrogen) and transferred to nitrocellulose membranes using the i-Blot dry transfer system (Invitrogen). Membranes were blocked with either 5% nonfat dry milk in TBST or 5% BSA in TBST, according to antibody instructions. Membranes were incubated overnight at 4° C with selective antibodies to: GluA1 (1:1000, Abcam), phosphorylated (p) GluA1-S831 (1:500, Millipore), CaMKII (1:1000, Millipore), pCaMKII-T286 (1:1000, Cell Signaling), and GAPDH (1:2000, Cell Signaling). Membranes were then incubated with fluorescent secondary antibodies (1:5000, IR-dye 680 or IR-dye 800), before being imaged on an Odyssey fluorescent scanner (Licor Biosciences). To ensure equal loading, GAPDH expression was used as a loading control.
Drug-paired and inactive lever responding was analyzed using a two-way ANOVA. The between subjects factors were pretreatment (cocaine self-administration or yoked-saline) and drug challenge injection (10 mg/kg cocaine or saline). Bonferroni post-tests were used to establish significant difference (p<0.05).
Quantification was performed by normalizing the intensity of all bands with protein-specific antibodies to the GAPDH intensity, followed by normalizing that value to saline-control (yoked-saline, i.p. saline) values. The immunoblot analyses were performed using a two-way ANOVA. The between-subjects factors were pretreatment (cocaine self-administration or yoked-saline) and drug challenge injection (10 mg/kg cocaine i.p. or saline). Bonferroni post-tests were used to establish significant difference (p<0.05).
Total drug-paired lever responding (mean +/− SEM) for the reinstatement test session are plotted in Fig 1. Responding was analyzed using a two-way ANOVA. Analyses revealed a significant main effect of pretreatment (F1,24=119.39, p<0.0001), a significant main effect of challenge injection (F1,24=92.14, p<0.0001), as well as a significant pretreatment × challenge interaction (F1,24=78.93, p<0.0001). Post-hoc tests revealed a significant difference of drug-paired lever responding due to cocaine challenge injection (Bonferroni p<0.001). There was no significant effect of pretreatment (F1,24=0.03, p=0.8543) or challenge (F1,24=3.68, p=0.0671) on inactive lever responding (data not shown).
The average fluorescent intensity for pGluA1-ser831 in the DL striatum was expressed as percent change from control and is plotted in Fig 2A. Percentages were analyzed by two-way ANOVA, which revealed a significant main effect of pretreatment (cocaine self-administration vs. yoked-saline, F1,12=14.03, p<0.005), a significant main effect of challenge (10 mg/kg cocaine or saline, F1,12=30.68, p<0.0001), as well as a significant pretreatment × challenge interaction (F1,12=20.75, p<0.0007). Post hoc tests revealed a significant difference in pGluA1-ser831 expression between yoked-saline and cocaine self-administering animals after cocaine challenge injection (Bonferroni p<0.001). There was no significant effect of pretreatment or challenge on the native GluA1 protein (Fig 2B).
The average fluorescent intensity for pCaMKII-thr286 in the DL striatum was expressed as percent change from control and plotted in Fig 2C. Percentages were analyzed by two-way ANOVA, which revealed a significant main effect of pretreatment (F1,11=15.73, p<0.0022), a significant main effect of challenge (F1,11=48.62, p<0.0001), as well as a significant pretreatment × challenge interaction (F1,11=9.78, p<0.0096). Post hoc tests revealed a significant difference in pCaMKII-thr286 expression between yoked-saline and cocaine self-administering animals after cocaine challenge injection (Bonferroni p<0.01). There was no significant effect of pretreatment or challenge on native CaMKII protein (Fig 2D).
The average fluorescent intensity for pGluA1-ser831 in the DM striatum was expressed as percent change from control and plotted in Fig 3A. Percentages were analyzed with a two-way ANOVA, which revealed no significant effect of pretreatment (p=0.3542) and no pretreatment × challenge interaction (p=0.9403), but a significant main effect of challenge (F1,24=4.33, p<0.05). There was no significant effect of pretreatment or challenge on the average fluorescent intensity of native GluA1 protein in the DM striatum (Fig 3B).
The average fluorescent intensity for pCaMKII-thr286 in the DM striatum was expressed as percent change from control and is plotted in Fig 3C. Percentages were analyzed with a two-way ANOVA, which revealed no significant effect of pretreatment (p=0.1615) or challenge (p=0.2906). There was no significant effect of pretreatment or challenge on the average fluorescent intensity of native CaMKII protein in the DM striatum (Fig 3D).
The present findings indicate that an acute injection of cocaine increased expression of pCaMKII-thr286 and pGluA1-ser831 in the DL, but not the DM, striatum of drug-naïve rats. Since the phosphorylation of CaMKII at threonine 286 is associated with increased phosphorylation of GluA1 at serine 831 and subsequent trafficking of these subunits [4], these results suggest that an acute cocaine injection may increase the surface expression of AMPA receptors in the DL striatum. However, following cocaine self-administration and extinction, a cocaine challenge injection had no effect on DL striatal pCaMKII-thr286 or pGluA1-ser831 expression. Taken together, these results indicated that tolerance to cocaine-induced increases in the phosphorylation of DL striatal CaMKII and GluA1 develops following cocaine self-administration.
Our findings are consistent with previous work that showed increased pCaMKII-thr286 and pGluA1-ser831 in the DL striatum following acute cocaine [10, 15]. A single exposure to cocaine also increases immediate early gene expression in the dorsal striatum [14, 17, 24]. Additionally, a single microinjection of cocaine into the dorsal striatum enhances locomotor activity [9]. Taken together, these results indicate that cocaine-induced hyperlocomotion is associated with increased neuronal activation in the DL striatum and increased phosphorylation of CaMKII and GluA1. Future studies are required to determine the functional significance of increased phosphorylation of CaMKII and GluA1 following acute cocaine exposure.
There also is evidence of altered glutamatergic transmission in the DL striatum following prolonged cocaine administration [26]. Animals identified as vulnerable to cocaine relapse following cocaine self-administration showed down-regulation of genes for GluA1 and activity-regulated cytoskeletal protein, both of which are involved in synaptic plasticity and AMPA receptor trafficking [7]. While we did not investigate alternate phosphorylation sites of GluA1 subunits in this study, there is also evidence of decreased expression of phospho-GluA1 ser845, a known target of protein kinase A, in the dorsal striatum in animals with chronic cocaine self-administration experience [10, 20]. Functionally, pharmacological inactivation of the DL striatum attenuates cocaine seeking in rats with a history of short- or long-access to cocaine self-administration [23]. Moreover, an intra-striatal infusion of an AMPA/kainate receptor antagonist attenuates cue-controlled cocaine seeking [23, 30]. Furthermore, overexpression of a dominant negative peptide that prevents trafficking of GluA1 to the membrane surface in the DL striatum attenuated cocaine-induced behavioral sensitization in juvenile mice [15]. It is important to note, however, that unlike adult mice, the juveniles exhibited increased GluA1 surface expression following repeated cocaine, suggesting that age at time of cocaine exposure influences DL striatal glutamatergic plasticity. Transgenic mice that express constitutively active CaMKII, where threonine 286 is replaced with aspartic acid, in the dorsal striatum exhibit impairments in goal-directed behaviors, with impairments in both cue-and reward-primed reinstatement [32]. As a whole, these results indicate that the changes in AMPA receptor transmission in adult rodents that contribute to cocaine sensitization and reinstatement are not likely due to increases in CaMKII or GluA1 phosphorylation in the DL striatum. Our data showed changes the phosphorylation state of CaMKII and GluA1 in DL striatum of drug naïve rats following acute cocaine only, offering novel biochemical support for such conclusions.
The present findings suggest differential involvement of the glutamatergic system in the DL striatum relative to the ventral striatum, particularly after cocaine self-administration and the reinstatement of cocaine seeking. It is well known that CaMKII plays a key role in phosphorylating GluA1 subunits at serine 831 and promotes AMPA receptor trafficking to the membrane surface [4]. In fact, plasticity of the glutamatergic system in the ventral striatum, including increased phosphorylation of CaMKII and GluA1 as well as increased GluA1 surface expression occurs in a variety of paradigms involving chronic cocaine exposure [5, 8, 13]. More specifically, during cocaine priming-induced reinstatement, there is increased phosphorylation of CaMKII and GluA1 AMPA receptor subunits, as well as increased trafficking of GluA1 to the plasma membrane in the accumbens shell [1]. Inhibiting CaMKII or preventing the trafficking of GluA1 subunits in the shell is sufficient to block reinstatement of cocaine seeking [1]. These findings coupled with the data presented here suggest that phosphorylation of CaMKII and GluA1 in the ventral, but not DL, striatum is required for reinstatement of cocaine seeking. Thus, changes in phospho-CaMKII and –GluA1 in the DL striatum likely reflect a compensatory mechanism in response to acute cocaine.
Our results indicate that acute cocaine increases expression of pCaMKII-thr286 and pGluA1-ser831 in the DL, but not DM, striatum and that these effects are not associated with the reinstatement of cocaine seeking. Therefore, changes in phospho-CaMKII and –GluA1 most likely reflect a compensatory adaptation in the DL striatum during early stages of drug use. It is possible that acute cocaine promotes transient biochemical modifications in the DL striatum that reverse with additional input from areas like the ventral striatum and/or prefrontal cortex as cocaine experience is extended [10].
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