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North American animals have numerous strategies for surviving the winter. Some migrate to warmer climes, others head for a long winter's nap, and some stay where they are and rely on fur, speed, fat, teeth, and claws to keep them safe, warm, and well fed. How much do you know about the winter behavior of North America's animals? Test your winter wildlife knowledge by taking our quiz. (Scroll down to see the answers.)
1. Which animal has the densest, warmest fur on earth?
A. Beaver
B. Sea Otter
C. Polar Bear
D. Mink
E. Muskox
2. How much weight might an adult Grizzly Bear gain before retiring for the winter?
A. 50 pounds
B. 100 pounds
C. 200 pounds
D. 300 pounds
E. 400 pounds
F. All of the above
3. Which animal remains active all winter, and doesn't retire to a den for a sustained period?
A. Eastern Chipmunk
B. White-tailed Prairie Dog
C. Red Squirrel
D. Black Bear
E. Polar Bear
4. Choose the animal species that does NOT regularly appear in a white-furred form.
A. Eastern Gray Squirrel
B. Arctic Fox
C. Grizzly Bear
D. Least Weasel
E. Gray Wolf
F. Black Bear
5. Which of the following species will travel to a new area for the winter?
A. Manatee
B. Caribou
C. Bison
D. Snow Goose
E. Snowy Owl
F. All of the above
6. Animals generally put on fat for two reasons: warmth and food reserves. Which of the following species reaches the heaviest weight?
A. Grizzly Bear
B. Polar Bear
C. Northern Elephant Seal
D. Walrus
|7. Match the predator (1-7) with the prey (a-g) that it commonly eats.
1. Arctic Fox
2. Lynx
3. Northern River Otter
4. Mink
5. Polar Bear
6. Walrus
7. Northern Elephant Seal
a) fish
b) squid
c) clams
d) seal
e) snowshoe hare
f) lemming
g) muskrat
Question 1: If you guessed the Sea Otter, you're right! Its fur is denser than the fur of any other mammal. One square inch of Sea Otter fur contains as many as 1 million hairs -- that's about the same number of hairs on the heads of ten humans. This water-loving mammal eats, sleeps, mates, and gives birth at sea. It lacks the layer of blubber that keeps many other marine mammals warm, and so has only its fur coat to protect it from chilly North Pacific waters. Oil spills can devastate Sea Otter populations, because the otters' coats lose their insulating properties when saturated with oil, and many affected otters die of exposure.
Question 2: All of the above! A Grizzly Bear can put on as much as 400 pounds to prepare for its winter sleep. This omnivore will eat just about anything in its quest to fatten up for the lean times. Grizzlies eat not just large mammals (elk, moose, deer) and fish, they'll eat roots, plant sprouts, berries, mushrooms, and any smaller critters, including insects, that come their way.
Question 3: The answer is the Red Squirrel. These gregarious creatures are active year-round, although they may hole up for a few days in inclement weather. The Red Squirrel's relatives the Eastern Chipmunk and White-tailed Prairie Dog are true hibernators, meaning they enter a state of dormancy during the winter in which body temperature drops to only a few degrees above air temperature, and all bodily processes greatly slow down. The hibernating animal thus conserves energy and stored fat, and is able to sleep through much of the winter.
Bears enter a protected area and sleep away the harshest part of the winter, but they do not truly hibernate, as their sleep is not deep, and their temperature falls only a few degrees below normal. Even Polar Bears retire to a den for part of the winter. Females den from November to March, during which time they give birth, while males usually den from late November to late January.
Question 4: The answer is the Grizzly Bear, which ranges in color from tawny to dark brown but is never white (except in the case of a rare albino individual). Arctic Foxes and Least Weasels vary seasonally, growing a white coat for winter camouflage (in northern populations). The Gray Wolf is a species that varies individually, ranging from white to black and any shade in between. Most Eastern Gray Squirrels are gray, but there are populations of white gray squirrels in several areas, with the largest concentration found in Olney, Illinois. And rarest of all of these white mammals are the cream-colored Black Bears that live in the coastal rain forests of British Columbia. These bears belong to a subspecies of Black Bear officially named the Kermode Bear but often referred to as the Spirit Bear.
Question 5: If you guessed all of the above you are right! At least some individuals of all of these species relocate seasonally. Even though Manatees live only in the South, they head for warmer water in the winter, some migrating tens to hundreds of miles and others merely congregating around the heated discharge from power plants. The Caribou of Alaska and Canada famously go on long mass migrations between winter and summer grounds. The Bison of the Great Plains once undertook mass migrations by the millions and ranged nearly from coast to coast. Few of the populations left are free-ranging, but some Canadian Bison still migrate about 150 miles between winter and summer grounds.
The Snow Goose is named for its white color rather than its love of snow; it summers in the Arctic but heads to the coastal United States and southward for the winter. The Snowy Owl, on the other hand, can and does survive the winter in our most northerly climes. In some years, however, if the owl's prey species (mainly lemmings) decline in population, large numbers of Snowy Owls will move into regions south of their normal range; this kind of migration is called an irruption or invasion.
Question 6: The Polar Bear, with its warm coat to keep it warm, is the svelte member of this group, reaching only about 1,100 pounds. A Grizzly Bear can weigh as much as 1,700 pounds. The largest Grizzlies are those of the southern Alaska coast and islands that are known as Alaskan Brown Bears. A Walrus can weigh nearly twice as much, topping out at 3,300 pounds. It lives in Arctic seas and needs a good solid layer of blubber to keep warm. Wisely, it spends a lot of time sunbathing on beaches or ice floes. When it does go on a prolonged dive (for up to 30 minutes) its blood flow decreases to the skin, thus conserving body heat, and increases to its vital internal organs.
The record holder among these four creatures is the Northern Elephant Seal: A male can reach 4,400 pounds! He doesn't stay that hefty, however: this seal can lose 50 percent of its weight during the mating season, when it is too busy to eat, and also fasts during the molt, when it grows a new coat. This Pacific Coast species does not live in waters as cold as the Walrus, but it spends more time in the water. It can stay underwater for nearly an hour and a half, pop up for a few minutes and dive again, so clearly it needs insulating blubber to help retain body heat.
Question 7: Lemmings are an important food source for the Arctic Fox. These small rodents undergo cyclical variations in population, increasing in number enormously when food is plentiful, then decreasing when the population outgrows the food supply. Arctic Fox populations follow these cycles, usually peaking a year after the lemmings. There is a similar relationship between the Lynx and the Snowshoe Hare, which makes up three-quarters of the Lynx's diet.
Northern River Otters mainly eat fish, and Minks fish too but their preferred prey in many areas is muskrat. Polar Bears stalk seals mainly, but also eat young Walruses, whales, fish, birds, eggs, shellfish, kelp whatever they can find. Walruses like clams and other mollusks so much that they can eat 3,000 to 6,000 in a single feeding. Northern Elephant Seals eat enormous quantities of squid (and they eat fish, too).
Click on a species below to learn more about its winter habits:
Arctic Fox
Snowshoe Hare
Snow Goose
Polar Bear
Eastern Chipmunk


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Have you ever purchased from PartSelect.com before?
Thank you for helping make our site better.
Below is a list of all parts for the Maytag Dryer DE24CM - Electric. To find the repair part you need, you can browse the list below or you can narrow your choices by searching for a part by name or description.
This drum support roller includes one drum roller and two bearings. Most dryers use two drum rollers, and is recommended to change both at once.
At 100 inches in length, this dryer drum belt is 3/8 of an inch wide and has five ridges. This belt is all black in color and it is made entirely of rubber.
This roller shaft acts as the axel for the drum roller. Also known as drum roller axle.
This front glide kit comes with a blue plastic glide, a brown cork pad, and two metallic rivets. Two kits are required per appliance. Also, a rivet tool is required for this kit - purchase locally.
This comes with a metallic hub clamp to hold it firmly in place.
This lint filter screen is of a slide in style type of Maytag filter screen. It is most typically used for the 08 and later series of Maytag gas or electric clothes dryers that appear before 1990.
This metallic retaining ring is less than an inch in diameter. It is made entirely of metal.
This roller shaft washer is just under an inch in diameter and is used for some of Whirlpool's brands of clothes dryers.Sold individually.
This part is used to help the belt to rotate the drum. It is white and made of plastic with a metallic bearing in the center.
This idler pulley shaft washer is used with many clothes dryers. It is less than an inch long, and it is black in color.
This part replaces older two terminal switches.
This high limit/safety thermostat is used on many of Maytag's brands of gas and electric clothes dryers. With this thermostat, a dryer's heating element will cut out if the internal temperature reaches 200 degrees Fahrenheit. The element will kick back in when the temperature drops to 170 degrees Fahrenheit.
This front or rear dryer drum felt seal is seventy-nine and a half inches long.
This dryer door latch kit comes in two pieces: a door latch, and a door strike. It's used to help hold your door shut to your dryer.
All brand logos are trademarks of their respective owners.
The PartSelect logo is a Registered Trademark of Atlantic Laundry Centres, Ltd.
Copyright © 1999-2017 , Eldis Group Partnership. All rights reserved.

subset: commoncrawl


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When it comes to the sheer variety of scenery on offer, the Goldsteig is hard to beat. Its route through the Upper Palatinate Forest and Bavarian Forest runs through five nature reserves and one national park.
Starting in Marktredwitz, the Goldsteig Trail passes through the Steinwald Forest before continuing to the wild and romantic ruins of Weissenstein Castle and on to Friedenfels. A thousand little lakes line the onward leg to mighty Falkenberg Castle, before the route follows the part gentle, part dramatic Waldnaab valley to the lead crystal town of Neustadt an der Waldnaab. Shortly after Oberviechtach, the Goldsteig Trail divides into two equally attractive routes. Walkers with a good level of fitness should choose the ridge route, which takes them past Furth im Wald, up the Grosser Arber and over the highest peaks of the Bavarian Forest National Park. The alternative option heading south offers glorious alpine panoramas from Mount Pröller, Mount Hirschenstein and Mount Brotjacklriegel. Descending to more level ground, it meets up with the mountain route in Passau, the town on three rivers. For those seeking accommodation, a warm welcome awaits at any one of the 'Goldsteig-Ge(h)nuss' accommodation providers.
Tip: ruins of Weissenstein Castle
Weissenstein Castle, now a ruin, was first mentioned in the 13th century. It has been undergoing reconstruction and archaeological examination since 1992. The castle keep affords fine panoramic views of the surrounding region and is accessed via an alarminglooking set of stairs.
At a glance:
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Hey you two don't you no where home is.
Have fun and be safe.
Terry Jeanette
Colombians Turn Out To Vote.
Trip Start Feb 01, 2005
435Trip End Dec 31, 2020
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Where I stayed
La Portada
What I did
Archaeological Park of Tierradentro Colombia
Read my review - 4/5 stars
Read my review - 4/5 stars
Election Day!
We find ourselves stranded for a day longer than planned in a little village, San Andres de Pisimbala (population 800) for Election Day. Hardly anyone apart from Colombians ever visits. Of no small interest is the fact that this area has been closed to tourism for some time, due to its reputation as a guerrilla stronghold and has only just begun to see tourists. There are only two other non-Colombians in town, a Melbourne Author updating for Lonely Planet and his Nicaraguan partner.
We had been warned several days ago that there may be no buses on Sunday, Election Day, but still we hoped this was not going to be so. The buses have been diverted for bringing in voters and usual schedules, all cancelled
A festive air abounds in this little village today. Bright colourful buses and 4 wheel drives, loaded impossibly full inside and rooftops full as well, are bringing laughing and excited Colombians in from remote communities to vote. A huge pot of meat and potato stew sits on a fire and is free for voters. We have just eaten tasty deep fired empanadas with unknown fillings (some sort of meat and vegetables but not recognizable) and are now back at our little hostel room hoping that election festivities do not get out of hand later.
But why are we here? Well, we came to see Tierradentro. UNESCO listed (of course!), the site is comprised of dozens of intricately and colourfully decorated sacred burial sites hewn out of rock, deep in the ground, and left behind from an era of around the 7th century by indigenous Colombians. Our journey has been long and arduous with the last 3 hours of travel from La Plata undertaken in a 4 wheel drive dual cab over rough roads - with 22 people! Yes 22! We travelled in the ute back with 6 others in our small space. Passengers clung to the outsides, sat on the roof above us (where our packs were) and squashed in the dual cab. Opposite us sat an attractive young woman with a massive scar down her eye and cheek that looked like it was just roughly stitched up (after whatever serious accident befell her), and was never seen by a plastic surgeon
We unwound ourselves from the tangled mass of bodies at San Andres de Pisimbala and looked forlornly at the state of our dust caked backpacks and ourselves. As we gingerly loaded them on our backs we felt the pull of cramped muscles from our enforced squashed positioning. A hostel was close by to where we got off, so we took a room and the set menu lunch of a delicious soup and salad before heading straight down the road for the 2 km walk to the entrance of the Tierradentro Site. About 500 metres on our way we were startled by a voice and looked up to see 2 gun toting soliders sitting on the bank. They literally pop out of the woodwork in Colombia which is sort of reassuring but also worrying at the same time!
As we paid our entry fee and signed the book requiring our details, we realized we were the only ones at all there that day and only 2 people, Colombians, had visited the day before. This would have to the first UNESCO World Heritage Site EVER that we have been the only visitors for the day
The tombs required a pretty strenuous hike up a rough path but on arrival at the top, we were met by a guard, who took us to each tomb and unlocked the protective mesh gate at the entry. Steep rock hewn steps led down to into each tomb and inside we found them to be highly decorated with colour still quite vivid in the decorations. Various artefacts and giant pottery urns were found in the tombs and one of the tombs had been left with all artifacts insitu. Two museums are on site, one has many other artifacts and urns recovered from the excavations. Many tombs are yet to be excavated but there is insufficient funding to continue.
Our hostel served us an evening meal and it was while at dinner, the other Australian who has excellent Spanish, confirmed for us that we would not be able to catch a bus out the next day to our next destination, due to the elections. An enforced rest is not too hard to take when Avan is just recovering from a cold and I seem to be coming down with one.
So today has been a "right time right place" moment to see Colombian elections undertaken in a tiny indigenous community and then tomorrow we'll brave transport again, with a 6.30am bus back to 'civilization".
Footnote: National Archaeological Park of Tierradentro is UNESCO World Heritage listed.
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My Review Of The Place I've Seen
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Toggle: English / Spanish
The creatinine blood test measures the level of creatinine in the blood. This test is done to see how well your kidneys work.
Creatinine can also be measured with a urine test.
Serum creatinine
A blood sample is needed.
The health care provider may tell you to temporarily stop taking certain medicines that can affect the test. Medicines include:
When the needle is inserted to draw blood, some people feel moderate pain. Others feel only a prick or stinging sensation. Afterward, there may be some throbbing or slight bruising. These soon go away.
Creatinine is a chemical waste product of creatine. Creatine is a chemical made by the body and is used to supply energy mainly to muscles.
This test is done to see how well your kidneys work. Creatinine is removed from the body entirely by the kidneys. If kidney function is not normal, creatinine level increases in your blood. This is because less creatinine is released through your urine.
The creatinine level also varies according to a person's size and muscle mass.
A normal result is 0.7 to 1.3 mg/dL for men and 0.6 to 1.1 mg/dL for women.
Women usually have a lower creatinine level than men. This is because women usually have less muscle mass than men.
The examples above are common measurements for results of these tests. Normal value ranges may vary slightly among different laboratories. Some labs use different measurements or test different samples. Talk to your doctor about the meaning of your specific test results.
Higher than normal level may be due to:
Lower than normal level may be due to:
There are many other conditions for which the test may be ordered, such as high blood pressure, diabetes or medication overdose. Your health care provider will tell you more, if needed.
There is very little risk involved with having your blood taken. Veins and arteries vary in size from one patient to another and from one side of the body to the other. Taking blood from some people may be more difficult than from others.
Other risks associated with having blood drawn are slight but may include:
Israni AK, Kasiske BL. Laboratory assessment of kidney disease: filtration rate, urinalysis, and proteinuria. In: Taal MW, Chertow GM, Marsden PA, et al., eds. Brenner and Rector's The Kidney. 9th ed. Philadelphia, Pa: Elsevier Saunders; 2011:chap 25.
A.D.A.M., Inc. is accredited by URAC, also known as the American Accreditation HealthCare Commission (www.urac.org). URAC's accreditation program is an independent audit to verify that A.D.A.M. follows rigorous standards of quality and accountability. A.D.A.M. is among the first to achieve this important distinction for online health information and services. Learn more about A.D.A.M.'s editorial policy, editorial process and privacy policy. A.D.A.M. is also a founding member of Hi-Ethics and subscribes to the principles of the Health on the Net Foundation (www.hon.ch)
The information provided herein should not be used during any medical emergency or for the diagnosis or treatment of any medical condition. A licensed medical professional should be consulted for diagnosis and treatment of any and all medical conditions. Call 911 for all medical emergencies. Links to other sites are provided for information only -- they do not constitute endorsements of those other sites. © 1997- 2013 A.D.A.M., Inc. Any duplication or distribution of the information contained herein is strictly prohibited.


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Microsoft Virtual Academy: Monitor & Operate Private Cloud Infrastructure with System Center 2012 SP1 - Part 2 - Control Access
About This Video:
In this module you will learn how System Center 2012 Service Pack 1 (SP1) has been enhanced to support Windows Server 2012 and SQL Server 2012 through simplified monitoring and operating of the private cloud infrastructure. You will see how the concepts of optimization, control and standardization can be used to delegate capacity and provide cloud abstraction across a diverse infrastructure. This will help ensure that the services running in your private cloud get the resources they need, and that your cloud dynamically reallocates resources as demand changes.
System Center 2012 SP1 can help protect your private cloud infrastructure by setting limits on your clouds while ensuring the correct owners have access to their cloud resources with role-based access. You will learn about how standardization can be applied to application and OS updating through service templates. You will be able to report on the current utilization of your private cloud, trend usage over time, and forecast future needs.
Finally you will see how you can optimize cloud utilization through dynamic optimization & power optimization, as well as reporting improvements to VMs running Linux distributions in the guest OS.
After watching this video, visit:
Published Date: January 07, 2013
Video: WMV (ZIP) (no longer available)
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This Smilodon is Not Smiling
Credit: Phil Scoville
Source: http://www.flickr.com/photos/philscoville/122990117
License: CC BY-NC 3.0
For 30,000 years, untold numbers of animals got stuck in the La Brea Tar Pits in Los Angeles, California. The museum there displays the fossils of some of these amazing Ice Age creatures!
Amazing But True!
- The Los Angeles basin is loaded with petroleum. Many of the oil fields have been exhausted and are no longer in production.
- The La Brea Tar Pits tap into these rich deposits. Crude oil seeps up from underground through an earthquake fault line. Some of the oil biodegrades to make thick tar that collects in pools.
- In the past, water, dust and leaves covered the surface of the pool. The pool became a death trap for animals that would go in for a drink. Predators would attack these vulnerable animals. Few of these animals got out alive.
- The tragedy of all these animals is a boon for us since the La Brea Tar Pits are a fabulous fossil locale.
Can You Apply It?
With the link below, learn more about fossils at the La Brea Tar Pits. Then answer the following questions.
- Why are fossilized pollen grains important?
- Why are the fossils at the La Brea Tar Pits so well preserved?
- Why is a Smilodon no longer called a saber-toothed tiger? What is a saber tooth?
- Paleontologists gain lots of information beyond just bones from good fossil sites. What sorts of information can they gain beyond just what the animal looked like?


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I was just taking a look at the pics of the Royal Starship playset. If you go to the picture archive and take a look at "Close Up 2", I think it is, the R2 unit in the elevator has a white body with red detailing. Is this a mock up, prototype, or did they not have a clue what they were gonna pack in at the time of the photograph?
Also... Did the final version of the Red Astromech even have a name. The box only reads: "Exclusive Red R2 unit" or something to that end. Anyone? Anyone? Bueller?


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Great decoration idea! This creative ball bowl is perfect for Christmas celebrations.
Christmas ball ornaments also make charming place settings. Write the name of each guest on an ornament with a metallic marking pen. Set one ornament at each place setting and steady it on a vinyl hose washer--that's right, the kind you find at your local hardware store. The washers work great for making the ornaments stay put, and you can spray paint them to match. Remove the tops from the ornaments, and fill about half full with water. Place flowers in ornaments and tie with a decorative ribbon.
© 2017 ®/TM General Mills All Rights Reserved


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ShopKo To Go Private
April 11, 2005,
Green Bay, Wis. — ShopKo's chairman and, possibly, other members of the retailer's senior management will participate in the acquisition of the chain by a Minneapolis-based private equity firm for slightly more than $1 billion, including cash.
The agreement, with buyout company Goldner, Hawn, Johnson & Morrison, will pay $24 in cash for each outstanding share of ShopKo common stock. The net transaction price is about $715 million plus the assumption of $330 million in debt.
Chairman Jack Eugster, who is an investor in the buy out, has resigned his position as chairman and will step down from board committee positions in order to avoid any conflicts of interest, according to spokesman John Vigeland. Eugster will retain his board seat during the transaction, however, and will become chairman and acting CEO following the close, until a new president and CEO is named.
GHJ&M, whose portfolio currently numbers eight firms, has no current retail holdings. It was outbid several years ago in an effort to acquire Elder-Beerman department stores.
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The Countdown to the ICON Honors Continues featuring Christophe Pourny


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The Waterfront Apartments afford magnificent views of the Marlborough Sounds. Being centrally positioned to Picton and overlooking the marina, the beautifully furnished, spacious apartments are the perfect place for a family holiday.
Secluded peaceful 2 stylish coastal cottages, Qualmark 5stars king beds/ensuites overlooking large ornamental lake, countryside, mountains. Includes gourmet hamper breakfast. Central to Abel Tasman Nat. Park beaches golf trout rivers art wineries
Across from Piha beach set in native bush: a modern self contained cottage. Queen bed, ensuite bathroom, kitchenette (microwave, hotplate, electric appliances, dishes). Lounge with sofa bed. Private & peaceful, perfect for couples. No smoking/pets.
Luxury 1, 2 & 3 bedroom serviced apartment accommodation with fabulous harbour views at Prince's Wharf in the centre of downtown Auckland. Short stay or long term rental - the perfect hotel alternative.
Find a Northland, New Zealand vacation rental home for your vacation. Stay at one of our rustic Northland, New Zealand cabins, cozy cottages, or authentic condos. We have everything from large rental homes to small villas. Rent for a weekend, week, month, or entire season! Vacation rentals are great accommodations for families and large groups. Enjoy the convenience of a full kitchen and save money by eating in. Living like the locals at a vacation rental assures you an authentic travel experience. Browse our Northland, New Zealand vacation rentals and plan your vacation today!


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Monoprice 109927 Enhanced Bass Hi-Fi Noise Isolating Earphones, Black
"I purchased these a long time ago after reading reviews that these were a good budget option in the earphone category. To save time, i am just going to jump in to the sound quality section.. Bass: A little bass light for my tastes, but i can live with it. I doubt people are going to complain with these at the price they are offered at. It has decent punch to it, but like i said, a..." --zach8278
Monoprice 8320 IEM
"I bought my first set of 8320 two years or more back. They were very flat in the beggining then someone sugested these should be burned in for a few days with white noise. An old cell phone and white noise generator helped break these things in. Man o man what a difference. Made my bose IEM look like dollar store headphones. Sound is a relative thing IMHO just like taste in food. Some like..." --engineerd3d
Monoprice 8322
Get your bling on with these stylish Monoprice earphones! With a sensitivity of 98 dB and a noise-isolating design, these earphones will provide plenty of volume to keep the world outside and the music inside your ears. The 47" cord is long enough to give you the freedom to move about and to keep your iPod safely in your pocket when you're mobile. It isn't enough to just have...
Ying Yang Earbud Design Clip
Budclicks patented the ability to click charms to earbuds. They allow kids, tweens, and adults to style up those bland white earbuds. The Budclick mount works on iPod and iPhone style white earbuds created by Apple. With over 150 million of these earbuds on the market Budclicks have appeal to just about everyone. Budclicks has found the largest audience in music listeners on the go. Whether on...


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Parvovirus (Parvoviral Enteritis or "Parvo," for short) is a virus causing severe infection in puppies and dogs. It invades and destroys rapidly growing cells in the intestine, bone marrow and lymphoid tissue resulting in nausea, vomiting and severe hemorrhagic (bloody) diarrhea in dogs. The invasion of the bone marrow cells causes a decrease in the white blood cell count leading to increased susceptibility to bacterial infections and sometimes to a shock-like condition called endotoxemia. The disease can vary from mild to fatal (if not properly treated).
Infection is generally attributed to ingestion of material contaminated by dog feces (stool or bowel movement) and can occur when a dog smells or licks the ground; direct contact with another dog is not necessary for infection. Parvovirus is shed in the feces of infected dogs for approximately two weeks after initial ingestion and can live in the environment for years.


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Do you think they were BBQ'ing the dead? Is that where the food was coming from?
In any case:
1) Never thought they'd go into suggested child rape (which Rick prevented) but I liked being shocked by this even darker turn for TWD. Yes, it was very uncomfortable to watch. That's the idea. You don't forget!
2) Rick and his "appetite!" Wow. All I can say is that was awesome!
3) Terminus - BigB, I think you understand it better than I do (Your insight is welcome!) But those people will not be able to handle themselves against TWD's "A-Team." I wonder if they'll use a new "war" to kill some characters next season? I don't think the zombies can ever kill our group now - after Darryl escaped that funeral home 2 episodes ago?!
4) Now I think Beth (instead of being kidnapped for rape) might have been rescued from the funeral home as it could have been a trap for people to be taken to Terminus.
5) Terminus - with all its signs out there - and it being a definite bad place (did you hear the screams for help coming from the cattle cars?) - needs to be destroyed. To me it represents the Sodom & Gomorrah of TWD world. (I doubt I'm wrong.)
6). I still think Judith is a key.
She was conceived after the zombie apocalypse. Maybe it only affected those that breathed something in the air at that particular time? Like a bio-weapon gone haywire?
There is no cure for those that turn. But maybe Judith and those conceived afterward never will? (A Child Will Lead.... - WaterWorld, but also Christian mythology - not to mention SW did it with The Chosen One.) There's always that...and deferring to a child to lead is always a safe way for other "apostle types" to gather power for themselves as the caretaker-types (China and Japan's empires for example - even led us into WWII with Japan).
Anyway, Carl's role will eventually be his sister's protector. My prediction. Everyone else will someday turn.


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We don't have any offers to host the 2013 British Open currently, so we're inviting people to submit offers to run this annual tournament.
In the last few years we've been to Edinburgh, Malmesbury and this year to Durham, so it would be nice to have a location in the South East for 2013; but we're prepared to consider any offers.
If you're interested please talk to any Council member.


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By The Associated Press
GULFPORT — If all professional wrestler Jack Swagger had were tickets for speeding and misdemeanor marijuana possession, he could pay them by mail or online.
Gulfport city prosecutor Kirk Clark tells The Sun Herald that it is the driving under the influence charge that requires Swagger to return to Gulfport.
Swagger, whose real name is Donald Jacob "Jake" Hager Jr., was arrested just before midnight Tuesday in Gulfport on charges of speeding, misdemeanor marijuana possession and DUI.
Hager, 30, had just completed taping the TV show "WWE SmackDown" in Biloxi at the Mississippi Coast Coliseum.
Police Lt. Craig Petersen said police stopped his northbound car on Lorraine Road about six miles south of Interstate 10. He said Swagger's car was towed and he was released from custody to a friend who picked him up at the police station.
Swagger's municipal court date is March 12. The charges are misdemeanors.
If convicted, records show Swagger faces fines of $188 for driving 10 mph over the speed limit, $437 for a small amount of marijuana and $1,300 if it's his first DUI offense. The DUI charge carries a mandatory court appearance and he could lose his driver's license for 90 days.
Clark said the court has allowed some defendants in limited circumstances to plead guilty to DUI without a court appearance. That only happens, he said, when an attorney enters a plea by affidavit and the defendant signs a notarized waiver of rights.
"We have never recommended mere fines and fees only on a DUI," Clark said. "We always recommend the maximum sentence of two days in jail for first offenders. That time can be suspended but only on completion of certain court-ordered conditions like probation, classes, AA meetings, etc."
Kirk said a conviction includes an order to complete a Victim Impact Panel class, he said.
Swagger, who lives with his wife and child in in Boca Raton, Fla., is a native of Perry, Okla. The 6-foot-7, 260-pound Swagger played football and wrestled at the University of Oklahoma before he became a pro wrestler.
WWE said in a statement that "Mr. Swagger is responsible for his own personal actions."


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The Tigertones, pictured above in 1970, are Princeton's second oldest all-male a cappella group. A cappella ("without instrumental accompaniment") performances have become an iconic Princeton tradition for the students who sing and the many fans who enjoy their music. Today, there are 13 active a cappella ensembles on campus that encompass a range of genres and groups.
Photo courtesy of the Alumni Council
Play the "Sounds of Princeton arch sings" video featuring student a cappella singers.
Video still courtesy of Susan Fou
Featured Story
A tradition of voice: A cappella at Princeton
Posted June 23, 2011; 12:00 p.m.
Walk through Princeton's campus at night, and it is not unusual to hear sweet melodies and perfect-pitched harmonies resonate from one of the University's Gothic archways. The waves of music emanate from Princeton's student a cappella singers, who entertain their classmates at arch sings, in addition to concerts and informal shows, throughout the year.
A cappella, which means "without instrumental accompaniment," relies on the power of voice alone. Since the University's first a cappella ensemble, the Nassoons, was formed in 1941, the groups' performances have become an iconic Princeton tradition for the students who sing and the many fans who enjoy their music.
Princeton a cappella originated in 1941 when eight students broke away from the Glee Club and started the Nassoons, according to historical records. The all-male group is shown here in 1970. (Photo courtesy of Princeton University Archives)
"Singing with Koleinu has been and continues to be one of the highlights of my time here at Princeton. It's not just about music -- the group has become my on-campus family," said Rivka Cohen, a member of the class of 2012 and president of that coed Jewish a cappella group.
"We spend almost as much time laughing and supporting each other as we spend singing," she added.
Chris Palermo, president of the all-male Nassoons, said being in an a cappella group has defined his time at Princeton so far.
"More than the music and the tours, being a member of the Nassoons has given me the opportunity to meet a great group of people that make up the arts scene on campus," said Palermo, a member of the class 2013.
Sweet melodies and perfect-pitched harmonies often resonate across campus at night as student a cappella groups gather for informal arch sings. (Photo by Bentley Drezner)
According to historical records, Princeton a cappella originated when eight students broke away from the Glee Club and started the Nassoons. Just as many groups do now, the Nassoons' gigs quickly extended beyond campus, with concerts scheduled around the country, even catching the attention of legendary jazz vocalist Ella Fitzgerald during a 1942 event at the Plaza Hotel in New York, according to Nassoons' history.
Since then, Princeton a cappella has grown to encompass a range of genres and groups. Today, there are 13 active groups on campus.
The Tigertones and Footnotes emerged in 1946 and 1959, respectively, as all-male groups. With undergraduate coeducation, the all-female Tigerlilies was established in 1971 and was followed in 1973 by the Katzenjammers, which is considered to be the oldest coed a cappella group in the Ivy League. The Tigressions and Wildcats joined the Tigerlilies in the 1980s as all female groups; and coed a cappella expanded with Roaring 20 in 1983 and Shere Khan in 1994. Recent decades also have seen the formation of groups focused on particular musical genres or cultures, such as Koleinu, Christian a cappella singers Kindred Spirit, the South Asian music ensemble Tarana and Old NasSoul, whose repertoire is old soul and rhythm and blues tunes.
Students in 1942 enjoy Step Sing, which began in 1760 when graduating seniors gathered on the steps of Nassau Hall to sing their favorite songs. Seniors today assemble for Step Sings on the steps of Blair Arch, while a cappella groups use Blair Arch for informal arch sing performances throughout the year. (Photo courtesy of Princeton University Archives)
"I attended a musical showcase during freshman orientation and I froze in my seat as Old NasSoul's baritone section chimed in after the basses at the beginning of Marvin Gaye's 'I Heard It Through the Grapevine,'" said Osei Wilks, a member of the class of 2012. "I could not believe that a group of such quality and caliber was only a few years old, and I've dedicated myself to helping them grow even more."
No matter what the musical style of the various groups, almost all students extol the a cappella performances in arch sings as their most memorable experience. The outdoor performances are as old as a cappella at Princeton and are connected with the tradition of Step Sings, which began in 1760 when graduating seniors gathered on the steps of Nassau Hall to sing their favorite songs.
"Arch sings are really the backbone of a cappella at Princeton. The groups get to show off their music, and students can drop by any time to hear their favorite songs," said Tadesh Inagaki, a member of the class of 2014 and singer with the Katzenjammers.
Palermo said he believed the informal gatherings make Princeton unique compared with other collegiate a cappella experiences.
"The fact that the vast majority of groups on campus come together every few weeks to sing and listen to each other is one of my favorite aspects of singing at Princeton," he said.
The Katzenjammers, pictured here in 1994, are considered to be the oldest coed a cappella group in the Ivy League. (Photo courtesy of Princeton University Archives)
Lights shine on Nassau Hall for an evening Step Sing performance in 1947. A cappella arch sing performances are connected to the tradition of Step Sings, which sometimes feature musical accompaniment. (Photo courtesy of Princeton University Archives)
The all-female a cappella ensemble the Wildcats were formed in 1987. Here, the group is pictured in 1994. (Photo courtesy of Princeton University Archives)
Founded in 2006, Old NasSoul is one of Princeton's newest a cappella groups. Its repertoire focuses on old soul and rhythm and blues tunes. (Photo courtesy of Old NasSoul)
The Tigerlilies sing during a practice in 2011. The Lils, as they are known, became Princeton's first all-female a cappella group in 1971 following undergraduate coeducation. (Video still by Nick Barberio)
Students circa 1960 give an a cappella performance in Blair Arch. Arch sings have been taking place in the University's Gothic archways since a cappella first emerged on campus in 1941. (Photo courtesy of Princeton University Archives)


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Underscoring its commitment to audio excellence and ongoing support for the art and craft of recorded music, the Producers & Engineers Wing of The Recording Academy will kick off a yearlong celebration of its 10th anniversary with a GRAMMY SoundTables panel discussion at the 129th Audio Engineering Society Convention, taking place Nov. 4–7 in San Francisco.
The event, titled Sonic Imprints: Songs That Changed My Life, will take place on Nov. 6 with a panel discussion led by producer Sylvia Massy, featuring producer/engineer Joe Barresi, GRAMMY-winning producer/engineers Bob Clearmountain, Jimmy Douglass and Nathaniel Kunkel, and DJ Khalil as they explore tracks from their careers that have left lifelong impressions on them.
"When The Producers & Engineers Wing was established a decade ago, an essential voice was created for music makers: the voice of those 'behind the glass' that work so diligently to ensure that the quality and integrity of recorded music is captured and preserved," said Neil Portnow, President/CEO of The Recording Academy. "Protecting and advocating for the rights of this group, bringing awareness to new technologies and working to improve the overall landscape of recording practices will continue to be priorities for the P&E Wing. We have made great strides and achieved many victories on behalf of this constituency, and we look forward to what the next decade will bring."
"The Producers & Engineers Wing provides a voice for those involved in every aspect of the recording process," said Maureen Droney, Senior Executive Director of the P&E Wing since 2005. "The music production industry has faced enormous challenges in the 10 years since the Wing was created, and it continues to face new technical, legislative and economic tests today. The P&E Wing's leaders, and its membership as a whole, are a dedicated body of creative professionals that has provided The Academy and the industry with a unique and invaluable resource with which to address these challenges."
"The Producers & Engineers Wing is about supporting the profession and preserving the beauty and timelessness of music and artistry," said GRAMMY-winning producer/engineer Ed Cherney. "It's about ensuring that music distributed over the Internet retains the integrity of sound that producers and engineers work so hard to create; the quest for fair and equitable intellectual property rights for our profession; thinking of ourselves as part of a group with common goals and ideas; nurturing our skills; and sharing our experience and knowledge."
Currently more than 6,000 professionals comprise The Recording Academy Producers & Engineers Wing. Key P&E Wing achievements over the past decade include:
These are the most read, shared and discussed articles on GRAMMY.com right now.


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A: You can get a monthly or yearly subscription to Office 365 Personal or Home at Microsoft Store or other retailers or resellers. If you are a qualifying student, you can get a 4-year subscription to Office 365 University.
A: Sign in to your Microsoft account page and you'll be able to see which version of Office 365 you're subscribed to, and how much time you have left on your subscription.
A: As the end of your subscription approaches, you'll get renewal notices within Office and via email.
A: If you're renewing your subscription to the same version of Office 365 that you already have, then the time you have remaining on your current subscription will be added to your new one. So if you have two months left on your current subscription to Office 365 Home and you renew for one year, your subscription will run for 14 months.
A: You can only have one active subscription at a time, so you'd have to buy a new one. Any time you may have left on your existing subscription will be added to your new one. For instance, if you have two months left on your current subscription to Office 365 Home, which allows installations on up to 5 PCs or Macs plus 5 tablets or smartphones, and you buy a new one-year subscription to Office 365 Personal, which allows installation on only one PC or Mac and one tablet or smartphone, then you will immediately have to delete any additional installations beyond the one that your new license entitles you to, and your new yearly subscription to Office 365 Personal will run for 14 months.
A: You can wait for your current subscription to expire and then buy your new subscription after that.
A: Once your subscription expires, you'll be able to view or print documents, but you won't be able to edit existing ones or create new ones. You could use the free Office Web Apps on OneDrive for basic editing until you get a new subscription. Any documents you saved to OneDrive will remain, and you'll still be able to get to them.
A: At Microsoft Store, your subscription starts when you complete your purchase. If you purchased from another retailer or reseller, your subscription starts when you activate your subscription.


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Diary of a Wimpy Kid: The Long Haul
Diary of a Wimpy Kid: Dog Days
Diary of a Wimpy Kid: Rodrick Rules
Astro Boy
Flushed Away
Shark Tale
FernGully: The Last Rainforest
The Road to El Dorado
Movies.com, the ultimate source for everything movies, is your destination for new movie trailers, reviews, photos, times, tickets + more! Stay in the know with the latest movie news and cast interviews at Movies.com.
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"Save America's Treasures" has been totally eliminated—the largest Federal program supporting preservation of such treasures as the original Star Spangled Banner and George Washington's tent.
65% of Americans don't know what happened at the Constitutional Convention, according to a recent survey by Newsweek.
The "Teaching American History" grants—the largest Federal program supporting history education—have been completely eliminated.
Visits to the Top 20 Civil War battlefields have dropped in half from 1970 to 2009 according to official National Park Service statistics.
40% of Americans can't identify whom we fought in World War II, according to a recent survey by Newsweek.
A quarter of Americans believe Congress shares power over U.S. foreign policy with the United Nations, according to a recent Annenberg survey.
"There is little that is more important for an American citizen to know than the history and traditions of his country," John F. Kennedy wrote in American Heritage.
The "We the People Program," which touched some 30 million students and 90,000 teachers over 25 years, has been completely eliminated.
Two-thirds of Americans could not correctly name Yorktown as the last major military action of the American Revolution, according to a recent national Gallup survey.
The National Heritage Areas and Scenic Byways program, the only major Federal program encouraging visits to historic places, has been completely eliminated in Congressional committee.
The third in a series on TIMES OF TRIAL IN AMERICAN STATECRAFT
Old Hickory's attack on Biddle's bank had some unexpected consequences
Editor's Note: Bray Hammond wrote this essay for American Heritage in 1956 and developed it into Banks and Politics in America from the Revolution to the Civil War, for which he won the Pulitzer Prize for history in 1958.
"Relief, sir!" interrupted the President. "Come not to me, sir! Go to the monster. It is folly, sir, to talk to Andrew Jackson. The government will not bow to the monster. … Andrew Jackson yet lives to put his foot upon the head of the monster and crush him to the dust." Read more »
© Copyright 1949-2016 American Heritage Publishing. All Rights Reserved.


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SunTouch has announced the SunTouch TapeMat with several product enhancements making installation even easier than the classic SunTouch mat design.
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SunTouch Mats patented dual wire design is engineered with Safety built-in to eliminate electromagnetic fields (EMF) and uses the highest quality materials for durability and longevity.
High heat output of 41 BTU/SF can replace many baseboard heaters as SunTouch uses only a 12 Watts/SF.
Installation is easy with our how-to videos online and adetailed instruction manual is included too. When replacing your flooring the addition of a SunTouch Heated Floor may be your best decision and the temperature is easily controlled with 1 of our SunStat thermostats.
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- Made in the USA by Watts Radiant a leader in safety and comfort since 1874 (NYSE:WTS).


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Web Stories
Lecture examines Chilean presidency, Nov. 27
Posted November 15, 2006; 11:51 p.m.
Eugenio Lahera, a Princeton alumnus and senior adviser to former Chilean President Ricardo Lagos, will speak on "The Presidency in Developing Countries: The Case of Chile, 2000-2006" at 4:30 p.m. Monday, Nov. 27, in 16 Robertson Hall.
Lahera, who holds an MPA and Ph.D. from the Woodrow Wilson School of Public and International Affairs, is the John L. Weinberg/Goldman Sachs and Company Visiting Professor at the Wilson School and an associate professor at the Universidad de Chile. He also is executive director of the Fundacion Chile 21 in Santiago, a social and economic think tank.
Lahera served as senior adviser and chief of public policies for Lagos, who was Chile's president from 2000 to 2006. Lahera previously worked for the United Nations Economic Commission for Latin America and Chile's foreign ministry.
The event is sponsored by the Wilson School and the Program in Latin American Studies.


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So polls drive me crazy. A few weeks ago Texas Tech beats the heck out of WV but in the polls they were 10 places higher the next day. The worst thing I think about them is that they make these random guesses a month before the season starts and that initial ranking impacts them all season long. I am from NC and a Georgia Tech Alumni but I admit that the ACC is awful. How is Florida St. #9 and Clemson #13. Clemson only wins against winning records is against Ball St. and a 4-4 Wake Forest team. Florida St. will end up winning the rest of their games until Florida St. and will end the year 10-2 but the only good win (against a team with a winning record) will be against Clemson but they will end up in the top 10. FSU would be a .500 team in the SEC and would lose 3-4 games in the PAC.
these are the 2 biggest pretenders in the top 15 I think. Anyone think there are any others?


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NAKAMARRA, Doreen Reid
Marrapinti is the rock hole site west of the Pollock Hills in Western Australia.
Ancestral women of the Nangala and Napangati subsections camped at Marrapinti during their travels east. There, the women made nose bones, also known as marrapinti. During ceremonies relating to Marrapinti, the older women pierced the nasal septums of the younger women who were participating in the ceremony. Now, nose bones are only used by the older generation for ceremonies.
Upon completion of the ceremonies at Marrapinti, the women continued their travels east, passing through Wala Wala, Ngaminya and Wirrulnga, before heading north east to Wilkinkarra [Lake Mackay].
Statement courtesy of Papunya Tula Artists Pty Ltd 2007.


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Media reactions
You can filter the media reactions by subject and/or by date.
Europe dragging its feet on tax dodging
Today EU leaders recognised the need to ramp up efforts to combat tax dodging, but failed to agree on a
Oxfam reaction to EU Court of Auditors report on EU climate finance
Oxfam reaction to EU Court of Auditors report on EU climate finance
$52 Billion for IDA, the World Bank's Fund for the Poorest
Donors have replenished the World Bank's fund for the world's poorest countries, the International Development Association, to the tune of
Oxfam strongly condemns the use of violent force against civilians, in Juba, South Sudan
Emma Jane Drew, Country Director (Acting) for Oxfam in South Sudan, said from Juba:
Oxfam reaction to EU biofuels debate by Energy Ministers
Commenting on the failure of EU Energy Ministers to reach agreement on EU biofuel policy today, Marc Olivier Herman, Oxfam's biofuels expert said:
Oxfam's reaction to the 9th WTO Ministerial Conference in Bali
In reaction to the new trade facilitation agreement approved in Bali on Saturday, Romain Benicchio, Senior Policy Advisor with Oxfam, said:
Oxfam's response to the death of Nelson Mandela
Winnie Byanyima, Executive Director of Oxfam International
"My direct contact with President Nelson Mandela was in the context of helping Burundian women to participate in the Arusha peac
Leaked EU document proves financial transaction tax is legal
A leaked European Commission document dispels earlier warnings that the fin
Geneva peace conference on Syria conflict must not be talks for the sake of talks
Nigel Timmins, who is heading up Oxfam's response to the Syria Crisis, said: "This announcement is long overdue.
Massive public pressure needed to rescue climate deal after Warsaw farce and fiasco
Only massive public pressure can rescue a climate deal, warned Oxfam today, as the Warsaw negotiations drew to a close after a fortnight of farce and fiasco.


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Modern Style
The lovely Xylophone Collection is made of cool interlocking minded segments of wood that invite children to explore. The lines are simple, yet intricate and bring a really cool puzzle to mind. The crisp simple colors NUMI NUMI is showcasing within the collection will look amazing in any child's space. Plus, if brought out of the bedroom or playroom, these pieces are sophisticated enough to look good in adult spaces too.
Sustainable Design
Like other NUMI NUMI collections, the Xylophone Collection is made of eco-friendly, but highly durable NAUF (no added urea formaldehyde) multi-ply birch wood. All finishes are non-toxic, water-based wood stains. The company only uses FSC (The Forest Stewardship Council – Monitored forest management) wood products and NUMI NUMI furniture contains zero veneer or plastic laminated materials.
The Xylophone Rocker
One of the most unique pieces in this collection is the Xylophone Rocker. This rocker is stylish and offers a wonderful place to relax, wonder, or read while rocking away.
The Xylophone Chair
This clever chair fits perfectly with the play table and offers more swooping style than a traditional four-legged chair. This chair, with it's single continuous shape that is reminiscent of a xylophone will work well even without the table, offering kids a place to sit and read or relax.
Easy Assembly
With simplicity in mind, NUMI NUMI, designed the entire Xylophone Collection with a zero hardware assembly feature, meaning you won't need tools to put these pieces together. Pieces in the collection snap together with a basic snug fitting design. The entire Xylophone Collection is coming soon. Visit the NUMI NUMI website to see more of their wonderful designs and to keep track of new product release dates.
+ Xylophone Collection – coming soon


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I can answer questions concerning physical and historical geology, environmental geology/hydrology, environmental consulting, remote sensing/aerial photo interpretation, G&G computer applications, petroleum exploration, drilling, geochemistry, geochemical and microbiological prospecting, 3D reservoir modeling, computer mapping and drilling.I am not a geophysicist.
I have 24 years experience split between the petroleum and environmental industries. I have served as an expert witness in remote sensing, developmental geologist, exploration geologist, enviromental project manager, and subject matter expert in geology and geophysical software development.
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Manufacturer: Nakajima Manufacturer: EarFuze 2 Results
FUZE Custom Earphones by EarFuze
Fuze Custom Earphones are the ultimate upgrade to your iPod/mp3 player. Perfect fit, comfortable, great sound and incredibly secure. Perfect for sports, helmet and motorsports use! Youll never have the annoying problem of earbuds that shift and fall out. Your music will always be in the sweet spot. Fuze Custom Earphones come packaged in an easy do-it-yourself system. By following the simple...
Kuromi Glam Earphones
Enjoy your favorite music the super stylish way with this neat Kuromi glam earphone. The ideal accessory for your iPod or other audio players. The cute design features Kuromi with shimmery pink metallic highlights. Digital stereo sound. Super lightweight.


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Web Stories
Quipfire! improv troupe to perform, April 29-May 1
Posted April 28, 2004; 05:14 p.m.
The Quipfire! student improvisational comedy troupe will perform at 9:30 p.m. Thursday through Saturday, April 29-May 1, at Murray-Dodge Theater. Audience members are encouraged to bring props.
Tickets are $5. They can be reserved by calling (609) 25-UCALL and entering IMPROV or by visiting the Quipfire! Web site .
Contact: Lauren Robinson-Brown (609) 258-3601


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You are here
Those suffering from post-Spoleto art withdrawl took great delight in Art For Charity's finale on Friday. The Michael Mitchell Gallery drew a large crowd of art aficionados, all of whom were more than happy to support the evening's cause, MUSC's Children's Hospital and Hollings Cancer Center.
The King Street space displayed works from more than 30 regional artists, including Ester Melton, Lee Monts, Karen Keene Day, and Marlies Williams, all of which were available for sale. Guests perused pieces while sipping glasses of wine and munching on incredible sushi from Oku. All the while strains of Michael Jackson pulsed through the gallery, lending a fresh, young feel to the event.
People wandered in and out throughout the evening, but the gallery always seemed to be comfortable crowded. Several people purchased work, and several artists were on hand to mingle with the partygoers.
Art For Charity was a wonderful way to kick off a Friday night on the town and pick up a couple new pieces all in one stop.
Art For Charity took place between May 20 and June 20, 2011 at Michael Mitchell Gallery. The event featured works in a variety of mediums from over 30 featured artists and benefited six local non-profit organizations.


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Neurointerventional Radiology
The quick treatment of patients suffering from aneurysms, strokes or serious back injury can have a dramatic impact on the recovery of the patient. The Eisenhower Neuroscience Institute team utilizes a number of intervention techniques to treat the patient within the brain or vertebrae before they experience debilitating brain or spinal injury. In the brain, these treatments include dissolving clots, placing coils in aneurysms and inserting angioplasty or stents in narrowed vessels. In the spine, Eisenhower interventional physicians perform vertebroplasty and kyphoplasty, procedures that inject a cement-like material into the vertebrae to stabilize a fracture and alleviate pain.
Neurointerventional Radiology Team
Mehran Elly, MD
- Radiology*
- Interventional Radiology
Brian Herman, MD
- Radiology*
- Interventional Radiology
Jerry Chang, MD
- Radiology
- Interventional Radiology


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Inferring the kinetics of stochastic gene expression from single-cell RNA-sequencing data
- Jong Kyoung Kim1 and
- John C Marioni1Email author
DOI: 10.1186/gb-2013-14-1-r7
© Kim and Marioni licensee Springer. 2013
Received: 7 November 2012
Accepted: 28 January 2013
Published: 28 January 2013
Genetically identical populations of cells grown in the same environmental condition show substantial variability in gene expression profiles. Although single-cell RNA-seq provides an opportunity to explore this phenomenon, statistical methods need to be developed to interpret the variability of gene expression counts.
We develop a statistical framework for studying the kinetics of stochastic gene expression from single-cell RNA-seq data. By applying our model to a single-cell RNA-seq dataset generated by profiling mouse embryonic stem cells, we find that the inferred kinetic parameters are consistent with RNA polymerase II binding and chromatin modifications. Our results suggest that histone modifications affect transcriptional bursting by modulating both burst size and frequency. Furthermore, we show that our model can be used to identify genes with slow promoter kinetics, which are important for probabilistic differentiation of embryonic stem cells.
We conclude that the proposed statistical model provides a flexible and efficient way to investigate the kinetics of transcription.
Keywordsgene regulation RNA-seq single-cell statistics transcriptional burst
RNA-sequencing (RNA-seq) is a recently developed approach that allows an unbiased examination of the transcriptome to be performed using high-throughput DNA sequencing [1–3]. Compared to gene expression microarrays, the previous gold standard for genome-wide quantification of gene expression levels, RNA-seq has some specific advantages: it allows splicing to be assayed in an unbiased manner [4], it better enables the measurement of expression levels over a wide dynamic range [1], and it allows allele-specific expression to be interrogated [5, 6].
Until recently, most RNA-sequencing experiments began with a large population of cells (> 105), and, as a result, the gene expression counts obtained can be viewed as an average across that population. However, recent developments in sequencing technology have enabled the use of much smaller volumes of starting material, and several groups have described protocols for assaying the transcriptome of single cells [7–11]. This is vital in many biological contexts, such as early embryonic development and tumor etiology, where it is expected that different cells will have distinctive expression profiles. Furthermore, even in tissues that are typically considered to consist of homogeneous populations of cells, inter-cellular variability in gene expression levels can be considerable. For example, the cells of a genetically identical population grown in the same environment have been shown to display substantial variability in the total number of mRNA molecules that they contain [12–14]. This variability can be partially explained by noting that gene expression levels are regulated by combinatorial interactions between numerous cellular components, where these interactions involve random biochemical reactions [12, 13, 15].
More generally, single-cell imaging methods (e.g., RNA fluorescence in situ hybridization or FISH) have been widely applied to elucidate the principles of gene expression regulation in vivo [16]. These studies have observed that: i) gene expression is heterogeneous [12–14]; ii) genes fluctuate between an 'on' and 'off' promoter state and transcripts are produced in bursts [17–19]; and iii) the transition to the 'on' state requires multiple rate-limiting steps that are determined by many sequential interactions between regulators and chromatin, but the transition to the 'off' state can be determined by a single rate-limiting step [16]. Some examples of the stochastic processes that play a role in the transition to the 'on' state are the recruitment of nucleosome remodelers and histone-modifying enzymes by activators, the rate at which RNA polymerase II (PolII) escapes from the core promoter to produce short RNA molecules prior to pausing, and the rate at which PolII leaves pausing and enters productive elongation [15].
One situation where stochastic fluctuation in gene expression levels plays an important role is in the regulation of mouse embryonic stem (ES) cells [14]. Mouse ES cells are derived from the inner cell mass (ICM) or the epiblast of the pre-implantation blastocyst [20], and they can proliferate in the same undifferentiated state indefinitely whilst retaining the ability to differentiate into all adult cell lineages. These two hallmarks of ES cells are conferred by tightly controlled gene regulatory networks [21]. However, growing evidence suggests that the ability of an individual ES cell to differentiate into an adult cell type at a specific time is determined stochastically [14, 22]. In particular, the expression levels of key regulatory genes, such as Nanog, Stella, and Rex1, which are markers of pluripotency, are heterogeneous in ES cells even though the cells are cultured in the same condition [23]. This implies that ES cells exist in a dynamic equilibrium between states that show different propensities for differentiation [22–24].
Here, we develop a statistical framework motivated by a kinetic model for transcriptional bursting to model the biological variability present in single-cell RNA-seq data. The framework derived makes it easy to perform parameter fitting and allows the kinetics of transcription to be investigated. We apply our model to single-cell RNA-sequencing data generated from mouse ES cells and demonstrate that the estimated parameters are consistent with promoter kinetics inferred from RNA polymerase II binding and chromatin state profiles.
A kinetic model for stochastic gene expression
Moreover, when the gene is in the active promoter state, it is assumed to be transcribed at a rate, s, per unit time and the number of mRNA molecules of the gene is assumed to decay at a rate, d, per unit time. Subsequently, transcriptional bursting can be characterized by two parameters: the average number of synthesized mRNA molecules while a gene remains in an active state (burst size or transcriptional efficiency, s/koff) and the frequency at which bursts occur per unit time (burst frequency, kon) [18, 26–28].
Given these four kinetic parameters, a set of differential equations has been derived describing how the number of mRNA molecules of a given gene within a cell, x, changes over time (Additional file 1; [17]). The steady state distribution of these equations has been shown to take the form [17–19]:
As noted previously, the four kinetic parameters are all measured in units of time. However, since the inverse of the decay rate, 1/d, denotes the average lifetime of an mRNA molecule, it can be used to normalize the other kinetic parameters so that they are independent of time [16, 17]. This is equivalent to setting d = 1 in (2), and we do this henceforth.
A Poisson-beta model
The parameters of the steady-state solution (2) have previously been estimated from observed data using two different approaches. The first approach is to match the first three moments to their empirical values [17]. Although this method is straightforward and computationally efficient, it does not guarantee that the estimates are within the parameter space. To overcome this problem the maximum likelihood estimates of the parameters can be found using a numerical optimization approach [18]. However, the computation of the confluent hypergeometric function, 1F1, is difficult, because there is no numerical method for its accurate, fast and reliable computation within all parameter values [29]. Furthermore, when the number of observations is small (less than 100), the maximum likelihood approach sometimes gives unrealistically large estimates of the kinetic parameters [18].
where p is an auxiliary variable following a beta distribution. The marginal distribution P(x|s, kon, koff), which is known as the Poisson-beta distribution (PoBe) [30], takes the same form as the steady-state distribution described in equation (2).
Interestingly, the mean of the auxiliary variable p is equal to the fraction of time that a gene spends in the active state (1). Further, Smiley and Proulx [31] showed that if a gene's expression level oscillates between 0 and s/d and the maximum expression level of the gene (s/d) is set to 1, then given the two-state model, its stationary distribution takes the same form as that of p.
Given count measurements from RNA-sequencing data, we assume that the number of reads mapped to a gene is proportional to the expression of the relevant mRNA molecule in the cell under study, and thus the parameters of the kinetic model can be inferred using a Bayesian hierarchical approach, such as a Gibbs sampler.
One of the most significant challenges in applying the kinetic model for gene expression is interpreting the parameters. As noted in a recent review by Munsky et al. [25], when kon and koff are large the transitions between the promoter states are rapid, resulting in a Poisson or negative binomial-like distribution of the number of mRNA molecules [18, 19]. In the context of fitting the kinetic model to real data this corresponds to areas of the parameter space where the three parameters are not identifiable (Method; Figure 1B). By contrast, when kon and koff are small, there are relatively few transitions between the two promoter states and the resulting distribution of gene expression molecules between different cells is bimodal - here all three parameters are identifiable (Figure 1B-E).
In practice, to ensure that the parameters are statistically identifiable, we suggest fitting three models (Poisson, negative binomial and Poisson-beta) to each gene before using a goodness-of-fit statistic to determine whether there is evidence that the parameters of the Poisson-beta model can be identified unambiguously (Methods). An alternative approach would be to fit a hierarchical Bayesian model to each gene and to use this to determine the best fitting distribution.
Assessing the reliability of the Poisson-beta model
Single-cell RNA-sequencing was recently used to assay the transcriptome of 12 mouse ES cells derived from the ICM at embryonic day 3.5 (E3.5) [8]. To explore the transcriptional kinetics of ES cells, we fitted the Poisson-beta model to these data (Methods). Before interpreting the inferred kinetic parameters, it is necessary to: i) account for the high amount of technical variability present in single-cell RNA-seq data; ii) consider whether the parameter estimates are statistically identifiable; and iii) assess whether we can draw meaningful inferences about transcriptional kinetics based on gene expression measurements from 12 cells.
Accurately quantifying the technical variability present in single-cell RNA-seq data is challenging. While experimental approaches vary, most suggest that when replicate libraries are generated from small quantities of RNA (taken from the same, large, population of RNA), the resulting read counts display more technical variability, especially for lowly expressed genes, than is observed in population-based RNA-sequencing analyses [7, 10, 11]. This is likely due to experimental factors such as the efficiency of the RT step and the PCR amplification when small quantities of starting material are considered [7, 10, 11]. Some attempts have been made to characterize the technical variation using spike-ins [11] but evidence for the efficacy of such approaches is still limited. Given these challenges and the limitations of current experimental approaches, we instead removed lowly expressed genes that are most likely to display high technical variability [7, 10, 11]. We considered a gene as lowly expressed if the maximum normalized read count was less than 50. This cutoff was determined using technical replicate data generated using the same protocol applied to the 12 ES cells [8] or oocytes [7] (Additional file 1, Figure S2). Across the set of 18,735 genes that were expressed in at least one cell, 12,551 genes had an expression level above this cutoff, and we fitted the Poisson-beta model separately to each of these genes (Additional file 1, Figure S3).
- 1.
Genes with relatively large values of koff,iand low values of kon,i. This corresponds to genes that have a low expression count in most cells and high expression in a small number of cells (typically one). When we simulated data from the Poisson-beta model with parameter values in this range (Methods), we found that kon,iwas estimated accurately, but that both koff,iand s i were underestimated (Additional file 1, Figures S6, S11).
- 2.
Genes with large values of koff,iand kon,i(Additional file 1, Figure S3). This set of genes are typically highly expressed (Additional file 1, Figure S12C, F) with a relatively low amount of variability across cells, as evidenced by the large values of koff,iand kon,i. While it is possible that this set of genes do have very fast promoter kinetics, statistically it is impossible to distinguish this from their being permanently in an active state (that is, koff,iis equal to or very close to zero). Hence, it is impossible to interpret either the raw or the derived parameters in this situation (Additional file 1, Figure S12). More generally, this explains why we do not observe many low values of koff,iin our set of parameter estimates. Moreover, it helps to explain some of the identifiability problems that other approaches have encountered when estimating koff,i.
Given these observations, we focus henceforth on the 10,298 genes that have identifiable estimates of the kinetic parameters. However, before going on to make biological inferences based upon these parameters it is first necessary to assess whether any meaningful conclusions can be drawn from fitting the Poisson-beta model to data from only 12 independent ES cells.
To do this, we fitted the Poisson-beta model to data simulated using the estimated parameters by increasing the number of cells from 3 to 100 (Methods). As expected, the correlation between the parameter estimates and the true values improved as the number of cells increased (Additional file 1, Figure S4-S8), with a good agreement when 12 cells were considered (Additional file 1, Figure S6-S8). Our simulations also displayed a tendency to underestimate s i ; the extent of the underestimation decreased as the number of cells increased (Additional file 1, Figure S8). One effect of this is a slight bias in the estimated values of kon,iand koff,i(Additional file 1, Figures S6-S8). This is not unexpected since s i can be considered to represent the 'maximum' rate of transcription and, especially when the number of cells is small, a cell where a gene is expressed at this 'maximal' value will not be simulated. Nevertheless, our simulations do provide confidence in the fit of the Poisson-beta model when a moderate number of cells (greater than or equal to 12) are considered. However, it is important to acknowledge that drawing strong biological inferences about the kinetic parameters of individual genes from only 12 cells is difficult - hence, in what follows we consider properties of sets of genes with specific values of the kinetic parameters. This will also help mitigate any effect that technical noise in the measurement of gene expression levels will have upon our interpretation of the data.
Transcriptional kinetics of mouse ES cells
To explore how the kinetic parameters provide information about the regulation of transcription we utilized independent information collected from v6.5 mouse ES cells (derived from the ICM at E3.5) on RNA polymerase II (PolII) occupancy and various histone modifications [32–34]. At the global level, the rates of transcription and gene activation are strongly correlated with the average expression level while the rate of gene inactivation displays a more modest correlation (Additional file 1, Figure S13).
The Poisson-beta model provides a convenient statistical framework for modeling single-cell RNA-seq data and for studying the kinetics of stochastic gene expression. Since the kinetic parameters of individual genes inferred from the small number of cells are likely to be noisy and may be influenced by technical variability, we focused on the summary properties of genes. Importantly, we confirmed that the kinetic parameters derived from the Poisson-beta model are consistent with PolII binding and chromatin modifications using single-cell RNA-seq data generated from mouse ES cells. Our results suggest that the chromatin state of genes, defined by H3K4me3, H3K27me3 and H3K36me3 modifications, affects transcriptional bursting by modulating both burst size and frequency, consistent with a recent study that suggested chromosomal location affected these kinetic characteristics [40].
However, while our model has clear advantages, it also has a number of limitations. First, in this manuscript, we do not address the modeling of the technical variability directly, primarily because our understanding of how different experimental characteristics (RT efficiency, PCR amplification, etc.) might contribute to the noise is very limited. Instead, we focus only on genes that are moderately to highly expressed, since previous single-cell RNA-seq studies have shown that such genes display less technical variability. However, as our understanding of the technical variability inherent to single-cell RNA-seq increases, it will be important to adapt the model presented herein.
Second, in common with most other biochemical models of gene expression, we assume that the rate of transitions to the 'on' state is governed by a single rate-limiting step. While this assumption facilitates the derivation of a closed-form solution for the master equations, and thus the implementation of the Poisson-beta model described in this paper, in higher eukaryotes activation requires many sequential steps [15]. However, the limited experimental data about the relative contribution of the different steps justifies the simplified model presented herein.
Third, the three kinetic parameters are currently measured in units of 'per mRNA average lifetime' since they are normalized by the decay rate. To estimate them in units of 'per second', we should directly measure the decay rates of all genes. This can be done by metabolic labeling of RNA with 4-thiouridine coupled with massively parallel sequencing [41]. Another improvement would be to measure the number of mRNA molecules directly rather than using the number of reads as a surrogate, which can be done by accurate digital quantification of transcriptome via digital RNA-seq [42].
Finally, our model assumes that the transition times and kinetic parameters are identical for the two alleles of each gene. A recent study established that 39% to 51% of heterozygous loci show allele-specific expression when expression patterns are measured in single cells of a two-cell embryo [43]. This suggests that the kinetic parameters and transition times of the underlying Markov chain might differ significantly between the two alleles of a gene. Further, Miyanari et al. [44] showed that Nanog is largely expressed from a single allele in ES cells and can transition between alleles randomly. Such variability can be incorporated into our model by measuring the expression of each allele independently (for example by using MMSEQ [45]), and using these measures as the input to the model. However, the mouse ES cell data we analyzed were generated from an inbred population of mice (C57BL/6J) and, as a result, we could not apply this approach. Examining allele-specific variation using the Poisson-beta model provides an interesting avenue for future research.
To summarize, as the single-cell field progresses towards analyzing the transcriptome of large numbers of individual cells in parallel, it will become increasingly important to develop statistical methods that accurately model stochastic gene expression. In this context, we anticipate that the Poisson-beta model presented here, and other similar approaches, will be vital in maximizing the amount of biological insight that can be obtained from these data.
Materials and methods
Properties of the steady-state solution of the master equation
A hierarchical Bayesian model
- 1.Draw s i for gene i from a gamma distribution:
- 2.Draw kon,ifor gene i from a gamma distribution:
- 3.Draw koff,ifor gene i from a gamma distribution
- 4.Draw p ij for gene i and cell j from a beta distribution
- 5.Draw x ij from a Poisson distribution
where t i is the length of gene i (the length of the transcripts measured in bp) and t j is the normalization factor for cell j. We used the scale normalization method of [46] to estimate the normalization factor for each cell. We make an implicit assumption that the number of reads is proportional to the number of mRNA molecules present in a cell.
The graphical model representing this generative process is shown in Additional file 1, Figure S1.
Learning by collapsed Gibbs sampling
Let be a set of observed read counts, and be a set of p ij . We treat the top-level variables, shown in the graphical model in Additional file 1, Figure S1, , as fixed hyperparameters. We derive a collapsed Gibbs sampler to infer all unknown variables given . In the following, a subscript with a minus sign that is attached to a set of variables means that the variables indexed by the subscript are excluded from the set.
- 1.Sampling p ij
- 2.Sampling kon,i
- 3.Sampling koff,i
- 4.Sampling s i
For the hyperparameters, we used the following settings: , , , , and We chose the empirical Bayes prior on s i so that it becomes almost uniform across all realistic ranges for the parameter. The priors on kon,iand koff,iwere chosen to place substantial probability across the identifiable parameter space. When we used different priors on kon,iand koff,i(, , , for more diffuse priors and , , , for more concentrated priors), the inferred kinetic parameters remained similar, except for the large values of the parameters that were penalized by the concentrated prior. These results suggest that our model is relatively insensitive to the choice of priors (Additional file 1, Figure S9, S10).
Bootstrap-based goodness-of-fit test
- 1.
Given the maximum likelihood estimates , generate n bootstrap samples from .
- 2.
Compute the maximum likelihood estimates from the bootstrap samples.
- 3.Estimate the empirical distribution function of the bootstrap samples
- 4.Compute the KS test statistic such that
For this study, we set n = 1,000 and B = 1,000.
Estimating the kinetic parameters from synthetic data
Given the posterior means of s i , kon,iand koff,ifor each gene, we generated 3, 6, 12, 20 or 100 independent samples from the Poisson-beta distribution. We run the Gibbs sampling algorithm by setting the total number of Gibbs iterations to 10,000, and computed the posterior means by discarding the first half of the samples in each chain as a burn-in period.
Gene activity and pause index for GRO-Seq
To quantify PolII activity at the promoters and gene body regions, we defined three measures based on the number of mapped reads of GRO-Seq data [49]. First, gene body activity is defined as N/L where N is the number of GRO-Seq reads mapped from +1 kb of the transcription start site to the end of a gene, and L is the length of the region. Second, gene promoter activity is defined as the maximum count of reads in a 50 bp window, where we took the maximum among all the windows within the ± 1 kb region of transcription start sites. Finally, we defined the pause index as the ratio of the gene promoter activity (divided by 50) to the gene body activity.
Gene ontology analysis using DAVID
To examine whether particular classes of GO biological processes (GOTERM_BP_FAT) are enriched in the top or bottom 3,000 genes sorted by koff,i/ kon,i, we used the DAVID functional annotation clustering tool (the classification stringency was set to 'medium') [36]. By setting up the 10,298 genes as a background, we chose a representative GO term from each annotation cluster with the Benjamini-corrected P value less than 0.05, providing 18 GO terms in total. The results are in Additional file 2, Table S1 (top 3,000) and Additional file 3, Table S2 (bottom 3,000).
Code availability
The MATLAB source code and a compiled version of the same are available in Additional file 4.
- bp:
base pair
- ES cell:
embryonic stem cell
gene ontology
inner cell mass
polymerase chain reaction
- PoBe:
RNA fluorescence in situ hybridization
- RNA PolII:
RNA polymerase II
reverse transcriptase.
We thank Simon Anders, Wolfgang Huber, Anestis Touloumis, Nuno Fonseca and all members of the Marioni group for helpful comments.
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