Editor's note: This story about college football's bulldog mascots was originally published on Jan. 22, 2020. Wednesday, Aug. 26 is National Dog Day.

LOW TO THE ground and panting, Bully XXI sways hip to hip down the Mississippi State sideline. After four years on the job, the English bulldog is used to 87,000-seat stadiums like Auburn's and might even sense that MSU's first possession on this September night -- a terrible three-and-out that gives Auburn great field position -- pleases no one on the MSU bench. He and his owner, Lisa Pritchard, saunter beyond the team and pull even with the Auburn end zone as the Tigers' JaTarvious Whitlow takes a handoff. A gap widens off tackle and Whitlow explodes through it, cutting to the MSU sideline. He races toward the pylon as Bulldogs safety C.J. Morgan closes on him. Whitlow gets one foot into the end zone before Morgan pushes him out of bounds, and as Whitlow falls and slides -- he hits Bully XXI hard.

The collision throws the dog into the air. When Bully lands on his paws a couple of feet away, he tenses, as if wondering whether Whitlow will do this again, and if he should attack him. Whitlow runs off to celebrate the TD, though, and Pritchard tugs Bully's leash and brings him close. She is a certified veterinary technician who works in MSU's Department of Clinical Sciences. She is also a loving and pampering mother who lets Bully sleep in her bed, takes him with her on vacation and manicures his nails before home games. She quickly inspects Bully and then, even more quickly, because something isn't right, walks him to the locker room. ESPN, broadcasting the game, produces a sideline report in which viewers see the dog, in slow motion, flying through the air.

What's Wrong With Bully becomes a plot point in an otherwise embarrassing loss for the Bulldogs. Pritchard gets concerned tweets from fans and has the dog fully examined over the weekend. On Monday, Sept. 30, Mississippi State tweets that Bully has "bruising to his chin and right hind leg" but will make a full recovery and "return to all his mascot duties on October 13." The following day, People for the Ethical Treatment of Animals writes a letter to Mississippi State president Mark E. Keenum, urging the school to "retire" Bully because the collision "could easily have left [him] severely injured or even dead." There is much guffawing over this in Starkville: Bully has been a live mascot since the 1930s, which means no one is retiring anyone. Says one local columnist: "PETA, you're barking up the wrong tree."

What everybody misses, though, is the reason Pritchard walked Bully beyond the MSU bench and toward the Auburn goal line in the first place.

It was hot that day, with a heat index at the 6 p.m. kickoff around 100 degrees. That kind of heat is among the many dangers a modern bulldog faces, bred to be stouter and more flat-faced than its forebears and, as a result, with a weaker constitution and a tiny nose that prevents it from breathing properly. Pritchard knew all this. She could tell she needed to act by the way Bully moved. He was slowly swaying hip to hip down the sideline and heading to the locker room before he got hit because Pritchard was trying to address a larger and more chronic problem. She wanted to make sure Bully didn't overheat. She wanted to make sure Bully didn't die.

There are more bulldog mascots in Division I college football than any other animal, the oldest of which is Yale's Handsome Dan. Bettmann via Getty Images

THE ENGLISH BULLDOG is a peculiar breed. It is objectively, almost aggressively, ugly, that tiny snout and wide jowls drawing themselves into a wrinkled mass, as if it is forever puzzled by a smell it doesn't like. Its frame is short and stocky, less a dog's than a pig's, but with a chest that's disproportionately broad, as if it were auditioning for the role of canine superhero. There is a listless roll to the bulldog's gait, implying it'll move only out of obligation.

Its flaws are also the features that endear the dog to us: a face so ugly it's cute, a body so compact it's powerful, a gait so slow it's confident. An English bulldog is not as much a companion as a statement, and what that statement says, for the dog and owner alike, is I am comfortable with myself and my place in the world. It has a vestigial athleticism, the residue of earlier eras when the dog looked fiercer and was the reason college after college -- 40 in all, according to the American Kennel Club -- chose it as a mascot. There are more bulldog mascots in Division I than any other animal. Many of these are still live bulldogs, like at Mississippi State, Yale and Georgia. Yale's first Handsome Dan stalked the sidelines in 1889, making it the oldest live mascot in college football. The dog's popularity in sports, coupled with its playful demeanor and sedentary habits, translates to something close to ubiquity on Instagram and among the celebrity set: John Legend, David Beckham, Reese Witherspoon and Adam Sandler all own bulldogs. The breed climbed from the 14th most popular in 2004 to the fourth most popular today, according to the AKC.

Auburn's JaTarvious Whitlow didn't just score a touchdown against Mississippi State. He collided with MSU's beloved mascot, Bully. Michael Chang/Getty Images

Why is an ugly dog so beloved? One reason, according to Dr. Niels Pedersen of the University of California at Davis, is anthropomorphism, the term for when animals acquire human attributes (see: nearly every Disney movie). The bulldog's small nostrils, its drowsy eyes, its rolls of flesh -- it kind of looks like an ugly baby. People love bulldogs, Pedersen argued in a 2016 paper in the journal Canine Genetics and Epidemiology, because they literally see part of themselves in these pets. This leads to a desire to see more of themselves in their dogs, and with each generation, breeders accentuate the dog's already comical dimensions.

And therein lies the problem: As the dog has acquired more human attributes, it has also gotten sicker. The bulldog ranks first among all canines in respiratory disease and second in puppy mortality. According to Pedersen's research, the average bulldog now lives just six to eight years, down from 10 to 12 a few decades ago. The high risk of heart, lung, skin or severe joint problems further dwindles the pool of healthy bulldogs who can pass on their genes to the next generation. By examining the DNA of 102 bulldogs from seven countries, 87 of which came from the U.S., Pedersen found inbreeding so ubiquitous that the only path forward for the dog was to "outcross": find another breed with which to mate so that healthy genes might be passed on. Bulldogs "have been bred into a corner," Pedersen says.

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Some people, like Annette Nobles, a spokeswoman for the Bulldog Club of America, take issue with this claim. "The BCA does not believe the small number of samples used in Dr. Pedersen's study represent the breed," she and other BCA executives wrote in response to Pedersen's research. But Nobles does not dispute that due to the bulldog's compact frame and accompanying weight issues, the average male no longer mates. Females tend to be artificially inseminated. The average female doesn't deliver pups naturally -- the head is now too wide -- and 80% of the time vets perform cesarean sections. In other words, without modern reproductive science, the English bulldog would risk extinction.

Bulldogs were once so virile that after World War I they were named the mascot of the Marines. In Britain, the bulldog was used to symbolize Winston Churchill's tenacity and his country's endurance in World War II. Today, however, the British Veterinary Association, concerned about the breed's suffering, has urged breeders to outcross the dog and has called for a ban on bulldog advertisements, as if the dog were a toxic cigarette company. The dog's respiratory problems are now so pronounced that many American airlines forbid it from traveling with its owners. The airlines fear that flat-faced canines like the bulldog might die in flight.

"If the genetic mutations we see in bulldogs occurred in humans, it would be terrible, horrible," Pedersen says. "And we take these mutations and we make them a feature of the [bulldog]."

Experts believe the bulldog's decline has accelerated over the past 40 years. Some vets and academics blame breeders for it, but breeders respond to what the public wants, which means there is another, potentially harder truth behind the bulldog's demise: It might have to do with how college sports fans spend their Saturdays.

A bulldog named Sergeant Major Jiggs became the official mascot of the Marines in 1922, chosen for its toughness and virility. Underwood & Underwood/Corbis via Getty Images

FOURTEEN MINUTES BEFORE kickoff, he emerges from the tunnel and moves past the goalpost to roars loud enough to feel in the chest. The weather in Athens this Saturday night is shockingly, comically nasty: The tropical storm that moved over Georgia in the morning never left, and to call the playing surface at Sanford Stadium a field is to indulge in euphemism. The rain is relentless, hitting everyone sideways, pushed along by gusts of wind that stiffen exposed fingers. Still, Bulldog Nation roars, the sound getting louder, because its king has begun his deliberate stroll to his castle.

UGAAAAAAAAAAA!

He is as white as milk and as stocky as a gallon jug. His owner, Charles Seiler, guides Uga into his doghouse on the Georgia sideline and then, on hands and knees, crawls hip-deep into the home, drying the dog with a large white towel. Fans crowd around, cellphones out, crouching and smiling in the furious rain, hoping to get a shot of the most famous bulldog in the country. Uga stares back, mute, an aristocrat bored by the ways of his commoners.

It's a demeanor centuries in the making. While certain enthusiasts obsess over the English bulldog's bloodlines just as certain Europeans obsess over noblemen's, the breed, in fact, owes its existence to the nobility. Bulldogs descend from mastiffs, large and regal dogs with powerful jaws, and the term "bulldog" arose from a long-lived amusement called bull baiting, in which British lords, among others, pitted a dog against a leashed bull and bet on whether the dog could tear off the bull's nostrils before the bull gored or kicked the dog to death. The most successful dogs clenched the bull's nostrils and didn't let go, even as the bull violently shook its head, even as the dog slapped against the bull's shoulders. Accounts vary, but bull baiting was an English pastime for the better part of 600 years. Owners learned to breed a dog particularly skilled for the sport, a lighter and quicker and more compact dog than a mastiff but with the same fearlessness and confidence. The best bull baiters were a breed ultimately known as bulldogs.

The British government banned bull baiting in 1835, and the bulldog, without a purpose, came close to extinction. It was saved by the Industrial Revolution and the insecurities of the nouveau riche capitalists in the United States and England who wanted to imitate the old-line aristocracy by adopting their trappings. These capitalists wanted bulldogs of their own, but they had to breed out the dog's meanness, which meant changing its form. By the 1890s, the bulldog looked more like the boxer -- the same ugly mug but with a strong chest and sleek body.

That kind of bulldog -- athletic, alert -- first roamed the Yale sidelines in 1889, when a student, the son of a New York industrialist, christened him Handsome Dan. Like luxury cars and expensive watches, the bulldog began to spread to other classes. In 1956, Sonny Seiler, a law school student at the University of Georgia, decided after many drinks at the Sigma Chi frat house to take the white English bulldog he and his wife, Cecelia, had received as a wedding present to Georgia's football game against Florida State. The dog sat with Sonny and Cecelia in the stands wearing a red shirt with a block-letter "G" Cecelia had sewn. He was a huge hit, Georgia beat FSU, and soon the Seilers found themselves the owners of Georgia's official mascot, which everybody started to call Uga.

In photos, he resembled the original Handsome Dan -- lithe through the body, strong through the chest -- and appeared at Georgia games for 10 years, until 1966. In successive decades, two things happened: Uga got shorter and fatter, a wrinkled furball with half-opened eyes and a pancaked face; and Uga became famous, a brand unto himself. Uga IV dressed in black tie and accompanied Herschel Walker to New York when the running back won the Heisman Trophy in 1982. Uga V appeared in the movie "Midnight in the Garden of Good and Evil" in 1997. Uga X, and the line of Ugas preceding him, was named by Sports Illustrated in 2019 as the greatest mascot in college football history. These days, Uga has his own hotel suite when he and Charles Seiler, Sonny's son, travel the four hours from Savannah for home games. These days, Uga's red shirt with the block-letter "G" comes courtesy of Nike.

Uga remains proudly purebred, and Charles Seiler takes issue with the idea that inbreeding has destroyed the bulldog. He says he and his family search diligently for healthy purebreds and keep tabs on the lineage of Uga's potential mates. "We make sure there's no inbreeding or line breeding going back eight generations," he says. (The American Kennel Club and the Bulldog Club of America, unlike the British Veterinary Association, have not recommended outcrossing.) The implication that owners would rather have a cute bulldog than a healthy one irritates Seiler. He and his family love Uga too much to see him suffer. His father has talked to the media about the surgeries the family pays for to help Uga breathe better. The family makes sure he exercises and even bathes him before games with two different shampoos, to help his skin.

But consider that pampering in another light: The Seilers have to pay for the surgeries, have to buy multiple shampoos. They even have to put an air conditioner in Uga's doghouse at Sanford Stadium because of the mounting evidence that shows the breed to be in a terrible way. It seems Uga cannot escape the fate of bulldogs everywhere. There have been 10 Ugas in 63 years, but five of those dogs have served as the mascot in just the past 20 years. The reigns are getting shorter. Uga VII made it two seasons, in 2008 and 2009, before dying from heart-related issues. Uga VIII served just six games in 2010 before a diagnosis of lymphoma forced him to retire. He died the next year.

Uga, the University of Georgia mascot, travels in style. He has his own hotel suite during home games as well as a doghouse on the Georgia sideline. Erik Lesser/Getty Images

THE CAMERAMAN SEES him first: in the corner of the Yale end zone, already so tired three minutes into the Harvard-Yale game that he's splayed out, paws before him, his belly flat against the turf, panting. The cameraman walks quickly toward Handsome Dan because an exhausted bulldog is an adorable one. He holds his camera low to the Yale Bowl's FieldTurf, as if hoping for a sweeping and cinematic shot. Some still photographers see the commotion and rush to Handsome Dan as well. His caretaker, Kevin Discepolo, knowing these images won't flatter Handsome Dan, tickles the dog's belly. Handsome Dan begrudgingly sits on his hind legs. The dynamic, and shot, changes. Slightly annoyed, the surrounding media's attention returns, one by one, to the game.

Some iteration of this scene plays out every time a live bulldog appears on the sideline of a college football broadcast: a pan away from the game to the slobbering dog and then back to the action. That simple sequence might explain the bulldog's decline better than anything else.

Satellite and cable broadband packages have skyrocketed over the past 40 years, moving from 16 million American households with cable in 1980 to more than 66 million households with cable broadband today, or 90% of American homes. This means the number of broadcast college football games has skyrocketed too. In Los Angeles, for example, the average number of college football games broadcast on a fall Saturday went from five games in 1979 to 36 games in 2017. The more college football games air, the higher the chance of someone in the U.S. seeing a bulldog, the most popular mascot in Division I sports. And "the more people ... see him on TV," Sonny Seiler once told The New York Times, "the more they want a bulldog."

For the 90 years preceding the 1980s -- from roughly the first Handsome Dan in the 1890s to Uga II in the early 1970s -- the breed looked more or less the same, tall and muscled and alert. After that, something happened. When scholars and vets look at photos of bulldogs from the 1980s until today, they see a breed that has grown ever stouter, ever fatter and ever more wrinkled. For that kind of rapid transformation, the breed would need constant exposure to the public, says Hal Herzog, a professor of psychology at Western Carolina University who has written extensively about why certain breeds of dogs become popular. There are "boom and bust" breeds, he says, dogs that grow in popularity around some cultural event, like when Dalmatian ownership spiked 710% after the 1996 live-action remake of "101 Dalmatians," according to one of Herzog's studies. But the bulldog is something altogether different. Herzog's research shows U.S. bulldog registration remaining flat for decades and then climbing slowly but steadily from 1986 onward, from a little shy of 7,000 dogs registered to a little more than 20,000 by 2005. A rise like that suggests sustained cultural exposure to the bulldog. "It's certainly possible," Herzog says, that college football games on cable provided that exposure and led to the increase in popularity -- and in turn the bulldogs' health problems.

The camera is drawn to what is cute, and what is cute in the bulldog is what gets passed on, says James A. Serpell, professor at the University of Pennsylvania's School of Veterinarian Medicine and a leading scholar in bulldog anthropomorphism. "TV could absolutely be helping to drive this downward trend," he says. College football games on TV gave the bulldog a fame the breed had never known. And when that lasting exposure tied itself to how we are "hardwired to respond to these babylike features," Serpell says, the extreme and comical attributes of one generation of bulldogs became the norm of the next. Which then became the extreme of the third, which then became the norm of the fourth -- and downward the breed spiraled. With the average bulldog living just six to eight years, perhaps as many as eight generations of bulldogs have roamed the sidelines of college football games since cable television became the national norm. Every time you watch a game, "you're indulging yourself at the bulldog's expense," Serpell says.

On game days, fans crowd around Uga hoping for photos with the most famous bulldog in the country. Kevin D. Liles for ESPN

This is what bothers Sandra Sawchuk. She's a veterinarian at the University of Wisconsin's Department of Medical Sciences and unique in the field in that she rails against the plight of bulldogs even as she owns them. "I get them as rejects," she says, from owners or breeders who can't keep caring for them because of the high costs of vet bills. "Of course I love the dogs," she says, but she's fully aware of the research and "totally" thinks college football broadcasts have harmed the breed. The dog's features in these games, she says, are "cartoons" of their former selves. She sees the damage this causes in her clinic. "When I started out in the '80s, you never saw bulldogs as clients," she says. "Now I get four or five a week." After a bulldog won the National Dog Show in November, Sawchuk told a local public radio broadcaster: "I have such misgivings about this because any time a dog becomes that popular, that they're now the national champion, a lot of people want one. And I just want people to know that although I have bulldogs, I'm a veterinarian. ... I have pet insurance for all three of my dogs. Because even as a veterinarian they have so many health issues." She began raising bulldogs 18 years ago because she thought her expertise could lessen their suffering. She now fears she's endorsing it. "This bulldog will be my last," she tells me.

Annette Nobles, the Bulldog Club of America spokeswoman, says views like Sawchuk's are part of the "radical extremists who hide behind social media with their non-confirmed insults." She says "extreme" breeders may have led to some unhealthy lines but that "since 2012, we have more than doubled the number of dogs participating in BCA-recommended [health] testing." The bulldog's win at the National Dog Show proves her point that the breed can be quite healthy. The winning dog, Thor, "is a BCA Platinum Health Award recipient, one of our highest health achievements." Nobles, a bulldog owner herself, says, "My dogs very rarely go to the veterinarian for anything other than yearly checkups. The bulldog is not fundamentally unhealthy."

Pedersen's research seems to rebut that claim. When his study compared the DNA of 102 "typical" bulldogs with the genes of 37 "unhealthy" bulldogs who'd come to UC-Davis' veterinary hospital for various ailments, it found no genetic difference between the two groups. To say the bulldog is healthy is to be in denial, Pedersen says. Lisa Pritchard, the veterinary technician at Mississippi State who owns Bully, says she reckons with the breed's popularity and problems every Saturday. When she and Bully roam the tailgates in Starkville, "I'm probably asked 20 times about getting a bulldog," she says. It places her in an uncomfortable position. She feels compelled to be honest with MSU fans about the bulldog's health risks, about its vet bills. In other words, the steward of nothing less than Mississippi State's identity must caution fans against owning a piece of it for themselves. "It hurts me," she says. The bulldog is a "great pet" and Pritchard loves Bully like a son, but "you can see a huge change in the way they look and the way they've evolved or devolved depending on your perspective."

Yale's current Handsome Dan, its 18th, is an outcrossed version of the traditional bulldog, bred in the hopes of restoring the dog's health and athleticism. AP Photo/Charles Krupa

WHICH BRINGS US back to Handsome Dan, sitting on his hind legs at the Harvard-Yale game. He looks slightly different from other modern-day bulldogs. His face isn't as wide, his jaw and snout more pronounced, his build sleeker, his gait faster. He is not modeled after purebreds like Uga or Bully. Kevin Discepolo, Handsome Dan's owner, oversees facilities within Yale's athletic department, and he inherited the responsibility in 2016 of caring for a new Handsome Dan. As Discepolo researched breeders, he also researched the breed and began to read a lot of articles discussing the bulldog's poor health. Discepolo shared his findings, and soon throughout Yale, "there was this push for a healthier version of the breed," he says. Discepolo discovered an outcrossed iteration of the traditional bulldog called the Olde English Bulldogge, which, much as its name implies, seeks to return the dog to its former athleticism and vigor, but without its bared-teeth aggression. Discepolo found an Olde English Bulldogge breeder in Maine, and the new Handsome Dan, the school's 18th, made his debut in 2017.

He was panting just a minute ago, paws out before him on the Yale sideline, because he'd spent half an hour before the game playing catch. "Everyone who comes in contact with him says he's the most athletic bulldog we've had," Discepolo says. Yesterday afternoon, he and Discepolo walked all over campus and then through the library, where Handsome Dan jumped onto a study table and licked the cheeks of students who posed for selfies. The night before, he and Discepolo went to a black-tie dinner honoring a few alumni. During the cocktail hour, the former attorney general of Maryland wrestled with Handsome Dan on the floor. Moments before dinner began, former U.S. Secretary of State John Kerry lumbered over. "Handsome Dan!" Kerry said, bending down to rub the dog's chin. "I've never met this one." He looked up to Discepolo. "Is he No. 18?" Discepolo nodded, and Kerry petted him until his name was called by the emcee.

No one knows or, rather, cares that Handsome Dan is a slight genetic break from the bulldog or that the American Kennel Club doesn't recognize Olde English Bulldogges as an official breed. Handsome Dan and Discepolo spend the first half of today's game prowling the sidelines and the Yale Bowl bleachers. Everywhere they go, they're stopped. Everyone wants pictures. Everyone wants to pet Handsome Dan. By halftime the dog needs to poop and release some pent-up energy. Discepolo takes him to Yale's soccer field, where Handsome Dan does his business on some fallen red leaves. Discepolo finds a lacrosse ball in a storage shed and tosses it deep onto the pitch. Handsome Dan takes off at a sprint, with the speed and eagerness of a Labrador, and chomps down on the bouncing ball. He races back to Discepolo.

"Good boy!" Discepolo says. Handsome Dan drops the ball for him to pick up and throw again, and when Discepolo doesn't do it fast enough, Handsome Dan jumps up and onto Discepolo, too excited to wait.

"Siiit," Discepolo says, then scoops up the ball with his stick and rolls it deep. Handsome Dan takes a wide but furious path toward the ball and Discepolo says with a half laugh, "Uga doesn't move like that, right?" Probably not: In a "CBS Sunday Morning" segment on Uga, Charles Seiler picked up the dog to move him down a flight of stairs.

As the second half approaches, Discepolo and Handsome Dan walk back to the stadium and come into view of a statue of the original Handsome Dan. Discepolo sees this statue every game day and is no longer struck by it, but it bears the same vigilant facial features as today's Handsome Dan, the same powerful body.

ESPN is broadcasting today's game, which means the second half, like the first, will feature a few cuts away from the action and to this new Handsome Dan -- which looks like the first Handsome Dan, which, "who knows," Discepolo says, might just be the Handsome Dan that Americans want their bulldogs to look like from now on.

It's happened once already.