Cats have 276 facial expressions. Why?
Famously aloof felines may be more expressive than we thought. A new study looks at how cats interact with each other, and with us.
JOEL SARTORE
We have this very deep history with cats, and for that reason there is “growing interest” in how to decipher our cats’ emotions.
Most previous research on facial signaling in cats has focused on interactions between cats and humans or assessing when a cat is in pain. The new study takes that a step further by analyzing how cats interact with one another.
And what better place to do that, than a cat café? After 150 hours at the CatCafe Lounge in Los Angeles, California, a nonprofit shelter where visitors can meet up to 30 cats for potential adoption. (What do cats think about us? You may be surprised.)
Their observations of 53 domestic shorthair cats revealed a wide variety of expressions—which combine various movements of the eyes, ears, and lips—and that most of these were friendly, not aggressive. Many cat people would instantly recognize some of these as friendly, such as ears and whiskers forward and eyes closed.
But some, like what the researchers call “play face,” with ears and whiskers forward and mouth pulled back at the corners, don’t have such obvious meanings.
What’s more, a lip lick might suggest a cat is anticipating a treat, but when combined with narrowed pupils and flattened ears, it’s an unfriendly cue.
Whiskers overall are surprisingly telling: Content or happy cats almost always point their whiskers forward.
Is that a fake smile?
After hours at the cat café, Scott recorded 194 minutes of cat interactions, excluding non-communicative behaviors such as yawning or chewing.
Back at the lab, the team used a coding system for facial expressions called Facial Action Coding System, or CatFACS. Trained users can identify “even the most subtle of muscle movements, and each of these movements, called an “action unit,” is noted and given a number using a video software, ELAN. This allows users to create and edit a video-footage timeline, which can read expressions at the millisecond level.
The analysis revealed 26 unique muscle movements that, in various combinations, created 276 facial expressions. The results categorized 46 percent of expressions as friendly, while 37 were unfriendly, and 17 fell into both categories.
Examining the facial expressions at such a granular level can catch nuances. A good example is the real vs. fake smile in humans: Though in both situations you’re smiling, there may be small differences, say in eye movements, that can indicate whether you’re actually happy. (Read surprising things you never knew about your cat.)
For some feline expressions, though, the meanings are still a mystery, and the team is planning follow-up studies to figure them out.
A tool for improving cat welfare
It will be really interesting to see what groupings are more common in different situations and contexts.
Cats don’t need eye contact to make their intentions known. Body posture and other behaviors, such as guarding a food bowl, sends a signal that “this is mine,” without needing to make any facial expressions.
Research could inspire scientists to observe facial expressions in wild cat species, which are more solitary. Knowing whether wild cats share specific meaningful expressions would discern whether domestication led to a greater variety of expressions in domestic cats. (Learn why cats domesticated themselves.)
The research also has a practical application, says Bennett, who consults with shelter organizations. For instance, the study could help develop a standardized tool of facial expressions, allowing animal shelter staff and volunteers to identify certain cues and act accordingly.
In one scenario, if two cats are brought into the shelter together, they may be considered to be bonded. But recognizing differences in their facial expressions could suggest otherwise, and point the staff toward finding a more suitable feline friend.
“I’m excited about this, They definitely think they’re on the right track.”
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How Siamese cats change their colours
A genetic mutation known as the Himalayan gene has made this breed’s fur extra sensitive to temperature—but also leaves them vulnerable to certain health problems.
PHOTOGRAPH BY WILLARD CULVER, NAT GEO IMAGE COLLECTION
Most of us, though, can pick out a Siamese cat. Known for their dark eye masks, socks, and tail—called “points”—that stand out against their cream-colored bodies and sparkling blue eyes, the Siamese is one of the world’s easily recognized breeds.
Those trademark points stem from a genetic mutation known as the Himalayan gene, which was discovered in the breed in 2005. This recessive gene, passed down from both parents, also gives the felines temperature-sensitive fur that can change color.
In the womb, Siamese kittens develop at temperatures of about 101 Fahrenheit, a cat’s normal temperature. Once born, all-white kittens eventually begin to cool at their extremities, such as their tail, legs, ears ,and face. These lower temperatures cause the Himalayan gene to activate the body’s melanin, a pigment also found in people. That’s why a Siamese cat’s warmer bodies stay light, but their cooler parts become darker as they mature. (Read how tabbies get their stripes.)
You can see them change. At about two weeks old, dark hues begin to spread to their limbs. By about a month old, their final color emerges, which may be one of several varieties, among them blue point, lilac point, chocolate point, and seal point, which has a light, cream-colored coat with dark brown hues on its paws, tail, nose, and ears.
A lifelong influence
The Himalayan mutation naturally occurs in South Asian domestic cats, says Leslie Lyons, a feline geneticist at the University of Missouri who first identified the mutation in Siamese cats.
Over time, as people selectively bred cats with the striking pale color and dark points, they passed the gene to different breeds, such as Siamese, Birmans, Ragdolls, and Burmese. Mixed breeds can also carry the genetic mutation.
These cats’ vibrant blue eyes are also due to the Himalayan gene, which influences their eye pigment. (See vintage pictures of pampered pets.)
The Himalayan mutation causes fur to darken as the cat ages, though any sort of trauma or change to the cat’s fur can alter its hues.
If a Siamese cat has surgery, for example, an area of fur that was shaved may grow back darker because the impacted area is cooler while the hair grows back. Eventually, the fur may become lighter again, Arnold says.
In the 1920s, a Siamese cat living in Moscow that wore a jacket over a shaved shoulder had its fur grow back all white, since that area was kept warmer. The fur later turned darker again, according to the University of Alaska.
Health concerns
Many other species have the Himalayan gene, including Himalayan rabbits, says Lyons.
Domestic rats, color-pointed gerbils, and American minks can also carry the mutation. In 2021 the journal Gene reported a dachshund with a different mutation on the same Himalayan gene that produced Siamese-type coloration, a rare occurrence in dogs.
Each species has its own mutation in the same gene, but then have been bred to have the specific coloration. (Read surprising things you never knew about your cat.)
While there’s no overarching advantage or disadvantage to the mutation in these domestic animals, it does come with some health concerns, too, including eye issues.
Siamese often have crossed or misaligned eyes, also called strabismus, which can compromise vision as well as depth perception.
Even if the cat’s eyes are aligned, the animal may still suffer from nystagmus, or shaking eyes, a condition in which the eye sometimes dart slightly from side to side in repetitive, uncontrolled movements that can cause vision or even balance problems, says Arnold. She adds she hasn’t seen the condition in her practice.
Both Lyons and Arnold recommend pet owners get their Siamese pets from reputable breeders, or adopt them from a rescue. Wherever they’re from, though, both experts agree these masked felines will steal anyone’s heart.
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Why Street Cats Are Taking Over Urban Neighborhoods
Jane and her husband loved the house they bought in central Los Angeles in 2022, until they noticed the cats. Lots and lots of cats. At any given time, more than a dozen would be lounging in their yard, peeing on their packages, leaving dead birds in the grass and fleas on their outdoor furniture. “The amount of cat feces we’re exposed to is truly insane,” says Jane, who asked that her real name or the name of her neighborhood not be used because of the ensuing battle over feeding the cats.
A family in the neighborhood had left their pet cat behind when they moved, neighbors told her. That cat had multiplied into two, then four, then more. Now, a cat colony rules the neighborhood. Jane tried planting lavender bushes, which are supposed to repel cats, only to find the cats sleeping under the plants. She tried adding sprinklers to scare them away, but they nimbly dodged the water. She tried to capture a few to bring them to a vet to be spayed and neutered, but inevitably ended up capturing the few cats that had already been fixed.
When she asked neighbors who left food out for the cats to stop, tensions rose. Though many people in the community complain about the cat problem, some are still leaving out cat food for the animals. Jane learned the situation wasn’t unusual. In cities like Los Angeles and New York, the population of street cats is booming. “It’s something people have come to expect in L.A.,” she says. “Cats are in everybody’s yards.”
Los Angeles is home to an estimated 1 to 3 million “community cats” that don’t have owners, according to the city. Some animal advocates believe the population is nearing 4 million, which is about how many people live in the city of Los Angeles. New York City’s community cat population could be as high as one million, says Will Zweigart, the executive director of Flatbush Cats, a nonprofit formed to reduce the outdoor cat population in BrooklynThese aren’t just feral cats that have never been exposed to humans, says Zweigart. Many of these cats are “friendly cats,” who once lived in a home before being put out on the streets. A few years ago, around 80% of the cats Zweigart’s organization encountered were feral. Now around 50% are friendly. “We’re seeing more people unable to care for their pets,” he says.
It’s not just cats. As the costs of pet ownership increases because of inflation and veterinary care gets more expensive, people are surrendering animals to shelters or leaving them on the street. Around 32% of all pets that ended up in shelters in 2024 were owner surrenders, up from 30.5% in 2019, according to Shelter Animals Count, a national database for animal rescues and shelters. People are also surrendering their pets because they are downsizing to new housing that doesn’t allow pets or can’t afford basic supplies like pet food.
“With inflation and the downturn in the economy, people are having to make very tough choices about whether to feed their families or feed their pets,” says Lauree Simmons, the founder and CEO at Big Dog Ranch Rescue, the largest no-kill dog rescue in the country. Owner surrenders are at an all-time high at her organization, which is based in Florida. The organization screens people who want to surrender their dogs, and the number one reason is that owners can’t afford them, Simmons says. The second most common reason is that they can’t find affordable housing that allows dogs, and the third is that they can’t afford veterinary care.
As would-be pet owners look at the costs and responsibilities of pet ownership, adoptions have fallen. In the first half of 2024, there were 82,000 fewer dog and cat adoptions than in 2023, a 4% drop from 2023, according to Shelter Animals Count. Nearly 700,000 dogs and cats were euthanized last year at shelters, according to the organization.
“This is the worst it’s been in the decade I’ve been in animal welfare,” says Katy Hansen, director of marketing and communications for Animal Care Centers of NYC, the city’s largest shelter. ACC is an open door shelter, which means it accepts any animal that people bring in, and Hansen says many people dropping off their pets say they just can’t afford to have animals anymore. Others say they’re moving for financial reasons and aren’t allowed to have a pet in their new place, or that the cost of a pet deposit is prohibitive. The shelter is at double capacity. “It’s a combination of people surrendering their pets and people not adopting because they’re not sure they can take on the financial commitment,” she says.
Historically, veterinarians in America focused on horses and farm animals; only in the last few decades has there been more of a focus on pets like cats and dogs. As pet owners became more willing to spend money on their animals, companies obliged, opening fancy veterinary hospitals and services like oncology. The private equity industry has taken notice, seeing how much money some pet owners are willing to fork over. As big corporations took over veterinary practices, prices rose. The cost of veterinary care is up 38% since 2019, according to the Bureau of Labor Statistics.
At Veterinary Care Group, a private equity-owned practice in Brooklyn, the cost of spaying or neutering a cat has soared to $850 per animal. By contrast, at the nonprofit veterinary clinic Zweigart recently founded in Brooklyn, the cost of spaying or neutering a cat is $225 and a mid-sized dog is $300. “Pet ownership,” says Zweigart, is now “out of reach for the average American.”.
Porsche Jones, who lives in Flatbush and takes animals to Zweigart’s clinic, says she tries to help neighbors who feel forced to abandon their pets for financial reasons. She encourages them to spay and neuter their pets, but the cost makes some owners balk, she says. “Two people on my block put their cats outside because they couldn’t afford to spay and neuter them,” she says. “Even $80 is a lot for people who don’t have the money for their electric bill.” Some pet owners are turning to sites like Waggle, which helps people raise money for veterinary care, but not everyone is successful at raising funds.“We’ve really skyrocketed in listings this year,” says Steven Mornelli, the founder of Waggle.
Meanwhile, some overwhelmed cities are turning animals back to the streets. The San Diego Humane Society started a program in 2021 to sterilize and vaccinate cats brought to the shelter, and then return them back to the streets if staff determine they can successfully live outdoors. The practice is currently being challenged in a lawsuit filed in San Diego Superior Court by the Pet Assistance Foundation, which argues the shelter should make more of an effort to find owners for cats that it would otherwise release into the wild.
Jane, the Los Angeles resident, is sick of waiting for the city to do anything about the cats that roam her yard. The city expects owners of private property to trap the cats themselves, she says, and then pay for them to be spayed and neutered. The hope is that if everyone does this, the cat population will start to shrink. Jane says that catching the cats is next to impossible, before even getting to the question of sterilizing costs.
She and her husband are now looking to move. She’s been subtly trying to discern if the new homes they’re looking to rent have cat problems by telling the real estate agent she loves cats and wonders if there are any roaming around. In some of the homes they’ve looked at, the answer has been yes.