In a project that has all the makings of a Roald Dahl classic, scientists have hit on

an answer to the mystery of how man’s best friend got its puppy dog eyes.

The sad, imploring expression held such power over humans during 33,000 years
of canine domestication that the preference for dogs that could pull off the look
steered the evolution of their facial muscles, researchers have said.

The result is that dogs gradually acquired a new forehead muscle named the
levator anguli oculi medialis, or LAOM, and have used it to deploy the doleful
look to devastating effect ever since.

“They are very powerful animals in how they capture our hearts,” said Prof
Bridget Waller, the director of the Centre for Comparative and Evolutionary
Psychology at the University of Portsmouth. “We pay a lot of attention to faces,
they are meaningful to us, and this expression makes dogs look juvenile and sad.
It induces a nurturing response. It’s a cute factor.”

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Puppy dog eyes are achieved by the LAOM raising the inner eyebrows, in some
cases quite dramatically. The movement makes the eyes look larger and the face
more babyish. Humans use different muscles to produce a similar expression
when they are sad, which may explain why it brings out the caregiver in people.

To investigate how the look developed in dogs, the UK-US research team
acquired wolf and dog cadavers from taxidermists and US state organisations and
dissected their heads to compare the facial muscles. No animals were killed for
the research.

Dissections of six dogs – a chihuahua, a labrador, a bloodhound, a German

shepherd, a Siberian husky and a mongrel – found all had the LAOM muscle. But
in the four grey wolves studied, the muscle was missing, save for a few scant
muscle fibres. Since all dogs are derived from wolves, the comparison suggests
the LAOM arose in the domestication process.

Only one other difference was noted in the head dissections. A muscle called the
retractor anguli oculi lateralis (RAOL), which pulls the eyelids out towards the
ears, was less prominent in wolves than dogs. The Siberian husky, one of the
most ancient breeds, was the only dog found to lack the RAOL muscle, according
to the report in the journal Proceedings of the National Academy of Sciences.

After establishing that dogs and wolves have different muscles around the eyes,
the researchers filmed the animals to see how their expressions varied. They
filmed nine wolves in two different animal parks, and 27 dogs, mostly
Staffordshire bull terriers, in shelters across the UK. The footage was reviewed by
a trained specialist who was not told about the scientists’ hypothesis. The
specialist recorded when the animals made the puppy dog eyes expression, and
rated its intensity on a five-point scale.

Dogs pulled the doleful face far more frequently than wolves, but the most
striking finding was the intensity of the expressions. While dogs and wolves both
produced “low intensity” expressions, only dogs appear to have weaponised the
look and achieved what the scientists classified as “high intensity expressions”.

The look has a real impact. In a previous study, Waller showed that the more
dogs deployed the expression, the faster they were rehomed from shelters. In that
regard, puppy dog eyes were more effective than tail wagging or the speed at
which dogs bounded over to visiting humans.

Waller does not believe dogs originally produced the expression to win humans
over. More likely, she said, is that animals that happened to deploy puppy dog
eyes tapped into a response humans had evolved over millennia of living in large
groups, where reading facial expressions was crucial.

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