Tabby Cats, UnfoldedThe Science of a Striped Coat

Tabby cats wear a dark letter M on their foreheads, drawn right in their fur, like a name tag they never take off.³

The cat registries agree about the letter more firmly than about almost anything else. An article on tabby patterns by a judge emeritus of the Cat Fanciers' Association calls the M "the characteristic marking on the forehead of every tabby-patterned cat." The association's pages for young readers put it more plainly: all tabby cats, whatever their pattern, have an M on the forehead. Alley Cat Allies, a cat-welfare group, calls the M the hallmark of the tabby. Mackerel, classic, spotted, ticked or rosetted, the body may carry stripes, swirls, spots or almost nothing, and the letter stays.⁴

Because the M belongs to the pattern, it marks a tabby, not a breed. "Tabby is a pattern, not a color, and not a breed," the association tells its young readers. Some pedigreed breeds are defined by a tabby pattern, the Egyptian Mau and the Ocicat by spots and the Abyssinian and Singapura by ticking, and others such as the Maine Coon and the Bengal allow or require tabby markings. Tabby patterns are also everywhere among ordinary feral and non-pedigree cats, and that is where geneticists collected many of the tabbies whose DNA appears later in this book.⁵

That fact shaped the science in the chapters that follow. The Taqpep work drew mostly on feral cats from Northern California, and in the study of ticked coats the clearest result came from non-breed cats rather than from show animals. The genetics of the striped coat was worked out as much on alley cats as on pedigreed ones, which suits a pattern that belongs to no breed in particular.⁶

A striped cat with no recorded ancestry is not a lesser tabby. Cornell University's Feline Health Center explains that mixed-breed cats are generally grouped as domestic shorthairs or domestic longhairs, so a mixed-breed cat with short fur is usually called a domestic shorthair. That name describes a coat length and a mixed family tree; like tabby, it is not a breed. A domestic shorthair with a forehead M is a tabby in exactly the same sense as a show cat with one.⁷

A word borrowed from silk

The word tabby did not begin with cats. Dictionaries trace it through the French tabis and the Arabic 'attābī to al-'Attābīya, a quarter of Baghdad where a particular fabric was first made. In English the word first named a watered silk, a cloth finished so that its surface ripples with wavy, striped light. Only later was it borrowed for cats whose coats show the same kind of striping, so a tabby cat is named, in effect, after a bolt of cloth.⁸

Dictionaries also record an informal sense of tabby meaning a domestic cat, especially a female, and that usage has led many people to assume that a tabby is a she-cat. Smithsonian Magazine noted that tabby may be a colloquial term for a female cat but is more properly the name for the common striped pattern on a domestic cat's coat. In cat genetics and registry use, tabby names the pattern, whether striped, swirled, spotted or ticked, and nothing else.⁹

What the letter is drawn in

Look closely enough and the M turns out to be drawn with two kinds of hair. Geneticists describe tabby markings as "a composite of two features": a light background in which individual hairs carry extensive light bands, and a superimposed darker component in which hairs have little or no banding. The registry article describes the light part as agouti hair, each hair carrying alternating bands of yellow and black along its length.¹⁰

Seen that way, the M is not ink laid on a plain cat. It is a place where the banding falls away and the hairs run dark, set against a background of banded hairs that, on a brown tabby, read from a distance as a speckled brown. The stripes, swirls and spots of every other tabby are made the same way. Each dark mark is a patch of solid-colored hairs replacing the banded ones, so a tabby's pattern is really a map of which hairs band and which do not.¹¹

The best evidence that the letter belongs to the pattern comes from cats whose bodies carry almost no pattern at all. In a ticked tabby every body hair is banded, so the body looks evenly flecked instead of striped. Yet the registry article notes that the face, legs and tail generally still show distinct tabby markings, the forehead M among them, along with mascara lines beside the eyes, bracelets on the legs and rings on the tail. The Abyssinian and the Singapura are ticked breeds, and the M on their faces is the same letter a striped alley cat wears.¹²

The M, then, is the single mark every tabby shares, from the patterned show cat to the stray on the fence. Below the forehead the patterns go their separate ways, and the oldest of them, the one every other tabby coat has to be measured against, is the narrow stripe.

A mackerel tabby has thin stripes down its sides, rings around its legs and tail, and striped necklaces across its chest, as if the cat were dressed in a striped sweater with bracelets to match.¹³

The registry description reads like a tailor's notes. Most of the markings are "dense, narrow pencilings," running from the spine down toward the belly. "The legs are barred with bracelets coming up to meet the body markings, and the tail is barred with tail rings. The necklaces on the neck and chest are distinct." Pencils, bracelets, rings and necklaces: the vocabulary the registry reaches for is that of a striped outfit and its jewelry.¹⁴

Geneticists describe the same coat in a single careful phrase. In mackerel cats, Kaelin and colleagues wrote in 2012, the dark component is organized "into narrow vertical stripes with a constant and regular spacing." The important word is regular. Stripes that are evenly spaced down the whole length of an animal need something to space them, and finding that something is the business of the next three chapters. Real cats vary, of course, and some mackerel tabbies wear crisp pencil lines while others wear softer, broken ones.¹⁵

The oldest coat

The mackerel stripe was not bred into house cats as a decoration. It is the coat they started with. The Old World wildcats from which domestic cats arose show a mackerel-like pattern, and in their 2021 paper Kaelin, McGowan and Barsh state that "the mackerel stripe pattern represents the ancestral state." When a mackerel tabby stretches on a windowsill, it is showing off the oldest tabby coat there is.¹⁶

Which wildcat, and where, was the question behind a 2007 study in Science by Driscoll and colleagues, which compared the genetics of domestic cats with those of wildcats. It concluded that cats were domesticated in the Near East, "probably coincident with agricultural village development in the Fertile Crescent." The ancestor is the Near Eastern wildcat, Felis silvestris lybica, also called the African wildcat, and house cats, striped or not, descend from it.¹⁷

Ancient DNA later filled in the travels. A 2017 study in Nature Ecology & Evolution by Ottoni and colleagues read genetic material from the remains of cats across the ancient world. It found that both the Near Eastern and the Egyptian populations of Felis silvestris lybica contributed to the domestic cat's gene pool, at different historical times. The house cat's family tree, in other words, has two wild roots in the same species rather than one.¹⁸

Neither of those studies was about stripes; what they supply is the family tree on which the stripe hangs. Put them together with the coat descriptions and the line runs straight: a striped wildcat in the Near East, cats probably living alongside early farming villages, and a mackerel tabby on a modern windowsill wearing much the same coat.¹⁹

Wildcat or house cat?

The coat has changed so little that it causes practical trouble. The 2007 Science paper notes that "a feral domestic cat with a 'wild-type' mackerel tabby pattern is difficult to distinguish visually from a 'true' wildcat," and that ongoing admixture, interbreeding between the two, makes the problem worse. A striped feral cat can pass for a wildcat by eye, which is one reason researchers turn to DNA to tell them apart.²⁰

All of this makes the name of the swirled pattern misleading. Blotched tabbies are often called classic tabbies, but Smithsonian Magazine pointed out that the name has nothing to do with being older or more common: the blotched pattern is the more recent mutation. Alley Cat Allies turns the point around and calls the mackerel tabby the true classic. The next chapter finds that mutation, and then asks when the swirls it made began to spread.²¹

Change just one gene, called Taqpep, and a tabby cat's thin stripes turn into big swirls, with a round blotch circled by rings on each side, like a bullseye. That is a classic tabby.²²

The registry description is precise. A classic, or blotched, tabby carries broad, swirling markings, with a butterfly shape across the shoulders, and "a large, solid blotch on each side of the body that is encircled by one or more unbroken rings." The association's page for children says simply that the cat will have swirls and a circle on its side. Geneticists, less poetic, describe the dark component of a blotched coat as "expanded into a less organized structure with wide whorls."²³

Finding Taqpep

In 2012 Kaelin and colleagues reported in Science that they had identified "the gene responsible for tabby pattern variation in domestic cats" as Transmembrane aminopeptidase Q, which they shortened to Taqpep. In their 2021 follow-up the same group summed up what it does. Loss-of-function mutations in Taqpep, changes that leave the gene unable to do its usual job, cause the ancestral pattern of narrow dark stripes to expand into large, less organized whorls.²⁴

That makes the classic tabby, strictly speaking, a mackerel tabby whose stripe-keeping gene has stopped working. A cat with a working copy of Taqpep keeps its stripes narrow and evenly spaced. When the gene is knocked out, the narrow stripes spread and merge into the swirls and the ringed blotch. One broken part changes the look of the whole coat, while the letter M on the forehead stays where it was.²⁵

A gene is a set of instructions for building a protein, and a loss-of-function change is one that leaves that protein missing or unable to do its work. The name Transmembrane aminopeptidase Q describes the kind of protein Taqpep makes, but the visible result of losing it is easier to describe than its chemistry: the same dark and light hairs as before, arranged in broad whorls instead of narrow, regularly spaced bands.²⁶

Dominant stripes, hidden swirls

Every cat carries two copies of Taqpep, one from each parent, and the two versions do not count equally. Mackerel is dominant to blotched, so a cat needs only one mackerel copy to wear mackerel stripes. A blotched tabby has two blotched copies. The registry article reaches the same conclusion from breeding experience, calling the mackerel pattern "the genetically dominant pattern."²⁷

The 2012 study checked that rule against real cats. Its tabbies came mostly from feral cats in Northern California, along with shelter cats. In the study, every blotched tabby carried a changed Taqpep gene on both of its copies, while every mackerel tabby had at least one unchanged copy. That is exactly the pattern expected when mackerel is dominant and blotched is recessive, found in cats that had never been bred for their coats at all.²⁸

Dominance explains a small family surprise, which is easiest to see with an imaginary pair of mackerel tabbies that each carry one hidden blotched copy. Each parent passes on one of its two copies at random, so a kitten can inherit a blotched copy from both, and that kitten will grow up swirled. Two striped parents can therefore raise a classic tabby, while two blotched parents, with no mackerel copy between them to pass on, would not be expected to raise a striped one.²⁹

Such hidden copies are far from unusual. Kaelin, McGowan and Barsh note that blotched versions of the gene are common in most feral cat populations, even though mackerel is the ancestral pattern. Because blotched is recessive, many cats that look striped carry a blotched copy they never show, so the swirl travels quietly through the striped cats of the world and surfaces whenever two copies meet.³⁰

Taqpep explains one difference, mackerel against blotched, and only that one. Other genes shape other tabby patterns. The spotted tabby is a case in point: its body stripes are broken into spots, yet the genetics that separates a spotted tabby from a mackerel one is not yet known. A later chapter meets one of those other genes, the one behind the ticked coat.³²

When the swirls arrived

If mackerel is the old coat, when did the swirls appear? The 2017 ancient-DNA study of cats across the ancient world found that the blotched coat variant reached high frequency only after the Middle Ages. Its authors read that late rise as a sign that directed breeding of cats began later than it did with most other domesticated animals. Reporting on the study, ABC News wrote that the researchers placed the emergence of the blotched mutation in the Ottoman Empire.³³

Pictures from the period tell the same story as the bones. Kaelin and colleagues note that depictions of tabby markings from the Middle Ages are mostly mackerel, so the cats that medieval artists drew were, for the most part, striped ones.³⁴

Once the swirls appeared, they spread fast. By 1758 the blotched pattern had become common enough for the naturalist Carl Linnaeus to describe it as characteristic of the domestic cat, a date that comes before the formation of most modern breeds. ABC News reported that the pattern became more frequent in southwest Asia, Africa and Europe and was quite common by the 18th century. The newcomer had become the pattern a naturalist reached for to describe the domestic cat.³⁵

Set side by side, the dates describe a strikingly quick change. Medieval artists drew mostly striped cats; the ancient DNA shows the blotched variant rising to high frequency only after the Middle Ages; and by 1758 a naturalist could treat the blotched coat as the characteristic look of the domestic cat. For most of their long history alongside people, house cats wore the wildcat's stripes, and the swirls on so many sofas today are a comparatively recent arrival.³⁶

The very gene that makes a classic tabby's swirls also gives the rare king cheetah its coat, with spots that join into stripes and blotches.³⁷

The 2012 paper that found Taqpep in tabby cats did not stop at house cats. Its authors also reported that they had identified Taqpep as the cause of the rare king cheetah phenotype, "in which spots coalesce into blotches and stripes." On a king cheetah the usual scattering of round spots runs together into larger blotches, and dark stripes run along the back.³⁸

The coat is striking enough that it once fooled biologists. Cheetahs with the king pattern were originally described as a distinct species. They were later recognized as ordinary cheetahs with a monogenic trait, one controlled by a single gene, with an autosomal recessive mode of inheritance. Autosomal means the gene sits on one of the chromosomes that do not determine sex, and recessive means the coat appears only in an animal carrying two copies of the changed gene.³⁹

That inheritance is the same shape as the blotched tabby's. A cheetah with one changed copy looks like any other cheetah and can pass the copy on unseen, and only when two carriers produce a cub with two changed copies does the king coat appear. A king cheetah is therefore neither a separate species nor a hybrid but a cheetah with two copies of one changed gene. Wild king cheetahs have been sighted only in a small area of sub-Saharan Africa.⁴⁰

Smithsonian Magazine summed up the parallel in one sentence: a normal Taqpep gene produced the regular spotted cheetahs, but a mutated Taqpep merged the spots into stripes, "just as the gene had merged the tabby stripes into blotches." Stanford's news office noted the different treatment the two animals received. The rare cheetahs became known as king cheetahs, while the affected tabby cats were given the less regal name blotched.⁴¹

Taken together, the two animals make a single point. In a cheetah and in a house cat, a working Taqpep keeps dark markings small and orderly, whether those markings are spots or narrow stripes. Without it, in both, the dark areas spread and run together. One gene, two very different cats and the same kind of change: that is why a rare cheetah belongs in a book about the tabby on the sofa.⁴²

From map to pigment

Knowing which gene changes the pattern is not the same as knowing how a hair turns dark. The 2012 study followed that question into the hair follicle. Pigment cells in each follicle make either yellow or black pigment, and which one they make depends on a signal called Endothelin3, or Edn3. In the paper's words, yellow and black pigment are made in hair follicles "that produce low and high levels of Edn3, respectively."⁴³

The signal comes from the dermal papilla, a small cluster of cells at the base of each hair follicle. In newborn tabby skin, the researchers measured Edn3 activity at about twice the level in dark areas as in light ones. They then tested the signal directly in mice, where adding extra Edn3 converted yellow hair to dark brown. Smithsonian Magazine's summary is hard to improve on: lots of Edn3 results in darker hair.⁴⁴

Edn3 explains how a follicle colors its hair, but not where the dark follicles sit. That job falls earlier. Taqpep is thought to help set up the pre-pattern that decides where Edn3 will be high, so that one gene lays out the map and another reads it, follicle by follicle. The evidence for that division of labor came mostly from tabby cats, with supporting experiments in cheetahs and mice, and the next chapter goes back to the moment the map is drawn.⁴⁵

A tabby kitten's stripe pattern is laid out in its skin before it is born, even before a single hair has grown.⁴⁷

The evidence comes from a 2021 study in Nature Communications by Kaelin, McGowan and Barsh, the same group that had found Taqpep. They looked at the skin of cat embryos, measuring which genes were active in single cells. In the study, they found "stripe-like alterations in epidermal thickness preceded by a gene expression pre-pattern." The epidermis is the outer layer of the skin, and in these embryos it was thicker in some bands and thinner in others.⁴⁸

The timing is what makes the finding remarkable. The stripes in the skin appeared before melanocytes, the pigment-making cells, had entered the epidermis, and before hair follicles had developed. Barsh called this step establishment and told Stanford's news office that it "happens long before color appears and long before hair follicles are mature." The pattern exists as a plan before there is anything to color.⁴⁹

The plan is easy to read once you know the key. As the Stanford report put it, the thick area marks the patches of fur that will later be darker, and the thin area marks the patches that will be lighter. Long before birth, the kitten's skin is already divided into future stripes and future background, like a drawing on a balloon that has not yet been blown up.⁵⁰

The study also checked that the embryonic pattern matched the real coats of real cats. Embryos with mackerel and with blotched Taqpep genotypes showed thick-skin layouts that, in the study, resembled the adult patterns of cats with those genotypes. The difference between stripes and swirls, in other words, is already visible in the skin of an embryo.⁵¹

One gene stood out in that early map. Dickkopf 4, or Dkk4, makes a secreted signal that blocks the Wnt family of signals, and in the study the researchers found that it plays a central role in laying out the pre-pattern. The Stanford report describes Dkk4 as involved in marking the areas that will eventually have darkly pigmented hairs. Its full story belongs to the ticked tabby, two chapters on.⁵²

Three stages of a pattern

From these results Kaelin, McGowan and Barsh built a working model, proposing that periodic color patterns in mammals arise in three stages. The first is establishment, in which each element of the pattern acquires its identity during fetal development. The second is implementation, in which signaling molecules made within individual hair follicles give the pattern its shape. The third is maintenance, in which each element keeps its identity through hair cycles and the animal's growth. They offer it as a working model, not a complete account.⁵³

The earlier chapters fit into that frame. The thickened bands of embryonic skin, and the Dkk4 and Taqpep genes that help shape them, belong to establishment. Edn3, made in the dermal papilla of each follicle and turning hair dark where it is high, is the clearest example of implementation. The third stage, maintenance, is the one any cat owner can watch, because it is what happens as a kitten grows.⁵⁴

A working model also shows where the gaps are. It names the stages without claiming that every gene acting in them has been found, and some patterns remain unexplained: how a spotted tabby's coat differs genetically from a mackerel one is not yet known. The frame is useful precisely because it tells researchers which stage a new result belongs to, and which stages still need explaining.⁵⁵

Bigger, not more

A tabby kitten does not add stripes as it grows. The 2012 paper notes that "tabby markings change in size but not number during organismal growth," and the 2021 paper adds that individual spots or stripes of hair apparent at birth "enlarge proportionally during postnatal growth." The balloon from earlier in this chapter is now being blown up: the drawing on it stretches, but no new lines appear.⁵⁶

Maintenance, the third stage of the model, is the reason. Each patch of hair keeps its light or dark identity as the hair is shed and regrows, so the same follicles go on making the same color through the cat's life. Geneticists contrast this with many fish, whose stripes can change in number as the fish grows, whereas a tabby's coat is laid down once and then kept.⁵⁷

That is also why the answer had to be sought in the embryo rather than in the kitten. A newborn tabby already wears its full set of stripes, each one destined only to stretch, so by the time a kitten can be photographed the decisions about where its stripes go have all been made. To watch them being made, researchers had to look earlier, at skin that had not yet grown a single hair.⁵⁸

The striped kitten asleep in its basket, then, is wearing the coat it will wear for life, only smaller. While it is still nursing, it can purr as it feeds, and its mother can purr while nursing it. A veterinary behavior expert at NC State University describes exactly that pairing: female cats purr when nursing their young, and kittens purr while nursing.⁵⁹

Tabby is a pattern, and a pattern can be printed in many colors. The registry article illustrates its tabby types with a brown mackerel tabby, a red classic tabby and a chocolate spotted tabby, and the association's page for children shows a silver classic tabby. Alley Cat Allies lists tabbies in brown, orange, silver and buff. The color is the color of the coat; the pattern is what makes the cat a tabby.⁶⁰

Any tabby pattern can also be combined with patches of red or cream, making what the registry calls a patched tabby, or torbie. The red patches come from a sex-linked red gene, and because of that link most patched tabbies are female. The patches may sit apart from the tabby markings or run through them, so a torbie can look like a tabby coat with a second coat laid over it in places.⁶¹

Color, patches and pattern combine freely. The same mackerel stripes can be printed in brown, silver or red, and the association notes that all of the tabby patterns can be combined with red or cream to make a patched tabby. Describing a tabby properly therefore takes at least two words, a color and a pattern, as in brown mackerel tabby, the coat of the cat in this book's pictures.⁶²

Spotted

In the spotted tabby the body markings are spots, which, as the registry puts it, "may vary in size, shape, and arrangement." The face still carries the usual tabby markings, the M included, and the legs and tail are still barred. The Egyptian Mau and the Ocicat are breeds selected for spots. How a spotted coat differs genetically from a mackerel one is not yet known, which makes the spotted tabby one of the open questions of the subject.⁶³

Ticked, and a gene that shrinks the map

The ticked tabby met in the first chapter, with its banded body and marked face, has a known cause. Kaelin, McGowan and Barsh found that Dkk4, the signal-blocking gene that helps lay out the embryonic pre-pattern, is mutated in cats with the ticked pattern, including the Abyssinian. Stanford's report described the result as identifying the mutations behind the Abyssinian's apparent lack of tabby markings.⁶⁴

Apparent is the right word, as Barsh explained to Stanford's news office: if you remove Dkk4, "the dark areas don't go away entirely, but they become smaller and more packed together." A ticked cat is therefore not a cat without a pattern. Its pattern has been shrunk and crowded until, across most of the body, the eye reads it as an even flecking of banded hairs.⁶⁵

The researchers then tested the idea across breeds. They genotyped cats for two Dkk4 variants, known by their protein-change codes as p.Ala18Val and p.Cys63Tyr. In the study, cats of ticked breeds such as the Abyssinian and the Singapura carried those variants, the vast majority of them in two copies. By contrast, none of the cats from breeds whose standards require tabby markings, the Egyptian Mau, the Ocicat and the Bengal, carried a variant at all.⁶⁶

The non-breed cats in the study gave the cleanest result. Among them both looks occur, ticked and not ticked, and the ticked look matched the presence of the p.Ala18Val variant perfectly. The same paper notes that the Ticked trait has been selected for in breeds with a uniform appearance, such as the Abyssinian, Burmese and Singapura, so breeders choosing that even look were, in effect, choosing Dkk4 variants.⁶⁷

Stripes in the shadows

Even a solid black cat is patterned underneath. Stanford's 2021 report notes that all-black cats, too, carry a pattern beneath their dark fur. A study of black coats in wild cats observed that in black domestic cats, faint "ghost" tabby markings are mostly visible in the young and become indistinguishable from the dark background in the adult. Some black cats can still show their stripes in the right light; Alley Cat Allies suggests looking at a black cat's fur in sunlight.⁶⁸

Black leopards show the same effect on a grander scale. In a melanistic leopard, one with a black coat, the rosettes remain visible as even darker markings against a darkened background. The researchers concluded that the rosettes are not obliterated by the mutation that makes the coat black, a change in the gene named in the paper's title, ASIP. The black coat darkens the background, and the pattern stays where it was.⁶⁹

Every coat in this chapter is a variation on the same arrangement: a map of dark and light, laid out early and kept for life, printed in one color or another, sometimes shrunk, sometimes patched, sometimes painted over. The letter M runs through nearly all of them. The next three chapters leave the map and turn to the animal that wears it.

Cats differ in how much contact they like, and tabby cats are no exception: some love cuddles and some like more space. So sit quietly nearby and let the cat choose to come to you.⁷⁰

That advice comes almost word for word from the feline environmental needs guidelines published in 2013 by two veterinary bodies, the American Association of Feline Practitioners and the International Society of Feline Medicine. "Do not force interaction with a cat," the guidelines say. "Let the cat initiate, choose and control the type of human contact." They add practical detail: get down to the cat's level, avoid staring and give the cat time to approach.⁷¹

The reason for that advice is individual variation. "Social preferences among cats vary widely," the guidelines continue, "and are influenced by factors such as genetics, early rearing conditions, and life experiences." A coat pattern appears nowhere on that list. The stripes, swirls and spots of the earlier chapters say a great deal about a cat's ancestry and development, and nothing in this book's sources ties them to how sociable a particular cat will be.⁷²

When a cat does choose company, it often says so with its face. Cats use the head, cheeks and chin for scent and touch communication, and the guidelines note that cats generally rub against their caregivers to mark their scent and as a sign of familiarity. A cat can rub its face against a person it knows, and those same places, the head, cheek and chin, are where most cats prefer to be petted, rather than the belly.⁷³

That preference gives a newcomer a simple guide. When a cat approaches and rubs against a hand, a gentle stroke along the cheek or under the chin can follow the cat's own lead, while the belly, which most cats prefer not to have petted, is best left alone.⁷⁴

The slow blink

A 2020 study by Humphrey and colleagues asked whether a simple facial signal could help. Owners slowly narrowed their eyes at their cats, and the researchers compared the cats' faces with their faces when the owner was present but not blinking. In the study, cats made more eye-narrowing movements of their own after their owners slow-blinked at them.⁷⁵

A second experiment replaced the owner with an unfamiliar experimenter, which removed the long shared history that might explain a friendly response on its own. In the study, cats were more likely to approach the experimenter after a slow-blink interaction than after the experimenter had adopted a neutral expression. The authors suggest that slow blinking may work as a form of positive communication between cats and people.⁷⁶

What the study supports is modest and useful. A slow blink is a calm signal that the cats in the study responded to, and it fits the guidelines' advice to avoid staring and give a cat time, but it is not a password. A cat that blinks back has not agreed to be picked up, and the choice to come closer still belongs to the cat.⁷⁷

A name among words

Whether cats know their own names was the question behind a 2019 study by Saito and colleagues, which played recordings of spoken words to cats. Household cats first heard a list of general nouns or the names of cats they lived with, and their reactions faded as the list went on; then they heard their own name. In the study, the cats reacted again, and they did so even when the words were spoken by people they did not know.⁷⁸

One group of cats behaved differently. Cats living in a cat café did not distinguish their own names from the names of the other café cats they lived with in the study. The result shows discrimination, telling one familiar sound from others, rather than an understanding of a name as a name, and it does not show that a cat will come when called.⁷⁹

Quiet voices and mixed purrs

Sound matters to a cat meeting someone new. The guidelines warn that loud and unfamiliar sounds, whether at home or at a veterinary clinic, can arouse fear in a cat, and a Texas A&M veterinary article adds that background noise people barely notice may stress one. A quiet voice and a calm room are part of letting the cat choose.⁸⁰

A purr is not a signed permission slip either. Purring is linked with friendly contact between cats and with people, and with drowsy rest before sleep. But cats can also purr when they are stressed or hurt: Leslie Lyons of the UC Davis veterinary school notes that cats often purr at the veterinarian or while recovering from injury, and that not every purring cat is content.⁸¹

Put together, the evidence turns the episode's advice into a small routine. Sit quietly, low down and a little apart, rather than looming over the cat, and keep your voice soft, because loud, unfamiliar sounds can frighten a cat. Look away now and then instead of staring, try a slow blink, the signal the cats in the study answered in kind, and then wait. If the cat comes over and rubs its cheek against your hand, it has made its choice, and the choice was always its to make.⁸²

Under the striped coat is a hunter, and play is how a house cat hunts without prey. The feline environmental guidelines advise letting a cat express as many parts of the predatory sequence as possible through play and feeding. Toy play can let a cat stalk, chase, pounce and catch: feather and fur toys to pounce on and toss into the air, soft toys to rake and bite, kibble tossed for it to chase and food hidden for it to find.⁸³

The guidelines are just as specific about safety. All toys with string or other parts that a cat could swallow should be put away after play, toys with small parts or bells should be avoided or have those parts removed before a cat plays alone, and kittens need toys of a suitable size. The cat in this book's pictures pounces on a paper mouse, never on loose string.⁸⁴

Senses tuned to small prey

A cat's whiskers do more than frame its face. The Texas A&M veterinary article explains that cats use their whiskers to enhance their senses of touch and balance, and that whiskers carry information such as whether the cat can fit through a passageway and where obstacles lie. Whiskers grow in several places, especially on the face.⁸⁵

Cats can see far better than people in dim light. Large pupils widen in low light, the retina is rich in light-collecting rod cells, and a reflective layer behind it, the tapetum lucidum, gives light a second chance to be detected. Cornell's Feline Health Center puts a cat's sensitivity to light at about six times a person's, and the Merck Veterinary Manual gives the same figure. Seeing in total darkness is a myth: a cat still needs some light.⁸⁶

Hearing is tuned the same way. Cats are especially good at high-frequency sounds and can hear sounds that people may miss entirely, such as the squeak of a mouse, and the feline guidelines note that cats can hear the ultrasonic chatter of rodents, which helps them locate prey. The same sharp hearing, the guidelines add, helps them notice potential danger.⁸⁷

Taken together, these senses describe an animal equipped to find small prey in poor light. Whiskers report on gaps and obstacles at close range, the eyes make far better use of dim light than a person's can, and the ears can pick up high-pitched rodent calls that people may not hear at all. A toy mouse twitched across a dim rug engages every one of them at once.⁸⁸

A meat eater's tongue

Anyone licked by a cat knows the rough feel of its tongue. A 2018 study of grooming by Noel and Hu describes the cat tongue as "covered in sharp, rear-facing spines called papillae," which the cat uses as it grooms its fur. Every lick of a tabby's flank combs through the same striped map that was laid down before birth.⁸⁹

That same tongue cannot taste sweetness at all. The Texas A&M article explains that cats lack the sweet taste receptor, so a cat that seems drawn to a sweet food is probably responding to its fat and texture instead.⁹⁰

The diet behind the hunt is meat. Cornell's Feline Health Center explains that cats are obligate carnivores, which means that they rely on nutrients found only in animal products, and that they evolved eating prey high in protein, moderate in fat and low in carbohydrate. A complete, balanced cat food, chosen with a veterinarian's advice, meets those needs.⁹¹

A home suits a cat best when it is laid out the way a cat uses space. The feline environmental guidelines recommend that each key resource, the toileting area, food and water bowls, play area, resting or sleeping area and scratching area, be in a separate location. Spreading them out enlarges the cat's territory and gives each cat choices, such as more than one place to rest, eat and use a litter box.⁹²

Drinking water belongs in that plan as well. Cornell's Feline Health Center says that cats should have access to clean, fresh water at all times, and the guidelines add that food and water should be kept apart from each other rather than side by side.⁹³

Height counts as territory too. A high perch can let a cat watch everything around it: the guidelines describe access to an elevated area as increasing a cat's vertical space and allowing it to monitor its surroundings. A perch set on top of a hiding box can help a cat feel safe, because it combines a lookout with a retreat.⁹⁴

Hiding belongs to the same picture. A cat can hide in a safe place when it feels scared, and the guidelines treat hiding as a normal coping behavior, used when a cat is stressed or wants to avoid other animals or people. They recommend giving each cat its own hiding places, such as a cardboard box turned on its side, and a hiding place works only if the cat stays free to come out of it.⁹⁵

Where several cats share a home, that advice applies to each of them. The guidelines recommend separate hiding places for each cat and choices for every cat, such as at least two resting, feeding and toileting areas. Each cat can then eat, rest and retreat on its own terms rather than queuing for a single bowl or box.⁹⁶

A carrier can be one of those hiding places. The guidelines describe a cat carrier as "a transportable, safe place that smells familiar to the cat." A carrier that lets the cat hide, or a blanket over part of it, helps, and the cat's own bedding inside adds familiar scent. Left out at home, it can become a retreat the cat already knows before it is ever used for a trip.⁹⁷

The first of those trips should come early. Cornell's Feline Health Center advises that any new cat be checked by a veterinarian as soon as possible after it comes home, so that a vet can help look after the cat from the start. That first visit is also the right time to ask about the cat's nutrition and grooming history.⁹⁸

The striped cat of the opening has now been unfolded. Its letter M is the signature of a pattern, not a breed. Its stripes are the wildcat's coat, its swirls a broken gene shared with a rare cheetah, and its whole design a map drawn in the skin before birth and kept for life. None of that tells you what kind of company a particular tabby wants. For that, sit quietly nearby and let the cat choose to come to you.