The reason, perhaps, why most animated cartoons are funny is because it’s in their pedigree; they came right out of the funny papers.

Windsor McCay was one of the most successful and technically brilliant newspaper cartoonists of the early twentieth century. His baroque inventive Little Nemo in Slumberland series galvanized readers across the country as the title hero’s adventures sprawled over an entire page in the Sunday morning funnies. Influenced by his son’s “flip book,” McCay tried his hand at animating Little Nemo for a brief cartoon—it took him four years of painstaking work. After a second attempt, he decided to create a character specifically for the movie screen, a gentle giant named Gertie the Dinosaur, whom McCay unveiled in 1914. It took 10,000 separate drawings, each hand-drawn by McCay on onion-skin, to bring Gertie to life in a one-reeler. Audiences didn’t know what hit them, but clearly the animated short was going to be an essential part of film vocabulary.

The early days of silent cartoons are filled with as many different experiments as their live action counterparts. Animators had to find ways to get beyond the sheer novelty of the process and create something amusing and fluid, a particularly difficult chore, considering the huge manpower required to churn out cartoons at a time when all the technology was done by hand. Transparent celluloid, or “cels”, allowed for a more efficient way of reproducing each frame and pioneer animators Max and Dave Fleischer invented a process called the Rotoscope in 1916, which projected live-action clowning directly onto a drawing board. Their Koko the Clown character was initially created by having one of the Fleischer brothers act out Koko’s antics, while the other rendered the action frame-by-frame. Many early characters were simply transfers from newspaper strips—Krazy Kat, Mutt and Jeff—but in 1919, an original character appeared, a cat who was easy to draw because he was all black. His name was Felix, and his plucky resourcefulness led critics to compare him with Chaplin (who briefly appeared as an animated character himself in the 1910s). In 1923’s “Felix in Hollywood,” our hero actually unscrews his own tail and uses it as a cane for a Chaplin imitation. Felix was also the first animated character to be licensed for commercial products.

According to animation historian Leonard Maltin, as the 1920s ended, “more and more artists and cartoonists got the hang of animation and began to explore more and more with each passing year how you could do things in animation you couldn’t do in a live action film. They created a whole new language for the animated cartoon which gave those characters abilities even beyond Charlie Chaplin and Buster Keaton–and that’s really saying something.” But cartoon animation might have been stopped in its tracks without the invention of sound synchronization in the mid-1920s. There had been some crude “follow-the-bouncing-ball” shorts, which used music, but the industry was revolutionized in 1928, when producer Walt Disney reintroduced a character named Mickey Mouse in a short called “Steamboat Willie.” Mickey’s previous two silent adventures had not even been picked up for distribution, but his new short subject had a synchronized sound score (and was delightfully illustrated by Ub Iwerks). Audiences loved it and Disney built his empire on the little mouse—to whom he always gave credit.

Exaggerated sound fit exaggerated motion to a T—and it was just the element that cartoons needed to lift off the ground. Sound also accelerated the need for tempo; now it was even more crucial for a successful cartoon to have the right timing; it was a difficult trick to master. Pioneer animator Chuck Jones said that “animation is the art of timing. . . . the difference between a huge laugh and a flop can be one frame.” With the success of Walt Disney’s short subjects (Silly Symphonies), movie studios in Hollywood set up full-time animation divisions (Disney’s short cartoons were distributed by RKO); as every feature presentation at the time included several short subjects and cartoons, animation became crucial moneymakers for the studios. Also, as with the creation of real-live movie stars, studios needed to create characters with whom the audience could identify and welcome back week after week. The creators of cartoons now had to expand their canvas dramatically—and that meant becoming part of the world of drama. As another pioneer, Walter Lantz (Woody Woodpecker) put it:

An animator is like an actor before the camera, only he has to act out his feelings and interpret that scene with his pencil; he also has to know how to space characters because the spacing of their movements determines the tempo; he must know expression; he must know feeling; he has to know the character and make him walk with a funny action.

Life as an animator for a studio division was chaotic, intense, frustrating, and often heck of a lot of fun. Executives—other than Disney—rarely knew how to handle these strange men who drew cartoons and animators, frequently fed up with one slight or another, shuttled among the various studios with an alarming frequency; even the most ardent animation fan would have trouble keeping the scorecard straight. Warner Bros started its own division in 1930 by animating songs from their vast musical catalogue with the rip-off title of Looney Tunes and soon added another series of one-offs called Merrie Melodies. But these were initially conceived without successful characters; other studios were having better luck drawing their own stable of stars. Fleischer created Betty Boop in 1930 and brought the comic strip hero Popeye to the screen in 1931. Disney added Donald Duck to Mickey’s menagerie in 1934 and the cantankerous waterfowl soon outstripped his friend’s popularity. Disney outpaced all of his rivals during the Depression years, adding professional voice talent and Technicolor to his cartoons. In 1937, he offered his competitors the greatest challenge of all: a feature-length adaptation of Snow White and the Seven Dwarfs, which used 750 artists who created a quarter of a million drawings at the cost of $1.5 million dollars. It earned six times that amount in its initial release.

Yet for many aficionados, the most exciting animation of the period (and into the 1950s) was to be found at Warner Bros. In the years 1936-1937, the studio (which stuck their animators in a remote bungalow they dubbed “Termite Terrace”) assembled the All-Star Team of cartoonists: directors Tex Avery, Chuck Jones, Bob Clampett, and Frank Tashlin, as well as voice professional extraordinaire Mel Blanc and musical director Carl Stalling. Working in deranged concert with each other, they demonstrated that classic comedy animation was a rare combination of design, voice, effect, character, timing, and point of view. Any combination of those would be amusing; to have all six at once, as Warner Bros often did, was exhilarating. Their stable of two-dimensional celebrities was impressive: Porky Pig, Daffy Duck, Bugs Bunny, and, joining them after WWII, Sylvester and Tweety, Yosemite Sam, Pepe le Pew, and Wile E. Coyote and the Road Runner. What the animators did with them was even better.

Each director had his own take on the material and although the differences might be subtle for the average viewer, they were clear to the animators. Tex Avery is widely created for taking a stock character, Bugs Bunny, and investing him with a definitive character for “A Wild Hare” in 1940. According to his biographer Joe Adamson, “Avery said, ‘How about a character who just isn’t fazed by anything? He comes up out of his rabbit hole, and he’s got a gun in his face, and he just chews his carrot and says, “What’s up, Doc?”’ Avery was known as a keen gag man who wanted his cartoons to be louder, faster, funnier and he was the first animator to have his characters talk to the audience

Chuck Jones, on the other hand, was interested in subtlety and the release of a quiet moment or humorous aside. He saw Bugs Bunny as a cool cookie, a character who only reacted when provoked. The creator of Wile E. Coyote and Road Runner, Jones was a comedy purist and the Stanislavksy of animation, for whom motivation was key: “It was important to give the characters some disciplines—rules that defined the limits of the game,” he once wrote. “Everyone I’ve ever respected used restricted tools. The greatest comedians were the ones who wore the simplest costumes and worked in prescribed areas like Chaplin.” Tashlin experimented in camera angles, blackouts, pace, and absurdity and went on to become a highly valued live-action comedy director, working with stars like Bob Hope and Jerry Lewis. When, on his first non-animated assignment, he created some dueling gags for a Bob Hope period comedy, Tashlin recalled that “It was full of cartoon jokes. I remember Bob saying—they must have told him I came from the cartoon business—things like, ‘Jesus Christ, now I’m a rabbit!’”

However innovative the gang at Warner Bros was, the fact remains that as the 1950s began, short cartoons for the movies were on the wane. The studio system had broken down and executives could no longer insist that exhibitors take two or three cartoons as part of the presentation package. Animation, which was always labor-intensive, became more and more expensive to produce; even Disney was cutting back. New animation units were sprouting up, looking for new ways to do things. UPA was created in 1944, and cut costs by simplifying the visual depth of their characters and abstracting backgrounds. They called the process “limited animation” and their new characters were Gerald McBoing-Boing and Mr. Magoo.

Television soon opened up a whole new market for cartoons; the Faustian bargain was a huge drop in quality and detail. It did little good for animators to wring their hands over the decline of artistry—the small black-and-white rectangular screen mercilessly dictated what looked good and what didn’t. William Hanna and Joseph Barbera had burnished their reputation with the Academy Award-winning Tom and Jerry series at MGM; when they bailed out of movies to work in television, they were shocked by the meager budgets offered by the networks. To make the numbers work, they created an even more limited animation style, often pixilating only the moving elements of a character: the mouth, the hands (or paws). Beginning in 1958, Hanna-Barbera introduced new characters directly to television: Huckleberry Hound, Yogi Bear, and, in 1960, created the first half-hour animated comedy series, The Flintstones, which was a huge ratings hit.

If you grew up in the 1960s, you had a nearly unlimited supply of cartoons available to you; every weekday afternoon, some local affiliate was repackaging old Bugs Bunny or Popeye cartoons, and every Saturday morning, there was nearly three hours of animated programming on each of the three major networks. Not to mention an entire menagerie of tigers, tunas, and toucans pitching products during the commercials. The quality of these candy-colored entertainments varied enormously—both as art and as literature—but they were omnipresent. Some shows broke out of the pack, such as Jay Ward’s various Rocky and Bullwinkle incarnations, which were absurdist, witty indictments of Cold War pieties (although Chuck Jones dismissed their visual crudity as “illustrated radio”).

While television often lacked the virtuosity to create first-rate animated comic characters, they borrowed an astonishing amount of real-life comedy to prime the pump. The Flintstones were Stone Age versions of The Honeymooners and another Hanna-Barbera prime-time show, Top Cat, was a feline adaptation of The Phil Silvers Show. The creators of Amos ‘n’ Andy resurfaced after a decade-long absence to create an animated rip-off called Calvin and the Colonel. Bargain-basement animated versions of Laurel and Hardy, Abbott and Costello, Laverne and Shirley, Jerry Lewis, the Fonz, the Mask, and the Three Stooges (once as human beings, the second time as robots) made the Saturday morning rounds, but even the best cartoons borrowed from their three-dimensional progenitors—who doesn’t recognize a shot of W.C. Fields in Mr. Magoo, a hint of Groucho in Bugs Bunny, a slather of Senator Claghorn in Foghorn Leghorn?

A combination of boredom with the limitations of television animation, computer technology, the freedom of cable networks, and some bright entrepreneurial thinking all help to, well, reanimate the field at the beginning of the 1990s. Disney found a way to recapture its old feature-length magic by tapping into Broadway-style musicals with The Little Mermaid and The Lion King, two major blockbusters that spawned countless imitations both at Disney and other, often brand-new, studios. Cartoon Network was added to the cable system in 1992, eventually acquiring the Hanna-Barbera and Warner Bros catalogues while developing some immensely creative cartoons, such as Dexter’s Laboratory and Powerpuff Girls. MTV took their freedom in another direction, using crudely conceived animation to deliver such counter-culture slackers as Beavis and Butthead and Daria directly to the Generation X audience. First-rate comic talent was no longer reduced to Saturday morning purgatory; comedians such as Nathan Lane, Robin Williams, Eddie Murphy, Ellen Degeneres, Billy Crystal, Woody Allen, Tim Allen, Mike Myers, Roseanne Barr, and Jerry Seinfeld gleefully leaped upon the voice-over bandwagon, providing signature personalities of such dimension in animated cartoons that they were often billed above the title for a film in which they never actually appeared. In 1997, The Simpsons beat The Flintstones’ record for most consecutive prime-time cartoon episodes, and they keep on ticking. . . .

In the 21st Century, animation has transformed, as Aladdin’s cartoon couple would put it, into a whole new world. Cartoons are more popular than ever, and they reach Americans in more ways, in more forms, with more resonance than ever before. It’s a medium that achieves hair-raising technical brilliance on one level—say, Shrek—while making countless viewers giggle uncontrollably with products that could be just as easily created with construction paper and school paste—say, South Park. Cartoon now exist in a wide, tense universe bracketed by alpha and omega, and it makes some purists wistful for the glory days. As Leonard Maltin put it, “Television has reinvigorated animation, has created a new audience for animation, but sadly, has forgotten the history of animation. Modern cartoons couldn’t exist without those [classic] cartoons, and yet they don’t have the heart or originality or the organic humor and originality of those cartoons.” Cartoons remain a comic medium; no matter how detailed a computer can pixilate the green fuzz on an ogre’s nose, it all begins with an artist brandishing a sharp gag and a sharp pencil.