Tiny Kitten Teeth has collaborated with a number of highly successful comics and artists, including videogame cultural icons Penny Arcade. Tiny Kitten Teeth is currently exhibiting at Comic-Con in San Diego, an event that has now become well-established on videogaming’s annual calendar.

Even the most cursory glance around Comic-Con will confirm that comics and videogames go hand in hand. That’s them, just over there. Neon lights and loudspeakers hurriedly dismantled at the close of E3 in Los Angeles and lovingly reassembled 200 kilometres south, here in San Diego.

Why exactly games and comics have always been such happy bedfellows is a little harder to pinpoint. My own best guess is that both mediums simultaneously crept out of the cultural basement – pale, limp-wristed and wheezing – over the past twenty years. Maybe it’s as simple as that.

But whatever the root of their symbiosis we're much richer for their collaboration.

The first comic book to videogame adaptation, or at the very least the first prominent one, was Spiderman for the Atari, back in 1981. It’s a good thing it was called that too, because without the titular cue, there was no way to know that the tiny cluster of pixels on the screen was meant to be your Friendly Neighbourhood Web-Slinger.

We've moved on from such modest beginnings. Showcased extensively both here at Comic-Con and at E3 is Marvel vs. Capcom 3. The damned thing is like crack for a videogame and comic enthusiast (read: nerd) like myself. The art direction endeavours to replicate the way a comic book is inked through the creative application of shadows and cell-shading. The stylised, super-fast motions emulate the jolted pacing of a comic book's panel-to-panel fight scenes. They even use comic book fonts created by professional comic letterers Blambot.

I've heard some trumpet that DC Universe vs. Mortal Kombat was more their cup of tea. Their tea needs to be spat back into the saucer.

It's not the first time Capcom has gone this loyal route, not even the third. The first Capcom Marvel fighting game, Marvel Superheroes, aped the unique style of recently artist deceased Jack Kirby, co-creator of Spiderman, Fantastic Four and many others.

Of all videogame publishers, Capcom has the most significant and storied past with comics. Witness, for example, the side-scrolling beat ‘em up Cadillacs & Dinosaurs, based on the late '80s independent comic sensation and designed to capitalise on the success of games such as SEGA's Streets of Rage. Of course it’s also possible this one was green lit because dinosaurs were pretty hot at the time. Everyone was searching for that next indie comic Ninja Turtle to ride all the way to the bank. See also: Street Sharks.

Shooters like Team Fortress 2 and Battlefield: Heroes are the happy exceptions to their genre. Team Fortress 2 opted – after nearly ten years in development – to throw realism out the window and go instead for a cartoonish look based loosely on the art of vintage illustrators like Norman Rockwell.

The marketing and characterisation of Team Fortress 2 has a distinctly comic book flair, drawing on some of the most absurd and dark humour present in the work of adult comic creators like Garth Ennis and Warren Ellis.

Nintendo has made a surfeit of games employing a cartoonish art style that many more serious gamers have dismissed as childish, titles perceived as neutered by an underpowered, over-gimmicked platform. But the wide appeal of these games is proven their sales figures. They have less in common with superheroes than they do with the more exaggerated comic art of Astroboy creator Osamu Tezuka.

In spite of Nintendo's limited online offerings next to the Xbox 360 and PlayStation 3, the developer has leveraged both their own back catalogue of intellectual property – and that of former rival SEGA – through emulation. Their flush of E3 title announcements trended toward long-neglected franchises such as Kid Icarus, demonstrating their ability to harness nostalgia and expand on simple gameplay mechanics creatively.

They’re the kinds of genres and mechanics in which comic games have previously thrived. Now they’re seeing a resurgence through online distribution models – Xbox LIVE Arcade, PlayStation Network and Steam, for example – low development costs and maturing gamer demand for simpler retro thrills.

Case in point is ultimate homage to retro gaming, Scott Pilgrim vs. The World: The Game. Scott Pilgrim was (is) a six volume, best-selling independent comic book by Brian Lee O’Malley that intertwines indie rock and a relationship story with videogame motifs.

Oni Press, which publishes Scott Pilgrim, originally billed itself within the industry as the “real mainstream.” What they meant is that superhero comics are the mainstream of comics, but non-superhero is the mainstream of wider entertainment media. They've shoe-horned that into a successful company, a new movie directed by Edgar Wright (Shaun of the Dead) and a sassy-looking 8-bit side-scrolling beat ‘em up, replete with arcade-style synths. It will be released on the 10th of August via Xbox LIVE Arcade and the PlayStation Network.

It’s also worth mentioning that while the Scott Pilgrim is under construction at perennial licensee developer Ubisoft, the game is in the style of the comic book rather than the movie, so you don't have to worry about a sprite version of Michael Cera ruining your Juno mad-on (just as they don't have to worry about paying him likeness rights).

The game is very much a throwback to classic comic crossovers such as Captain America & The Avengers and X-Men: The Arcade Game.

That era, defined by arcade fighters, was very good to comic games. It was at the cusp of the ‘90s comic boom with the ridiculous and ill-informed collector market snapping up any and all issue #1s. Generally this involved over-drawn, over-wrought and over-muscled superheroes that no one will ever remember. Even if the comics were dross, the games were entertaining.

More under “It Came From E3”: Telltale Games announced both Jurassic Park and Back to the Future adventure titles. They’re both exceptions to Telltale’s portfolio. Telltale's business is built around downloadable content, with their Strong Bad (of Homestar Runner fame) games at the forefront. They have dealt with comic books a great deal.

Downloadable content is a happy middle ground for comic book adaptations: It gives the name recognition of a licensed work to promising smaller studios without the baggage of a current movie license.

Originally employees of LucasArts, Telltale’s founding members entered the escape pod when the adventure game division at Lucas was dissolved. Telltale are responsible for a well-intentioned adaptation of the fantasy epic graphic novel Bone, as well as a new Sam & Max game based on the comic of the same name by Steve Purcell.

Also created from the dissolution at Lucas was Tim Schafer’s Double Fine, creators of Psychonauts and Brütal Legend. Double Fine’s official website is home to five webcomics, all drawn by members of the studio’s art team.

All of these games are a welcome stylistic diversion from the mode of contemporary videogame graphics. As games strive more earnestly for realism, as they carry on the pilgrimage toward “uncanny valley” – bowed under the weight of greater resolutions, increased draw distances and skyrocketing polygon counts – the visual cues adapted to games from comic books have become an important, necessary counterpoint.

Today, the comic industry is no more immune to the decline of print than any other “paper” medium – newspapers, magazines or books. Curiously, it didn’t necessarily have to be that way for comics. If this industry hadn’t cannibalised its collector culture in the mid to late ‘90s, it may have proved a little more resistant than others.

But as comics increasingly move online, their cross-pollination with videogames strengthens. Just as many games have leaned on comics for inspiration, comics are beginning to borrow from games. Already, some are performing curious digital experiments in partial animation. The defining feature of videogames – user interaction – is the only small step left to make.

You can read Tiny Kitten Teeth here and see their guest pieces on Penny Arcade here, and here.