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Inside the mind of the octopus | Orion Magazine

provided by EOL authors

ON AN UNSEASONABLY WARM day in the middle of March, I traveled from New Hampshire to the moist, dim sanctuary of the New England Aquarium, hoping to touch an alternate reality. I came to meet Athena, the aquarium’s forty-pound, five-foot-long, two-and-a-half-year-old giant Pacific octopus.

For me, it was a momentous occasion. I have always loved octopuses. No sci-fi alien is so startlingly strange. Here is someone who, even if she grows to one hundred pounds and stretches more than eight feet long, could still squeeze her boneless body through an opening the size of an orange; an animal whose eight arms are covered with thousands of suckers that taste as well as feel; a mollusk with a beak like a parrot and venom like a snake and a tongue covered with teeth; a creature who can shape-shift, change color, and squirt ink. But most intriguing of all, recent research indicates that octopuses are remarkably intelligent.

Many times I have stood mesmerized by an aquarium tank, wondering, as I stared into the horizontal pupils of an octopus’s large, prominent eyes, if she was staring back at me—and if so, what was she thinking?

Not long ago, a question like this would have seemed foolish, if not crazy. How can an octopusknowanything, much less form an opinion? Octopuses are, after all, “only” invertebrates—they don’t even belong with the insects, some of whom, like dragonflies and dung beetles, at least seem to show some smarts. Octopuses are classified within the invertebrates in the mollusk family, and many mollusks, like clams, have no brain.

Only recently have scientists accorded chimpanzees, so closely related to humans we can share blood transfusions, the dignity of having a mind. But now, increasingly, researchers who study octopuses are convinced that these boneless, alien animals—creatures whose ancestors diverged from the lineage that would lead to ours roughly 500 to 700 million years ago—have developed intelligence, emotions, and individual personalities. Their findings are challenging our understanding of consciousness itself.

I had always longed to meet an octopus. Now was my chance: senior aquarist Scott Dowd arranged an introduction. In a back room, he would open the top of Athena’s tank. If she consented, I could touch her. The heavy lid covering her tank separated our two worlds. One world was mine and yours, the reality of air and land, where we lumber through life governed by a backbone and constrained by jointed limbs and gravity. The other world was hers, the reality of a nearly gelatinous being breathing water and moving weightlessly through it. We think of our world as the “real” one, but Athena’s is realer still: after all, most of the world is ocean, and most animals live there. Regardless of whether they live on land or water, more than 95 percent of all animals are invertebrates, like Athena.

The moment the lid was off, we reached for each other. She had already oozed from the far corner of her lair, where she had been hiding, to the top of the tank to investigate her visitor. Her eight arms boiled up, twisting, slippery, to meet mine. I plunged both my arms elbow deep into the fifty-seven-degree water. Athena’s melon-sized head bobbed to the surface. Her left eye (octopuses have one dominant eye like humans have a dominant hand) swiveled in its socket to meet mine. “She’s looking at you,” Dowd said.

As we gazed into each other’s eyes, Athena encircled my arms with hers, latching on with first dozens, then hundreds of her sensitive, dexterous suckers. Each arm has more than two hundred of them. The famous naturalist and explorer William Beebe found the touch of the octopus repulsive. “I have always a struggle before I can make my hands do their duty and seize a tentacle,” he confessed. But to me, Athena’s suckers felt like an alien’s kiss—at once a probe and a caress. Although an octopus can taste with all of its skin, in the suckers both taste and touch are exquisitely developed. Athena was tasting me and feeling me at once, knowing my skin, and possibly the blood and bone beneath, in a way I could never fathom.

When I stroked her soft head with my fingertips, she changed color beneath my touch, her ruby-flecked skin going white and smooth. This, I learned, is a sign of a relaxed octopus. An agitated giant Pacific octopus turns red, its skin gets pimply, and it erects two papillae over the eyes, which some divers say look like horns. One name for the species is “devil fish.” With sharp, parrotlike beaks, octopuses can bite, and most have neurotoxic, flesh-dissolving venom. The pressure from an octopus’s suckers can tear flesh (one scientist calculated that to break the hold of the suckers of the much smaller common octopus would require a quarter ton of force). One volunteer who interacted with an octopus left the aquarium with arms covered in red hickeys.

Occasionally an octopus takes a dislike to someone. One of Athena’s predecessors at the aquarium, Truman, felt this way about a female volunteer. Using his funnel, the siphon near the side of the head used to jet through the sea, Truman would shoot a soaking stream of salt water at this young woman whenever he got a chance. Later, she quit her volunteer position for college. But when she returned to visit several months later, Truman, who hadn’t squirted anyone in the meanwhile, took one look at her and instantly soaked her again.

Athena was remarkably gentle with me—even as she began to transfer her grip from her smaller, outer suckers to the larger ones. She seemed to be slowly but steadily pulling me into her tank. Had it been big enough to accommodate my body, I would have gone in willingly. But at this point, I asked Dowd if perhaps I should try to detach from some of the suckers. With his help, Athena and I pulled gently apart.

I was honored that she appeared comfortable with me. But what did she know about me that informed her opinion? When Athena looked into my eyes, what was she thinking?


WHILE ALEXA WARBURTON was researching her senior thesis at Middlebury College’s newly created octopus lab, “every day,” she said, “was a disaster.”

She was working with two species: the California two-spot, with a head the size of a clementine, and the smaller, Florida species,Octopus joubini. Her objective was to study the octopuses’ behavior in a T-shaped maze. But her study subjects were constantly thwarting her.

The first problem was keeping the octopuses alive. The four-hundred-gallon tank was divided into separate compartments for each animal. But even though students hammered in dividers, the octopuses found ways to dig beneath them—and eat each other. Or they’d mate, which is equally lethal. Octopuses die after mating and laying eggs, but first they go senile, acting like a person with dementia. “They swim loop-the-loop in the tank, they look all googly-eyed, they won’t look you in the eye or attack prey,” Warburton said. One senile octopus crawled out of the tank, squeezed into a crack in the wall, dried up, and died.

It seemed to Warburton that some of the octopuses were purposely uncooperative. To run the T-maze, the pre-veterinary student had to scoop an animal from its tank with a net and transfer it to a bucket. With bucket firmly covered, octopus and researcher would take the elevator down to the room with the maze. Some octopuses did not like being removed from their tanks. They would hide. They would squeeze into a corner where they couldn’t be pried out. They would hold on to some object with their arms and not let go.

Some would let themselves be captured, only to use the net as a trampoline. They’d leap off the mesh and onto the floor—and then run for it. Yes,run. “You’d chase them under the tank, back and forth, like you were chasing a cat,” Warburton said. “It’s soweird!”

Octopuses in captivity actually escape their watery enclosures with alarming frequency. While on the move, they have been discovered on carpets, along bookshelves, in a teapot, and inside the aquarium tanks of other fish—upon whom they have usually been dining.

Even though the Middlebury octopuses were disaster prone, Warburton liked certain individuals very much. Some, she said, “would lift their arms out of the water like dogs jump up to greet you.” Though in their research papers the students refer to each octopus by a number, the students named them all. One of thejoubiniwas such a problem they named her The Bitch. “Catching her for the maze always took twenty minutes,” Warburton said. “She’d grip onto something and not let go. Once she got stuck in a filter and we couldn’t get her out. It was awful!”

Then there was Wendy. Warburton used Wendy as part of her thesis presentation, a formal event that was videotaped. First Wendy squirted salt water at her, drenching her nice suit. Then, as Warburton tried to show how octopuses use the T-maze, Wendy scurried to the bottom of the tank and hid in the sand. Warburton says the whole debacle occurred because the octopus realized in advance what was going to happen. “Wendy,” she said, “just didn’t feel like being caught in the net.”

Data from Warburton’s experiments showed that the California two-spots quickly learned which side of a T-maze offered a terra-cotta pot to hide in. But Warburton learned far more than her experiments revealed. “Science,” she says, “can only say so much. I know they watched me. I know they sometimes followed me. But they are so different from anything we normally study. How do you prove the intelligence of someone so different?”


MEASURING THE MINDS OF OTHER creatures is a perplexing problem. One yardstick scientists use is brain size, since humans have big brains. But size doesn’t always match smarts. As is well known in electronics, anything can be miniaturized. Small brain size was the evidence once used to argue that birds were stupid—before some birds were proven intelligent enough to compose music, invent dance steps, ask questions, and do math.

Octopuses have the largest brains of any invertebrate. Athena’s is the size of a walnut—as big as the brain of the famous African gray parrot, Alex, who learned to use more than one hundred spoken words meaningfully. That’s proportionally bigger than the brains of most of the largest dinosaurs.

Another measure of intelligence: you can count neurons. The common octopus has about 130 million of them in its brain. A human has 100 billion. But this is where things get weird. Three-fifths of an octopus’s neurons are not in the brain; they’re in its arms.

“It is as if each arm has a mind of its own,” says Peter Godfrey-Smith, a diver, professor of philosophy at the Graduate Center of the City University of New York, and an admirer of octopuses. For example, researchers who cut off an octopus’s arm (which the octopus can regrow) discovered that not only does the arm crawl away on its own, but if the arm meets a food item, it seizes it—and tries to pass it to where the mouth would be if the arm were still connected to its body.

“Meeting an octopus,” writes Godfrey-Smith, “is like meeting an intelligent alien.” Their intelligence sometimes even involves changing colors and shapes. One video online shows a mimic octopus alternately morphing into a flatfish, several sea snakes, and a lionfish by changing color, altering the texture of its skin, and shifting the position of its body. Another video shows an octopus materializing from a clump of algae. Its skin exactly matches the algae from which it seems to bloom—until it swims away.

For its color palette, the octopus uses three layers of three different types of cells near the skin’s surface. The deepest layer passively reflects background light. The topmost may contain the colors yellow, red, brown, and black. The middle layer shows an array of glittering blues, greens, and golds. But how does an octopus decide what animal to mimic, what colors to turn? Scientists have no idea, especially given that octopuses are likelycolorblind.

But new evidence suggests a breathtaking possibility. Woods Hole Marine Biological Laboratory and University of Washington researchers found that the skin of the cuttlefishSepia officinalis, a color-changing cousin of octopuses, contains gene sequences usually expressed only in the light-sensing retina of the eye. In other words, cephalopods—octopuses, cuttlefish, and squid—may be able to see with their skin.

The American philosopher Thomas Nagel once wrote a famous paper titled “What Is It Like to Be a Bat?” Bats can see with sound. Like dolphins, they can locate their prey using echoes. Nagel concluded it was impossible to know what it’s like to be a bat. And a bat is a fellow mammal like us—not someone who tastes with its suckers, sees with its skin, and whose severed arms can wander about, each with a mind of its own. Nevertheless, there are researchers still working diligently to understand what it’s like to be an octopus.


JENNIFER MATHER SPENT MOST of her time in Bermuda floating facedown on the surface of the water at the edge of the sea. Breathing through a snorkel, she was watchingOctopus vulgaris—the common octopus. Although indeed common (they are found in tropical and temperate waters worldwide), at the time of her study in the mid-1980s, “nobody knew what they were doing.”

In a relay with other students from six-thirty in the morning till six-thirty at night, Mather worked to find out. Sometimes she’d see an octopus hunting. A hunting expedition could take five minutes or three hours. The octopus would capture something, inject it with venom, and carry it home to eat. “Home,” Mather found, is where octopuses spend most of their time. A home, or den, which an octopus may occupy only a few days before switching to a new one, is a place where the shell-less octopus can safely hide: a hole in a rock, a discarded shell, or a cubbyhole in a sunken ship. One species, the Pacific red octopus, particularly likes to den in stubby, brown, glass beer bottles.

One octopus Mather was watching had just returned home and was cleaning the front of the den with its arms. Then, suddenly, it left the den, crawled a meter away, picked up one particular rock and placed the rock in front of the den. Two minutes later, the octopus ventured forth to select a second rock. Then it chose a third. Attaching suckers to all the rocks, the octopus carried the load home, slid through the den opening, and carefully arranged the three objects in front. Then it went to sleep. What the octopus was thinking seemed obvious: “Three rocks are enough. Good night!”

The scene has stayed with Mather. The octopus “must have had some concept,” she said, “of what it wanted to make itself feel safe enough to go to sleep.” And the octopus knew how to get what it wanted: by employing foresight, planning—and perhaps even tool use. Mather is the lead author ofOctopus: The Ocean’s Intelligent Invertebrate, which includes observations of octopuses who dismantle Lego sets and open screw-top jars. Coauthor Roland Anderson reports that octopuses even learned to open the childproof caps on Extra Strength Tylenol pill bottles—a feat that eludes many humans with university degrees.

In another experiment, Anderson gave octopuses plastic pill bottles painted different shades and with different textures to see which evoked more interest. Usually each octopus would grasp a bottle to see if it were edible and then cast it off. But to his astonishment, Anderson saw one of the octopuses doing something striking: she was blowing carefully modulated jets of water from her funnel to send the bottle to the other end of her aquarium, where the water flow sent it back to her. She repeated the action twenty times. By the eighteenth time, Anderson was already on the phone with Mather with the news: “She’s bouncing the ball!”

This octopus wasn’t the only one to use the bottle as a toy. Another octopus in the study also shot water at the bottle, sending it back and forth across the water’s surface, rather than circling the tank. Anderson’s observations were reported in theJournal of Comparative Psychology.“This fit all the criteria for play behavior,” said Anderson. “Only intelligent animals play—animals like crows and chimps, dogs and humans.”

Aquarists who care for octopuses feel that not only can these animals play with toys, but they may need to play with toys. AnOctopus Enrichment Handbookhas been developed by Cincinnati’s Newport Aquarium, with ideas of how to keep these creatures entertained. One suggestion is to hide food inside Mr. Potato Head and let your octopus dismantle it. At the Seattle Aquarium, giant Pacific octopuses play with a baseball-sized plastic ball that can be screwed together by twisting the two halves. Sometimes the mollusks screw the halves back together after eating the prey inside.

At the New England Aquarium, it took an engineer who worked on the design of cubic zirconium to devise a puzzle worthy of a brain like Athena’s. Wilson Menashi, who began volunteering at the aquarium weekly after retiring from the Arthur D. Little Corporation sixteen years ago, devised a series of three Plexiglas cubes, each with a different latch. The smallest cube has a sliding latch that twists to lock down, like the bolt on a horse stall. Aquarist Bill Murphy puts a crab inside the clear cube and leaves the lid open. Later he lets the octopus lift open the lid. Finally he locks the lid, and invariably the octopus figures out how to open it.

Next he locks the first cube within a second one. The new latch slides counterclockwise to catch on a bracket. The third box is the largest, with two different locks: a bolt that slides into position to lock down, and a second one like a lever arm, sealing the lid much like the top of an old-fashioned glass canning jar.

All the octopuses Murphy has known learned fast. They typically master a box within two or three once-a-week tries. “Once they ‘get it,’” he says, “they can open it very fast”—within three or four minutes. But each may use a different strategy.

George, a calm octopus, opened the boxes methodically. The impetuous Gwenevere squeezed the second-largest box so hard she broke it, leaving a hole two inches wide. Truman, Murphy said, was “an opportunist.” One day, inside the smaller of the two boxes, Murphy put two crabs, who started to fight. Truman was too excited to bother with locks. He poured his seven-foot-long body through the two-inch crack Gwenevere had made, and visitors looked into his exhibit to find the giant octopus squeezed, suckers flattened, into the tiny space between the walls of the fourteen-cubic-inch box outside and the six-cubic-inch one inside it. Truman stayed inside half an hour. He never opened the inner box—probably he was too cramped.

Three weeks after I had first met Athena, I returned to the aquarium to meet the man who had designed the cubes. Menashi, a quiet grandfather with a dark moustache, volunteers every Tuesday. “He has a real way with octopuses,” Dowd and Murphy told me. I was eager to see how Athena behaved with him.

Murphy opened the lid of her tank, and Athena rose to the surface eagerly. A bucket with a handful of fish sat nearby. Did she rise so eagerly sensing the food? Or was it the sight of her friend that attracted her? “She knows me,” Menashi answered softly.

Anderson’s experiments with giant Pacific octopuses in Seattle prove Menashi is right. The study exposed eight octopuses to two unfamiliar humans, dressed identically in blue aquarium shirts. One person consistently fed a particular octopus, and another always touched it with a bristly stick. Within a week, at first sight of the people, most octopuses moved toward the feeders and away from the irritators, at whom they occasionally aimed their water-shooting funnels.

Upon seeing Menashi, Athena reached up gently and grasped his hands and arms. She flipped upside down, and he placed a capelin in some of the suckers near her mouth, at the center of her arms. The fish vanished. After she had eaten, Athena floated in the tank upside down, like a puppy asking for a belly rub. Her arms twisted lazily. I took one in my hand to feel the suckers—did that arm know it had hold of a different person than the other arms did? Her grip felt calm, relaxed. With me, earlier, she seemed playful, exploratory, excited. The way she held Menashi with her suckers seemed to me like the way a long-married couple holds hands at the movies.

I leaned over the tank to look again into her eyes, and she bobbed up to return my gaze. “She has eyelids like a person does,” Menashi said. He gently slid his hand near one of her eyes, causing her to slowly wink.


BIOLOGISTS HAVE LONG NOTED the similarities between the eyes of an octopus and the eyes of a human. Canadian zoologist N. J. Berrill called it “the single most startling feature of the whole animal kingdom” that these organs are nearly identical: both animals’ eyes have transparent corneas, regulate light with iris diaphragms, and focus lenses with a ring of muscle.

Scientists are currently debating whether we and octopuses evolved eyes separately, or whether a common ancestor had the makings of the eye. But intelligence is another matter. “The same thing that got them their smarts isn’t the same thing that got us our smarts,” says Mather, “because our two ancestors didn’t have any smarts.” Half a billion years ago, the brainiest thing on the planet had only a few neurons. Octopus and human intelligence evolved independently.

“Octopuses,” writes philosopher Godfrey-Smith, “are a separate experiment in the evolution of the mind.” And that, he feels, is what makes the study of the octopus mind so philosophically interesting.

The octopus mind and the human mind probably evolved for different reasons. Humans—like other vertebrates whose intelligence we recognize (parrots, elephants, and whales)—are long-lived, social beings. Most scientists agree that an important event that drove the flowering of our intelligence was when our ancestors began to live in social groups. Decoding and developing the many subtle relationships among our fellows, and keeping track of these changing relationships over the course of the many decades of a typical human lifespan, was surely a major force shaping our minds.

But octopuses are neither long-lived nor social. Athena, to my sorrow, may live only a few more months—the natural lifespan of a giant Pacific octopus is only three years. If the aquarium added another octopus to her tank, one might eat the other. Except to mate, most octopuses have little to do with others of their kind.

So why is the octopus so intelligent? What is its mindfor? Mather thinks she has the answer. She believes the event driving the octopus toward intelligence was the loss of the ancestral shell. Losing the shell freed the octopus for mobility. Now they didn’t need to wait for food to find them; they could hunt like tigers. And while most octopuses love crab best, they hunt and eat dozens of other species—each of which demands a different hunting strategy. Each animal you hunt may demand a different skill set: Will you camouflage yourself for a stalk-and-ambush attack? Shoot through the sea for a fast chase? Or crawl out of the water to capture escaping prey?

Losing the protective shell was a trade-off. Just about anything big enough to eat an octopus will do so. Each species of predator also demands a different evasion strategy—from flashing warning coloration if your attacker is vulnerable to venom, to changing color and shape to camouflage, to fortifying the door to your home with rocks.

Such intelligence is not always evident in the laboratory. “In the lab, you give the animals this situation, and they react,” points out Mather. But in the wild, “the octopus is actively discovering his environment, not waiting for it to hit him. The animal makes the decision to go out and get information, figures out how to get the information, gathers it, uses it, stores it. This has a great deal to do with consciousness.”

So what does it feel like to be an octopus? Philosopher Godfrey-Smith has given this a great deal of thought, especially when he meets octopuses and their relatives, giant cuttlefish, on dives in his native Australia. “They come forward and look at you. They reach out to touch you with their arms,” he said. “It’s remarkable how little is known about them . . . but I could see it turning out that we have to change the way we think of the nature of the mind itself to take into account minds with less of a centralized self.”

“I think consciousness comes in different flavors,” agrees Mather. “Some may have consciousness in a way we may not be able to imagine.”


IN MAY, I VISITED Athena a third time. I wanted to see if she recognized me. But how could I tell? Scott Dowd opened the top of her tank for me. Athena had been in a back corner but floated immediately to the top, arms outstretched, upside down.

This time I offered her only one arm. I had injured a knee and, feeling wobbly, used my right hand to steady me while I stood on the stool to lean over the tank. Athena in turn gripped me with only one of her arms, and very few of her suckers. Her hold on me was remarkably gentle.

I was struck by this, since Murphy and others had first described Athena’s personality to me as “feisty.” “They earn their names,” Murphy had told me. Athena is named for the Greek goddess of wisdom, war, and strategy. She is not usually a laid-back octopus, like George had been. “Athena could pull you into the tank,” Murphy had warned. “She’s curious about what you are.”

Was she less curious now? Did she remember me? I was disappointed that she did not bob her head up to look at me. But perhaps she didn’t need to. She may have known from the taste of my skin who I was. But why was this feisty octopus hanging in front of me in the water, upside down?

Then I thought I might know what she wanted from me. She was begging. Dowd asked around and learned that Athena hadn’t eaten in a couple of days, then allowed me the thrilling privilege of handing her a capelin.

Perhaps I had understood something basic about what it felt like to be Athena at that moment: she was hungry. I handed a fish to one of her larger suckers, and she began to move it toward her mouth. But soon she brought more arms to the task, and covered the fish with many suckers—as if she were licking her fingers, savoring the meal.


A WEEK AFTER I LAST VISITED ATHENA, I was shocked to receive this e-mail from Scott Dowd: “Sorry to write with some sad news. Athena appears to be in her final days, or even hours. She will live on, though, through your conveyance.” Later that same day, Dowd wrote to tell me that she had died. To my surprise, I found myself in tears.

Why such sorrow? I had understood from the start that octopuses don’t live very long. I also knew that while Athena did seem to recognize me, I was not by any means her special friend. But she was very significant to me, both as an individual and as a representative from her octopodan world. She had given me a great gift: a deeper understanding of what it means to think, to feel, and to know. I was eager to meet more of her kind.

And so, it was with some excitement that I read this e-mail from Dowd a few weeks later: “There is a young pup octopus headed to Boston from the Pacific Northwest. Come shake hands (x8) when you can.”

Octopus

provided by wikipedia EN

The octopus (plural octopuses) is a soft-bodied, eight-limbed mollusc of the order Octopoda (/ɒktˈɑːpədə/, ahk-TAH-pə-də). Around 300 species are recognised, and the order is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like other cephalopods, the octopus is bilaterally symmetric with two eyes and a beak, with its mouth at the center point of the eight limbs ("tentacle" is used as an umbrella term for cephalopod limbs; however, within a teuthological context, "arm" is used to refer to such limbs while "tentacle" is reserved for feeding appendages not found on octopuses). The soft body can rapidly alter its shape, enabling octopuses to squeeze through small gaps. They trail their eight appendages behind them as they swim. The siphon is used both for respiration and for locomotion, by expelling a jet of water. Octopuses have a complex nervous system and excellent sight, and are among the most intelligent and behaviourally diverse of all invertebrates.

Octopuses inhabit various regions of the ocean, including coral reefs, pelagic waters, and the seabed; some live in the intertidal zone and others at abyssal depths. Most species grow quickly, mature early, and are short-lived. In most species, the male uses a specially adapted arm to deliver a bundle of sperm directly into the female's mantle cavity, after which he becomes senescent and dies, while the female deposits fertilised eggs in a den and cares for them until they hatch, after which she also dies. Strategies to defend themselves against predators include the expulsion of ink, the use of camouflage and threat displays, the ability to jet quickly through the water and hide, and even deceit. All octopuses are venomous, but only the blue-ringed octopuses are known to be deadly to humans.

Octopuses appear in mythology as sea monsters like the Kraken of Norway and the Akkorokamui of the Ainu, and probably the Gorgon of ancient Greece. A battle with an octopus appears in Victor Hugo's book Toilers of the Sea, inspiring other works such as Ian Fleming's Octopussy. Octopuses appear in Japanese erotic art, shunga. They are eaten and considered a delicacy by humans in many parts of the world, especially the Mediterranean and the Asian seas.

Etymology and pluralisation

The scientific Latin term octopus was derived from Ancient Greek ὀκτώπους, a compound form of ὀκτώ (oktō, "eight") and πούς (pous, "foot"), itself a variant form of ὀκτάπους, a word used for example by Alexander of Tralles (c. 525–605) for the common octopus.[3][4][5] The standard pluralised form of "octopus" in English is "octopuses";[6] the Ancient Greek plural ὀκτώποδες, "octopodes" (/ɒkˈtɒpədz/), has also been used historically.[7] The alternative plural "octopi" is considered grammatically incorrect because it wrongly assumes that octopus is a Latin second declension "-us" noun or adjective when, in either Greek or Latin, it is a third declension noun.[8][9] Fowler's Modern English Usage states that the only acceptable plural in English is "octopuses", that "octopi" is misconceived, and "octopodes" pedantic;[10][11][12] the latter is nonetheless used frequently enough to be acknowledged by the descriptivist Merriam-Webster 11th Collegiate Dictionary and Webster's New World College Dictionary. The Oxford English Dictionary lists "octopuses", "octopi", and "octopodes", in that order, reflecting frequency of use, calling "octopodes" rare and noting that "octopi" is based on a misunderstanding.[13] The New Oxford American Dictionary (3rd Edition, 2010) lists "octopuses" as the only acceptable pluralisation, and indicates that "octopodes" is still occasionally used, but that "octopi" is incorrect.[14]

Anatomy and physiology

Size

"Captured
A Giant Pacific octopus at Echizen Matsushima Aquarium, Japan.

The giant Pacific octopus (Enteroctopus dofleini) is often cited as the largest known octopus species. Adults usually weigh around 15 kg (33 lb), with an arm span of up to 4.3 m (14 ft).[15] The largest specimen of this species to be scientifically documented was an animal with a live mass of 71 kg (156.5 lb).[16] Much larger sizes have been claimed for the giant Pacific octopus:[17] one specimen was recorded as 272 kg (600 lb) with an arm span of 9 m (30 ft).[18] A carcass of the seven-arm octopus, Haliphron atlanticus, weighed 61 kg (134 lb) and was estimated to have had a live mass of 75 kg (165 lb).[19][20] The smallest species is Octopus wolfi, which is around 2.5 cm (1 in) and weighs less than 1 g (0.035 oz).[21]

External characteristics

The octopus is bilaterally symmetrical along its dorso-ventral axis; the head and foot are at one end of an elongated body and function as the anterior (front) of the animal. The head includes the mouth and brain. The foot has evolved into a set of flexible, prehensile appendages, known as "arms", that surround the mouth and are attached to each other near their base by a webbed structure.[22] The arms can be described based on side and sequence position (such as L1, R1, L2, R2) and divided into four pairs.[23][22] The two rear appendages are generally used to walk on the sea floor, while the other six are used to forage for food; hence some biologists refer to the animals as having six "arms" and two "legs".[24][25] The bulbous and hollow mantle is fused to the back of the head and is known as the visceral hump; it contains most of the vital organs.[26][27] The mantle cavity has muscular walls and contains the gills; it is connected to the exterior by a funnel or siphon.[22][28] The mouth of an octopus, located underneath the arms, has a sharp hard beak.[27]

"Schematic
Diagram of octopus from side, with gills, funnel, eye, ocellus (eyespot), web, arms, suckers, hectocotylus and ligula labelled.

The skin consists of a thin outer epidermis with mucous cells and sensory cells, and a connective tissue dermis consisting largely of collagen fibres and various cells allowing colour change.[22] Most of the body is made of soft tissue allowing it to lengthen, contract, and contort itself. The octopus can squeeze through tiny gaps; even the larger species can pass through an opening close to 2.5 cm (1 in) in diameter.[27] Lacking skeletal support, the arms work as muscular hydrostats and contain longitudinal, transverse and circular muscles around a central axial nerve. They can extend and contract, twist to left or right, bend at any place in any direction or be held rigid.[29][30]

The interior surfaces of the arms are covered with circular, adhesive suckers. The suckers allow the octopus to anchor itself or to manipulate objects. Each sucker is usually circular and bowl-like and has two distinct parts: an outer shallow cavity called an infundibulum and a central hollow cavity called an acetabulum, both of which are thick muscles covered in a protective chitinous cuticle. When a sucker attaches to a surface, the orifice between the two structures is sealed. The infundibulum provides adhesion while the acetabulum remains free, and muscle contractions allow for attachment and detachment.[31][32]

"A
A finned Grimpoteuthis species with its atypical octopus body plan.

The eyes of the octopus are large and are at the top of the head. They are similar in structure to those of a fish and are enclosed in a cartilaginous capsule fused to the cranium. The cornea is formed from a translucent epidermal layer and the slit-shaped pupil forms a hole in the iris and lies just behind. The lens is suspended behind the pupil and photoreceptive retinal cells cover the back of the eye. The pupil can be adjusted in size and a retinal pigment screens incident light in bright conditions.[22]

Some species differ in form from the typical octopus body shape. Basal species, the Cirrina, have stout gelatinous bodies with webbing that reaches near the tip of their arms, and two large fins above the eyes, supported by an internal shell. Fleshy papillae or cirri are found along the bottom of the arms, and the eyes are more developed.[33][34]

Circulatory system

Octopuses have a closed circulatory system, in which the blood remains inside blood vessels. Octopuses have three hearts; a systemic heart that circulates blood around the body and two branchial hearts that pump it through each of the two gills. The systemic heart is inactive when the animal is swimming and thus it tires quickly and prefers to crawl.[35][36] Octopus blood contains the copper-rich protein haemocyanin to transport oxygen. This makes the blood very viscous and it requires considerable pressure to pump it around the body; octopuses' blood pressures can exceed 75 mmHg (10 kPa).[35][36][37] In cold conditions with low oxygen levels, haemocyanin transports oxygen more efficiently than haemoglobin. The haemocyanin is dissolved in the plasma instead of being carried within blood cells, and gives the blood a bluish colour.[35][36]

The systemic heart has muscular contractile walls and consists of a single ventricle and two atria, one for each side of the body. The blood vessels consist of arteries, capillaries and veins and are lined with a cellular endothelium which is quite unlike that of most other invertebrates. The blood circulates through the aorta and capillary system, to the vena cavae, after which the blood is pumped through the gills by the auxiliary hearts and back to the main heart. Much of the venous system is contractile, which helps circulate the blood.[22]

Respiration

"An
Octopus with open siphon. The siphon is used for respiration, waste disposal and discharging ink.

Respiration involves drawing water into the mantle cavity through an aperture, passing it through the gills, and expelling it through the siphon. The ingress of water is achieved by contraction of radial muscles in the mantle wall, and flapper valves shut when strong circular muscles force the water out through the siphon.[38] Extensive connective tissue lattices support the respiratory muscles and allow them to expand the respiratory chamber.[39] The lamella structure of the gills allows for a high oxygen uptake, up to 65% in water at 20 °C (68 °F).[40] Water flow over the gills correlates with locomotion, and an octopus can propel its body when it expels water out of its siphon.[39][37]

The thin skin of the octopus absorbs additional oxygen. When resting, around 41% of an octopus's oxygen absorption is through the skin. This decreases to 33% when it swims, as more water flows over the gills; skin oxygen uptake also increases. When it is resting after a meal, absorption through the skin can drop to 3% of its total oxygen uptake.[41]

Digestion and excretion

The digestive system of the octopus begins with the buccal mass which consists of the mouth with its chitinous beak, the pharynx, radula and salivary glands.[42] The radula is a spiked, muscular tongue-like organ with multiple rows of tiny teeth.[27] Food is broken down and is forced into the oesophagus by two lateral extensions of the esophageal side walls in addition to the radula. From there it is transferred to the gastrointestinal tract, which is mostly suspended from the roof of the mantle cavity by numerous membranes. The tract consists of a crop, where the food is stored; a stomach, where food is ground down; a caecum where the now sludgy food is sorted into fluids and particles and which plays an important role in absorption; the digestive gland, where liver cells break down and absorb the fluid and become "brown bodies"; and the intestine, where the accumulated waste is turned into faecal ropes by secretions and blown out of the funnel via the rectum.[42][43]

During osmoregulation, fluid is added to the pericardia of the branchial hearts. The octopus has two nephridia (equivalent to vertebrate kidneys) which are associated with the branchial hearts; these and their associated ducts connect the pericardial cavities with the mantle cavity. Before reaching the branchial heart, each branch of the vena cava expands to form renal appendages which are in direct contact with the thin-walled nephridium. The urine is first formed in the pericardial cavity, and is modified by excretion, chiefly of ammonia, and selective absorption from the renal appendages, as it is passed along the associated duct and through the nephridiopore into the mantle cavity.[22][44]

A common octopus (Octopus vulgaris) moving around. Its nervous system allows the arms to move with some autonomy.

Nervous system and senses

The octopus (along with cuttlefish) has the highest brain-to-body mass ratios of all invertebrates; it is also greater than that of many vertebrates.[45][46] It has a highly complex nervous system, only part of which is localised in its brain, which is contained in a cartilaginous capsule.[47] Two-thirds of an octopus's neurons are found in the nerve cords of its arms, which show a variety of complex reflex actions that persist even when they have no input from the brain.[48] Unlike vertebrates, the complex motor skills of octopuses are not organised in their brain via an internal somatotopic map of its body, instead using a nonsomatotopic system unique to large-brained invertebrates.[49]

Like other cephalopods, octopuses can distinguish the polarisation of light. Colour vision appears to vary from species to species, for example being present in O. aegina but absent in O. vulgaris.[50] Researchers believe that opsins in the skin can sense different wavelengths of light and help the creatures choose a coloration that camouflages them, in addition to light input from the eyes.[51] Other researchers hypothesise that cephalopod eyes in species which only have a single photoreceptor protein may use chromatic aberration to turn monochromatic vision into colour vision, though this sacrifices image quality.[52] This would explain pupils shaped like the letter U, the letter W, or a dumbbell, as well as explaining the need for colourful mating displays.[53]

Attached to the brain are two special organs called statocysts (sac-like structures containing a mineralised mass and sensitive hairs), that allow the octopus to sense the orientation of its body. They provide information on the position of the body relative to gravity and can detect angular acceleration. An autonomic response keeps the octopus's eyes oriented so that the pupil is always horizontal.[22] Octopuses may also use the statocyst to hear sound. The common octopus can hear sounds between 400 Hz and 1000 Hz, and hears best at 600 Hz.[54]

Octopuses also have an excellent sense of touch. The octopus's suction cups are equipped with chemoreceptors so the octopus can taste what it touches. Octopus arms do not become tangled or stuck to each other because the sensors recognise octopus skin and prevent self-attachment.[55]

The arms contain tension sensors so the octopus knows whether its arms are stretched out, but this is not sufficient for the brain to determine the position of the octopus's body or arms. As a result, the octopus does not possess stereognosis; that is, it does not form a mental image of the overall shape of the object it is handling. It can detect local texture variations, but cannot integrate the information into a larger picture. The neurological autonomy of the arms means the octopus has great difficulty learning about the detailed effects of its motions. It has a poor proprioceptive sense, and it knows what exact motions were made only by observing the arms visually.[56]

Ink sac

The ink sac of an octopus is located under the digestive gland. A gland attached to the sac produces the ink, and the sac stores it. The sac is close enough to the funnel for the octopus to shoot out the ink with a water jet. Before it leaves the funnel, the ink passes through glands which mix it with mucus, creating a thick, dark blob which allows the animal to escape from a predator.[57] The main pigment in the ink is melanin, which gives it its black colour.[58] Cirrate octopuses lack the ink sac.[33]

Lifecycle

Reproduction

Octopuses are gonochoric and have a single, posteriorly-located gonad which is associated with the coelom. The testis in males and the ovary in females bulges into the gonocoel and the gametes are released here. The gonocoel is connected by the gonoduct to the mantle cavity, which it enters at the gonopore.[22] An optic gland creates hormones that cause the octopus to mature and age and stimulate gamete production. The gland may be triggered by environmental conditions such as temperature, light and nutrition, which thus control the timing of reproduction and lifespan.[59][60]

When octopuses reproduce, the male uses a specialised arm called a hectocotylus to transfer spermatophores (packets of sperm) from the terminal organ of the reproductive tract (the cephalopod "penis") into the female's mantle cavity.[61] The hectocotylus in benthic octopuses is usually the third right arm, which has a spoon-shaped depression and modified suckers near the tip. In most species, fertilisation occurs in the mantle cavity.[22]

The reproduction of octopuses has been studied in only a few species. One such species is the giant Pacific octopus, in which courtship is accompanied, especially in the male, by changes in skin texture and colour. The male may cling to the top or side of the female or position himself beside her. There is some speculation that he may first use his hectocotylus to remove any spermatophore or sperm already present in the female. He picks up a spermatophore from his spermatophoric sac with the hectocotylus, inserts it into the female's mantle cavity, and deposits it in the correct location for the species, which in the giant Pacific octopus is the opening of the oviduct. Two spermatophores are transferred in this way; these are about one metre (yard) long, and the empty ends may protrude from the female's mantle.[62] A complex hydraulic mechanism releases the sperm from the spermatophore, and it is stored internally by the female.[22]

"A
Female giant Pacific octopus guarding strings of eggs

About forty days after mating, the female giant Pacific octopus attaches strings of small fertilised eggs (10,000 to 70,000 in total) to rocks in a crevice or under an overhang. Here she guards and cares for them for about five months (160 days) until they hatch.[62] In colder waters, such as those off of Alaska, it may take as much as 10 months for the eggs to completely develop.[63]:74 The female aerates the eggs and keeps them clean; if left untended, many eggs will not hatch.[64] She does not feed during this time and dies soon afterwards. Males become senescent and die a few weeks after mating.[65]

The eggs have large yolks; cleavage (division) is superficial and a germinal disc develops at the pole. During gastrulation, the margins of this grow down and surround the yolk, forming a yolk sac, which eventually forms part of the gut. The dorsal side of the disc grows upwards and forms the embryo, with a shell gland on its dorsal surface, gills, mantle and eyes. The arms and funnel develop as part of the foot on the ventral side of the disc. The arms later migrate upwards, coming to form a ring around the funnel and mouth. The yolk is gradually absorbed as the embryo develops.[22]

"A
Octopus paralarva, a planktonic hatchling

Most young octopuses hatch as paralarvae and are planktonic for weeks to months, depending on the species and water temperature. They feed on copepods, arthropod larvae and other zooplankton, eventually settling on the ocean floor and developing directly into adults with no distinct metamorphoses that are present in other groups of mollusc larvae.[22] Octopus species that produce larger eggs – including the southern blue-ringed, Caribbean reef, California two-spot, Eledone moschata[66] and deep sea octopuses – do not have a paralarval stage, but hatch as benthic animals similar to the adults.[63]:74–75[67]

In the argonaut (paper nautilus), the female secretes a fine, fluted, papery shell in which the eggs are deposited and in which she also resides while floating in mid-ocean. In this she broods the young, and it also serves as a buoyancy aid allowing her to adjust her depth. The male argonaut is minute by comparison and has no shell.[68]

Lifespan

Octopuses have a relatively short life expectancy; some species live for as little as six months. The giant Pacific octopus, one of the two largest species of octopus, may live for as much as five years. Octopus lifespan is limited by reproduction: males can live for only a few months after mating, and females die shortly after their eggs hatch. The larger Pacific striped octopus is an exception, as it can reproduce multiple times over a life of around two years.[69] Octopus reproductive organs mature due to the hormonal influence of the optic gland but result in the inactivation of their digestive glands, typically causing the octopus to die from starvation.[70]:276–277 Experimental removal of both optic glands after spawning was found to result in the cessation of broodiness, the resumption of feeding, increased growth, and greatly extended lifespans.[71]

Distribution and habitat

Octopus cyanea in Kona, Hawaii

Octopuses live in every ocean, and different species have adapted to different marine habitats. As juveniles, common octopuses inhabit shallow tide pools. The Hawaiian day octopus (Octopus cyanea) lives on coral reefs; argonauts drift in pelagic waters. Abdopus aculeatus mostly lives in near-shore seagrass beds. Some species are adapted to the cold, ocean depths. The spoon-armed octopus (Bathypolypus arcticus) is found in abyssal plains at depths of 1,000 m (3,300 ft), and Vulcanoctopus hydrothermalis lives near hydrothermal vents at 2,000 m (6,600 ft).[26] The cirrate species are often free-swimming and live in deep-water habitats.[34] No species are known to live in fresh water.[72]

Behaviour and ecology

Most species are solitary when not mating,[73] though a few are known to occur in high densities and with frequent interactions, signaling, mate defending and eviction of individuals from dens. This is likely the result of abundant food supplies combined with limited den sites.[74] The larger Pacific striped octopus however is social, living in groups of up to 40 individuals that share dens.[69] Octopuses hide in dens, which are typically crevices in rocky outcrops or other hard structures, though some species burrow into sand or mud. Octopuses are not territorial but generally remain in a home range; they may leave the area in search of food. They can use navigation skills to return to a den without having to retrace their outward route.[75] They are not known to be migratory.[76]

Octopuses bring captured prey back to the den where they can eat it safely. Sometimes the octopus catches more prey than it can eat, and the den is often surrounded by a midden of dead and uneaten food items. Other creatures, such as fish, crabs, molluscs and echinoderms, often share the den with the octopus, either because they have arrived as scavengers, or because they have survived capture.[77]

Feeding

"An
Veined octopus eating a crab

Nearly all octopuses are predatory; bottom-dwelling octopuses eat mainly crustaceans, polychaete worms, and other molluscs such as whelks and clams; open-ocean octopuses eat mainly prawns, fish and other cephalopods.[78] Major items in the diet of the giant Pacific octopus include bivalve molluscs such as the cockle Clinocardium nuttallii, clams and scallops and crustaceans such as crabs and spider crabs. Prey that it is likely to reject include moon snails because they are too large and limpets, rock scallops, chitons and abalone, because they are too securely fixed to the rock.[77]

A benthic (bottom-dwelling) octopus typically moves among the rocks and feels through the crevices. The creature may make a jet-propelled pounce on prey and pull it towards the mouth with its arms, the suckers restraining it. Small prey may be completely trapped by the webbed structure. Octopuses usually inject crustaceans like crabs with a paralysing saliva then dismember them with their beaks.[78][79] Octopuses feed on shelled molluscs either by forcing the valves apart, or by drilling a hole in the shell to inject a nerve toxin.[80][79] It used to be thought that the hole was drilled by the radula, but it has now been shown that minute teeth at the tip of the salivary papilla are involved, and an enzyme in the toxic saliva is used to dissolve the calcium carbonate of the shell. It takes about three hours for O. vulgaris to create a 0.6 mm (0.024 in) hole. Once the shell is penetrated, the prey dies almost instantaneously, its muscles relax, and the soft tissues are easy for the octopus to remove. Crabs may also be treated in this way; tough-shelled species are more likely to be drilled, and soft-shelled crabs are torn apart.[81]

Some species have other modes of feeding. Grimpoteuthis has a reduced or non-existent radula and swallows prey whole.[33] In the deep-sea genus Stauroteuthis, some of the muscle cells that control the suckers in most species have been replaced with photophores which are believed to fool prey by directing them towards the mouth, making them one of the few bioluminescent octopuses.[82]

Locomotion

"An
Octopuses swim with their arms trailing behind.

Octopuses mainly move about by relatively slow crawling with some swimming in a head-first position. Jet propulsion or backwards swimming, is their fastest means of locomotion, followed by swimming and crawling.[83] When in no hurry, they usually crawl on either solid or soft surfaces. Several arms are extended forwards, some of the suckers adhere to the substrate and the animal hauls itself forwards with its powerful arm muscles, while other arms may push rather than pull. As progress is made, other arms move ahead to repeat these actions and the original suckers detach. During crawling, the heart rate nearly doubles, and the animal requires ten or fifteen minutes to recover from relatively minor exercise.[29]

Most octopuses swim by expelling a jet of water from the mantle through the siphon into the sea. The physical principle behind this is that the force required to accelerate the water through the orifice produces a reaction that propels the octopus in the opposite direction.[84] The direction of travel depends on the orientation of the siphon. When swimming, the head is at the front and the siphon is pointed backwards, but when jetting, the visceral hump leads, the siphon points towards the head and the arms trail behind, with the animal presenting a fusiform appearance. In an alternative method of swimming, some species flatten themselves dorso-ventrally, and swim with the arms held out sideways, and this may provide lift and be faster than normal swimming. Jetting is used to escape from danger, but is physiologically inefficient, requiring a mantle pressure so high as to stop the heart from beating, resulting in a progressive oxygen deficit.[83]

"Three
Movements of the finned species Cirroteuthis muelleri

Cirrate octopuses cannot produce jet propulsion and rely on their fins for swimming. They have neutral buoyancy and drift through the water with the fins extended. They can also contract their arms and surrounding web to make sudden moves known as "take-offs". Another form of locomotion is "pumping", which involves symmetrical contractions of muscles in their webs producing peristaltic waves. This moves the body slowly.[33]

In 2005, Adopus aculeatus and veined octopus (Amphioctopus marginatus) were found to walk on two arms, while at the same time mimicking plant matter.[85] This form of locomotion allows these octopuses to move quickly away from a potential predator without being recognised.[83] A study of this behaviour led to the suggestion that the two rearmost appendages may be more accurately termed "legs" rather than "arms".[86] Some species of octopus can crawl out of the water briefly, which they may do between tide pools while hunting crustaceans or gastropods or to escape predators.[87][88] "Stilt walking" is used by the veined octopus when carrying stacked coconut shells. The octopus carries the shells underneath it with two arms, and progresses with an ungainly gait supported by its remaining arms held rigid.[89]

Intelligence

"A
Octopus opening a container by unscrewing its cap

Octopuses are highly intelligent; the extent of their intelligence and learning capability are not well defined.[90][91][92][93] Maze and problem-solving experiments have shown evidence of a memory system that can store both short- and long-term memory. It is not known precisely what contribution learning makes to adult octopus behaviour. Young octopuses learn nothing from their parents, as adults provide no parental care beyond tending to their eggs until the young octopuses hatch.[63]:75

In laboratory experiments, octopuses can be readily trained to distinguish between different shapes and patterns. They have been reported to practise observational learning,[94] although the validity of these findings is contested.[90][91] Octopuses have also been observed in what has been described as play: repeatedly releasing bottles or toys into a circular current in their aquariums and then catching them.[95] Octopuses often break out of their aquariums and sometimes into others in search of food.[87][96][97] They have even boarded fishing boats and opened holds to eat crabs.[92] The veined octopus collects discarded coconut shells, then uses them to build a shelter, an example of tool use.[89][98][99]

Camouflage and colour change

Video of Octopus cyanea moving and changing its colour, shape and texture

Octopuses use camouflage when hunting and to avoid predators. To do this they use specialised skin cells which change the appearance of the skin by adjusting its colour, opacity, or reflectivity. Chromatophores contain yellow, orange, red, brown, or black pigments; most species have three of these colours, while some have two or four. Other colour-changing cells are reflective iridophores and white leucophores.[100] This colour-changing ability is also used to communicate with or warn other octopuses.[101]

Octopuses can create distracting patterns with waves of dark coloration across the body, a display known as the "passing cloud". Muscles in the skin change the texture of the mantle to achieve greater camouflage. In some species, the mantle can take on the spiky appearance of algae; in others, skin anatomy is limited to relatively uniform shades of one colour with limited skin texture. Octopuses that are diurnal and live in shallow water have evolved more complex skin than their nocturnal and deep-sea counterparts.[101]

A "moving rock" trick involves the octopus mimicking a rock and then inching across the open space with a speed matching the movement in the surrounding water, allowing it to move in plain sight of a predator.[93][102]

Defence

"An
Warning display of greater blue-ringed octopus (Hapalochlaena lunulata)

Aside from humans, octopuses may be preyed on by fishes, seabirds, sea otters, pinnipeds, cetaceans, and other cephalopods.[103] Octopuses typically hide or disguise themselves by camouflage and mimicry; some have conspicuous warning coloration (aposematism) or deimatic behaviour.[101] An octopus may spend 40% of its time hidden away in its den. When the octopus is approached, it may extend an arm to investigate. 66% of Enteroctopus dofleini in one study had scars, with 50% having amputated arms.[103] The blue rings of the highly venomous blue-ringed octopus are hidden in muscular skin folds which contract when the animal is threatened, exposing the iridescent warning.[104] The Atlantic white-spotted octopus (Callistoctopus macropus) turns bright brownish red with oval white spots all over in a high contrast display.[105] Displays are often reinforced by stretching out the animal's arms, fins or web to make it look as big and threatening as possible.[106]

Once they have been seen by a predator, they commonly try to escape but can also use distraction with an ink cloud ejected from the ink sac. The ink is thought to reduce the efficiency of olfactory organs, which would aid evasion from predators that employ smell for hunting, such as sharks. Ink clouds of some species might act as pseudomorphs, or decoys that the predator attacks instead.[107]

When under attack, some octopuses can perform arm autotomy, in a manner similar to the way skinks and other lizards detach their tails. The crawling arm may distract would-be predators. Such severed arms remain sensitive to stimuli and move away from unpleasant sensations.[108] Octopuses can replace lost limbs.[109]

Some octopuses, such as the mimic octopus, can combine their highly flexible bodies with their colour-changing ability to mimic other, more dangerous animals, such as lionfish, sea snakes, and eels.[110][111]

Pathogens and parasites

The diseases and parasites that affect octopuses have been little studied, but cephalopods are known to be the intermediate or final hosts of various parasitic cestodes, nematodes and copepods; 150 species of protistan and metazoan parasites have been recognised.[112] The Dicyemidae are a family of tiny worms that are found in the renal appendages of many species;[113] it is unclear whether they are parasitic or are endosymbionts. Coccidians in the genus Aggregata living in the gut cause severe disease to the host. Octopuses have an innate immune system, and the haemocytes respond to infection by phagocytosis, encapsulation, infiltration or cytotoxic activities to destroy or isolate the pathogens. The haemocytes play an important role in the recognition and elimination of foreign bodies and wound repair. Captive animals have been found to be more susceptible to pathogens than wild ones.[114] A gram-negative bacterium, Vibrio lentus, has been found to cause skin lesions, exposure of muscle and death of octopuses in extreme cases.[115]

Evolution

The scientific name Octopoda was first coined and given as the order of octopuses in 1818 by English biologist William Elford Leach,[116] who classified them as Octopoida the previous year.[2] The Octopoda consists of around 300 known species[117] and were historically divided into two suborders, the Incirrina and the Cirrina.[34] However, more recent evidence suggests that Cirrina are merely the most basal species and are not a unique clade.[118] The incirrate octopuses (the majority of species) lack the cirri and paired swimming fins of the cirrates.[34] In addition, the internal shell of incirrates is either present as a pair of stylets or absent altogether.[119]

Fossil history and phylogeny

Cephalopods have existed for 500 million years and octopus ancestors were in the Carboniferous seas 300 million years ago. The oldest known octopus fossil is Pohlsepia, which lived 296 million years ago. Researchers have identified impressions of eight arms, two eyes, and possibly an ink sac.[121] Octopuses are mostly soft tissue, and so fossils are relatively rare. Octopuses, squids and cuttlefish belong to the clade Coleoidea. They are known as "soft-bodied" cephalopods, lacking the external shell of most molluscs and other cephalopods like the nautiloids and the extinct Ammonoidea.[122] Octopuses have eight limbs like other coleoids but lack the extra specialised feeding appendages known as tentacles which are longer and thinner with suckers only at their club-like ends.[123][124][125] The vampire squid (Vampyroteuthis) also lacks tentacles but has sensory filaments.[126]

The cladograms are based on Sanchez et al., 2018, who created a molecular phylogeny based on mitochondrial and nuclear DNA marker sequences.[118]

Cephalopods Nautiloids

Nautilus "A

  Coleoids  

Squids and cuttlefish "A

     

Vampyroteuthis "A

   

Octopods "A

       

The molecular analysis of the octopods shows that the suborder Cirrina (Cirromorphida) and the superfamily Argonautoidea are paraphyletic and are broken up; these names are shown in quotation marks and italics on the cladogram.

Octopoda

part of "Cirromorphida" (Cirroteuthidae, Stauroteuthidae)

    Octopodida      

part of "Argonautoidea" (Argonautidae, Ocythoidae) "Papierboot

Octopodoidea

Bathypolypodidae "Bathypolypus

       

Octopodidae "Octopus

   

Megaleledonidae "Graneledone

     

Enteroctopodidae "E

           

Bolitaenidae "Eledonella

     

Amphitretidae "Amphitretus

   

Vitreledonellidae "Vitreledonella

         

part of "Argonautoidea" (Tremoctopodidae, Alloposidae) "Pelagic

     

part of "Cirromorphida" (Opisthoteuthidae, Cirroctopodidae)"Opisthoteuthis

     

RNA editing

Octopuses and other coleoid cephalopods are capable of greater RNA editing (which involves changes to the nucleic acid sequence of the primary transcript of RNA molecules) than any other organisms. Editing is concentrated in the nervous system and affects proteins involved in neural excitability and neuronal morphology. More than 60% of RNA transcripts for coleoid brains are recoded by editing, compared to less than 1% for a human or fruit fly. Coleoids rely mostly on ADAR enzymes for RNA editing, which requires large double-stranded RNA structures to flank the editing sites. Both the structures and editing sites are conserved in the coleoid genome and the mutation rates for the sites are severely hampered. Hence, greater transcriptome plasticity has come at the cost of slower genome evolution. High levels of RNA editing do not appear to be present in more basal cephalopods or other molluscs.[127][128]

Relationship to humans

"An
Minoan clay vase with octopus decoration, c. 1500 BC

Cultural references

Ancient seafaring people were aware of the octopus, as evidenced by certain artworks and designs. For example, a stone carving found in the archaeological recovery from Bronze Age Minoan Crete at Knossos (1900–1100 BC) has a depiction of a fisherman carrying an octopus.[129] The terrifyingly powerful Gorgon of Greek mythology has been thought to have been inspired by the octopus or squid, the octopus itself representing the severed head of Medusa, the beak as the protruding tongue and fangs, and its tentacles as the snakes.[130] The Kraken are legendary sea monsters of giant proportions said to dwell off the coasts of Norway and Greenland, usually portrayed in art as a giant octopus attacking ships. Linnaeus included it in the first edition of his 1735 Systema Naturae.[131][132] One translation of the Hawaiian creation myth the Kumulipo suggests that the octopus is the lone survivor of a previous age.[133][134][135] The Akkorokamui is a gigantic octopus-like monster from Ainu folklore.[136]

"A
The NROL-39 mission patch, depicting the National Reconnaissance Office as an octopus with a long reach

A battle with an octopus plays a significant role in Victor Hugo's book Travailleurs de la mer (Toilers of the Sea), relating to his time in exile on Guernsey.[137] Ian Fleming's 1966 short story collection Octopussy and The Living Daylights, and the 1983 James Bond film were partly inspired by Hugo's book.[138]

Japanese erotic art, shunga, includes ukiyo-e woodblock prints such as Katsushika Hokusai's 1814 print Tako to ama (The Dream of the Fisherman's Wife), in which an ama diver is sexually intertwined with a large and a small octopus.[139][140] The print is a forerunner of tentacle erotica.[141] The biologist P. Z. Myers noted in his science blog, Pharyngula, that octopuses appear in "extraordinary" graphic illustrations involving women, tentacles, and bare breasts.[142][143]

Since it has numerous arms emanating from a common centre, the octopus is often used as a symbol for a powerful and manipulative organisation.[144]

Danger

"Coloured
Pen and wash drawing of an imagined colossal octopus attacking a ship, by the malacologist Pierre de Montfort, 1801

Octopuses generally avoid humans, but incidents have been verified. For example, a 2.4-metre (8 ft) Pacific octopus, said to be nearly perfectly camouflaged, "lunged" at a diver and "wrangled" over his camera before it let go. Another diver recorded the encounter on video.[145]

All species are venomous, but only blue-ringed octopuses have venom that is lethal to humans.[146] Bites are reported each year across the animals' range from Australia to the eastern Indo-Pacific Ocean. They bite only when provoked or accidentally stepped upon; bites are small and usually painless. The venom appears to be able to penetrate the skin without a puncture, given prolonged contact. It contains tetrodotoxin, which causes paralysis by blocking the transmission of nerve impulses to the muscles. This causes death by respiratory failure leading to cerebral anoxia. No antidote is known, but if breathing can be kept going artificially, patients recover within 24 hours.[147][148] Bites have been recorded from captive octopuses of other species; they leave swellings which disappear in a day or two.[149]

Fisheries and cuisine

Octopus fisheries exist around the world with total catches varying between 245,320 and 322,999 metric tons from 1986 to 1995.[150] The world catch peaked in 2007 at 380,000 tons, and fell by a tenth by 2012.[151] Methods to capture octopuses include pots, traps, trawls, snares, drift fishing, spearing, hooking and hand collection.[150] Octopus is eaten in many cultures and is a common food on the Mediterranean and Asian coasts.[152][153] The arms and sometimes other body parts are prepared in various ways, often varying by species or geography. Live octopuses are eaten in several countries around the world, including the US.[154][155] Animal welfare groups have objected to this practice on the basis that octopuses can experience pain.[156] Octopuses have a food conversion efficiency greater than that of chickens, making octopus aquaculture a possibility.[157]

In science and technology

"An
Octopus in captivity at Aquarium Finisterrae in Corunna, Galicia, Spain. These animals are known to escape from their tanks.

In classical Greece, Aristotle (384–322 BC) commented on the colour-changing abilities of the octopus, both for camouflage and for signalling, in his Historia animalium: "The octopus ... seeks its prey by so changing its colour as to render it like the colour of the stones adjacent to it; it does so also when alarmed."[158] Aristotle noted that the octopus had a hectocotyl arm and suggested it might be used in sexual reproduction. This claim was widely disbelieved until the 19th century. It was described in 1829 by the French zoologist Georges Cuvier, who supposed it to be a parasitic worm, naming it as a new species, Hectocotylus octopodis.[159][160] Other zoologists thought it a spermatophore; the German zoologist Heinrich Müller believed it was "designed" to detach during copulation. In 1856 the Danish zoologist Japetus Steenstrup demonstrated that it is used to transfer sperm, and only rarely detaches.[161]

"
Flexible biomimetic 'Octopus' robotics arm. The BioRobotics Institute, Scuola Superiore Sant'Anna, Pisa, 2011

Octopuses offer many possibilities in biological research, including their ability to regenerate limbs, change the colour of their skin, behave intelligently with a distributed nervous system, and make use of 168 kinds of protocadherins (humans have 58), the proteins that guide the connections neurons make with each other. The California two-spot octopus has had its genome sequenced, allowing exploration of its molecular adaptations.[162] Having independently evolved mammal-like intelligence, octopuses have been compared to hypothetical intelligent extraterrestrials.[163] Their problem-solving skills, along with their mobility and lack of rigid structure enable them to escape from supposedly secure tanks in laboratories and public aquariums.[164]

Due to their intelligence, octopuses are listed in some countries as experimental animals on which surgery may not be performed without anesthesia, a protection usually extended only to vertebrates. In the UK from 1993 to 2012, the common octopus (Octopus vulgaris) was the only invertebrate protected under the Animals (Scientific Procedures) Act 1986.[165] In 2012, this legislation was extended to include all cephalopods[166] in accordance with a general EU directive.[167]

Some robotics research is exploring biomimicry of octopus features. Octopus arms can move and sense largely autonomously without intervention from the animal's central nervous system. In 2015 a team in Italy built soft-bodied robots able to crawl and swim, requiring only minimal computation.[168] In 2017 a German company made an arm with a soft pneumatically controlled silicone gripper fitted with two rows of suckers. It is able to grasp objects such as a metal tube, a magazine, or a ball, and to fill a glass by pouring water from a bottle.[169]

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Octopus: Brief Summary

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The octopus (plural octopuses) is a soft-bodied, eight-limbed mollusc of the order Octopoda (/ɒktˈɑːpədə/, ahk-TAH-pə-də). Around 300 species are recognised, and the order is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like other cephalopods, the octopus is bilaterally symmetric with two eyes and a beak, with its mouth at the center point of the eight limbs ("tentacle" is used as an umbrella term for cephalopod limbs; however, within a teuthological context, "arm" is used to refer to such limbs while "tentacle" is reserved for feeding appendages not found on octopuses). The soft body can rapidly alter its shape, enabling octopuses to squeeze through small gaps. They trail their eight appendages behind them as they swim. The siphon is used both for respiration and for locomotion, by expelling a jet of water. Octopuses have a complex nervous system and excellent sight, and are among the most intelligent and behaviourally diverse of all invertebrates.

Octopuses inhabit various regions of the ocean, including coral reefs, pelagic waters, and the seabed; some live in the intertidal zone and others at abyssal depths. Most species grow quickly, mature early, and are short-lived. In most species, the male uses a specially adapted arm to deliver a bundle of sperm directly into the female's mantle cavity, after which he becomes senescent and dies, while the female deposits fertilised eggs in a den and cares for them until they hatch, after which she also dies. Strategies to defend themselves against predators include the expulsion of ink, the use of camouflage and threat displays, the ability to jet quickly through the water and hide, and even deceit. All octopuses are venomous, but only the blue-ringed octopuses are known to be deadly to humans.

Octopuses appear in mythology as sea monsters like the Kraken of Norway and the Akkorokamui of the Ainu, and probably the Gorgon of ancient Greece. A battle with an octopus appears in Victor Hugo's book Toilers of the Sea, inspiring other works such as Ian Fleming's Octopussy. Octopuses appear in Japanese erotic art, shunga. They are eaten and considered a delicacy by humans in many parts of the world, especially the Mediterranean and the Asian seas.

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