An octopus (: octopuses or octopodes) is a soft-bodied, eight-limbed Mollusca of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class with , cuttlefish, and . Like other cephalopods, an octopus is bilaterally symmetric with two eyes and a cephalopod beak mouth at the centre point of the eight limbs. An octopus can radically deform its shape, enabling it to squeeze through small gaps. They trail their appendages behind them as they swim. The siphon is used for respiration and locomotion (by water jet propulsion). Octopuses have a complex nervous system and excellent sight, and are among the most intelligent and behaviourally diverse .
Octopuses inhabit various ocean habitats, including , pelagic waters, and the seabed; some live in the intertidal zone and others at abyssal zone. Most species grow quickly, mature early, and are short-lived. In most species, the male uses a specially-adapted arm to deliver sperm directly into the female's mantle cavity, after which he becomes senescence and dies, while the female deposits fertilised eggs in a den and cares for them until they hatch, after which she also dies. They are predators and hunt , Bivalvia, and fish. Strategies to defend themselves against their own predators include expelling octopus ink, camouflage, and deimatic display, the ability to jet quickly through the water and hide, and 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 such as the kraken of Norway and the Akkorokamui of the Ainu people, and possibly the Gorgon of ancient Greece. A battle with an octopus appears in Victor Hugo's book Toilers of the Sea. Octopuses appear in Japanese shunga erotic art. They are eaten and considered a delicacy by humans in many parts of the world, especially the Mediterranean and Asia.
The standard form of octopus in English is octopuses; Relevant data at Google Ngram Viewer. the Ancient Greek plural ὀκτώποδες, octopodes (), has also been used historically. The alternative plural octopi is usually considered etymologically incorrect because it hypercorrection that octopus is a Latin second-declension noun or adjective when, in either Greek or Latin, it is a third declension noun.Peters, Pam (2004). The Cambridge Guide to English Usage. Cambridge: Cambridge University Press. , p. 388. Historically, the first plural to commonly appear in English language sources, in the early 19th century, is the Latinate form octopi, followed by the English form octopuses in the latter half of the same century. The Hellenic plural is roughly contemporary in usage, although it is also the rarest.
Fowler's Modern English Usage states that the only acceptable plural in English is octopuses, that octopi is misconceived, and octopodes pedantic; the last 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. The New Oxford American Dictionary (3rd Edition, 2010) lists octopuses as the only acceptable pluralisation, and indicates that octopodes is occasionally used, but that octopi is incorrect.
The skin consists of a thin epidermis with mucous cells and sensory cells and a fibrous dermis made of collagen and containing various cells that allow colour change. Most of the body is made of soft tissue, allowing it to squeeze through tiny gaps; even the larger species can pass through a gap little more than in diameter. Lacking skeletal support, the arms work as muscular hydrostats and feature longitudinal, transverse, and circular muscles around a central axial nerve. They can squash and stretch, coil at any place in any direction or stiffen.
The interior surfaces of the arms are covered with circular, adhesive suckers. The suckers allow the octopus to secure itself in place or to handle objects. Each sucker is typically circular and bowl-like and has two distinct parts: an outer disc-shaped Cephalopod limb and a inner cup-like acetabulum, both of which are thick muscles covered in connective tissue. A chitinous cuticle lines the outer surface. When a sucker attaches to a surface, the orifice between the two structures is sealed and the infundibulum flattens. Muscle contractions allow for attachment and detachment. Each of the eight arms senses and responds to light, allowing the octopus to control its limbs even if its head is obscured.
The cranium has two cartilage capsules each containing one large eye, which resembles those of fish. The cornea is formed from a translucent epidermal layer; the slit-shaped pupil forms a hole in the iris just behind the cornea. The lens hangs behind the pupil; photoreceptive Retina line the back. The pupil can expand and contract; a retinal pigment screens incident light in bright conditions.
Some species differ in form from the typical body shape. Basal species, the Cirrina, have gelatinous bodies with two Cephalopod fin located above the eyes, an cirrate shell and mostly webbed arms that are lined with fleshy papillae or cirri underneath.
The systemic heart has muscular contractile walls and consists of a single ventricle and two atria, which attach it to each of the two gills. The blood vessels consist of arteries, capillaries and veins and are lined with a cellular endothelium unlike that of most other . The blood circulates through the aorta and capillary system, to the Venae cavae, after which the blood is pumped through the gills by the branchial hearts and back to the main heart. Much of the venous system is contractile, which helps circulate the blood.
The thin skin absorbs additional oxygen. When resting, around 41% of oxygen absorption is through the skin, reduced to 33% when the octopus swims, despite the amount of oxygen absorption increasing as water flows over the body. When it is resting after a meal, skin absorption can drop to 3%.
During osmoregulation, fluid is added to the Pericardium of the branchial hearts. The octopus has two nephridium (equivalent to vertebrate kidneys) that are associated with the branchial hearts; these and their associated ducts connect the pericardial cavities with the mantle cavity. Each branch of the vena cava has renal appendages that pass over the thin-walled nephridium before reaching the branchial heart. Urine is created in the pericardial cavity, and is altered by excretion, of mostly ammonia, and absorption from the renal appendages, as it is passed along the associated duct and through the nephridiopore into the mantle cavity.
Like other cephalopods, octopuses have camera-like eyes. Colour vision appears to vary from species to species, for example, it is present in A. aegina but absent in Common octopus. in the skin respond to different wavelengths of light and help the animals choose a colouration that matches the surroundings and camouflages them; chromatophores in the skin can respond to light independently of the eyes. An alternative hypothesis is that in species that only have a single photoreceptor protein may use chromatic aberration to turn monochromatic vision into colour vision, though this lowers image quality. This would explain pupils shaped like the letter "U", the letter "W", or a dumbbell, as well as the need for colourful mating displays.
Attached to the optic capsules are two organs called (sac-like structures containing a mineralised mass and sensitive hairs), that allow the octopus to sense the orientation of its body, relative to both gravity and time (angular acceleration). An autonomic response keeps the octopus's eyes oriented so that the pupil is always horizontal. Octopuses may also use the statocyst to hear. The common octopus can hear sounds between 400 Hz and 1000 Hz, and hears best at 600 Hz.
Octopuses have an excellent somatosensory system. Their suction cups are equipped with chemoreceptors so they can taste what they touch. Octopus arms move easily because the sensors recognise octopus skin and prevent self-attachment. Octopuses appear to have poor proprioception sense and must see their arms to keep track of their position.
Reproduction has been studied in some species. In the giant Pacific octopus, courtship includes changes in skin texture and colour, mostly in the male. 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 in 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. A complex hydraulic mechanism releases the sperm from the spermatophore.
The eggs have large yolks; cleavage (division) is relatively shallow and a germinal disc develops at the pole. During gastrulation, the disc surrounds the yolk, forming a yolk sac, which eventually forms part of the gut. The embryo forms as the dorsal side of the disc grows upward, with a shell gland, gills, mantle and eyes on its dorsal side. The arms and funnel form on the ventral side of the disc, with the former moving upward to surround the mouth. The embryo consumes the yolk during development.
Over a month after mating, Giant Pacific octopuses lay eggs. The species can lay 180,000 eggs in a single clutch, while O. rubescens clutches host up to 45,000 eggs while O. vulgaris clutches can include 500,000 eggs. Fertilised octopus eggs are laid as strings within a shelter. Female giant Pacific octopuses nurture and protect their eggs for five months (160 days) until they hatch. In colder waters, such as those off Alaska, it may take up to ten months for the eggs to completely develop. In the argonaut (paper nautilus), the female is much larger than the male. She secretes a thin shell shaped like a cornucopia, in which the eggs are deposited and in which she also resides and broods the young while swimming.
Most young octopuses hatch as , Octopus larvae in particular are for weeks or months. Larvae feed on shrimp, isopods and , eventually settling on the ocean floor to mature. Species that produce larger eggs instead hatch as benthic animals similar to the adults. These include the southern blue-ringed, Caribbean reef, California two-spot and Eledone moschata.
The Cirrina species are often free-swimming and live in deep-water habitats. Although several species live at bathyal and abyssal depths, only a single indisputable record documents their presence in the hadal zone; a species of Grimpoteuthis (dumbo octopus) photographed at .
Octopuses bring captured prey to the den to eat. Dens are often surrounded by a midden of dead and uneaten food items. These middens may attract scavengers such as fish, molluscs, and . On rare occasions, octopuses hunt cooperatively with other species, with fish as their partners. They regulate the species composition of the hunting and the behavior of their by punching them.
Octopuses typically locate prey by feeling through their environment; some species hide and ambush their target. When prey tries to escape, the octopus jets after it. Octopuses may drill into the shells of crustaceans, bivalves and gastropods. It used to be thought that drilling was done 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. This can take hours and once the shell is penetrated, the prey dies almost instantaneously. With crabs, tough-shelled species are more likely to be drilled, and soft-shelled crabs are torn apart.
Some species have other modes of feeding. Grimpoteuthis either lacks or has a small radula and swallows prey whole. In the deep-sea genus Stauroteuthis, the suckers in most species have been altered into which are believed to fool prey by directing them to the mouth, making them one of the few bioluminescent octopuses.
Cirrate octopuses cannot produce jet propulsion and swim using their fins. Their neutrally buoyant bodies float along while the fins are spread. 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 peristalsis, moving them slowly.
Octopuses can create distracting patterns with waves of dark colouration 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 bumpy appearance of algae-covered rocks. Diurnal, shallow water octopuses have more complex skin than their nocturnal and deep-sea counterparts. In the latter species, skin anatomy is limited to one colour or pattern.
Octopus' "moving rock" trick involves mimicking a rock and then inching across the open space with a speed matching that of the surrounding water.
Octopus try to escape from a predator by ejecting an ink cloud, which acts as a "smoke-screen" or a decoy, as well as to interfere with the attacker's sense of smell. When severed by a predator, some octopuses can autotomy, which can grow back. Some octopuses, such as the mimic octopus, can combine their flexible bodies with their colour-changing ability to mimic other, more dangerous animals, such as Pterois, sea snakes, and .
The are based on Sanchez et al., 2018, who created a molecular phylogeny based on and nuclear DNA marker sequences. The position of the Eledonidae is from Ibáñez et al., 2020, with a similar methodology. Divergence dates are from Kröger et al., 2011 and Fuchs et al., 2019.
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.
The octopus genome is unremarkably bilaterian except for large developments of two gene families: , which regulate the development of neurons; and the C2H2 zinc-finger transcription factors. Many novel genes in both cephalopods generally and octopus specifically manifest in the animals' skin, suckers, and nervous system.
In the Asuka period Japanese legend Taishokan, a female diver battles an octopus to recover a stolen jewel, which became the inspiration for woodblock printings. Similarly, in the 1973 novel Gravity's Rainbow an octopus named Grigori attacks a woman on the beach. A battle with an octopus plays a significant role in Victor Hugo's 1866 book Travailleurs de la mer ( Toilers of the Sea). The octopus continues to be depicted as antagonistic in films such as Wake of the Red Witch (1948).
In political cartoons, octopuses have been used to symbolise empires and large organizations, the arms representing long reach. Octopus also have an erotic appeal. 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 a woman is sexually intertwined with a large and a small octopus. This art style would inspire Pablo Picasso's 1903 drawing An Erotic Drawing: Woman and Octopus. Some individual octopuses gained celebrity status, notably Paul the Octopus who predicted the winners of the 2010 FIFA World Cup.
Octopus is eaten in many cultures, such as those on the Mediterranean and Asian coasts. The arms and other body parts are prepared in ways that vary by species and geography. Live octopuses or their wriggling pieces are consumed as san-nakji in Korean cuisine. If not prepared properly, however, the severed arms can choke the diner with their suction cups, causing at least one death in 2010. Animal welfare groups have objected to the live consumption of octopuses on the basis that they can experience pain.
Octopuses offer many model organism; the California two-spot octopus had its genome sequenced, allowing exploration of its molecular adaptations. Having independently evolved mammal-like intelligence, octopuses were compared by the philosopher Peter Godfrey-Smith, who studied the nature of intelligence, to hypothetical intelligent extraterrestrials. Their intelligence and flexible bodies enable them to escape from supposedly secure tanks in .
Due to their intelligence, many argue that octopuses should be given protections when used for experiments. In the UK from 1993 to 2012, the common octopus ( Octopus vulgaris) was the only invertebrate protected under the Animals (Scientific Procedures) Act 1986. In 2012, this legislation was extended to include all cephalopods in accordance with a general European Union directive.
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.
Circulatory system
Respiration
Digestion and excretion
Nervous system and senses
Ink sac
Life cycle
Reproduction
Lifespan
Distribution and habitat
Behaviour and ecology
Feeding
Locomotion
Intelligence
Camouflage and colour change
Defence
Pathogens and parasites
Evolution
Fossil history and phylogeny
RNA editing and the genome
Relationship to humans
Cultural significance
Danger to humans
As a food source
Science and technology
See also
Notes
External links
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