Box Octopus

[Varinia Swicegood]

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An octopus (pl.: octopuses or octopodes[a]) is a soft-bodied, eight-limbed mollusc of the order Octopoda (/ɒkˈtɒpədə/, ok-TOP-ə-də[3]). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like other cephalopods, an octopus is bilaterally symmetric with two eyes and a beaked mouth at the center point of the eight limbs.[b] The soft body can radically 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 possibly 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.

Historically, the first plural to commonly appear in English language sources, in the early 19th century, is the latinate form octopi,[12] 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.[13]

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).[19] The largest specimen of this species to be scientifically documented was an animal with a live mass of 71 kg (157 lb).[20] Much larger sizes have been claimed for the giant Pacific octopus:[21] one specimen was recorded as 272 kg (600 lb) with an arm span of 9 m (30 ft).[22] 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).[23][24] The smallest species is Octopus wolfi, which is around 2.5 cm (1 in) and weighs less than 1 g (0.035 oz).[25]

The octopus is bilaterally symmetrical along its dorso-ventral (back to belly) 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.[26] The arms can be described based on side and sequence position (such as L1, R1, L2, R2) and divided into four pairs.[27][26] The two rear appendages are generally used to walk on the sea floor, while the other six are used to forage for food.[28] 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.[29][30] The mantle cavity has muscular walls and contains the gills; it is connected to the exterior by a funnel or siphon.[26][31] The mouth of an octopus, located underneath the arms, has a sharp hard beak.[30]

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.[26] 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.[30] 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.[32][33]

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.[34][35] Each of the eight arms senses and responds to light, allowing the octopus to control the limbs even if its head is obscured.[36]

The eyes of the octopus are large and 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; the slit-shaped pupil forms a hole in the iris just behind the cornea. The lens is suspended behind the pupil; photoreceptive retinal cells cover the back of the eye. The pupil can be adjusted in size; a retinal pigment screens incident light in bright conditions.[26]

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.[37][38]

Octopuses have a closed circulatory system, in which the blood remains inside blood vessels. Octopuses have three hearts; a systemic or main heart that circulates blood around the body and two branchial or gill hearts that pump it through each of the two gills. The systemic heart becomes inactive when the animal is swimming. Thus the octopus tires quickly and prefers to crawl.[39][40] 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).[39][40][41] 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.[39][40]

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 branchial hearts and back to the main heart. Much of the venous system is contractile, which helps circulate the blood.[26]

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.[42] Extensive connective tissue lattices support the respiratory muscles and allow them to expand the respiratory chamber.[43] The lamella structure of the gills allows for a high oxygen uptake, up to 65% in water at 20 C (68 F).[44] Water flow over the gills correlates with locomotion, and an octopus can propel its body when it expels water out of its siphon.[43][41]

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