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Cephalopod ink is a dark-coloured or luminous ink released into water by most species of
cephalopod A cephalopod is any member of the molluscan class Cephalopoda ( Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
, usually as an escape mechanism. All cephalopods, with the exception of the
Nautilidae The nautilus (, ) is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in ...
and the Cirrina (deep-sea octopuses), are able to release ink to confuse predators. The ink is released from the ink sacs (located between the
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they ar ...
s) and is dispersed more widely when its release is accompanied by a jet of water from the
siphon A siphon (from grc, σίφων, síphōn, "pipe, tube", also spelled nonetymologically syphon) is any of a wide variety of devices that involve the flow of liquids through tubes. In a narrower sense, the word refers particularly to a tube in a ...
. Its dark color is caused by its main constituent,
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
. Each species of cephalopod produces slightly differently coloured inks; generally, octopuses produce black ink, squid ink is blue-black, and cuttlefish ink is a shade of brown. A number of other aquatic molluscs have similar responses to attack, including the
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
clade known as
sea hare The clade Anaspidea, commonly known as sea hares ('' Aplysia'' species and related genera), are medium-sized to very large opisthobranch gastropod molluscs with a soft internal shell made of protein. These are marine gastropod molluscs in the ...
s.


Types of ink shapes

The shapes taken by ink releases are classified as six types: * pseudomorphs; * pseudomorph series; * ink ropes; * clouds/smokescreens; * diffuse puffs; * mantle fills.


Inking behaviors


Escape strategies

Two distinct behaviors have been observed in inking cephalopods. The first is the release of large amounts of ink into the water by the cephalopod in order to create a dark, diffuse cloud (much like a
smoke screen A smoke screen is smoke released to mask the movement or location of military units such as infantry, tanks, aircraft, or ships. Smoke screens are commonly deployed either by a canister (such as a grenade) or generated by a vehicle (such as ...
) that can obscure the predator's view, allowing the cephalopod to make a rapid retreat by jetting away. The second response to a predator is to release pseudomorphs ("false bodies"), smaller clouds of ink with a greater
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
content, which allows them to hold their shape for longer. These are expelled slightly away from the cephalopod in question, which will often release several pseudomorphs and change colour ( blanch) in conjunction with these releases. The pseudomorphs are roughly the same volume as and look similar to the cephalopod that released them, and many predators have been observed attacking them mistakenly, allowing the cephalopod to escape (this behaviour is often referred to as the "blanch-ink-jet manoeuvre"). Thus, such capture avoidance method is analogous to fighter jet dogfights where the followed pilot releases countermeasures (such as flares) to misdirect the sensors in some guided missiles. Furthermore,
green turtle The green sea turtle (''Chelonia mydas''), also known as the green turtle, black (sea) turtle or Pacific green turtle, is a species of large sea turtle of the family Cheloniidae. It is the only species in the genus ''Chelonia''. Its range exten ...
(''Chelonia mydas'') hatchlings that have been observed mistakenly attacking pseudomorphs released by '' Octopus bocki'' have subsequently ignored
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organis ...
octopuses.


Hiding strategy

The spotty bobtail squid releases ropes of ink longer than itself and hides among them, maybe to be confused with floating
seagrass Seagrasses are the only flowering plants which grow in marine environments. There are about 60 species of fully marine seagrasses which belong to four families ( Posidoniaceae, Zosteraceae, Hydrocharitaceae and Cymodoceaceae), all in the ...
leaves.


Behavior around eggs

Octopuses have also been observed squirting ink at
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class ...
s or
crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all th ...
s approaching their
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
. Numerous cuttlefish species add a coat of ink to their eggs, presumably to camouflage them from potential predators.


Properties


Attack protection

Inking has been shown to protect species of squids against predatory fish attacks, as well during the capture phase as during the consummatory phase, due to visual and chemical effects of the ink releases.


Chemical effects

Many cephalopod predators (for instance
moray eel Moray eels, or Muraenidae (), are a family of eels whose members are found worldwide. There are approximately 200 species in 15 genera which are almost exclusively marine, but several species are regularly seen in brackish water, and a few are f ...
s) have advanced chemosensory systems, and some anecdotal evidenceMacGinitie, G.E. and MacGinitie, N. (1968) ''Natural History of Marine Animals'', pp. 395–397, 2nd ed. McGraw-Hill, New York. suggests that compounds (such as
tyrosinase Tyrosinase is an oxidase that is the rate-limiting enzyme for controlling the production of melanin. The enzyme is mainly involved in two distinct reactions of melanin synthesis otherwise known as the Raper Mason pathway. Firstly, the hydroxy ...
) found in cephalopod ink can irritate, numb or even deactivate such apparatus. Few controlled experiments have been conducted to substantiate this. Cephalopod ink is nonetheless generally thought to be more sophisticated than a simple "smoke screen"; the ink of a number of squid and cuttlefish has been shown to function as a conspecific chemical alarm.


Physical properties

'' Sepia officinalis'' ink forms a polydisperse suspension composed by spheric particles with a size between 80 and 150 nm (measured by
TRPS Tunable Resistive Pulse Sensing (TRPS) is a single-particle technique used to measure the size, concentration and zeta potential of particles as they pass through a size-tunable nanopore. The technique adapts the principle of resistive pulse sens ...
and SEM). The particles have a density of , which may be due to the amount of metals that the ink has in its composition (4.7% in weight). '' Heteroteuthis dispar'' is a cephalopod species known for releasing luminous ink. The light comes from a substance produced by a dedicated organ before being transferred into the ink sac.


Chemical composition

Cephalopod ink contains a number of chemicals in a variety of different concentrations, depending on the species. However, its main constituents are
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
and
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
. It can also contain, among others,
tyrosinase Tyrosinase is an oxidase that is the rate-limiting enzyme for controlling the production of melanin. The enzyme is mainly involved in two distinct reactions of melanin synthesis otherwise known as the Raper Mason pathway. Firstly, the hydroxy ...
,
dopamine Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic chemical of the catecholamine and phenethylamine families. Dopamine constitutes about 80% o ...
, and
L-DOPA -DOPA, also known as levodopa and -3,4-dihydroxyphenylalanine, is an amino acid that is made and used as part of the normal biology of some plants and animals, including humans. Humans, as well as a portion of the other animals that utilize -DOPA ...
, as well as small amounts of free
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
s, including
taurine Taurine (), or 2-aminoethanesulfonic acid, is an organic compound that is widely distributed in animal tissues. It is a major constituent of bile and can be found in the large intestine, and accounts for up to 0.1% of total human body weight. I ...
,
aspartic acid Aspartic acid (symbol Asp or D; the ionic form is known as aspartate), is an α-amino acid that is used in the biosynthesis of proteins. Like all other amino acids, it contains an amino group and a carboxylic acid. Its α-amino group is in the pro ...
,
glutamic acid Glutamic acid (symbol Glu or E; the ionic form is known as glutamate) is an α-amino acid that is used by almost all living beings in the biosynthesis of proteins. It is a non-essential nutrient for humans, meaning that the human body can synt ...
,
alanine Alanine (symbol Ala or A), or α-alanine, is an α-amino acid that is used in the biosynthesis of proteins. It contains an amine group and a carboxylic acid group, both attached to the central carbon atom which also carries a methyl group side ...
, and
lysine Lysine (symbol Lys or K) is an α-amino acid that is a precursor to many proteins. It contains an α-amino group (which is in the protonated form under biological conditions), an α-carboxylic acid group (which is in the deprotonated − ...
.


Use by humans

Cephalopod ink has, as its name suggests, been used in the past as ink for pens and quills; the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
name for cuttlefish, and the taxonomic name of a cuttlefish genus, ''
Sepia Sepia may refer to: Biology * ''Sepia'' (genus), a genus of cuttlefish Color * Sepia (color), a reddish-brown color * Sepia tone, a photography technique Music * ''Sepia'', a 2001 album by Coco Mbassi * ''Sepia'' (album) by Yu Takahashi * " ...
'', is associated with the brown colour of cuttlefish ink (for more information, see
sepia Sepia may refer to: Biology * ''Sepia'' (genus), a genus of cuttlefish Color * Sepia (color), a reddish-brown color * Sepia tone, a photography technique Music * ''Sepia'', a 2001 album by Coco Mbassi * ''Sepia'' (album) by Yu Takahashi * " ...
). Modern use of cephalopod ink is generally limited to
cooking Cooking, cookery, or culinary arts is the art, science and craft of using heat to prepare food for consumption. Cooking techniques and ingredients vary widely, from grilling food over an open fire to using electric stoves, to baking in vario ...
, primarily in
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the n ...
and the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on ...
, where it is used as a
food colouring Food coloring, or color additive, is any dye, pigment, or substance that imparts color when it is added to food or drink. They come in many forms consisting of liquids, powders, gels, and pastes. Food coloring is used in both commercial food ...
and flavouring, for example in
pasta Pasta (, ; ) is a type of food typically made from an unleavened dough of wheat flour mixed with water or eggs, and formed into sheets or other shapes, then cooked by boiling or baking. Rice flour, or legumes such as beans or lentils, ...
and
sauces In cooking, a sauce is a liquid, cream, or semi- solid food, served on or used in preparing other foods. Most sauces are not normally consumed by themselves; they add flavor, moisture, and visual appeal to a dish. ''Sauce'' is a French wor ...
, and '' calamares en su tinta''. For this purpose it is generally obtainable from
fishmonger A fishmonger (historically fishwife for female practitioners) is someone who sells raw fish and seafood. Fishmongers can be wholesalers or retailers and are trained at selecting and purchasing, handling, gutting, boning, filleting, displaying, ...
s, gourmet food suppliers, and is widely available in markets in Japan and Spain. The ink is extracted from the ink sacs during preparation of the dead cephalopod, usually cuttlefish, and therefore contains no mucus. While it is not commonly used in China, cephalopod ink is sometimes used to dye the dough of
dumpling Dumpling is a broad class of dishes that consist of pieces of dough (made from a variety of starch sources), oftentimes wrapped around a filling. The dough can be based on bread, flour, buckwheat or potatoes, and may be filled with meat, ...
s. Studies have shown that cephalopod ink is toxic to some
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
s, including
tumor cells A neoplasm () is a type of abnormal and excessive growth of tissue. The process that occurs to form or produce a neoplasm is called neoplasia. The growth of a neoplasm is uncoordinated with that of the normal surrounding tissue, and persists ...
. It is being researched in mice for its antitumor activity against Meth-A fibrosarcoma. It currently remains unclear however if any of the antitumor activity of squid ink can be obtained from oral consumption, and this is indicated as an area for future investigation.


References


External links


An article on harvesting squid ink
{{Cephalopod anatomy Antipredator adaptations Ink Mollusc products Inks