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Siphuncle

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A cross-section through a Nautilus shell, showing a narrow siphuncle connecting the chambers of the shell
A cross-section through a Nautilus shell, showing a narrow siphuncle connecting the chambers of the shell

The siphuncle is a strand of tissue passing longitudinally through the shell of a cephalopod mollusk. Only cephalopods with chambered shells have siphuncles, such as the extinct ammonites and belemnites, and the living nautiluses, cuttlefish, and Spirula. In the case of the cuttlefish, the siphuncle is indistinct and connects all the small chambers of that animal's highly modified shell; in the other cephalopods it is thread-like and passes through small openings in the septa (walls) dividing the camerae (chambers). Some older studies have used the term siphon for the siphuncle, though this naming convention is uncommon in modern studies to prevent confusion with a mollusc organ of the same name.[1]

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Mollusc shell

Mollusc shell

The mollusc shell is typically a calcareous exoskeleton which encloses, supports and protects the soft parts of an animal in the phylum Mollusca, which includes snails, clams, tusk shells, and several other classes. Not all shelled molluscs live in the sea; many live on the land and in freshwater.

Cephalopod

Cephalopod

A cephalopod is any member of the molluscan class Cephalopoda such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles modified from the primitive molluscan foot. Fishers sometimes call cephalopods "inkfish", referring to their common ability to squirt ink. The study of cephalopods is a branch of malacology known as teuthology.

Nautilus

Nautilus

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.

Cuttlefish

Cuttlefish

Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of buoyancy.

Spirula

Spirula

Spirula spirula is a species of deep-water squid-like cephalopod mollusk. It is the only extant member of the genus Spirula, the family Spirulidae, and the order Spirulida. Because of the shape of its internal shell, it is commonly known as the ram's horn squid or the little post horn squid. Because the live animal has a light-emitting organ, it is also sometimes known as the tail-light squid.

Septum (cephalopod)

Septum (cephalopod)

Septa are thin walls or partitions between the internal chambers (camerae) of the shell of a cephalopod, namely nautiloids or ammonoids.

Camera (cephalopod)

Camera (cephalopod)

Camerae are the spaces or chambers enclosed between two adjacent septa in the phragmocone of a nautiloid or ammonoid cephalopod molluscus. These can be seen in cross-sections of a nautilus shell and in the polished cross-sections of ammonites. In life these chambers are filled with gas, mediated by the siphuncle, and used to control buoyancy.

Siphon (mollusc)

Siphon (mollusc)

A siphon is an anatomical structure which is part of the body of aquatic molluscs in three classes: Gastropoda, Bivalvia and Cephalopoda.

Function

Simplified structure and mechanism of cephalopodic siphuncle.
Simplified structure and mechanism of cephalopodic siphuncle.

The siphuncle is used primarily in emptying water from new chambers as the shell grows.[2] To perform this task, the cephalopod increases the saltiness of the blood in the siphuncle, and the water moves from the more dilute chamber into the blood through osmosis. At the same time gasses, mostly nitrogen, oxygen, and carbon dioxide, diffuse from the blood in the siphuncle into the emptying chamber. This is not a form of active pumping: the gas moving into the chamber is a passive process. Most energy is expended through the absorption of water from the chamber.

Removing water from the chambers of the shell reduces the overall density of the shell, and thus the shell behaves as a flotation device comparable to the swim bladder in bony fish. Typically, cephalopods maintain a density close to that of sea water, allowing them to keep a stable buoyancy with minimal effort. In the geologic past, many cephalopods grew to an enormous size (perhaps approaching ten meters in length) thanks to this.

Generally, the siphuncle is unable to provide a way to change the density of shell rapidly and thus cause the animal to rise or sink at will; rather, the animal must swim up or down as required.

Cephalopods with a wider siphuncle have a higher rate of metabolic activity.[3]

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Blood

Blood

Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the circulatory system is also known as peripheral blood, and the blood cells it carries, peripheral blood cells.

Osmosis

Osmosis

Osmosis is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential to a region of low water potential, in the direction that tends to equalize the solute concentrations on the two sides. It may also be used to describe a physical process in which any solvent moves across a selectively permeable membrane separating two solutions of different concentrations. Osmosis can be made to do work. Osmotic pressure is defined as the external pressure required to be applied so that there is no net movement of solvent across the membrane. Osmotic pressure is a colligative property, meaning that the osmotic pressure depends on the molar concentration of the solute but not on its identity.

Nitrogen

Nitrogen

Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at seventh in total abundance in the Milky Way and the Solar System. At standard temperature and pressure, two atoms of the element bond to form N2, a colorless and odorless diatomic gas. N2 forms about 78% of Earth's atmosphere, making it the most abundant uncombined element. Nitrogen occurs in all organisms, primarily in amino acids (and thus proteins), in the nucleic acids (DNA and RNA) and in the energy transfer molecule adenosine triphosphate. The human body contains about 3% nitrogen by mass, the fourth most abundant element in the body after oxygen, carbon, and hydrogen. The nitrogen cycle describes the movement of the element from the air, into the biosphere and organic compounds, then back into the atmosphere.

Oxygen

Oxygen

Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as well as with other compounds. Oxygen is Earth's most abundant element, and after hydrogen and helium, it is the third-most abundant element in the universe. At standard temperature and pressure, two atoms of the element bind to form dioxygen, a colorless and odorless diatomic gas with the formula O2. Diatomic oxygen gas currently constitutes 20.95% of the Earth's atmosphere, though this has changed considerably over long periods of time. Oxygen makes up almost half of the Earth's crust in the form of oxides.

Carbon dioxide

Carbon dioxide

Carbon dioxide (chemical formula CO2) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature, and as the source of available carbon in the carbon cycle, atmospheric CO2 is the primary carbon source for life on Earth. In the air, carbon dioxide is transparent to visible light but absorbs infrared radiation, acting as a greenhouse gas. Carbon dioxide is soluble in water and is found in groundwater, lakes, ice caps, and seawater. When carbon dioxide dissolves in water, it forms carbonate and mainly bicarbonate (HCO−3), which causes ocean acidification as atmospheric CO2 levels increase.

Density

Density

Density is the substance's mass per unit of volume. The symbol most often used for density is ρ, although the Latin letter D can also be used. Mathematically, density is defined as mass divided by volume:

Swim bladder

Swim bladder

The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ that contributes to the ability of many bony fish to control their buoyancy, and thus to stay at their current water depth without having to expend energy in swimming. Also, the dorsal position of the swim bladder means the center of mass is below the center of volume, allowing it to act as a stabilizing agent. Additionally, the swim bladder functions as a resonating chamber, to produce or receive sound.

Fish

Fish

Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of living fish species are ray-finned fish, belonging to the class Actinopterygii, with around 99% of those being teleosts.

Cephalopod size

Cephalopod size

Cephalopods, which include squids and octopuses, vary enormously in size. The smallest are only about 1 centimetre (0.39 in) long and weigh less than 1 gram (0.035 oz) at maturity, while the largest—the giant and colossal squids—can exceed 10 metres (33 ft) in length and weigh close to half a tonne (1,100 lb), making them the largest living invertebrates. Living species range in mass more than three-billion-fold, or across nine orders of magnitude, from the lightest hatchlings to the heaviest adults. Certain cephalopod species are also noted for having individual body parts of exceptional size. The giant and colossal squids, for example, have the largest known eyes among living animals.

Morphology

A shell of Nautilus pompilius in cross section. Septal necks are preserved, but the thin connecting rings have been degraded and lost.
A shell of Nautilus pompilius in cross section. Septal necks are preserved, but the thin connecting rings have been degraded and lost.

The siphuncle of fossilised cephalopods is assumed to have worked in the same general way as in living nautiluses. The siphuncle itself is only rarely preserved, but its shape can be inferred from hardened structures which lie around it. Many fossils show the holes where the siphuncle passes through each septum. Around these holes, the rim of the septum is bent into a stout aragonitic tube known as a septal neck (or siphuncle notch).[1][4]

These fossilized Orthoceras shells have fully preserved septal necks and connecting rings outlining the shape of the siphuncle
These fossilized Orthoceras shells have fully preserved septal necks and connecting rings outlining the shape of the siphuncle

In each chamber of the shell, the siphuncle is encased by a tubular structure known as a connecting ring. In living nautiluses, the connecting ring is a simple, thin-walled cylinder, with organic or thinly calcitic layers secreted from the tissues of the siphuncle. This fragile and poorly-mineralized form is known as a nautilosiphonate morphology. Many extinct cephalopods have a much more prominent connecting ring, with a very thick and porous inner calcitic layer. This more strongly-mineralized form is known as a calciosiphonate connecting ring. Connecting rings are strongly variable in morphology, from narrow homogenous tubes to bulbous, segmented cavities. Some are infolded, sending lobes or blades of calcite into the siphuncle. Connecting rings are typically continuous with the septal necks, and are difficult to distinguish without close examination. However, their developmental origin is wholly separate from the shell and septa, and they utilize calcite rather than aragonite as a biomineralized reinforcement.[1][4]

Biomineralized structures which develop within the siphuncle are known as endosiphuncular deposits (or simply siphonal deposits). These may include horizontal partitions (diaphragms), stacked conical structures (endocones), longitudinal rods, and various other concretions. Endosiphuncular deposits are typically thin structures which may be homologous to parts of the septae or connecting rings.[1][4]

In most fossil nautiluses, the siphuncle runs more or less through the center of each chamber, but in ammonites and belemnites it usually runs along the ventral edge of the shell. In some fossil straight shelled nautiloids, cylindrical calcareous growths ("siphuncular deposits") around the siphuncle can be seen towards the apex of the shell. These were apparently counterweights for the soft body at the other end of the shell, and allowed the nautilus to swim in a horizontal position. Without these deposits, the apex of the buoyant shell would have pointed upwards and the heavier body downwards, making horizontal swimming difficult. The siphuncle of the Endocerida also contained much of the organisms' body organs.[5]

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Chambered nautilus

Chambered nautilus

The chambered nautilus, also called the pearly nautilus, is the best-known species of nautilus. The shell, when cut away, reveals a lining of lustrous nacre and displays a nearly perfect equiangular spiral, although it is not a golden spiral. The shell exhibits countershading, being light on the bottom and dark on top. This is to help avoid predators, because when seen from above, it blends in with the darkness of the sea, and when seen from below, it blends in with the light coming from above.

Fossil

Fossil

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

Nautilus

Nautilus

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.

Aragonite

Aragonite

Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, CaCO3. It is formed by biological and physical processes, including precipitation from marine and freshwater environments.

Orthoceras

Orthoceras

Orthoceras is a genus of extinct nautiloid cephalopod restricted to Middle Ordovician-aged marine limestones of the Baltic States and Sweden. This genus is sometimes called Orthoceratites. Note it is sometimes misspelled as Orthocera, Orthocerus or Orthoceros.

Calcite

Calcite

Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratch hardness comparison. Large calcite crystals are used in optical equipment, and limestone composed mostly of calcite has numerous uses.

Endocerida

Endocerida

Endocerida is an extinct nautiloid order, a group of cephalopods from the Lower Paleozoic with cone-like deposits in their siphuncle. Endocerida was a diverse group of cephalopods that lived from the Early Ordovician possibly to the Late Silurian. Their shells were variable in form. Some were straight (orthoconic), others curved (cyrtoconic); some were long (longiconic), others short (breviconic). Some long-shelled forms like Endoceras attained shell lengths close to 6 metres (20 ft). The related Cameroceras is anecdotally reported to have reached lengths approaching 9 metres (30 ft), but these claims are problematic. The overwhelming majority of endocerids and nautiloids in general are much smaller, usually less than a meter long when fully grown.

Source: "Siphuncle", Wikipedia, Wikimedia Foundation, (2022, October 22nd), https://en.wikipedia.org/wiki/Siphuncle.

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References
  1. ^ a b c d Flower, Rousseau H. (1964). "Nautiloid shell morphology" (PDF). New Mexico Bureau of Mines and Mineral Resources, Memoir. 13: 1–78.
  2. ^ Mutvei, Harry; Zhang, Yun-bai; Dunca, Elena (2007). "Late Cambrian Plectronocerid Nautiloids and Their Role in Cephalopod Evolution". Palaeontology. 50 (6): 1327–1333. doi:10.1111/j.1475-4983.2007.00708.x.
  3. ^ Kröger, Björn (2003). "The size of the siphuncle in cephalopod evolution". Senckenbergiana Lethaea. 83: 39. doi:10.1007/BF03043304.
  4. ^ a b c King, Andy H.; Evans, David H. (2019). "High-level classification of the nautiloid cephalopods: a proposal for the revision of the Treatise Part K". Swiss Journal of Palaeontology. 138 (1): 65–85. doi:10.1007/s13358-019-00186-4. ISSN 1664-2384.
  5. ^ Kroger, B; Yun-Bai, Zhang (2008). "Pulsed cephalopod diversification during the Ordovician". Palaeogeography, Palaeoclimatology, Palaeoecology. 273: 174. doi:10.1016/j.palaeo.2008.12.015.

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