
A byssus () is a bundle of filaments secreted by many species of
bivalve
Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
mollusc
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
that function to attach the mollusc to a solid surface. Species from several families of clams have a byssus, including pen shells (
Pinnidae), true mussels (
Mytilidae
The Mytilidae are a family (biology), family of small to large Marine life, marine and Brackish water, brackish-water bivalve molluscs in the order (biology), order Mytilida. One of the genera, ''Limnoperna fortunei, Limnoperna'', even inhabits f ...
), and
Dreissenidae.
Filaments
Byssus filaments are created by certain kinds of marine and freshwater
bivalve
Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
mollusk
Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The ...
s, which use the byssus to attach themselves to rocks,
substrates, or seabeds. In edible
mussel
Mussel () is the common name used for members of several families of bivalve molluscs, from saltwater and Freshwater bivalve, freshwater habitats. These groups have in common a shell whose outline is elongated and asymmetrical compared with other ...
s, the inedible byssus is commonly known as the "beard", and is removed before cooking.
Many species of mussels secrete byssus threads to anchor themselves to surfaces, with
families
Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
including the
Mytilidae
The Mytilidae are a family (biology), family of small to large Marine life, marine and Brackish water, brackish-water bivalve molluscs in the order (biology), order Mytilida. One of the genera, ''Limnoperna fortunei, Limnoperna'', even inhabits f ...
,
Arcidae,
Anomiidae
Anomiidae is a family of saltwater clams, marine bivalve molluscs related to scallops and oysters, and known as anomiids. It contains seven genera.
The family is known by several common names, including jingle shells, mermaid's toenails, and ...
,
Pinnidae,
Pectinidae
Scallop () is a common name that encompasses various species of Marine (ocean), marine bivalve mollusc, molluscs in the Taxonomy (biology), taxonomic Family (biology), family Pectinidae, the scallops. However, the common name "scallop" is also ...
,
Dreissenidae, and
Unionidae
The Unionidae are a Family (biology), family of freshwater mussels, the largest in the order Unionida, the bivalve molluscs sometimes known as river mussels, or simply as unionids.
The range of distribution for this family is world-wide. It is a ...
.
Mechanics
The byssus, or byssal complex, is composed of multiple extracellular collagenous threads that are placed radially by the mussel from a central stem. Each thread is composed of three regions: a corrugated proximal region close to the mussel body, a longer, smooth distal region connecting the proximal region to the ending plaque, and the adhesive plaque itself, which anchors the mussel to the surface.
The proximal region consists of a corrugated sheath enveloping loosely-arranged coiled fibers; these coils can unravel to extend the fiber under an applied force. The distal region is more ordered, consisting of aligned collagenous fiber bundles that give the fiber stiffness. The plaque consists of collagen-like fibers over a spongy matrix, in which the adhesive protein is deposited and hardens.
The purpose of the byssus is to keep the mussel attached to the desired surface, and to this end byssal threads must be able to withstand strong cyclic motion due to tidal action near the shorelines mussels inhabit. Mechanical testing of live mussels has shown that byssal threads can extend 39% before yield and 64% before breaking, at a nominal strain rate of 10 mm/min.
Tensile testing shows that threads exhibit three distinct phases: initial stiffness from both the distal and proximal regions, softening due to yield in the distal region, and finally stiffening directly preceding tensile failure.
The ability of the distal region to yield before breaking gives the mussels their characteristic hardiness even under strong tidal forces.
Many variables that influence the performance of byssal threads have been studied, including species variations,
seasonal variations,
temperature effects,
and ageing effects.
Temperature effects in particular have revealed a glass transition temperature of 6 °C.
The number of threads used by a mussel to attach is typically between 20 and 60; this can vary by the species, season, or age of the mussel. Under cyclic tidal conditions, the radial spread of fiber placement allows the mussel to dynamically align most of its fibers in the direction of applied force. This lowers the stress on any one thread, reducing the chances of failure and detachment.
Mussels are also capable of ejecting the entire byssal complex, including the central stem, without damaging themselves. The complex can simply be regenerated and with fibers placement resuming within 24 hours.
When a mussel's foot encounters a crevice, it creates a vacuum chamber by forcing out the air and arching up, similar to a plumber's
plunger
A plunger is a device driven by or against fluid pressure. In plumbing, the term plunger commonly refers to handheld tools used to clear blockages in drains and pipes. Plumbing plungers consist of a rubber suction cup attached to a stick (''sh ...
unclogging a drain. The byssus, which is made of
keratin
Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
,
quinone
The quinones are a class of organic compounds that are formally "derived from aromatic compounds benzene.html" ;"title="uch as benzene">uch as benzene or naphthalene] by conversion of an even number of –CH= groups into –C(=O)– groups with ...
-tanned proteins (
polyphenolic proteins), and other proteins, is spewed into this chamber in a liquid form similar to
injection moulding
Injection moulding (U.S. spelling: injection molding) is a manufacturing process for producing parts by injecting molten material into a mould, or mold. Injection moulding can be performed with a host of materials mainly including metals (for ...
in polymer processing, and bubbles into a sticky foam. By curling its foot into a tube and pumping the foam, the mussel produces sticky threads about the size of a human hair. The mussel then varnishes the threads with another protein, resulting in an adhesive.
The attachment dynamics of the plaque are studied both to imitate the strong adhesive and to create coatings to which the plaque cannot adhere. Foul release strategies such as fluoropolymer paints and lubricant-infused coatings are an active research area important to preventing the fouling of marine structures by invasive mussel species such as the zebra and quagga mussel.
Biomimetics
Byssus is a remarkable adhesive, one that is neither degraded nor deformed by water as many synthetic adhesives are.
The remarkable properties of this adhesive, specifically the mussel foot proteins (Mfps), has spurred many attempts to imitate the excellent adhesive capacity that mussels show, either by producing Mfps via other organisms or by creating synthetic polymers with similar properties. For instance,
genetic engineers have inserted mussel
DNA
Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
into
yeast
Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom (biology), kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are est ...
cells to translate the genes into the appropriate proteins. Synthetic approaches generally utilize
catechol
Catechol ( or ), also known as pyrocatechol or 1,2-dihydroxybenzene, is an organic compound with the molecular formula . It is the ''ortho'' isomer of the three isomeric benzenediols. This colorless compound occurs naturally in trace amounts. It ...
as a
cross-link
In chemistry and biology, a cross-link is a bond or a short sequence of bonds that links one polymer chain to another. These links may take the form of covalent bonds or ionic bonds and the polymers can be either synthetic polymers or natural ...
ing agent to produce wear-resistant polymer networks. Imitation of Mfp-3 to induce
coacervation is another key property, as this protects the material from partial dissolution in saltwater.
The protein structure of byssus is reminiscent of that of silk produced by insects. Other examples of biomimetic approaches for creating mussel-inspired adhesives use these polymers as a backbone.
Applications of biomimetic byssus adhesive include biomedical adhesives,
therapeutic applications,
and anti-fouling coatings.
Historical uses
"Byssus" often refers to the long, fine, silky threads secreted by the large
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 east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
pen shell, ''
Pinna nobilis''. The byssus threads from this ''
Pinna'' species can be up to in length and have historically been made into cloth.
Byssus cloth is a rare fabric, also known as
sea silk
Sea silk is an extremely fine, rare, and valuable fabric that is made from the long silky filaments or byssus secreted by a gland in the foot of pen shells (in particular '' Pinna nobilis''). The byssus is used by the mussel to attach itself ...
, that is made using the byssus of pen shells as the fiber source.
The byssus of ''
Atrina pectinata'', a shell of the same family, has been used in
Sardinia
Sardinia ( ; ; ) is the Mediterranean islands#By area, second-largest island in the Mediterranean Sea, after Sicily, and one of the Regions of Italy, twenty regions of Italy. It is located west of the Italian Peninsula, north of Tunisia an ...
as a substitute for
critically endangered
An IUCN Red List critically endangered (CR or sometimes CE) species is one that has been categorized by the International Union for Conservation of Nature as facing an extremely high risk of extinction in the wild. As of December 2023, of t ...
''
Pinna nobilis'', to weave sea silk.
''Byssus'' was used by
Carl Linnaeus
Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
as a genus of plants (some later known to be cyanobacteria). It has little relevance in current taxonomy, as most specimens are either lost or not identified. The ones that are identified have been either syonymized (''B. jolithus'', ''B. aurea'') or turned into ''nomen rejiciendum'' (''B. cryptarum'').
References
External links
*
{{Fibers
Bivalve anatomy
Mollusc products