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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.
168:
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.
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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.
176:
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.
167:
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
199:
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
171:
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%
208:
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,
175:
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
233:
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.
870:
Lo Presti, Marco; Ostrovsky-Snider, Nicholas; Rizzo, Giorgio; Portoghese, Marina; Blasi, Davide; Farinola, Gianluca M.; Omenetto, Fiorenzo G. (29 August 2023).
913:"Prospective randomized study evaluating a biodegradable polymeric sealant for sealing intraoperative air leaks that occur during pulmonary resection"
403:
348:
1184:
715:
Robert L. Strausberg; et al. (31 December 1989). "Development of a
Microbial System for Production of Mussel Adhesive Protein".
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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
89:
that function to attach the mollusc to a solid surface. Species from several families of clams have a byssus, including pen shells (
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748:
Simmons, Miriam; Horbelt, Nils; Sverko, Tara; Scoppola, Ernesto; Jackson, Daniel J.; Harrington, Matthew J. (28 November 2023).
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Applications of biomimetic byssus adhesive include biomedical adhesives, therapeutic applications, and anti-fouling coatings.
732:
959:"Polydopamine-enabled surface functionalization of gold nanorods for cancer cell-targeted imaging and photothermal therapy"
872:"The role of tyrosine in protein-dopamine based bioinspired adhesives: the stoichiometry that maximizes bonding strength"
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either by producing Mfps via other organisms or by creating synthetic polymers with similar properties. For instance,
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750:"Invasive mussels fashion silk-like byssus via mechanical processing of massive horizontally acquired coiled coils"
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Through the Jade Gate to Rome: A Study of the Silk Routes during the Later Han
Dynasty, 1st to 2nd centuries CE
1177:
1116:
517:"Tensile and dynamic mechanical analysis of the distal portion of mussel (Mytilus edulis) byssal threads"
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1052:
McKinley, Daniel (June 1998). "Pinna and Her Silken Beard: A Foray Into
Historical Misappropriations".
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912:
1980:
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cells to translate the genes into the appropriate proteins. Synthetic approaches generally utilize
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species can be up to 6 cm (2.4 in) in length and have historically been made into cloth.
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Lo Presti, Marco; Rizzo, Giorgio; Farinola, Gianluca M.; Omenetto, Fiorenzo G. (August 2021).
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Maeder, Felicitas (2002). "The project Sea-silk: Rediscovering an
Ancient Textile Material".
449:
280:
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Turner, Ruth; Rosewater, Joseph (June 1958). "The Family
Pinnidae in the Western Atlantic".
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1008:"Protein Resistance of Titanium Oxide Surfaces Modified by Biologically Inspired mPEG−DOPA"
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666:"Recent approaches in designing bioadhesive materials inspired by mussel adhesive protein"
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124:, the inedible byssus is commonly known as the "beard", and is removed before cooking.
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Many species of mussels secrete byssus threads to anchor themselves to surfaces, with
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815:"Bioinspired Biomaterial Composite for All-Water-Based High-Performance Adhesives"
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404:"Mechanical design of mussel byssus: material yield enhances attachment strength"
269:, that is made using the byssus of pen shells as the fiber source. The byssus of
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agent to produce wear-resistant polymer networks. Imitation of Mfp-3 to induce
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Byssus often refers to the long, fine, silky threads secreted by the large
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450:"Interspecific Comparison of the Mechanical Properties of Mussel Byssus"
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Byssus filaments are created by certain kinds of marine and freshwater
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719:. ACS Symposium Series. Vol. 385. pp. 453–464.
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82:) is a bundle of filaments secreted by many species of
349:"Seasonal variation in mussel byssal thread mechanics"
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Moeser, Gretchen M.; Carrington, Emily (15 May 2006).
116:, which use the byssus to attach themselves to rocks,
74:
68:
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Journal of
Polymer Science Part A: Polymer Chemistry
62:
448:Brazee, Shanna; Carrington, Emily (December 2006).
59:
397:
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664:Forooshani, Pegah; Lee, Bruce (11 October 2016).
183:unclogging a drain. The byssus, which is made of
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1054:Ars Textrina: A Journal of Textiles and Costumes
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754:Proceedings of the National Academy of Sciences
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275:, a shell of the same family, has been used in
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1094:(2nd ed.). Charleston, SC: Book Surge.
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621:Journal of Coatings Technology and Research
402:Bell, Emily; Gosline, John (1 April 1996).
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1126:. Geneva: Patek Philippe. pp. 35–39
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329:Biology: The Unity and Diversity of Life
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615:Verma, Shatakshi (20 February 2019).
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327:Starr, Cecie; Taggart, Ralph (2004).
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1006:Dalsin, Jeffrey (9 December 2004).
564:Peyer, Suzanne (23 December 2008).
31:), attached to a rock by its byssus
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1073:Archaeological Textiles Newsletter
717:Adhesives from Renewable Resources
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876:Frontiers in Biomaterials Science
515:Aldred, Nick (22 December 2007).
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331:. Belmont, CA: Thomson Learning.
265:is a rare fabric, also known as
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573:Journal of Experimental Biology
408:Journal of Experimental Biology
353:Journal of Experimental Biology
255:. The byssus threads from this
16:Fibre secreted by some molluscs
957:Black, Kvar (14 August 2012).
917:The Annals of Thoracic Surgery
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39:Illustration of the byssus of
1:
1153:The dictionary definition of
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45:, the freshwater zebra mussel
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7:
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1997:
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1117:"From the Soul of the Sea"
1115:Cubello, Stefania (2018).
889:10.3389/fbiom.2023.1184088
725:10.1021/bk-1989-0385.ch032
633:10.1007/s11998-018-00174-2
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120:, or seabeds. In edible
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454:The Biological Bulletin
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533:10.1098/rsif.2007.1026
420:10.1242/jeb.199.4.1005
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281:critically endangered
213:have inserted mussel
193:polyphenolic proteins
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279:as a substitute for
42:Dreissena polymorpha
1090:Hill, John (2009).
766:2023PNAS..12011901S
760:(48): e2311901120.
683:10.1002/pola.28368
586:10.1242/jeb.028688
197:injection moulding
191:-tanned proteins (
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579:(13): 2027–2036.
527:(17): 1159–1167.
366:10.1242/jeb.02234
359:(10): 1996–2003.
211:genetic engineers
93:), true mussels (
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285:Pinna nobilis
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252:Pinna nobilis
248:
247:Mediterranean
238:
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227:cross-linking
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1891:Polyethylene
1595:
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1256:Periostracum
1246:Pallial line
1155:
1128:. Retrieved
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969:(1): 17–28.
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963:Nanomedicine
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153:Dreissenidae
126:
108:
99:Dreissenidae
50:
48:
40:
26:
18:
1722:Regenerated
1666:Spider silk
1394:Trochophore
1389:Pseudofeces
1221:Hinge teeth
676:(1): 9–33.
249:pen shell,
204:Biomimetics
1970:Categories
1871:Modacrylic
1866:Microfiber
1783:Triacetate
1735:Milk fiber
1601:Camel hair
1533:Lotus silk
1384:Glochidium
1358:Nephridium
1331:Soft parts
1216:Hinge line
292:References
149:Pectinidae
118:substrates
1886:Polyester
1758:Diacetate
1713:Synthetic
1616:Chiengora
1343:Ctenidium
1276:Sculpture
1266:Resilifer
1130:17 August
898:2813-3749
839:2198-3844
782:0027-8424
641:139442176
307:Johnsonia
163:Mechanics
157:Unionidae
141:Anomiidae
133:Mytilidae
105:Filaments
95:Mytilidae
1945:Category
1854:Technora
1816:Metallic
1699:Asbestos
1641:Pashmina
1606:Cashmere
1379:Abductin
1322:Sea silk
1271:Resilium
1226:Ligament
1060:: 9–223.
1032:15641834
1012:Langmuir
993:22891865
937:15111188
857:34080324
800:37983489
791:10691215
702:27917020
595:19525429
551:17439859
490:24797335
482:17179385
375:16651564
277:Sardinia
267:sea silk
223:catechol
145:Pinnidae
129:families
114:mollusks
91:Pinnidae
1955:Commons
1913:Vinylon
1908:Vectran
1903:Spandex
1834:Acrylic
1826:Polymer
1793:Mineral
1773:Piñatex
1763:Lyocell
1753:Acetate
1691:Mineral
1621:Guanaco
1471:Bagasse
1449:Natural
1399:Veliger
1196:anatomy
1194:Bivalve
1079:: 8–11.
984:3544340
848:8373158
762:Bibcode
693:5132118
542:2396211
474:4134548
428:9318809
189:quinone
185:keratin
181:plunger
137:Arcidae
122:mussels
111:bivalve
97:), and
87:mollusc
84:bivalve
28:Mytilus
1918:Vinyon
1896:UHMWPE
1881:Olefin
1849:Kevlar
1844:Twaron
1839:Aramid
1811:Basalt
1806:Carbon
1676:Vicuña
1661:Tendon
1651:Rabbit
1646:Qiviut
1636:Mohair
1611:Catgut
1596:Byssus
1591:Angora
1586:Alpaca
1578:Animal
1558:Rattan
1548:Raffia
1491:Cotton
1476:Bamboo
1441:Fibers
1363:Siphon
1353:Mantle
1312:Byssus
1296:Annuli
1236:Lunule
1211:Callus
1156:byssus
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155:, and
51:byssus
1923:Zylon
1876:Nylon
1859:Nomex
1801:Glass
1778:Rayon
1768:Modal
1631:Llama
1563:Sisal
1553:Ramie
1528:Kenaf
1523:Kapok
1506:Linen
1496:Fique
1481:Bashō
1466:Abacá
1458:Plant
1372:Other
1317:Pearl
1286:Valve
1281:Umbo
1241:Nacre
1203:Shell
1120:(PDF)
1037:9 May
942:9 May
646:9 May
637:S2CID
600:9 May
569:(PDF)
495:9 May
486:S2CID
470:JSTOR
433:9 May
380:8 May
258:Pinna
225:as a
219:yeast
217:into
1671:Wool
1656:Silk
1626:Hair
1568:Wood
1543:Pine
1538:Piña
1518:Jute
1513:Hemp
1501:Flax
1486:Coir
1291:Beak
1231:Lira
1132:2024
1096:ISBN
1039:2021
1028:PMID
989:PMID
944:2021
933:PMID
894:ISSN
853:PMID
835:ISSN
796:PMID
778:ISSN
729:ISBN
698:PMID
648:2021
602:2021
591:PMID
547:PMID
497:2021
478:PMID
435:2021
424:PMID
382:2021
371:PMID
1681:Yak
1020:doi
979:PMC
971:doi
925:doi
884:doi
843:PMC
827:doi
786:PMC
770:doi
758:120
721:doi
688:PMC
678:doi
629:doi
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537:PMC
529:doi
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