title: Analysis of the behaviours mediating barnacle cyprid reversible adhesion.
abstract: When exploring immersed surfaces the cypris larvae of barnacles employ a tenacious and rapidly reversible adhesion mechanism to facilitate their characteristic 'walking' behaviour. Although of direct relevance to the fields of marine biofouling and bio-inspired adhesive development, the mechanism of temporary adhesion in cyprids remains poorly understood. Cyprids secrete deposits of a proteinaceous substance during surface attachment and these are often visible as 'footprints' on previously explored surfaces. The attachment structures, the antennular discs, of cyprids also present a complex morphology reminiscent of both the hairy appendages used by some terrestrial invertebrates for temporary adhesion and a classic 'suction cup'. Despite the numerous analytical approaches so-far employed, it has not been possible to resolve conclusively the respective contributions of viscoelastic adhesion via the proteinaceous 'temporary adhesive', 'dry' adhesion via the cuticular villi present on the disc and the behavioural contribution by the organism. In this study, high-speed photography was used for the first time to capture the behaviour of cyprids at the instant of temporary attachment and detachment. Attachment is facilitated by a constantly sticky disc surface - presumably due to the presence of the proteinaceous temporary adhesive. The tenacity of the resulting bond, however, is mediated behaviourally. For weak attachment the disc is constantly moved on the surface, whereas for a strong attachment the disc is spread out on the surface. Voluntary detachment is by force, requiring twisting or peeling of the bond - seemingly without any more subtle detachment behaviours. Micro-bubbles were observed at the adhesive interface as the cyprid detached, possibly an adaptation for energy dissipation. These observations will allow future work to focus more specifically on the cyprid temporary adhesive proteins, which appear to be fundamental to adhesion, inherently sticky and exquisitely adapted for reversible adhesion underwater.
authors.affiliation: School of Marine Science and Technology, Newcastle University, Newcastle upon Tyne, United Kingdom.
authors.forename: Nick
authors.initials: N
authors.lastname: Aldred
authors.forename: Jens T
authors.initials: JT
authors.lastname: Høeg
authors.forename: Diego
authors.initials: D
authors.lastname: Maruzzo
authors.forename: Anthony S
authors.initials: AS
authors.lastname: Clare
mesh_terms: D000818:Animals; D001522:Behavior, Animal / Q000502:physiology; D057219:Biofouling; D002448:Cell Adhesion / Q000502:physiology; D007814:Larva / Q000502:physiology; D009068:Movement / Q000502:physiology; D010781:Photography / Q000379:methods; D001468:Thoracica / Q000502:physiology*