Underwater adhesion: The barnacle way
Introduction
On immersion of a surface in the marine environment the fouling process is initiated instantly. Biofouling is one of the most serious problems and costs the US Navy an estimated $1 billion per annum [1]. In general, the first stage is conditioning of the surface by the adsorption of organic and inorganic compounds. Conditioned surfaces are then colonized by various microorganisms such as bacteria, diatoms, etc. [2], [3] followed by attachment of algal spores and invertebrate larvae [4]. These biofilms play an important role in mediating settlement and metamorphosis of invertebrate larvae [5], [6], [7], [8], [9], [10], [11], [12], [13], [14], [15], [16], [17], [18], [19]. Chemical cues such as exopolymers and other excreted products produced by microorganisms have been shown to be involved in settlement of macrofoulers, metamorphosis induction, growth and development of organisms [15], [16], [20], [21], [22], [23], [24], [25].
Section snippets
Attachment of barnacles
Among the macrofoulers, barnacles are the dominant fouling organisms found throughout the world. A lot of research on barnacles has been carried out with respect to its settlement and ecology [11], [12], [13], [14], [15], [16], [17], [20], [23], [26], [27], [28].
The larval development of these organisms includes six naupliar instars and a non-feeding presettling cyprid instar. The anatomy of the cyprid is different from the preceding naupliar instars (Fig. 1). The cypris larvae explore the
Process of secretion
Barnacle cement is recognized as the most durable and toughest connection in the living aquatic world [41]. The cement is secreted by a pair of ovoid or kidney shaped glands (Fig. 3), 60–90 μm thick and 150 μm long which are located behind the compound eyes of cyprids [31], [42]. If the glands are filled with secretion, the volume of this material would be 471 μm3 per gland. Each antennule has 20 glands [32], [33], [34]. Walker examined these glandular structures within the body of larva before
Properties of barnacle adhesive
The resistance to chemical breakdown by barnacle adhesive caused a major problem in its characterization. However, many investigators have been successful in partially characterizing the barnacle adhesive model proteins of Balanus hameri [52], Balanus nubilus [47], Balanus crenatus [52], [53] and B. balanoides [54] which has been synthesized by polycondensation [55]. The alkaline phosphatase activity in the cementing apparatus was found by histoenzymology [56]. Proteins, phenols and polyphenol
Adhesive strength
In addition to amino acid composition and molecular weight, the natural barnacle adhesive system operates on different physical factors of the surface. Adhesion is not instantaneous and appears to take a long time. Factors such as increasing adhesive area and enzymic actions which oxidize and insolubilize the secreted water-soluble proteins from the glands, must also be considered [55]. The influence of enzymatic actions was considered to prevent adhesion and thus biofouling. It was
Application of barnacle adhesive
Marine proteins secreted by the invertebrates such as mussels and barnacles have potential application as powerful adhesives as they insolubilize and adhere to variety of substrates in a aqueous environment. They provide an example of macroadhesion as they anchor themselves to any solid substrata in marine environment in exchange for habitat advantages. Barnacle cement as well as mussel and clam byssus, all of which are 99% protein [76] resist enzymatic as well as chemical degradation at
Acknowledgments
We thank Dr. S.R. Shetye, Director National Institute of Oceanography, Goa, for the facilities and encouragement. We thank US Naval Institute for permission to reproduce Figs. 14 and 17 from the article “The attachment of macrofouling invertebrates” by Elek Lindner from the book Marine Biodeterioration: An interdisciplinary study, in this review article. We thank The Royal Society of London’ for granting permission to reproduce the illustration from the article “Structure and function in
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