Chemical defence in Sea Hares - references

I have generated this list of scientific papers on aspects of chemical defence in Sea Hares and chemicals they appear to get from the algae they eat. There are probably papers I have missed. If you know of any let me know and I will add them to the list...Bill Rudman (Last updated 20 Sept 2000)

See also:
Sea Hares - attack & defence, ink glands
Sea Hares - collecting sites in Australasia
Sea Hares - mating chains
Sea Hares - Species List
Sea Hares - what are they?


    •Avila,C (1992): A preliminary catalogue of natural substances of opisthobranch molluscs from western Mediterranean and near Atlantic. Scientia Marina 56(4), 373-382.
    •Avila, C. (1995): Natural products of opisthobranch molluscs: a biological review. Oceanography and Marine Biology: an Annual Review 33, 487-559.
    •Bailey,CH; Chen,M; Keller,F; Kandel,ER (1992): Serotonin-mediated endocytosis of apCAM: an early step of learning-related synaptic growth in Aplysia. Science 256(1 May), 645-649.
    • Blankenship, J.E., Langlais, P.J. & Kittredge, J.S. (1975) Identification of a cholinomimetic compound in the digestive gland of Aplysia californica. Comparative Biochemistry and Physiology, 51C: 129-137.
    •Carefoot, T.H. (1967) Growth and nutrition of Aplysia punctata feeding on a variety of marine algae. Journal of the marine Biological Association, U.K., 47: 335-350.
    • Carefoot, T.H., S.C. Pennings, and J.P. Danko. 1999. A test of novel functions for the ink of sea hares. Journal of Marine Biology and Ecology, 234: 185-197.
    • Chapman, D.J. & Fox, D.L. (1969) Bile pigment metabolism in the sea hare Aplysia. Journal of experimental Marine Biology and Ecology, 4: 71-78.
    •Coelho, L., Prince, J. and Nolen, T. (1998). Processing of defensive pigment in Aplysia californica: Acquisition, modification and mobilization of the red algal pigment, r-phycoerythrin by the digestive gland. J. Exp. Biol. 201(3): 425-438.
    •Di Matteo,T (1982): The ink of Aplysia dactylomela (Rang, 1828) (Gastropoda: Opisthobranchia) and its role as a defensive mechanism. J. Exp. Mar. Biol. Ecol. 57, 169-180.
    •Fenical,W; Sleeper,HL; Paul,VJ; Stallard,MO; Sun,HH (1979): Defensive chemistry of Navanax and related opisthobranch molluscs. Pure and Applied Chemistry 51, 1865-1874.
    •Fiorito,G; Bergamo,P; Capuano,C; Lepore,C (1985): Considerazioni sul comportamento di emissione di inchiostro nel genre Aplysia (Mollusca, Gastropoda, Opisthobranchia). Boll Soc Natur Napoli 94, 25-42.
    • Hino K., Mitsui Y. & Hirano Y. (1994) Four cases of acute liver damage following the ingestion of a sea hare egg. Journal of Gastroenterology, 29: 679.
    •Imperato,F; Minale,L; Riccio,R (1977): Constituents of the digestive gland of molluscs of the genus Aplysia. Halogenated monoterpenes from Aplysia limacina. Exper. 33, 1273-1274.
    •Ireland,C; Faulkner,DJ (1977): Diterpenes from Dolabella californica. Scripps Institute Oceanography 47(1-2), 1177-1182.
    •Johnson, P.M & Willows, A.O.D, (1999) Defense in Sea Hares (Gastropoda, Opisthobranchia, Anaspidea): multiple layers of protection from egg to adult. Marine & Freshwater Behaviour & Physiology, 32: 147-180.
    •Kamiya,H; Muramoto,K; Yamazaki,M (1986): Aplysianin-A, an antibacterial and antineoplastic glycoprotein in the albumen gland of a sea hare, Aplysia kurodai. Exper. 42, 1065-1067.
    •Kamiya,H; Muramoto,K; Goto,R; Sakai,M; Endo,Y; Yamazaki,M (1989): Purification and characterization of an antibacterial and antineoplastic protein secretion of a sea hare, Aplysia juliana. Toxicon 27(12), 1269-1277.
    •Kandel,ER (1979): Behavioural Biology of Aplysia. San Francisco, W.H.Freeman&Co.463pp.
    •Kandel,ER (1976): Cellular Basis of Behaviour. An introduction to behavioural neurobiology. W.H.Freeman & Co., San Francisco. 727 pages.
    • MacColl, R., Galivan, J., Berns, D.S., Nimec, Z., Guard-Friar, D. & Wagoner, D. (1990)The chromophore and polypeptide composition of Aplysia ink. Biological Bulletin, 179: 326-331.
    •Martinez,E; Cimino,G; Crispino,A; Ortea,JA; Spinella,A (1992): A biochemical comparative study of Anaspidea molluscs from distinct geographical areas. Abstracts, 11th International Malacological Congress, Siena 1992, 283-284.
    •Mayford,M; Barzilai,A; Kellar,F; Schacher,S; Kandel,ER (1992): Modulation of an NCAM-related adhesion molecule with longterm synaptic plasticity in Aplysia. Science 256, 638-644.
    •Nolen, T.G.; Johnson, P.M.; Kicklighter, C.E.; Capo, T. (1995): Ink secretion by the marine snail Aplysia californica enhances its ability to escape from a natural predator. J. Comp. Physiol. A 176, 239-254.
    • Nolen, T.G. & Johnson, P.M. (2001) Defensive inking in Aplysia spp: Multiple episodes of ink secretion and the adaptive use of a limited chemical resource The Journal of Experimental Biology, 204: 1257–1268
    •Paul,VJ; Pennings,SC (1991): Diet-derived chemical defenses in the sea hare Stylocheilus longicauda (Quoy & Gaimard, 1824). J. Exp. Mar. Biol. Ecol. 151, 227-243.
    •Pennings,SC (1990): Multiple factors promoting narrow host range in the sea hare, Aplysia californica. Oecologia 82, 192-200.
    •Pennings,SC (1990): Predator-prey interactions in opisthobranch gastropods: effects of prey body size and habitat complexity. Mar. Ecol. Prog. Ser. 62, 95-101.
    •Pennings,SC; Paul,VJ (1993): Sequestration of dietary secondary meatabolites by three species of sea hares: location, specificity and dynamics. Marine Biology 117, 535-546.
    •Pennings,SC (1994): Interspecific variation in chemical defenses in the sea hares (Opisthobranchia: Anaspidea). J. Exp. Mar. Biol. Ecol. 180, 203-219.
    •Pennings,SC; Paul,VJ (1992): Effect of plant toughness, calcification, and chemistry on herbivory by Dolabella auricularia-. Ecol. 73(5), 1606-1619.
    •Pennings,SC; Weiss,AM; Paul,VJ (1996): Secondary metabolites of the cyanobacterium Microcoleus lynbyaceus and the sea hare Stylocheilus longicauda: palatability and toxicity. Marine Biology 126, 735-743.
    •Prince, J., Nolen, T. and Coelho, L. (1998). Defensive ink pigment processing and secretion in Aplysia californica: Concentration and storage of phycoerythrobilin in the ink gland. J. Exp. Biol. 201(10): 1595-1613.
    •Rogers,CN; Steinberg,PD; Nys,R de (1995): Factors associated with oligophagy in two species of sea hares (Mollusca: Anaspidea). J. Exp. Mar. Biol. Ecol. 192, 47-73.
    • Sorokin M. (1988) Human poisoning by ingestion of a sea hare. Toxicon, 26: 1095-1097.
    •Stallard,MO; Faulkner,DJ (1974): Chemical constituents of the digestive gland of the sea hare Aplysia californica - I. Importance of diet. Comp. Biochem. Physiol. 49 B, 25-35.
    •Stallard,MO; Fenical,W; Kittredge,JS (1978): The brasilenols, rearranged sesquiterpene alcohols isolated from the marine opisthobranch Aplysia brasiliana. Scripps Institute Oceanography 48(1), 2077-2081.
    •Tobach,E; Zaferes,A; Migenis-Lopez,L (1989): Aplysia ink and opaline: exploration of their relation to predation. Bull. Mar. Sci. 45(3), 664-670.
    •Trowbridge,CD (1994): Defensive Responses and palatability of specialist herbivores: predation on NE Pacific ascoglossan gastropods. Mar. Ecol. Prog. Ser. 105, 61-70.

Authorship details
Rudman, W.B., 1998 (October 14) Chemical defence in Sea Hares - references. [In] Sea Slug Forum. Australian Museum, Sydney. Available from

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