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

Related messages

Sea Hares - attack and defence

April 21, 1998
From: Bill Rudman

Following the recent flood of queries on Sea Hares I have generated the list, which appears at the top of this page, 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