Kleptoplasty

Kleptoplasty in Elysia chlorotica. The slug on the left sucks the sap from its food plant, Vaucheria, shcematized at top. Chloroplasts (red circles) are taken into the slug's cells at bottom, enabling the animal's cells to perform photosynthesis. From Rumpho et al., 2011, J. Exp. Biol. 214: 303.

Kleptoplasty in Elysia chlorotica. The slug on the left sucks the sap from its food plant, Vaucheria litorea., Chloroplasts (red circles) are taken from the algal cells at top into the slug’s cells at bottom, enabling the animal’s cells to perform photosynthesis. From Rumpho et al., 2011, J. Exp. Biol. 214: 303.

Based on the number of papers below and the number of articles that have appeared in the popular press, this aspect of Elysia biology gets the most attention.  When Elysia species feed on their host algae, they can retain the chloropasts, which are referred to as “kleptoplasts” after having been stolen from the plants.  Depending on the species of Elysia, these kelptoplasts can survive for many months in digestive diverticula that extend throughout the animals’ bodies.  Although the animals must still eat to survive, the kleptoplasts continue to perform photosynthesis in the animals’ bodies.

Scientific Literature

Barber K, Middlebrooks M, Bell S, Pierce S. (2021) The Specialist Marine Herbivore Elysia papillosa Grows Faster on a Less Utilized Algal Diet. Biol Bull. 241:158-167. [F] [K] [NH] [Pa]

Cartaxana P, Rey F, LeKieffre C, Lopes D, Hubas C, Spangenberg JE, Escrig S, Jesus B, Calado G, Domingues R, Kühl M, Calado R, Meibom A, Cruz S.(2021) Photosynthesis from stolen chloroplasts can support sea slug reproductive fitness Proc Biol Sci. 288: 20211779. [K] [Ti]

Frankenbach, S., Luppa, Q., Serôdio, J., Greve, C., Bleidissel, S., Melo Clavijo, J.  Laetz, E.M.J., Preisfeld, A., Christa, G. (2021) Kleptoplasts are continuously digested during feeding in the plastid-bearing sea slug Elysia viridis [K] [Vi]

Havurinne V, Handrich M, Antinluoma M, Khorobrykh S, Gould SB, Tyystjärvi E. (2021) Genetic autonomy and low singlet oxygen yield support kleptoplast functionality in photosynthetic sea slugs J Exp Bot. 72: 5553-5568. [K] [Ch]

Maeda T, Takahashi S, Yoshida T, Shimamura S, Takaki Y, Nagai Y, Toyoda A, Suzuki Y, Arimoto A, Ishii H, Satoh N, Nishiyama T, Hasebe M, Maruyama T, Minagawa J, Obokata J, Shigenobu S. (2021) Chloroplast acquisition without the gene transfer in kleptoplastic sea slugs, Plakobranchus ocellatus. Elife. 10: e60176.  [K] [Pl]

Cartaxana P, Cruz S. (2020) On the art of stealing chloroplasts. Elife 9: e64057. [K] [R]

Cruz S, LeKieffre C, Cartaxana P, Hubas C, Thiney N, Jakobsen S, Escrig S, Jesus B, Kühl M, Calado R, Meibom A. (2020) Functional kleptoplasts intermediate incorporation of carbon and nitrogen in cells of the Sacoglossa sea slug Elysia viridis Sci Rep. 10: 10548. [K] [Vi]

Donohoo, S.A., Wade, R.M., Sherwood, A.R.  (2020) Finding the Sweet Spot: Sub-Ambient Light Increases Fitness and Kleptoplast Survival in the Sea Slug Plakobranchus cf. ianthobaptus Gould, 1852. Biol. Bull 238: 154-166. [K] [Pl]

Havurinne V, Tyystjärvi E. (2020) Photosynthetic sea slugs induce protective changes to the light reactions of the chloroplasts they steal from algae. Elife 9:e57389. [K] [Ti]

Melo Clavijo J, Frankenbach S, Fidalgo C, Serôdio J, Donath A, Preisfeld A, Christa G. (2020)  Identification of scavenger receptors and thrombospondin-type-1 repeat proteins potentially relevant for plastid recognition in Sacoglossa. Ecol Evol. 10: 12348-12363. [K] [Co]

Rey F, Melo T, Cartaxana P, Calado R, Domingues P, Cruz S, Domingues MRM. (2020) Coping with Starvation: Contrasting Lipidomic Dynamics in the Cells of Two Sacoglossan Sea Slugs Incorporating Stolen Plastids from the Same Macroalga. Integr Comp Biol. 60: 43-56. [F] [K] [Vi]

Shiroyama H, Mitoh S, Ida TY, Yusa Y. (2020) Adaptive significance of light and food for a kleptoplastic sea slug: implications for photosynthesis. Oecologia 194: 455-463. [F] [K] [At]

Cai H, Li Q, Fang X, Li J, Curtis NE, Altenburger A, Shibata T, Feng M, Maeda T, Schwartz JA, Shigenobu S, Lundholm N, Nishiyama T, Yang H, Hasebe M, Li S, Pierce SK, Wang J. (2019) A draft genome assembly of the solar-powered sea slug Elysia chlorotica. Sci Data. 6: 190022. [HT] [K] [Ch]

Cartaxana P, Morelli L, Jesus B, Calado G, Calado R, Cruz S. (2019) The photon menace: kleptoplast protection in the photosynthetic sea slug Elysia timida. J Exp Biol. 222: 202580. [K] [Ti]

Middlebrooks ML, Curtis NE, Pierce SK. (2019) Algal Sources of Sequestered Chloroplasts in the Sacoglossan Sea Slug Elysia crispata Vary by Location and Ecotype. Biol Bull. 236: 88-96. [F] [K] [Cr]

Zan J, Li Z, Tianero MD, Davis J, Hill RT, Donia MS. (2019) A microbial factory for defensive kahalalides in a tripartite marine symbiosis. Science. 364: eaaw6732. [NP] [Ru]

Cartaxana P, Morelli L, Quintaneiro C, Calado G, Calado R, Cruz S. (2018) Kleptoplast photoacclimation state modulates the photobehaviour of the solar-powered sea slug Elysia viridis. J Exp Biol. 221:  jeb180463. [K] [NB] [Vi]

Chan CX, Vaysberg P, Price DC, Pelletreau KN, Rumpho ME, Bhattacharya D. (2018) Active Host Response to Algal Symbionts in the Sea Slug Elysia chlorotica. Mol Biol Evol. 35: 1706-1711. [K] [Ch]

Dionísio G, Faleiro F, Bispo R, Lopes AR, Cruz S, Paula JR, Repolho T, Calado R, Rosa R. (2018) Distinct Bleaching Resilience of Photosynthetic Plastid-Bearing Mollusks Under Thermal Stress and High CO(2) Conditions. Front Physiol. 9:1675. [K] [NH] [Cr]

Cartaxana P, Trampe E, Kühl M, Cruz S. (2017) Kleptoplast photosynthesis is nutritionally relevant in the sea slug Elysia viridis. Sci Rep. 7: 7714. [K] [Vi]

Laetz EMJ, Wägele H. (2017) Chloroplast digestion and the development of functional kleptoplasty in juvenile Elysia timida (Risso, 1818) as compared to short-term and non-chloroplast-retaining sacoglossan slugs. PLoS One 12: e0182910. [K] [Ti]

Laetz, E.M.J., Ruhr, P.T., Bartolomaeus, T., Preisfeld, A., Wagele, H. (2017) Examining the retention of functional kleptoplasts and digestive activity in sacoglossan sea slugs. Organisms Diversity & Evolution 17: 87-99 [K] [Ti] [Vi]

Laetz, E.M.J., Moris, V.C., Moritz, L., Haubrich, A.N., Wagele, H. (2017) Photosynthate accumulation in solar-powered sea slugs – starving slugs survive due to accumulated starch reserves. Frontiers In Zoology 14: 4 [K] [Ti]

Rauch, C., Jahns, P., Tielens, A.G.M., Gould, S.B., Martin, W.F. (2017) Being the Right Size as an Animal with Plastids. Frontiers In Plant Science 8: 1402 [K] Calculating energy production

Rauch, C., Christa, G., de Vries, J., Woehle, C., Gould, S.B. (2017) Mitochondrial Genome Assemblies of Elysia timida and Elysia cornigera and the Response of Mitochondrion-Associated Metabolism during Starvation. Genome Biology And Evolution 7: 1873-1879 [K] [Co] [Ti]

Rey F, Costa ED, Campos AM, Cartaxana P, Maciel E, Domingues P, Domingues MRM, Calado R, Cruz S. (2017) Kleptoplasty does not promote major shifts in the lipidome of macroalgal chloroplasts sequestered by the sacoglossan sea slug Elysia viridis. Sci Rep. 7:11502. [K] [Vi]

Laetz, E.M.J., Ruhr, P.T., Bartolomaeus, T., Preisfeld, A., Wagele, H. (2017) Examining the retention of functional kleptoplasts and digestive activity in sacoglossan sea slugs. Organisms Diversity & Evolution 17: 87-99 [K] [Ti] [Vi]

Laetz, E.M.J., Moris, V.C., Moritz, L., Haubrich, A.N., Wagele, H. (2017) Photosynthate accumulation in solar-powered sea slugs – starving slugs survive due to accumulated starch reserves. Frontiers In Zoology 14: 4 [K] [Ti]

Baumgartner, F.A., Pavia, H., Toth, G.B. (2015) Acquired phototrophy through retention of functional chloroplasts increases growth efficiency of the sea slug Elysia viridis. PLoS ONE, 10 (4), art. no. e0120874 [F] [K] [Vi]

de Vries, J., Rauch, C., Christa, G., Gould, S.B. (2014) A sea slug’s guide to plastid symbiosis. Acta Soc Bot Pol 83(4):415–421 [K] [R]

Curtis, N.E., Middlebrooks, M.L., Schwartz, J.A., Pierce, S.K. (2015) Kleptoplastic sacoglossan species have very different capacities for plastid maintenance despite utilizing the same algal donors. Symbiosis, 65 (1), pp. 23-31 [F] [K] [Cl] [Pp]

Christa, G., Händeler, K., Kück, P., Vleugels, M., Franken, J., Karmeinski, D., Wägele, H. (2014) Phylogenetic evidence for multiple independent origins of functional kleptoplasty in Sacoglossa (Heterobranchia, Gastropoda). Organisms Diversity and Evolution, 15 (1), pp. 23-36 [K] [F] [Cl] [Pp] [Pt] [Zu]

Pelletreau, K.N., Weber, A.P.M., Weber, K.L., Rumpho, M.E. (2014) Lipid accumulation during the establishment of kleptoplasty in Elysia chlorotica. PLoS ONE, 9 (5), art. no. e97477. [K] [Cl]

Serôdio, J., Cruz, S., Cartaxana, P., Calado, R. (2014) Photophysiology of kleptoplasts: Photosynthetic use of light by chloroplasts living in animal cells. Philosophical Transactions of the Royal Society B: Biological Sciences, 369 (1640), 20130242. [K] [R]

Schwartz, J.A., Curtis, N.E., Pierce, S.K. (2014) FISH labeling reveals a horizontally transferred algal (vaucheria litorea) nuclear gene on a sea slug (elysia chlorotica) chromosome. Biological Bulletin, 227 (3), pp. 300-312. [HT] [K] [Ch]

de Vries, J., Crista, G., Gould, S.B. (2014) Plastid survival in the cytosol of animal cells.  Trends in Plant Science 1168 [K] [R]

Christa, G., Händeler, K., Schäberle, T.F., König, G.M., Wägele, H. (2014) Identification of sequestered chloroplasts in photosynthetic and non-photosynthetic sacoglossan sea slugs (Mollusca, Gastropoda) Frontiers in Zoology, 11 (1), art. no. 15. Abstract PDF [F] [K]

Schmitt, V., Händeler, K., Gunkel, S., Escande, M.-L., Menzel, D., Gould, S.B., Martin, W.F., Wägele, H. (2014) Chloroplast incorporation and long-term photosynthetic performance through the life cycle in laboratory cultures of Elysia timida (sacoglossa, heterobranchia). Frontiers in Zoology, 11 (1), art. no. 5, . Abstract PDF [AQ] [F] [K] [Ti]

Middlebrooks, M.L., Bell, S.S., Curtis, N.E., Pierce, S.K. (2014) Atypical plant-herbivore association of algal food and a kleptoplastic sea slug (Elysia clarki) revealed by DNA barcoding and field surveys. Marine Biology, 161 (6), pp. 1429-1440.  Abstract [F] [K] [Cl]

Akimoto, A., Hirano, Y.M., Sakai, A., Yusa, Y. (2014) Relative importance and interactive effects of photosynthesis and food in two solar-powered sea slugs. Marine Biology, 161 (5), pp. 1095-1102. Abstract [F] [K] [At] [Tr]

Christa, G., Zimorski, V., Woehle, C., Tielens, A.G.M., Wägele, H., Martin, W.F., Gould, S.B. (2013) Plastid-bearing sea slugs fix CO2 in the light but do not require photosynthesis to survive. Proceedings of the Royal Society B: Biological Sciences, 281 (1774), art. no. 20132493. Abstract [K] [Ti] [contains phylogenetic analysis of Plakobranchia]

Martin, R., Walther, P., Tomaschko, K.-H. (2013) Phagocytosis of algal chloroplasts by digestive gland cells in the photosynthesis-capable slug Elysia timida (Mollusca, Opisthobranchia, Sacoglossa). Zoomorphology, 132 (3), pp. 253-259. Abstract [K] [Ti]

Bhattacharya, D., Pelletreau, K.N., Price, D.C., Sarver, K.E., Rumpho, M.E.(2013) Genome analysis of Elysia chlorotica egg DNA provides no evidence for horizontal gene transfer into the germ line of this kleptoplastic mollusc Molecular Biology and Evolution, 30 (8), pp. 1843-1852. PDF [HT] [K] [Ch]

Pelletreau, K.N., Worful, J.M., Sarver, K.E., Rumpho, M.E. (2012) Laboratory culturing of Elysia chlorotica reveals a shift from transient to permanent kleptoplasty Symbiosis, 58 (1-3), pp. 221-232. Page1 [AQ] [F] [K] [Ch]

Middlebrooks, M.L., Bell, S.S., Pierce, S.K. (2012) The kleptoplastic sea slug Elysia clarki prolongs photosynthesis by synthesizing chlorophyll a and b. Symbiosis, 57 (3), pp. 127-132. Page1 [HT] [K] [Cl]

Pierce, S.K., Fang, X., Schwartz, J.A., Jiang, X., Zhao, W., Curtis, N.E., Kocot, K.M., Yang, B., Wang, J. (2012) Transcriptomic evidence for the expression of horizontally transferred algal nuclear genes in the photosynthetic sea slug, Elysia chlorotica. Molecular Biology and Evolution, 29 (6), pp. 1545-1556. PDF [HT] [K] [Ch]

Soule, K.M., Rumpho, M.E. (2012) Light-regulated photosynthetic gene expression and phosphoribulokinase enzyme activity in the heterokont alga Vaucheria litorea (xanthophyceae) and its symbiotic molluskan partner Elysia chlorotica (gastropoda). Journal of Phycology, 48 (2), pp. 373-383. Abstract [HT] [K] [Ch]

Schmitt, V., Wägele, H. (2011) Behavioral adaptations in relation to long-term retention of endosymbiotic chloroplasts in the sea slug Elysia timida (Opisthobranchia, Sacoglossa). Thalassas, 27 (2), pp. 225-238. PDF [BN] [K] [Ti]

Middlebrooks, M.L., Pierce, S.K., Bell, S.S. (2011) Foraging behavior under starvation conditions is altered via photosynthesis by the marine gastropod, Elysia clarki. PLoS ONE, 6 (7), art. no. e22162.  PDF [BN] [K] [Cl]

Wägele, H., Deusch, O., Händeler, K., Martin, R., Schmitt, V., Christa, G., Pinzger, B., Gould, S.B., Dagan, T., Klussmann-Kolb, A., Martin, W. (2011) Transcriptomic evidence that longevity of acquired plastids in the photosynthetic slugs Elysia timida and Plakobranchus ocellatus does not entail lateral transfer of algal nuclear genes. Molecular Biology and Evolution, 28 (1), pp. 699-706. PDF [HT] [K] [Ti]

Rumpho, M.E., Pelletreau, K.N., Moustafa, A., Bhattacharya, D. (2011) The making of a photosynthetic animal. Journal of Experimental Biology, 214 (2), pp. 303-311. PDF [Aq] [HT] [K] [NH] [Ch]

Skulachev, V.P. (2010) Discovery of a photosynthesizing animal that can survive for months in a light-dependent manner. Biochemistry (Moscow), 75 (12), pp. 1498-1499. PDF [HT] [K] [R] [Ch]

Curtis, N.E., Schwartz, J.A., Pierce, S.K. (2010) Ultrastructure of sequestered chloroplasts in sacoglossan gastropods with differing abilities for plastid uptake and maintenance. Invertebrate Biology, 129 (4), pp. 297-308. Abstract [K] [Cl] [Ru] [Pt]

Schwartz, J.A., Curtis, N.E., Pierce, S.K. (2010) Using algal transcriptome sequences to identify transferred genes in the sea slug, Elysia chlorotica. Evolutionary Biology, 37 (1), pp. 29-37. PDF [HT] [K] [Ch]

Serôdio, J., Pereira, S., Furtado, J., Silva, R., Coelho, H., Calado, R. (2010) In vivo quantification of kleptoplastic chlorophyll a content in the “solar-powered” sea slug Elysia viridis using optical methods: Spectral reflectance analysis and PAM fluorometry. Photochemical and Photobiological Sciences, 9 (1), pp. 68-77. PDF [K] [Vi]

Pierce, S.K., Curtis, N.E., Schwartz, J.A. (2009) Chlorophyll a synthesis by an animal using transferred algal nuclear genes. Symbiosis, 49 (2), pp. 121-131.PDF [HT] [K] [Ch]

Händeler, K., Grzymbowski, Y.P., Krug, P.J., Wägele, H. (2009) Functional chloroplasts in metazoan cells – A unique evolutionary strategy in animal life. Frontiers in Zoology, 6 (1), art. no. 28. PDF [F] [K] [R]

Rumpho, M.E., Pochareddy, S., Worful, J.M., Summer, E.J., Bhattacharya, D., Pelletreau, K.N., Tyler, M.S., Lee, J., Manhart, J.R., Soule, K.M. (2009) Molecular characterization of the calvin cycle enzyme phosphoribulokinase in the stramenopile alga Vaucheria litorea and the plastid hosting mollusc Elysia chlorotica. Molecular Plant, 2 (6), pp. 1384-1396. PDF [HT] [K] [Ch]

Vieira, S., Calado, R., Coelho, H., Serôdio, J. (2009) Effects of light exposure on the retention of kleptoplastic photosynthetic activity in the sacoglossan mollusc Elysia viridis. Marine Biology, 156 (5), pp. 1007-1020. PDF [K] [Vi]

Evertsen, J., Johnsen, G. (2009) In vivo and in vitro differences in chloroplast functionality in the two north Atlantic sacoglossans (Gastropoda, Opisthobranchia) Placida dendritica and Elysia viridis. Marine Biology, 156 (5), pp. 847-859. PDF [K] [Vi]

Teugels, B., Bouillon, S., Veuger, B., Middelburg, J.J., Koedam, N. (2008) Kleptoplasts mediate nitrogen acquisition in the sea slug Elysia viridis. Aquatic Biology, 4 (1), pp. 15-21.PDF [K] [Vi]

Rumpho, M.E., Worful, J.M., Lee, J., Kannan, K., Tyler, M.S., Bhattacharya, D., Moustafa, A., Manhart, J.R. (2008) Horizontal gene transfer of the algal nuclear gene psbO to the photosynthetic sea slug Elysia chlorotica. Proceedings of the National Academy of Sciences of the United States of America, 105 (46), pp. 17867-17871. PDF [HT] [K] [Ch]

Gimenez Casalduero, F.G., Muniain, C. (2008) The role of kleptoplasts in the survival rates of Elysia timida (Risso, 1818): (Sacoglossa: Opisthobranchia) during periods of food shortage. Journal of Experimental Marine Biology and Ecology, 357 (2), pp. 181-187. PDF [K] [Ti]

Pierce, S.K., Curtis, N.E., Hanten, J.J., Boerner, S.L., Schwartz, J.A. (2007) Transfer, integration and expression of functional nuclear genes between multicellular species. Symbiosis, 43 (2), pp. 57-64. [HT] [K] [Ch]

Schmitt, V., Anthes, N., Michiels, N.K. (2007) Mating behaviour in the sea slug Elysia timida (Opisthobranchia, Sacoglossa): Hypodermic injection, sperm transfer and balanced reciprocity. Frontiers in Zoology, 4, art. no. 17. [B] PDF [BN] [NH] [Ti]

Evertsen, J., Burghardt, I., Johnsen, G., Wägele, H. (2007) Retention of functional chloroplasts in some sacoglossans from the indo-pacific and Mediterranean. Marine Biology, 151 (6), pp. 2159-2166. PDF [K] [Pu] [Ti] [To]

Curtis, N.E., Pierce, S.K., Massey, S.E., Schwartz, J.A., Maugel, T.K. (2007) Newly metamorphosed Elysia clarki juveniles feed on and sequester chloroplasts from algal species different from those utilized by adult slugs. Marine Biology, 150 (5), pp. 797-806. PDF [AQ] [F] [K] [Cl]

Giménez Casalduero, F., Muniain, C. (2006) Photosynthetic activity of the solar-powered lagoon mollusc Elysia timida (Risso, 1818) (Opisthobranchia: Sacoglossa). Symbiosis, 41 (3), pp. 151-158. PDF [K] [Ti]

Pierce, S.K., Curtis, N.E., Massey, S.E., Bass, A.L., Karl, S.A., Finney, C.M. (2006) A morphological and molecular comparison between Elysia crispata and a new species of kleptoplastic sacoglossan sea slug (Gastropoda: Opisthobranchia) from the Florida Keys, USA.  Molluscan Research, 26 (1), pp. 23-38. Abstract [F] [K] [NH] [Cr] [Cl]

Curtis, N.E., Massey, S.E., Pierce, S.K. (2006) The symbiotic chloroplasts in the sacoglossan Elysia clarki are from several algal species. Invertebrate Biology, 125 (4), pp. 336-345.HTML [F] [K] [Cl]

Green, B.J., Fox, T.C., Rumpho, M.E. (2005) Stability of isolated algal chloroplasts that participate in a unique mollusc/kleptoplast association. Symbiosis, 40 (1), pp. 31-40. Abstract [K] [Ch]

Mondy, W.L., Pierce, S.K. (2003) Apoptotic-like morphology is associated with annual synchronized death in kleptoplastic sea slugs (Elysia chlorotica). Invertebrate Biology, 122 (2), pp. 126-137. Abstract [K] [NH] [Ch]

Hanten, J.J., Pierce, S.K. (2001) Synthesis of several light-harvesting complex I polypeptides is blocked by cycloheximide in symbiotic chloroplasts in the sea slug, Elysia chlorotica (Gould): A case for horizontal gene transfer between alga and animal? Biological Bulletin, 201 (1), pp. 34-44.  PDF [K] [HT] [Ch]

Raven, J.A., Walker, D.I., Jansen, K.R., Handley, L.L., Scrimgeour, S.M., McInroy, S.G. (2001) What fraction of organic carbon in sacoglossans is obtained from photosynthesis by kleptoplastids? An investigation using the natural abundance of stable carbon isotopes.  Mar. Biol. 138: 537-545 [F] [K] [Au] [Ex] [Ma] [Pu]

Rumpho, M.E., Summer, E.J., Green, B.J., Fox, T.C., Manhart, J.R. (2001) Mollusc/algal chloroplast symbiosis: How can isolated chloroplasts continue to function for months in the cytosol of a sea slug in the absence of an algal nucleus? Zoology, 104 (3-4), pp. 303-312. Abstract [HT] [K] [Ch]

Muniain, C., Marín, A., Penchaszadeh, P.E. (2001) Ultrastructure of the digestive gland of larval and adult stages of the sacoglossan Elysia patagonica. Marine Biology, 139 (4), pp. 687-695. PDF [K] [Pa]

Trowbridge, C.D. (2000) The missing links: Larval and post-larval development of the ascoglossan opisthobranch Elysia viridis. Journal of the Marine Biological Association of the United Kingdom, 80 (6), pp. 1087-1094. Abstract [K] [NH] [Vi]

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Trench, R.K., Boyle, J.E., Smith, D.C. (1974) The association between chloroplast of Codium fragile and the mollusc Elysia viridis. III. Movement of photosynthetically fixed 14C in tissues of intact living E. viridis and in Tridachia crispata. Proceedings of the Royal Society of London – Biological Sciences, 185 (1081), pp. 453-464. JStor PDF [K] [Cr] [Vi]

Hinde, R., Smith, D.C. (1972) Persistence of Functional Chloroplasts in Elysia viridis (Opisthobranchia, Sacoglossa) Nat New Biol 239: 30-31 [F] [K] [Vi]

Trench, RK, ME Trench, L Muscatine (1972) Symbiotic chloroplasts; their photosynthetic products and contribution to mucus synthesis in two marine slugs The Biological Bulletin, 142 no. 2 335-349 [K] [Cr] [Di]

Taylor, D.L. (1971) Photosynthesis of symbiotic chloroplasts in Tridachia crispata (Bërgh). Comparative Biochemistry and Physiology — Part A: Physiology, 38 (1), pp. 233-236. PDF [K] [Cr]

Greene, R.W. (1970) Symbiosis in sacoglossan opisthobranchs: functional capacity of symbiotic chloroplasts. Marine Biology, 7 (2), pp. 138-142. PDF [K] [He]

Trench, R.K., Smith, D.C. (1970) Synthesis of pigments in symbiotic chloroplasts. Nature 227: 196 [F] [K] [Cr]

Trench, R.K. (1969) Chloroplasts as functional endosymbionts in the mollusc Tridachia crispata (Bërgh), (Opisthobranchia, Sacoglossa). Nature, 222 (5198), pp. 1071-1072. PDF [K] [Cr] [Paper chromatography of pigments]

Kawaguti, S., Yamasu, T. (1965) Electron microscopy on the symbiosis between an elysioid gastropod and chloroplasts of a green alga.  Biol. J. Okayama Univ. 11: 57-65.  [K] No abstract or PDF available.

Trench, RK, RW Greene, BG Bystrom (1969) Chloroplasts as functional organelles in animal tissues J. Cell Biol.  42: 404-417  [F] [K] [Cr] [Di]

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