Culturing echinoderm larvae through metamorphosis
Tài liệu tham khảo
Agudo, 2007
Allen, 2012, Effects of egg size reductions on development time and juvenile size in three species of echinoid echinoderms: Implications for life history theory, Journal of Experimental Marine Biology and Ecology, 422–423, 72, 10.1016/j.jembe.2012.04.015
Allen, 2007, Uncommon diversity in developmental mode and larval form in the genus Macrophiothrix (Echinodermata: Ophiuroidea), Marine Biology, 151, 85, 10.1007/s00227-006-0470-6
Amemiya, 2015, Development of ciliary bands in larvae of the living isocrinid sea lily Metacrinus rotundus, Acta Zoologica, 96, 36, 10.1111/azo.12049
Aronowicz, 2007, The “agitator”: An inexpensive device for culturing large numbers of fragile marine invertebrate larvae, Marine Biology, 151, 1137, 10.1007/s00227-006-0554-3
Balser, 1998, Cloning by ophiuroid echinoderm larvae, The Biological Bulletin, 194, 187, 10.2307/1543049
Balser, 2002, Phylum echinodermata: Crinoidea, 463
Bashevkin, 2016, Prior exposure to low salinity affects the vertical distribution of Pisaster ochraceus (Echinodermata: Asteroidea) larvae in haloclines, Marine Ecology Progress Series, 542, 123, 10.3354/meps11563
Becker, 2004, The skin ulceration disease in cultivated juveniles of Holothuria scabra (Holothuroidea, Echinodermata), Aquaculture, 242, 13, 10.1016/j.aquaculture.2003.11.018
Birkeland, 1971, Development, substratum selection, delay of metamorphosis and growth in the seastar, Mediaster aequalis, The Biological Bulletin, 141, 99, 10.2307/1539994
Bishop, 2007, Development of nitric oxide synthase-defined neurons in the sea urchin larval ciliary band and evidence for a chemosensory function during metamorphosis, Developmental Dynamics, 236, 1535, 10.1002/dvdy.21161
Brooks, 1899, Ophiura brevispina, Memoirs of the National Academy of Sciences, 8, 79
Burke, 1980, Development of pedicellariae in the pluteus larva of Lytechinus pictus (Echinodermata: Echinoidea), Canadian Journal of Zoology, 58, 1674, 10.1139/z80-229
Burke, 1984, Pheromonal control of metamorphosis in the pacific sand dollar, Dendraster excentricus, Science, 225, 442, 10.1126/science.225.4660.442
Byrne, 1991, Reproduction, development and population biology of the Caribbean ophiuroid Ophionereis olivacea, a protandric hermaphrodite that broods its young, Marine Biology, 111, 387, 10.1007/BF01319411
Byrne, 1995, Changes in larval morphology in the evolution of benthic development by Patiriella exigua (Asteroidea), a comparison with the larvae of Patiriella species with planktonic development, The Biological Bulletin, 188, 293, 10.2307/1542306
Byrne, 1996, Viviparity and intragonadal cannibalism in the diminutive asterinid sea stars Patiriella vivipara and P. parvivipara, Marine Biology, 125, 551, 10.1007/BF00353268
Byrne, 2005, Viviparity in the sea star Cryptasterina hystera (Asterinidae)—Conserved and modified features in reproduction and development, The Biological Bulletin, 208, 81, 10.2307/3593116
Byrne, 2008, Brooding of pelagic-type larvae in Ophiopeza spinosa: Reproduction and development in a tropical ophiodermatid brittle-star, Invertebrate Biology, 127, 98, 10.1111/j.1744-7410.2007.00110.x
Byrne, 2001, Ciliated band structure in planktotrophic and lecithotrophic larvae of Heliocidaris species (Echinodermata:Echinoidea): Conservation and change, Acta Zoologica, 82, 189
Byrne, 2002, Phylum Echinodermata: Ophiuroidea, 483
Cameron, 1989, The control of sea urchin metamorphosis: Ionic effects, Development, Growth & Differentiation, 31, 589, 10.1111/j.1440-169X.1989.00589.x
Carpizo-Ituarte, 2002, Induction of metamorphosis with KCl in three species of sea sea urchins and its implications in the production of juveniles, Ciencias Marinas, 28, 157, 10.7773/cm.v28i2.217
Castilla-Gavilan, 2018, Optimising microalgae diets in sea urchin Paracentrotus lividus larviculture to promote aquaculture diversification, Aquaculture, 490, 251, 10.1016/j.aquaculture.2018.02.003
Cavanaugh, 1975
Chen, 1992, Reproductive cycle, larval development, juvenile growth and population dynamics of Patiriella pseudoexigua (Echinodermata: Asteroidea) in Taiwan, Marine Biology, 113, 271, 10.1007/BF00347281
Chia, 1966, Development of a deep-sea cushion star, Pteraster tesselatus, Proceedings of the California Academy of Sciences, 34, 505
Chia, 1978, Echinoderm metamorphosis: Fate of larval structures, 219
Cisternas, 2005, Larval development in the ophiuroid Ophiarachnella gorgonia (Ophiodermatidae): Evidence for developmental heterochronies between species with type I and type II development, Canadian Journal of Zoology, 83, 1067, 10.1139/z05-092
Derbès, 1847, Observations sur le mechanism et les phénomènes qui accompagnent la formation de l'embryon chez l'oursin comestible, Annales des Sciences Naturelles III S'er. Zool., 8, 80
Dupont, 2010, Impact of near-future ocean acidification on echinoderms, Ecotoxicology, 19, 449, 10.1007/s10646-010-0463-6
Eeckhaut, 2004, Parasites and biotic diseases in field and cultivated sea cucumbers, Vol. 463, 311
Eaves, 2003, Reproduction—Widespread cloning in echinoderm larvae, Nature, 425, 146, 10.1038/425146a
Elphick, 2001, Neural control of muscle relaxation in echinoderms, The Journal of Experimental Biology, 204, 875, 10.1242/jeb.204.5.875
Emlet, 1988, Larval form and metamorphosis of a "primitive" sea urchin, Eucidaris thouarsi (Echinodermata, Echinoidea, Cidaroida), with implications for developmental and phylogenetic studies, The Biological Bulletin, 174, 4, 10.2307/1541754
Emlet, 2006, Direct development of the brittle star Amphiodia occidentalis (Ophiuroidea, Amphiuridae) from the northeastern Pacific Ocean, Invertebrate Biology, 125, 154, 10.1111/j.1744-7410.2006.00049.x
Emlet, 2010, Morphological evolution of newly metamorphosed sea urchins—a phylogenetic and functional analysis, Integrative and Comparative Biology, 50, 571, 10.1093/icb/icq073
Fadl, 2017, New biomarkers of post-settlement growth in the sea urchin Strongylocentrotus purpuratus, Heliyon, 3, 10.1016/j.heliyon.2017.e00412
Fell, 1946, The embryology of the viviparous ophiuroid Amphipholis squamata Delle Chiajie, Transactions of the Royal Society of New Zealand, 75, 419
Fenaux, 1969, Le developpement larvaire chez Ophioderma longicauda (Retzius), Cahiers de Biologie Marine, 10, 59
Gaylord, 2013, Turbulent shear spurs settlement in larval sea urchins, Proceedings of the National Academy of Sciences of the United States of America, 110, 6901, 10.1073/pnas.1220680110
George, 1994, Population differences in maternal size and offspring quality for Leptasterias epichlora (Brandt) (Echinodermata, Asteroidea), Journal of Experimental Marine Biology and Ecology, 175, 121, 10.1016/0022-0981(94)90179-1
Gianasi, 2018, Full allogeneic fusion of embryos in a holothuroid echinoderm, Proceedings of the Royal Society B: Biological Sciences, 285, 10.1098/rspb.2018.0339
Gianasi, 2018, Morphometric and behavioural changes in the early life stages of the sea cucumber Cucumaria frondosa, Aquaculture, 490, 5, 10.1016/j.aquaculture.2018.02.017
Gladfelter, 1978, General ecology of the cassiduloid urchin Cassidulus caribbearum, Marine Biology, 47, 149, 10.1007/BF00395636
Gosselin, 1998, From competent larva to exotrophic juvenile, a morphological study of the perimetamorphic period of Paracentrotus lividus (Echinodermata, Echinoida), Zoomorphology, 118, 31, 10.1007/s004350050054
Guillard, 1975, Culture of phytoplankton for feeding marine invertebrates, 26
Hadfield, 2011, Biofilms and marine invertebrate larvae: What bacteria produce that larvae use to choose settlement sites, Annual Review of Marine Science, 3, 453, 10.1146/annurev-marine-120709-142753
Hair, 2011, Sandfish culture in Fiji Islands, SPC Beche-de-mer Information Bulletin, 31, 3
Hamel, 1996, Early development, settlement, growth, and spatial distribution of the sea cucumber Cucumaria frondosa (Echinodermata, Holothuroidea), Canadian Journal of Fisheries and Aquatic Sciences, 53, 253, 10.1139/f95-186
Hamel, 2008, Population status, fisheries and trade of sea cucumbers in temperate areas of the northern hemisphere, Vol. 516, 257
Hamel, 2008, Precautionary management of Cucumaria frondosa in Newfoundland and Labrador, Canada, Vol. 516, 293
Hamel, 2001, The sea cucumber Holothuria scabra (Holothuroidea, Echinodermata), its biology and exploitation as beche-de-mer, Advances in Marine Biology, 41, 129, 10.1016/S0065-2881(01)41003-0
Hart, 1991, Particle captures and the method of suspension feeding by echinoderm larvae, The Biological Bulletin, 180, 12, 10.2307/1542425
Hendler, 1975, Adaptational significance of the patterns of ophiuroid development, American Zoologist, 15, 691, 10.1093/icb/15.3.691
Hendler, 1977, Development of Amphioplus abditus (Verrill) (Echinodermata, Ophiuroidea), I. Larval biology, The Biological Bulletin, 152, 51, 10.2307/1540726
Hendler, 1991, Echinodermata, Ophiuroidea, 355
Herrmann, 2003, Sterile polystyrene culture dishes induce transformation of polyps into medusae in Aurelia aurita (Scyphozoa, Cnidaria), Methods in Cell Science, 25, 135, 10.1007/s11022-004-5592-y
Heyland, 2004, Heterochronic developmental shift caused by thyroid hormone in larval sand dollars and its implications for phenotypic plasticity and the evolution of nonfeeding development, Evolution, 58, 524, 10.1111/j.0014-3820.2004.tb01676.x
Heyland, 2014, A detailed staging scheme for late larval development in Strongylocentrotus purpuratus focused on readily-visible juvenile structures within the rudiment, BMC Developmental Biology, 14, 22, 10.1186/1471-213X-14-22
Heyland, 2006, Thyroid hormone metabolism and peroxidase function in two non-chordate animals, Journal of Experimental Zoology. Part B, Molecular and Developmental Evolution, 306, 551, 10.1002/jez.b.21113
Heyland, 2004, Thyroid hormones determine developmental mode in sand dollars (Echinodermata, Echinoidea), Evolution & Development, 6, 382, 10.1111/j.1525-142X.2004.04047.x
Heyland, 2006, Endogenous thyroid hormone synthesis in facultative planktotrophic larvae of the sand dollar Clypeaster rosaceus, implications for the evolutionary loss of larval feeding, Evolution & Development, 8, 568, 10.1111/j.1525-142X.2006.00128.x
Highsmith, 1982, Induced settlement and metamorphosis of sand dollar (Dendraster excentnicus) larvae in predator-free sites, adult sand dollar beds, Ecology, 63, 329, 10.2307/1938950
Highsmith, 1986, Delayed metamorphosis, effect on growth and survival of juvenile sand dollars (Echinoidea, Clypeasteroida), Bulletin of Marine Science, 39, 347
Hill, 2013, Methods for disinfection of aquaculture establishments, 28
Ho, 2016, Perturbation of gut bacteria induces a coordinated cellular immune response in the purple sea urchin larva, Immunology and Cell Biology, 94, 861, 10.1038/icb.2016.51
Hodin, 2006, Expanding networks, signaling components in and a hypothesis for the evolution of metamorphosis, Integrative and Comparative Biology, 46, 719, 10.1093/icb/icl038
Hodin, 2018, Turbulence exposure recapitulates desperate behavior in late-stage sand dollar larvae, BMC Zoology, 3, 9, 10.1186/s40850-018-0034-5
Hodin, 2015, Rethinking competence in marine life cycles, ontogenetic changes in the settlement response of sand dollar larvae exposed to turbulence, Royal Society Open Science, 2, 150114, 10.1098/rsos.150114
Hoekstra, 2012, Novel insights into the echinoderm nervous system from histaminergic and FMRFaminergic-like cells in the sea cucumber Leptosynapta clarki, PLoS One, 7, 10.1371/journal.pone.0044220
Holland, 1991, Echinodermata, Crinoidea, 247
Huggett, 2005, Larval development and metamorphosis of the australian diadematid sea urchin Centrostephanus rodgersii, Invertebrate Reproduction and Development, 47, 197, 10.1080/07924259.2005.9652160
Hurst, 1967, The egg masses and veligers of thirty northeast Pacific opistobranchs, Veliger, 9, 255
Israel, 2016, Comparative developmental transcriptomics reveals rewiring of a highly conserved gene regulatory network during a major life history switch in the sea urchin genus Heliocidaris, PLoS Biology, 14, 10.1371/journal.pbio.1002391
Kacenas, 2018, Density-dependent expression of plasticity in larval morphology, effects of actual and apparent competitors, Marine Ecology Progress Series, 593, 1, 10.3354/meps12573
Kamya, 2014, Larvae of the coral eating crown-of-thorns starfish, Acanthaster planci in a warmer-high CO2 ocean, Global Change Biology, 20, 3365, 10.1111/gcb.12530
Kitazawa, 2012, Morphogenesis of adult traits during the early development of Mespilia globulus Linnaeus, 1758 (Echinodermata, Echinoidea), Zoological Studies, 51, 1481
Kumé, 1968
2013, Vol. 38
Lawrence, 2019, Large-scale production of sea urchin (Strongylocentrotus intermedius) seed in a hatchery in China, Aquaculture International, 10.1007/s10499-018-0319-2
Leahy, 1986, Laboratory culture of Strongylocentrotus purpuratus adults, embryos, and larvae, Vol. 27, 1
Leong, 1997, Metabolic importance of Na+/K+-atpase activity during sea urchin development, The Journal of Experimental Biology, 200, 2881, 10.1242/jeb.200.22.2881
Lizárraga, 2017, Low concentrations of large inedible particles reduce feeding rates of echinoderm larvae, Marine Biology, 164, 102, 10.1007/s00227-017-3134-9
Marsh, 2001, High macromolecular synthesis with low metabolic cost in Antarctic sea urchin embryos, Science, 291, 1950, 10.1126/science.1056341
Martinez, 2017, Filling in the grazing puzzle, a synthesis of herbivory in starfish, Oceanography and Marine Biology: An Annual Review, 55, 1
McAlister, 2018, Phenotypic plasticity of feeding structures in marine invertebrate larvae, 103
McDonald, 2010, Abrupt change in food environment induces cloning in plutei of Dendraster excentricus, The Biological Bulletin, 219, 38, 10.1086/BBLv219n1p38
McEdward, 1993, Life cycle evolution in asteroids, what is a larva?, The Biological Bulletin, 184, 255, 10.2307/1542444
McEdward, 2001, Larval and life-cycle patterns in echinoderms, Canadian Journal of Zoology, 79, 1125, 10.1139/z00-218
Mercier, 2013, Sea cucumber aquaculture, hatchery production, juvenile growth and industry challenges, 431
Mercier, 2012, Hatchery experience and useful lessons from Isostichopus fuscus in Ecuador and Mexico, 79
Mercier, 2004, Aquaculture of the Galapagos sea cucumber, Isostichopus fuscus, Vol. 463, 347
Miller, 1999, Development of newly metamorphosed juvenile sea urchins (Strongylocentrotus franciscanus and S. purpuratus), morphology, the effects of temperature and larval food ration, and a method for determining age, Journal of Experimental Marine Biology and Ecology, 235, 67, 10.1016/S0022-0981(98)00164-6
Milonas, 2010, Light influences feeding and growth of echinoplutei, Marine Ecology Progress Series, 404, 69, 10.3354/meps08488
Mladenov, 1979, Unusual lecithotrophic development of the Caribbean brittle star Ophiothrix oerstedi, Marine Biology, 55, 55, 10.1007/BF00391717
Mladenov, 1985, Development and metamorphosis of the brittle star Ophiocoma pumila, evolutionary and ecological implications, The Biological Bulletin, 168, 285, 10.2307/1541241
Montgomery, 2017, Ontogenetic shifts in swimming capacity of echinoderm propagules, a comparison of species with planktotrophic and lecithotrophic larvae, Marine Biology, 164, 43, 10.1007/s00227-017-3072-6
Montgomery, 2018, Ontogenetic variation in photosensitivity of developing echinoderm propagules, Journal of Experimental Marine Biology and Ecology, 500, 63, 10.1016/j.jembe.2017.12.003
Mortensen, 1920, On hermaphroditism in viviparous ophiuroids, Acta Zoologica, 1, 1, 10.1111/j.1463-6395.1920.tb00326.x
Mortensen, 1921
Mortensen, 1931
Mortensen, 1933, Papers from Dr. T. Mortensen's Pacific expedition 1914-16, Vol. 93, 401
Mortensen, 1937
Mortensen, 1938
Müller, 1846, Über die larven und die metamorphose der ophiuren und seeigel, Vorgetragen in der Königlichen Akademie der Wissenschaften am 29, 1846, 273
Olson, 1993, Larval development (with observations on spawning) of the pencil urchin Phyllacanthus imperialis: A new intermediate larval form?, The Biological Bulletin, 185, 77, 10.2307/1542131
Pearse, 1991, Echinodermata, Echinoidea, 514
Podolsky, 2005, Developmental plasticity in Macrophiothrix brittlestars, are morphologically convergent larvae also convergently plastic?, The Biological Bulletin, 209, 127, 10.2307/3593130
Purcell, 2016, Ecological roles of exploited sea cucumbers, Oceanography and Marine Biology: An Annual Review, 54, 367
Purcell, 2013, Sea cucumber fisheries, global analysis of stocks, management measures and drivers of overfishing, Fish and Fisheries, 14, 34, 10.1111/j.1467-2979.2011.00443.x
Purcell, 2014, The cost of being valuable, predictors of extinction risk in marine invertebrates exploited as luxury seafood, Proceedings of the Royal Society B, 9, e95075
Qiu, 2015, Development, settlement and post-settlement growth, 111
Raff, 2006, The active evolutionary lives of echinoderm larvae, Heredity, 97, 244, 10.1038/sj.hdy.6800866
Ren, 2016, Effects of mono-species and bi-species microalgal diets on the growth, survival and metamorphosis of auricularia larvae of the California sea cucumber, Parastichopus californicus (Stimpson, 1857), Aquaculture Nutrition, 22, 304, 10.1111/anu.12253
Ramofafia, 2003, Development of three commercial sea cucumbers, Holothuria scabra, H. fuscogilva and Actinopyga mauritiana: larval structure and growth, Marine and Freshwater Research, 54, 657, 10.1071/MF02145
Roman, 2001, Temporal and spatial patterns of zooplankton in the Chesapeake Bay turbidity maximum, Marine Ecology Progress Series, 213, 215, 10.3354/meps213215
Scholtz, 2013, Effects of different microalgal feeds and their influence on larval development in the white-spined sea urchin Tripneustes gratilla, African Journal of Marine Science, 35, 25, 10.2989/1814232X.2013.769902
Selvakumaraswamy, 2000, Reproduction, spawning and development in 5 ophiuroids from Australia and New Zealand, Invertebrate Biology, 119, 394, 10.1111/j.1744-7410.2000.tb00109.x
Selvakumaraswamy, 2006, Evolution of larval form in ophiuroids, insights from the metamorphic phenotype of Ophiothrix (Echinodermata, Ophiuroidea), Evolution & Development, 8, 183, 10.1111/j.1525-142X.2006.00088.x
Smith, 1997, Echinoderm larvae and phylogeny, Annual Review of Ecology and Systematics, 28, 219, 10.1146/annurev.ecolsys.28.1.219
Smith, 2008, The larval stages of the sea urchin, Strongylocentrotus purpuratus, Journal of Morphology, 269, 713, 10.1002/jmor.10618
Soars, 2015, Contrasting arm elevation angles of multi- and two-armed sea urchin echinoplutei supports Gru¨nbaum and Strathmann's hydromechanical model, Marine Biology, 162, 607, 10.1007/s00227-014-2608-2
Soars, 2009, Overview of phenotypic plasticity in echinoid larvae, 'Echinopluteus transversus' type vs. typical echinoplutei, Marine Ecology Progress Series, 383, 113, 10.3354/meps07848
Stancyk, 1973, Development of Ophiolepis elegans (Echinodermata, Ophiuroidea) and its implications in the estuarine environment, Marine Biology, 21, 7, 10.1007/BF00351186
Strathmann, 1971, The feeding behavior of planktotrophic echinoderm larvae, mechanisms, regulation, and rates of suspension feeding, Journal of Experimental Marine Biology and Ecology, 6, 109, 10.1016/0022-0981(71)90054-2
Strathmann, 1978, Larval settlement in echinoderms, 235
Strathmann, 1987
Strathmann, 1987, Larval feeding, 465
Strathmann, 2014, Culturing larvae of marine invertebrates, Vol. 1128, 1
Strathmann, 1992, Heterochronic developmental plasticity in larval sea urchins and its implications for the evolution of nonfeeding larvae, Evolution, 46, 972, 10.1111/j.1558-5646.1992.tb00613.x
Strathmann, 2006, Good eaters, poor swimmers, compromises in larval form, Integrative and Comparative Biology, 46, 312, 10.1093/icb/icj031
Strickland, 2004, Light microscopy of echinoderm embryos, Methods in Cell Biology, 74, 371, 10.1016/S0091-679X(04)74016-9
Suckling, 2018, Optimising stocking density for the commercial cultivation of sea urchin larvae, Aquaculture, 488, 96, 10.1016/j.aquaculture.2018.01.022
Sui, 1990, 107
Sutherby, 2012, Histamine is a modulator of metamorphic competence in Strongylocentrotus purpuratus (Echinodermata, Echinoidea), BMC Developmental Biology, 12, 14, 10.1186/1471-213X-12-14
Swanson, 2012, Dissolved histamine, a potential habitat marker promoting settlement and metamorphosis in sea urchin larvae, Marine Biology, 159, 915, 10.1007/s00227-011-1869-2
Toral-Granda, 2008, Sea cucumbers, A global review of fisheries and trade, Vol. 516
Turner, 1976, Fixation and preservation of molluscan zooplankton, 290
Vellutini, 2010, Embryonic, larval, and juvenile development of the sea biscuit Clypeaster subdepressus (Echinodermata, Clypeasteroida), PLoS One, 5, 10.1371/journal.pone.0009654
Wheeler, 2016, Ontogenetic changes in larval swimming and orientation of pre-competent sea urchin Arbacia punctulata in turbulence, The Journal of Experimental Biology, 219, 1303, 10.1242/jeb.129502
Wolfe, 2017, Superstars, assessing nutrient thresholds for enhanced larval success of Acanthaster planci, a review of the evidence, Marine Pollution Bulletin, 116, 307, 10.1016/j.marpolbul.2016.12.079
Wray, 1992, The evolution of larval morphology during the post-Paleozoid radiation of echinoids, Paleobiology, 18, 258, 10.1017/S0094837300010848
Wray, 1996, Parallel evolution of nonfeeding larvae in echinoids, Systematic Biology, 45, 308, 10.1093/sysbio/45.3.308
Wray, 1994, Several distinct factors drive the evolution of early development in echinoderms, Development, 97, 10.1242/dev.1994.Supplement.97
Wray, 1989, Evolutionary modification of cell lineage in the direct developing sea urchin Heliocidaris erythrogramma, Developmental Biology, 132, 458, 10.1016/0012-1606(89)90242-X
Wray, 1991, The evolution of developmental strategy in marine invertebrates, Trends in Ecology & Evolution, 6, 45, 10.1016/0169-5347(91)90121-D
Xing, 2007, The potential value of different species of benthic diatoms as food for newly metamorphosed sea urchin Strongylocentrotus intermedius, Aquaculture, 263, 142, 10.1016/j.aquaculture.2006.10.003
Yaguchi, J. (2019). Microinjection methods for sea urchin eggs and blastomeres. Methods in Cell Biology, 150, 171–188.
Yamashita, 1985, Embryonic development of the brittle-star Amphipholis kochii in laboratory culture, The Biological Bulletin, 169, 131, 10.2307/1541393
Yang, 2015
Zhang, 2015, Aquaculture, stock enhancement, and restocking, 289
Zhang, 2017, The sea cucumber genome provides insights into morphological evolution and visceral regeneration, PLoS Biology, 15, 10.1371/journal.pbio.2003790