Ocean EcoSystems Biology publications

Click here for publications from the Experimental Ecology - Food Webs (2004 - 2018).


Min MA, Needham DM, Sudek S, Truelove NK, Pitz K, Chavez GM, Poirier C, Gardeler B, von der Esch E, Ludwig A, Riebesell U, Worden AZ & FP Chavez. Ecological divergence of a mesocosm in an eastern boundary upwelling system assessed with multi-marker DNA metabarcoding. Biogeosciences. In press. Worden Lab Publication

Needham DM, Poirier C, Bachy C, George EE, Wilken S, Yung CCM, Limardo AJ, Morando M, Sudek L, Malstrom RR, Keeling PJ, Santoro AE & AZ Worden. The microbiome of a bacterivorous marine choanoflagellate contains a resource-demanding obligate bacterial associate. Nature Microbiology. Vol 7, 1466-1479. Worden Lab Publication

Wittmers F, Needham DM, Hehenberger E, Giovannoni SJ & AZ Worden. Genomes from uncultivated pelagiphages reveal multiple phylogenetic clades exhibiting extensive auxiliary metabolic genes and cross-family multigene transfers. mSystems. e01522-21. Worden Lab Publication

Bachy C, Sudek L, Choi C-J, Eckmann CA, Nöthig E-M, Metfies K & AZ Worden. Phytoplankton surveys in the arctic Fram Strait demonstrate the tiny eukaryotic alga Micromonas and other picoprasinophytes contribute to deep sea export. Microorganisms. Vol 10, 961 doi.org/10.3390/microorganisms10050961. Worden Lab Publication

Bachy C, Wittmers F, Muschiol J, Hamilton M, Henrissat B & AZ Worden. The land-sea connection: insights into the plant lineage from a green algal perspective. Annual Reviews of Plant Biology. Vol 73, 585–616. Worden Lab Publication


Hamilton M, Mascioni M, Hehenberger E, Bachy C, Yung C, Vernet M & AZ Worden. Spatiotemporal variations in Antarctic protistan communities highlight phytoplankton diversity and seasonal dominance by a novel cryptophyte lineage. mBio. Vol 12, Issue 6 e02973-21. Worden Lab Publication

Schön ME, Zlatogursky VV, Singh RP, Poirier C, Wilken S, Mathur V, Strassert JFH, Pinhassi J, Worden AZ, Keeling PJ, Ettema TJG, Wideman JG & F Burki. Single cell genomics reveals plastid-lacking Picozoa as close relatives of red algae. Nature Communications. Vol 12, 6651. doi.org/10.1038/s41467-021-26918-0. Worden Lab Collaboration

Bolaños LM, Choi CJ, Worden AZ, Baetge N, Carlson CA & S Gionvannoni. Seasonality of the Microbial Community Composition in the North Atlantic. Frontiers in Marine Science. 8(23). https://doi.org/10.3389/fmars.2021.624164.

Bachy C, Yung C-M, Needham DM, Gazitua C, Roux S, Limardo AJ, Choi CJ, Sullivan MB & AZ Worden. Viruses infecting a warm water picoeukaryote shed light on spatial co-occurrence dynamics of marine viruses and their hosts. The ISME Journal. doi.org/10.1038/s41396-021-00989-9. Worden Lab Publication

Hoadley KD, Hamilton M, Poirier CL, Choi CJ, Yung C-M & AZ Worden. Selective uptake of pelagic microbial community members by Caribbean reef corals. Applied and Environmental Microbiology.Vol 87, e03175-20. Worden Lab Publication.       


Bi R & U Sommer. Food Quantity and Quality Interactions at Phytoplankton–Zooplankton Interface: Chemical and Reproductive Responses in a Calanoid Copepod. Frontiers in Marine Science 7(274). http://dx.doi.org/10.3389/fmars.2020.00274.  

Bi R, Ismar-Rebitz SMH, Sommer U, Zhang H & M Zhao. Ocean-related global change alters lipid biomarker production in common marine phytoplankton. Biogeosciences (BG) 17(24), pp. 6287-6307. http://dx.doi.org/10.5194/bg-17-6287-2020.  

Breusing C, Schultz DT, Sudek S, Worden AZ & CR Young. High‐contiguity genome assembly of the chemosynthetic gammaproteobacterial endosymbiont of the cold seep tubeworm Lamellibrachia barhami. Molecular Ecology Resources, 20(5). pp. 1432-1444. http://dx.doi.org/10.1111/1755-0998.13220.  

Burian A, Nielsen JM, Hansen T, Bermudez, R & M Winder. The potential of fatty acid isotopes to trace trophic transfer in aquatic food-webs. Philosophical Transactions of the Royal Society B: Biological Sciences, 375(1804). p. 20190652. http://dx.doi.org/10.1098/rstb.2019.0652

Choi C, Jimenez V, Needham D, Poirier C, Bachy C, Alexander H, Wilken S, Chavez F, Sudek S, Giovannoni, SJ & AZ Worden. Seasonal and geographical transitions in eukaryotic phytoplankton community structure in the Atlantic and Pacific Oceans. Frontiers in Microbiology, 11. Art. Nr. 542372. http://dx.doi.org/10.3389/fmicb.2020.542372.  

Cooney EC, Okamoto N, Cho A, Hehenberger E, Richards TA, Santoro AE, Worden AZ, Leander BS & PJ Keeling. Single-Cell Transcriptomics of Abedinium Reveals a New Early-Branching Dinoflagellate Lineage. Genome Biology and Evolution, 12 (12). pp. 2417-2428. http://dx.doi.org/10.1093/gbe/evaa196.

Djeghri N, Pondaven P, Stockenreiter M, Behl S, Huang JYT, Hansen T, Patris S, Ucharm G & H Stibor. Isotopic and elemental compositions reveal density‐dependent nutrition pathways in a population of mixotrophic jellyfish. Ecosphere, 11 (11). Art.NR. e03295. http://dx.doi.org/10.1002/ecs2.3295.  

Engel FG, Dini-Andreote F, Eriksson BK, Salles JF, de Lima Brossi MJ  & B Matthiessen. Dispersal mitigates bacterial dominance over microalgal competitor in metacommunities. Oecologia, 193. pp. 677-687. http://dx.doi.org/10.1007/s00442-020-04707-8.  

Engel FG, Matthiessen B & BK Eriksson. A heatwave increases turnover and regional dominance in microbenthic metacommunities. Basic and Applied Ecology, 47. pp. 1-11. http://dx.doi.org/10.1016/j.baae.2020.03.003.  

Horn HG, Boersma M, Garzke J, Sommer U & N Aberle. High CO2 and warming affect microzooplankton food web dynamics in a Baltic Sea summer plankton community. Marine Biology, 167 (5). http://dx.doi.org/10.1007/s00227-020-03683-0.  

Hoving HJT, Neitzel P, Hauss H, Christiansen S, Kiko R, Robison BH, Silva P & A Körtzinger. In situ observations show vertical community structure of pelagic fauna in the eastern tropical North Atlantic off Cape Verde. Scientific Reports, 10 (1). Art.Nr. 21798. http://dx.doi.org/10.1038/s41598-020-78255-9.  

Huang PC, Liu TY, Hu MY, Casties I. & YC Tseng. Energy and nitrogenous waste from glutamate/glutamine catabolism facilitates acute osmotic adjustment in non-neuroectodermal branchial cells. Scientific Reports, 10 (1). Art.Nr. 9460. http://dx.doi.org/10.1038/s41598-020-65913-1.  

Ismar-Rebitz SMH, Sommer U & J Garzke. Zooplankton growth and survival differentially respond to interactive warming and acidification effects. Journal of Plankton Research, 42 (2). pp. 189-202. http://dx.doi.org/10.1093/plankt/fbaa005.  

Kiko R, Bianchi D, Grenz C, Hauss H, Iversen M, Kumar S, Maas A & C Robinson. Editorial: Zooplankton and Nekton: Gatekeepers of the Biological Pump. Frontiers in Marine Science, 7.   Art.Nr. 545. http://dx.doi.org/10.3389/fmars.2020.00545.  

Kiko R, Brandt P, Christiansen S, Faustmann J, Kriest I, Rodrigues E, Schütte F & H Hauss.  Zooplankton-Mediated Fluxes in the Eastern Tropical North Atlantic. Frontiers in Marine Science, 7. Art.Nr. 358. http://dx.doi.org/10.3389/fmars.2020.00358.  

Larsen T, Hansen T & J Dierking. Characterizing niche differentiation among marine consumers with amino acid δ 13 C fingerprinting. Ecology and Evolution, 10 (14). pp. 7768-7782. http://dx.doi.org/10.1002/ece3.6502.  

Lebrato M, Garbe-Schönberg D, Müller MN, Blanco-Ameijeiras S, Feely RA, Lorenzoni L, Molinero JC, Bremer K, Jones DOB, Iglesias-Rodriguez D, Greeley D, Lamare MD, Paulmier A, Graco M, Cartes J, Barcelos e Ramos J, de Lara A, Sanchez-Leal R, Jimenez P, Paparazzo FE, Hartman SE, Westernströer U, Küter M, Benavides R, da Silva AF, Bell S, Payne C, Olafsdottir S, Robinson K, Jantunen LM, Korablev A, Webster RJ, Jones EM, Gilg O, Bailly du Bois P,  Beldowski J, Ashjian C, Yahia ND, Twining B,  Chen XG, Tseng LC, Hwang JS, Dahms HU & A Oschlies. Global variability in seawater Mg:Ca and Sr:Ca ratios in the modern ocean.  PNAS Proceedings of the National Academy of Sciences of the United States of America, 117  (36). pp. 22281-22292. http://dx.doi.org/10.1073/pnas.1918943117.  

Listmann L,  Hattich GSI, Matthiessen B & TBH Reusch. Eco-Evolutionary Interaction in Competing Phytoplankton: Nutrient Driven Genotype Sorting Likely Explains Dominance Shift and Species Responses to CO2. Frontiers in Marine Science, 7. Art.Nr. 634. http://dx.doi.org/10.3389/fmars.2020.00634.  

Makareviciute K, Matthiessen B, Lotze HK & U Sommer. Decrease in diatom dominance at lower Si:N ratios alters plankton food webs. Journal of Plankton Research, 42 (4). pp. 411-424.  http://dx.doi.org/10.1093/plankt/fbaa032.  

Moustaka-Gouni M & U Sommer. Effects of Harmful Blooms of Large-Sized and Colonial Cyanobacteria on Aquatic Food Webs. Water, 12 (6). Art.Nr. 1587. http://dx.doi.org/10.3390/w12061587.  

Ory NC, Lehmann A, Javidpour J, Stöhr R, Walls GL & C Clemmesen. Factors influencing the spatial and temporal distribution of microplastics at the sea surface – A year-long monitoring case study from the urban Kiel Fjord, southwest Baltic Sea. Science of the Total Environment, 736 .   Art.Nr. 139493. http://dx.doi.org/10.1016/j.scitotenv.2020.139493.  

Ramírez-Romero E, Molinero JC, Sommer U, Salhi N, Kéfi-Daly Yahia O & MN Daly Yahia. Phytoplankton size changes and diversity loss in the southwestern Mediterranean Sea in relation to long-term hydrographic variability. Estuarine, Coastal and Shelf Science, 235. Art.Nr. 106574. http://dx.doi.org/10.1016/j.ecss.2019.106574.  

Stagličić N, Šegvić-Bubić T, Ezgeta-Balić D, Bojanić Varezić D, Grubišić L, Žuvić L, Lin Y & E Briski. Distribution patterns of two co-existing oyster species in the northern Adriatic Sea: The native European flat oyster Ostrea edulis and the non-native Pacific oyster Magallana gigas. Ecological Indicators, 113. Art.Nr. 106233.  http://dx.doi.org/10.1016/j.ecolind.2020.106233.  

Suffridge CP, Bolanos LM, Bergauer K, Worden AZ, Morré J, Behrenfeld MJ & SJ Giovannoni.  Exploring Vitamin B1 Cycling and Its Connections to the Microbial Community in the North Atlantic Ocean. Frontiers in Marine Science, 7. Art.Nr. 606342. http://dx.doi.org/10.3389/fmars.2020.606342.  

Tikhonenkov DV, Mikhailov KV, Hehenberger E, Karpov SA, Prokina KI, Esaulov AS, Belyakova OI, Mazei YA, Mylnikov AP, Aleoshin VV & PJ Keeling. New Lineage of Microbial Predators Adds Complexity to Reconstructing the Evolutionary Origin of Animals. Current Biology, 30 (22). 4500-4509.e5. http://dx.doi.org/10.1016/j.cub.2020.08.061.  

Trombetta T, Vidussi F, Roques C, Scotti M & B Mostajir. Marine Microbial Food Web Networks During Phytoplankton Bloom and Non-bloom Periods: Warming Favors Smaller Organism Interactions and Intensifies Trophic Cascade. Frontiers in Microbiology, 11. Art.Nr. 502336. http://dx.doi.org/10.3389/fmicb.2020.502336.  

Wilken S, Choi CJ & AZ Worden. Contrasting Mixotrophic Lifestyles Reveal Different Ecological Niches in Two Closely Related Marine Protists. Journal of Phycology, 56 (1). pp. 52-67. http://dx.doi.org/10.1111/jpy.12920.  

Dörner I, Hauss H, Aberle N, Lohbeck K, Spisla C, Riebesell U& SMH Ismar-Rebitz. Ocean acidification impacts on biomass and fatty acid composition of a post-bloom marine plankton community. MEPS Vol. 647 https://doi.org/10.3354/meps13390.

Bolaños LM, Karp-Boss L, Choi C-J, Worden AZ, Graff JR, Haëntjens N, Chase AP, Della Penna A, Gaube P, Morison F, Menden-Deuer S, Westberry TK, O’Malley RT, Boss E, Behrenfeld MJ & SJ Giovannoni (2020). Small phytoplankton dominate western North Atlantic biomass. The ISME Journalhttps://doi.org/10.1038/s41396-020-0636-0.

Faktorová D, Nisbet RER, Fernández JA, Casacuberta E, Sudek L, Allen AE, Ares M, Aresté C, Balestreri C, Barbrook AC, Beardslee B, Bender S, Booth DS, Bouget F-Y, Bowler C, Breglia SA, Brownlee C, Burger G, Cerutti H, Cesaroni R, Chiurillo MA, Clemente T, Coles DB, Collier JL, Cooney EC, Coyne K, Docampo R, Dupont CL, Edgcomb V, Einarsson E, Elustondo PA, Federici F, Freire-Beneitez V, Freyria NJ, Fukuda K, García PA, Girguis PR, Gomaa F, Gornik SG, Guo J, Hampl V, Hanawa Y, Haro-Contreras ER, Hehenberger H, Highfield A, Hirakawa Y, Hopes A, Howe CJ, Hu I, Ibañez J, Irwin NAT, Ishii Y, Janowicz NE, Jones AC, Kachale A, Fujimura-Kamada K, Kaur B, Kaye JZ, Kazana E, Keeling PJ, King N, Klobutcher LA, Lander N, Lassadi I, Li Z, Lin S, Lozano J-C, Luan F, Maruyama S, Matute T, Miceli C, Minagawa J, Moosburner M, Najle SR, Nanjappa D, Nimmo IC, Noble L, Novák Vanclová AMG, Nowacki M, Nuñez I, Pain A, Piersanti A, Pucciarelli S, Pyrih J, Rest JS, Rius M, Robertson D, Ruaud A, Ruiz-Trillo I, Sigg MA, Silver PA, Slamovits CH, Smith GJ, Sprecher BN, Stern R, Swart E, Tsaousis A, Tsypin L, Turkewitz A, Turnšek J, Valach M, Vergé V, von Dassow P, von der Haar T, Waller RF, Wang L, Wen X, Wheeler G, Woods A, Zhang H, Mock* T, Worden* AZ & J Lukeš* (2020). Genetic tool development in marine protists: Emerging model organisms for experimental cell biology. Nature Methods Vol 17:481-494. Co-first authors, *Co-corresponding.

Rodríguez-Martínez R, Leonard G, Milner DS, Sudek S, Conway M, Moore K, Hudson T, Mahé F, Keeling PJ, Santoro AE, Worden AZ & TA Richards (2020). Controlled sampling of ribosomally active protistan diversity in marine sediment-surface layers identifies putative players in the carbon sink. The ISME Journal Vol 14:984-998.

Wideman JG, Monier A, Rodríguez-Martínez R, Leonard G, Cook E, Poirier C, Maguire F, Milner D, Irwin NAT, Moore K, Santoro AE, Keeling PJ, Worden AZ & TA Richards (2020). Unexpected mitochondrial genome diversity revealed by targeted single-cell genomics of heterotrophic flagellated protists. Nature Microbiology Vol 5:154-165.

Zimmerman AE, Howard-Varona C, Needham DM, John S, Worden AZ, Sullivan MB, Waldbauer JR & ML Coleman (2020). Metabolic and biogeochemical consequences of viral infection in aquatic ecosystems. Nature Reviews MicrobiologyVol 18:21-34. https://doi.org/10.1038/s41579-019-0270-x

Hehenberger E, Eitel M, Fortunato SVA, Miller DJ, Keeling PJ and MA Cahill. Early eukaryotic origins and metazoan elaboration of MAPR family proteins. Molecular Phylogenetics and Evolution. Volume 148, July 2020, 106814. https://doi.org/10.1016/j.ympev.2020.106814

Tikhonenkov DV, Hehenberger E, Esaulov AS, Belyakova OI, Mazei YA, Mylnikov AP and PJ Keeling. Insights into the origin of metazoan multicellularity from predatory unicellular relatives of animals.  BMC Biology. 18:39. https://doi.org/10.1186/s12915-020-0762-1


Needham DM, Poirier C, Hehenberger E, Jimenez V, Swalwell JE, Santoro AE & Worden AZ. Targeted metagenomic recovery of four divergent viruses reveals shared and distinctive characteristics of giant viruses of marine eukaryotes. Philosophical Transactions  of the Royal Society B. 374: 20190086. https://doi:10.1098/rstb.2019.0086

Wilken S, Choi CJ &Worden AZ. Contrasting mixotrophic lifestyles reveal different ecological niches in two closely related marine protists. Journal of Phycology.https://doi:10.1111/jpy.12920

Wilken S, Yung C, Hamilton M, Hoadley K, Nzongo J, Eckmann C, Corrochano Luque M, Poirier C & Worden AZ. The need to account for cell biology on characterising predatory mixotrophs in aquatic environments. Philosophical Transactions of the Royal Society B.  Vol 374: 20190090. http://dx.doi.org/10.1098/rstb.2019.0090

Needham DM, Yoshizawa S, Hosaka T, Poirier C, Choi CJ, Hehenberger E, Irwin NTA, Wilken S, Yung C-M, Bachy C, Kurihara R, Nakajima Y, Kojima K, Kimura-Someya T, Leonard G, Malmstrom RR, Mende D, Olson DK, Sudo Y, Sudek S, Richards TA, DeLong EF, Keeling PJ, Santoro AE, Shirouzu M, Iwasaki W & Worden AZ. A distinct lineage of giant viruses brings a rhodopsin photosystem to unicellular marine predators. Proceedings of the National Academy of Sciences USA. https://doi.org/10.1073/pnas.1907517116.

Kolody BJ, McCrow J, Zeigler Allen L, Aylward FO, Fontanez KM, Moustafa A, Moniruzzaman M, Chavez FP, Scholin CA, Allen EE, Worden AZ, DeLong EF. Diel transcriptional response of a California Current plankton microbiome to light, low iron, and enduring viral infection. The ISME Journal. doi: 10.1038/s41396-019-0472-2.https://www.nature.com/articles/s41396-019-0472-2

Zimmerman AE, Bachy C, Ma X, Roux S, Jang HB, Sullivan MB, Waldbauer JR & Worden AZ. Closely-related viruses of the marine picoeukaryotic alga, ostreococcus lucimarinus,exhibit different ecological strategies. Environmental Microbiology. Vol 21:2148-2170.https://doi.org/10.1111/1462-2920.14608

Okamoto N, Gawryluk RMR, del Campo J, Strassert JFH, Lukeš J, Richards TA, Worden AZ, Santoro AE &PJ Keeling. A Revised Taxonomy of Diplonemids Including the Eupelagonemidae n. fam. and a Type Species, Eupelagonema oceanica n. gen. &sp. The Journal of Eukaryotic Microbiology. Vol 66:519-524. https://doi.org/10.1111/jeu.12679.

del Campo J, Heger TJ, Rodríguez-Martínez R, Worden AZ, Richards TA, Massana R & PJ Keeling. Assessing the diversity and distribution of apicomplexans in host and free-living environments using high-throughput amplicon data and a phylogenetically informed reference framework. Frontiers in Microbiology.https://doi.org/10.3389/fmicb.2019.02373

Wahl M , Werner FJ, Buchholz B, Raddatz S, Graiff A, Matthiessen B, Karsten U, Hiebenthal C, Hamer J, Ito M, Gülzow E, Rilov G & T Guy-Haim. Season affects strength and direction of the interactive impacts of ocean warming and biotic stress in a coastal seaweed ecosystem. Limnology and Oceanography. https://doi.org/10.1002/lno.11350

Ribalet F, Berthiaume C, Hynes A, Swalwell J, Carlson M, Clayton S, Hennon G, Poirier C, Shimabukuro E, White A, & Armbrust EV. SeaFlow data v1, high-resolution abundance, size and biomass of small phytoplankton in the North Pacific. Scientific Data.

Ito M, Scotti M, Franz M, Barboza FR, Buchholz B, Zimmer M, Guy-Haim T & M Wahl. Effects of temperature on carbon circulation in macroalgal food webs are mediated by herbivores. Marine Biology. 166:158.

Carcedo MC, Fiori SM, Scotti M, Ito M, Dutto MS & ME Carbone. Dominant bivalve in an exposed sandy beach regulates community structure through spatial competition. Estuaries and Coasts. 42, 1912-1923.

Kluger LC, Scotti M, Vivar I & M Wolff. Specialization of fishers leads to greater impact of external disturbance: evidence from a social-ecological network modelling exercise for Sechura Bay, northern Peru. Ocean & Coastal Management. 179:104861.

Olmo Gilabert R, Navia AF, De La Cruz-Agüero G, Molinero JC, Sommer U & M Scotti. Body size and mobility explain species centralities in the Gulf of California food web. Community Ecology. 20:149-160.

Franz M, Barboza FR, Hinrichsen HH, Lehmann A, Scotti M, Hiebenthal C, Molis M, Schütt R & M Wahl. Long-term records of hard-bottom communities in the southwestern Baltic Sea reveal the decline of a foundation species. Estuarine, Coastal and Shelf Science. 219:242-251.

Picq S, Scotti M & O Puebla. Behavioural syndromes as a link between ecology and mate choice: a field study in a reef fish population. Animal Behaviour. 150:219-237.

Kiko R & H Hauss. On the Estimation of Zooplankton-Mediated Active Fluxes in Oxygen Minimum Zone Regions. Frontiers in Marine Science. 6:741.

Lombard F, Boss E, Waite AM, Vogt M, Uitz J, Stemmann L, Sosik HM, Schulz J, Romagnan JB, Picheral M, Pearlman J, Ohman MD, Niehoff B, Möller KO, Miloslavich P, Lara-Lpez A, Kudela R, Lopes RM, Kiko R, Karp-Boss L, Jaffe JS, Iversen MH, Irisson JO, Fennel K, Hauss H, Guidi L, Gorsky G, Giering SLC, Gaube P, Gallager S, Dubelaar G, Cowen RK, Carlotti F, Briseño-Avena C, Berline L, Benoit-Bird K, Bax N, Batten S, Ayata SD, Artigas LF & W Appeltans. Globally Consistent Quantitative Observations of Planktonic Ecosystems. Frontiers in Marine Science. 6: 196.

Hoving HJT , Christiansen S, Fabrizius E, Hauss H, Kiko R, Linke P, Neitzel P, Piatkowski U &A Körtzinger. The Pelagic In situ Observation System (PELAGIOS) to reveal biodiversity, behavior, and ecology of elusive oceanic fauna. Ocean Science. 15 . pp. 1327-1340.

Hehenberger E, Gast RJ & PJ Keeling. A kleptoplastidic dinoflagellate and the tipping point between transient and permanent plastid endosymbiosis. Proc National Acad Sci USA. 116(36): 17934-17942.

Gawryluk RMR†, Tikhonenkov DV†, Hehenberger E, Husník F, Mylnikov A P & PJ Keeling. Non-photosynthetic predators are sisters to red algae. Nature. 572: 240-243.† These authors contributed equally.

Mathur V†, Kolisko M†, Hehenberger E, Irwin NAT, Leander BS, Kristmundsson Á, Freeman MA & PJ Keeling. Multiple independent origins of apicomplexan-like parasites. Curr Biol. 29(17): R836-R839. † These authors contributed equally.



Strassert JFH, Hehenberger E, del Campo J,  Okamoto N,  Kolisko M, Richards TA,  Worden AZ,  Santoro AE & PJ Keeling. Phylogeny, evidence for a cryptic plastid, and distribution of Chytriodinium parasites (Dinophyceae) infecting copepods. Journal of Eukaryotic Microbiology.https://doi.org/10.1111/jeu.12701

Joo S, Wang MH, Lui G, Lee J, Barnas A, Kim E, Sudek S, Worden AZ & JH Lee. Common ancestry of heterodimerizing TALE homeobox transcription factors across Metazoa and Archaeplastida. BMC Biology. 16:136. doi: 10.1186/s12915-018-0605-5

Bachy C, Charlesworth CJ, Chan AM, Finke JF, Wong C-H, Wei C-L, Sudek S, Coleman ML, Suttle CA & AZ Worden. Transcriptional responses of the marine green alga Micromonas pusilla and an infecting prasinovirus under different phosphate conditions. Environmental Microbiology. Vol 20:2898-2912.

Guo J, Wilken S, Jimenez V, Choi CJ, Ansong CK, Dannebaum R, Sudek L, Milner D, Bachy C, Reistetter EN, Elrod VA, Klimov D, Purvine SO, Wei C-L, Kunde-Ramamoorthy G, Richards TA, Goodenough U, Smith RD, Callister SJ & AZ Worden. Specialized proteomic responses and an ancient photoprotection mechanism sustain marine green algal growth during phosphate limitation. Nature Microbiology. Vol 3:781–790.

Orsi WD, Wilken S, del Campo J, Heger T, James E, Richards TA, Keeling PJ, Worden AZ & AE. Identifying protist consumers of photosynthetic picoeukaryotes in the surface ocean using stable isotope probing. Environmental Microbiology. Vol 20:815-827. http://doi.org/10.1111/1462-2920.14018

Strassert JFH, Karnkowska A, Hehenberger E, del Campo J, Kolisko M, Okamoto N, Burki F, Janouškovec J, Poirier C, Leonard G, Hallam S, Richards TA, Worden AZ, Santoro A & PJ Keeling. Single cell genomics of uncultured Marine Alveolates (MALVs) shows paraphyly of basal dinoflagellates. The ISME Journal. Vol 12:304-308.



Gutowska MA, Shome B, McRose D, Sudek S, Hamilton M, Giovannoni SJ, Begley TP & AZ Worden. Ecologically important marine haptophyte algae use exogenous pyrimidine compounds more efficiently than thiamin. mBio Vol 8: e01459-17.doi: 10.1128/mBio.01459-17

Limardo AJ, Sudek S, Choi CJ, Poirier C, Rii YM, Blum M, Roth R, Goodenough U, Church MJ & AZ Worden. Quantitative biogeography of picoprasinophytes establishes differential ecotype and significant contributions to marine phytoplankton. Environmental Microbiology19:3219-3234. doi: 10.1111/1462-2920

Caron DA, Alexander H, Allen AE, Archibald JM, Armbrust EV, Bachy C, Bharti A, Dyhrman ST, Guida S, Heidelberg KB, Kaye JZ, Metzner J, Smith SR & AZ Worden. Probing the evolution, ecology and physiology of marine protists using transcriptomics. Nature Reviews Microbiology15:6-20. early online doi:10.1038/nrmicro.2016.160

Choi CJ, Bachy C, Jaeger GS, Poirier C, Sudek L, Sarma VVSS, Mahadevan A, Giovannoni SJ & AZ Worden. Newly discovered deep-branching marine plastid lineages are numerically rare but globally distributed. Current Biology 27:R17–R18. Co-first authors.

Cuvelier ML, Guo J, Ortiz AC, van Baren MJ, Tariq MA, Partensky F & AZ Worden. Responses of the picoprasinophyte Micromonas commoda to light and ultraviolet stress. PLoS One. 27 pgs. doi: 10.1371/journal.pone.0172135

Mock T, Otillar RP, Strauss J, Allen AE, Dupont CL, Frickenhaus S, Maumus F, McMullan M, A, Sanges R, Schmutz J, Toseland A, Veluchamy A, Ward BJ, Wu T, Barry KW, Falciatore A, Ferrante MI, Fortunato AE, Glöckner G, Gruber A, Hipkin R, Janech MG, Kroth P, Leese F, Lindquist EA, Lyon BR, Martin J, Mayer C, Parker M, Quesneville H, Raymond JA, Uhlig C, Valas RE, Valentin KU, Worden AZ, Armbrust EV, Bowler C, Green BR, Moulton V, van Oosterhout C & IV Grigoriev. Evolutionary genomics of a cold-adapted diatom, Fragilariopsis cylindrus. Nature 541:536-540. doi: 10.1038/nature20803

Clayton SC, Lin Y-C, Follows MJ & AZ Worden. Co-existence of distinct Ostreococcus ecotypes at an oceanic front. Limnology & Oceanography62:75–88.doi: 10.1002/lno.10373



Gawryluk R, del Campo J, Okamoto N, Strassert JFH, Lukes J, Richards TA, Worden AZ, Santoro AE & PJ Keeling. Morphological identification and single-cell genomics of marine diplonemids. Current Biology26:3053–3059.

Gregory AC, Solonenko S, LaButti K, Copeland A, Sudek S, Maitland A, Chittick L, dos Santos F, Weitz JS, Worden AZ, Woyke T & MB Sullivan. Genomic differentiation amongst wild cyanophages despite widespread horizontal gene transfer. BMC Genomics 17:930 (13 pgs). doi: 10.1186/s12864-016-3286-x

Orsi WD, Smith JM, Liu S, Liu Z, Sakamoto CM, Wilken S, Poirier C, Richards TA, Keeling PJ, Worden AZ & AE Santoro. Diverse, uncultivated bacteria and archaea underlying the cycling of dissolved protein in the ocean. The ISME Journal10:2158-2173. doi: 10.1038/ismej.2016.20

Simmons MP, Sudek S, Monier A, Limardo AJ, Jimenez V, Perle C, Elrod VA, Pennington JT & AZ Worden. Abundance and biogeography of picoprasinophyte ecotypes and other phytoplankton in the Eastern North Pacific Ocean. Applied & Environmental Microbiology 82:1693-1705.

van Baren MJ, Bachy C, Reistetter EN, Purvine SO, Grimwood J, Sudek S, Yu H, Poirier C, Kuo A, Grigoriev IV, Wong C-H, Smith RD, Schmutz J, Wei C-L, Callister SJ & AZ Worden. Evidence-based green algal genomics reveals marine diversity and ancestral characteristics of land plants. BMC Genomics 17:267 (22 pgs). doi: 10.1186/s12864-016-2585-6

Monier A, Worden AZ & TA Richards. Phylogenetic diversity and biogeography of the Mamiellophyceae lineage of eukaryotic marine phytoplankton across the oceans. Environmental Microbiology Reports 8:461-469. doi: 10.1111/1758-2229.12390

Waltman PH, Guo J, Reistetter EN, Purvine S, Ansong CK, van Baren MJ, Wong C-H, Wei C-H, Smith RD, Callister SJ, Stuart JM & AZ Worden. Identifying aspects of the post-transcriptional program governing the proteome of the green alga Micromonas pusilla. PLoS One. doi: 10.1371/journal.pone.0155839

Worden AZ & Wilken S. A plankton bloom shifts as the ocean warms. Science. Vol 354:287-288. doi: 10.1126/science.aaj1751.



RibaletF, SwalwellJE, ClaytonS, JimenezV, SudekS, LinY, JohnsonZI, WordenAZ & EV Armbrust.    Light-driven synchrony of Prochlorococcus growth and mortality in the north Pacific gyre. Proceedings of the National Academy of Sciences, USA 112:8008–8012.

Simmons MP, Bachy C, Sudek S, Baren MJ, Sudek L, Ares M & AZ Worden. Intron invasions trace algal speciation and reveal nearly identical Arctic and Antarctic Micromonas populations. Molecular Biology and Evolution 32:2219-2235.

Worden AZ, Follows MJ, Giovannoni SJ, Wilken S, Zimmerman AE & PJ Keeling. Rethinking the marine carbon cycle: factoring in multifarious lifestyles of microbes. Science 347. doi: 10.1126/science.1257594

Derelle E, Monier A, Cooke R, Worden AZ, Grimsely NH & H Moreau. Diversity of viruses infecting the green micro-alga Ostreococcus lucimarinus. Journal of Virology 89:5812-21.

OrsiWD, SmithJM, WilcoxHM, SwalwellJE, CariniP, WordenAZ & AE Santoro. Ecophysiology of uncultivated marine euryarchaea is linked to particulate organic matter. The ISME Journal 9: 1747-1763.

Sudek S, Everroad RC, GehmanA, SmithJM, Poirier C, ChavezFP & AZ Worden. Cyanobacterial distributions along a physico-chemical gradient in the northeastern Pacific Ocean. Environmental Microbiology 17:3692-3707.

ŠevčíkováT, HorákA, KlimešV, ZbránkováV, Demir-HiltonE, SudekS, JenkinsJ, SchmutzJ, PřibylP, FousekJ, VlčekC, LangBF, OborníkM, Worden*AZ & M Eliáš*. Updating algal evolutionary relationships through plastid genome sequencing: did alveolate plastids emerge through endo-symbiosis of an ochrophyte? Scientific Reports 5: doi: 10.1038/srep10134. *co-corresponding.

Limardo AJ &Worden AZ, News and Views, Microbiology: Exclusive networks in the sea. Nature. Vol 522:36-37. doi 10.1038/nature14530.



McRose D, Guo J, Monier A, Sudek S, Wilken S, Yan S, Mock T, Archibald JM, Begley TP, Reyes-A & AZ Worden. Alternatives to Vitamin B1 uptake revealed with discovery of riboswitches in multiple marine unicellular eukaryotes. The ISME Journal 17:3692-3707. Co-first authors.

Duanmu D, Bachy C, SudekS, WongCH, JimenezV, RockwellNC, MartinSS, NganCY, ReistetterE, van BarenMJ, PriceDC, WeiCL, Reyes-PrietoA, Lagarias* JC & AZ Worden*. Marine algae and plants share conserved phytochrome signaling systems. Proceedings of the National Academy of Sciences, USA doi: 111:15827–15832. Co-first authors, *co-corresponding.

Keeling* PJ, Burki F, Wilcox HM, Allam B, Allen EE, 75 others & AZ Worden*. The Marine Microbial Eukaryote Transcriptome Sequencing Project (MMETSP): Illuminating the Functional Diversity of Eukaryotic Life in the Oceans through Transcriptome Sequencing. PLoS Biology 12(6): e1001889. *co-corresponding.

Rockwell, NC, Duanmu D, Martin SS, Bachy C, Price DC, Bhattacharya D, Worden AZ & JC Lagarias. Eukaryotic algal phytochromes span the visible spectrum. Proceedings of the National Academy of Sciences, USA 11:3871-3876.

Worden AZ. Women in Oceanography - A Decade Later. Autobiographical Sketches - Alexandra Z. Worden. TOS Oceanography. Vol. 27, No.4 Supplement, pg. 251.

Bachy, C & Worden AZ. Microbial ecology: Finding structure in the rare biosphere. Current biology. Vol 24:R315-R317.



ReadBA, Kegel J, Klute MJ, Kuo A, Lefebvre SC, Maumus F, Mayer C, Miller J, Monier A, Salamov A, Emiliania huxleyi Annotation Consortium, Dacks JB, Delwiche CF, Dyhrman ST, Glöckner G, John U, Richards TA, Worden AZ, Zhang X and IV Grigoriev. Emiliania’s pan genome drives the global distribution. Nature 499: 209-213.Co-senior authors.

Monier A, Sudek S, Fast N & AZ Worden. Gene invasion in distant eukaryotic lineages: discovery of mutually exclusive genetic elements reveal marine biodiversity. The ISME Journal 7:1764–1774.

Santoro AE, Sakamoto CM, Smith JM, Plant JN, Gehman AL, Worden AZ, Johnson KS, Francis CA & KL Casciotti. Measurements of nitrite production and nitrite-producing organisms in and around the primary nitrite maximum in the central California Current. Biogeosci. Disc. 10:5803-5840.

Vergin KL, Beszteri B, Monier A, Thrash JC, Treusch AH, Kilpert F, Worden AZ & SJ Giovannoni. High resolution SAR11 ecotype dynamics at the Bermuda Atlantic Time-series Study site as determined by phylogenetic placement of pyrosequences. The ISME Journal 7:1322-1332.



Worden AZ, JanouskovecJ, McRose D, Engman A, WelshRM, MalfattiS, TringeSG & PJ Keeling. Global distribution of a wild alga revealed by targeted metagenomics. Current Biology 22:R675-R677.

Curtis BA, and 72 others (including McRose D & AZ Worden). Algal genomes reveal evolutionary mosaicism and the fate of nucleomorphs. Nature 492:59–65.

Lin YC, Campbell T, Chung CC, Gong G, Chiang KP & AZ Worden. Distribution patterns and phylogeny of novel marine stramenopiles (MAST) in the North Pacific Ocean.Applied & Environmental Microbiology 78:3387-3399.

Monier A, Welsh RM, Gentemann C, Weinstock G, Sodergren E, Armbrust EV, Eisen JA & AZ. Phosphate transport in marine phytoplankton and their viruses: cross-domain commonalities in viral-host gene exchanges. Environmental Microbiology14:162–176.

Treusch AH, Demir-Hilton E, Vergin KL, Worden AZ, Carlson CA & SJ Giovannoni. Phytoplankton distribution patterns in the northwestern Sargasso Sea revealed by small subunit rRNA genes from plastids. The ISME Journal6:481-492.



Paerl RW, JohnsonKS, WelshRM, WordenAZ, Chavez FP & JP Zehr. Differential distributions of Synechococcus subgroups across the California Current System. Frontiers in Microbiology. doi: 10.3389/fmicb.2011.00059

Demir-Hilton E, Sudek S, Cuvelier ML, Gentemann C, Zehr JP & AZ Worden. Global distribution patterns of distinct clades of the photosynthetic picoeukaryote Ostreococcus. The ISME Journal5:1095–1107.

Kim E, Harrison J, Sudek S, Jones MDM, Wilcox HM, Richards T*, Worden AZ* & JM Archibald*. Newly identified and diverse plastid-bearing branch on the eukaryotic tree of life. Proceedings of the National Academy of Sciences, USA 108:1496-1500. *co-corresponding.

Worden AZ, Dupont C & AE Allen. GEnomics of uncultured eucaryotes: sorting FACS from fiction. Genome Biology. 12:117. (3 pages).



Cuvelier ML, Allen AE, Monier A, McCrow JP, Messié M, Tringe SG, Woyke T, Welsh RM, Ishoey T, Lee JH, Binder BJ, DuPont CL, Latasa M, Guigand C, Buck KR, Hilton J, Thiagarajan M, Caler E, Read B, Lasken RS, Chavez FP & AZ Worden. Targeted metagenomics and ecology of globally important uncultured eukaryotic phytoplankton. Proceedings of the National Academy of Sciences, USA 107:14679-14684.

McDonald SM, PlantJN & AZ Worden. The mixed lineage nature of nitrogen transport and assimilation in marine eukaryotic phytoplankton: a case study of Micromonas. Molecular Biology and Evolution27:2268-2283.

Worden AZ & AE Allen. The voyage of he microbial eukaryote. Current Opinion in Microbiology. Vol 13:652-660.



Caron DA, Worden AZ, Countway PD, Demir E & KB Heidelberg. Protists are microbes too: a perspective. The ISME Journal 3:4-12.

Worden AZ, Lee JH, Mock T, Rouze P, Simmons MP, Aerts AL, Allen AE, Cuvelier ML, Derelle E, Everett MV, Foulon E, Grimwood J, Gundlach H, Henrissat B, Napoli C, McDonald SM, Parker MS, Rombauts S, Salamov A, Von Dassow P, Badger JH, Coutinho PM, Demir E, Dubchak I, Gentemann C, Eikrem W, Gready JE, John U, Lanier W, Lindquist EA, Lucas S, Mayer KFX, Moreau H, Not F, Otillar R, Panaud O, Pangilinan J, Paulsen I, Piegu B, Poliakov S, Robbens S, Schmutz J, Toulza E, Wyss T, Zelensky A, Zhou K, Armbrust EV, Bhattacharya D, Goodenough UW, Van de Peer Y & IV Grigoriev. Green evolution and dynamic adaptations revealed by genomes of the marine picoeukaryote Micromonas. Science 324:268-272.

Worden AZ & D Mc Rose. the otherness of the oceans. Nature. Vol 459:166-167.



Guillou L, Viprey M, Chambouvet A, Welsh RM, Kirkham AR, Massana R, Scanlan DJ & AZ Worden. Widespread occurrence and genetic diversity of marine parasitoids belonging to Syndiniales (Alveolata). Environmental Microbiology 10:3349-3365.

Deschamps P, Moreau H, Worden AZ, Dauvillée D & SG Ball. Early gene duplication within chloroplastida and its correspondence with relocation of starch metabolism to chloroplasts. Genetics178:2373-2387.

Cuvelier ML, Ortiz A, Kim E, Moehlig H, Richardson DE, Heidelberg JF, Archibald JM & AZ Worden. Widespread distribution of a unique marine protistan lineage. Environmental Microbiology 10:1621–1634.

Worden AZ & F Not.Ecology and diversity of picoeukaryotes. In: Microbial Ecology of the Ocean, 2nd Edition. Ed. D. Kirchman. Wiley.



Robbens S, Rouze P. Cock JM, Spring J, Worden* AZ & Y Van de Peer*. The FTO gene, implicated in human obesity, is found only in vertebrates and marine algae. Journal of Molecular Evolution 66:80-84. *co-corresponding.

Not F, Gausling R, Azam F, Heidelberg, JF & AZ Worden. Vertical distribution of picoeukaryotic  diversity in the Sargasso Sea. Environmental Microbiology 9:1233-1252.

Sherr BF, Sherr EB, Caron DA, Vaulot D & AZ Worden. Oceanic protists. Oceanography20:130-134.



Worden AZ, Cuvelier ML & DH Bartlett. In depth marine microbial community genomics. Trends in Microbiology 14:331-336.

Derelle E, Ferraz C, Rombauts S, Rouzé P, Worden AZ, Robbens S, Partensky F, Degroeve S, Echeynié S, Cooke R, Wuyts J, Saeys Y, Jabbari K, Bowler C, Ball S, Ral JP, Bouget FY, Piganeau G, De Baets B, Picard A, Delseny M, Demaille J, Van de Peer Y & H Moreau (2006). Genome analysis of the smallest free-living eukaryote Ostreococcus tauri unveils many unique features. Proceedings of the National Academy of Sciences, USA 103:11647-11652.

Worden AZ. Picoeukaryote diversity in coastal waters of the Pacific Ocean. Aquatic Microbial Ecology 43:165-175.

Worden AZ, Seidel M, Smriga S, Wick A, Malfatti F, Bartlett D & F Azam. Trophic regulation of Vibrio cholerae in coastal marine waters.Environmental Microbiology 8:21-29.

Worden AZ, Cuvelier ML & DH Bartlett. In depth marine microbial community genomics. Trends in Microbiology. Vol. 14:331-336.



Six C, Worden AZ, Rodriguez F, Moreau H & F Partensky. New insights into the nature and of prasinophyte antennae proteins: Ostreococcus tauri, a case study. Molecular Biology and Evolution 22:2217-2230. Co-first authors.



Azam F & AZ Worden. Microbes, molecules and marine ecosystems. Science 303:1622-1624.

Worden AZ, Nolan JK & B Palenik. Assessing the dynamics and ecology of marine picophyto-plankton: the importance of the eukaryotic component. Limnology & Oceanography 49:168-179.



Mouriño RR, Worden AZ & F Azam. Growth of Vibrio cholerae in Red Tide waters off California. Applied and Environmental Microbiology 69:6923-6931.

Worden AZ & BJ Binder. Growth regulation of rRNA content of Prochlorococcus and Synechococcus (marine cyanobacteria) measured by whole-cell hybridization of rRNA targeted peptide nucleic acids. Journal of Phycology 39:527-534.

Worden AZ & BJ Binder. Application of dilution experiments for measuring growth and mortality rates among Prochlorococcus and Synechococcus populations in oligotrophic environments. Aquatic Microbial Ecology 30:159-174.



Worden AZ, Chisholm SW & BJ Binder. In situ hybridization of Prochlorococcus and Synechococcus (Marine Cyanobacteria) with rRNA-targeted Peptide Nucleic Acid Probes. Applied and Environmental Microbiology 66:284-289.



Binder BJ, Chisholm SW, Olson RJ, Frankel SL & AZ Worden. Dynamics of pico-phytoplankton, ultra-phytoplankton, and bacteria in the central equatorial Pacific. Deep-Sea Research Part II 43:907-931.