{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,11,11]],"date-time":"2025-11-11T22:38:50Z","timestamp":1762900730581,"version":"3.41.2"},"reference-count":46,"publisher":"The Company of Biologists","issue":"15","license":[{"start":{"date-parts":[[2025,7,30]],"date-time":"2025-07-30T00:00:00Z","timestamp":1753833600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100006260","name":"Technion \u2013 Israel Institute of Technology","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100006260","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2025,8,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n               <jats:p>Sessile marine organisms rely on water flows to enhance mass transfer and facilitate key physiological processes. Pulsating corals exhibit rhythmic pulsation of their tentacles to generate flow that removes excess oxygen produced during photosynthesis, thereby enhancing the process. The ejected flow is counterbalanced by an incoming flow directed toward the polyp, potentially delivering essential nutrients. This mechanism may be crucial for these corals, which rely on epidermal nutrient uptake rather than zooplankton predation. The characteristics of the incoming flow and its interactions with the coral tissue, where mass transfer occurs, are largely unknown. Here, we characterize the origin of new water approaching the polyp, the pathway it takes, where on the polyp and when during the pulsation period this interaction occurs. We used particle image velocimetry on single polyps to measure the flow field around the polyp and reconstruct the trajectories of the incoming water. We found that incoming water primarily originates from below the polyp. Eighty percent of the new water interacts with the polyp during the downward stroke of the tentacle motion, and 75% of the new water contacts the aboral face. We used a conservation of mass analysis to estimate the flow between the tentacles and found significant bidirectional flow through the gaps. Pulsation draws in \u223c26,800 polyp volumes of new water daily, containing sufficient nitrogen to fully meet the polyp's estimated daily demand for growth. Increased nutrient uptake may explain the persistence of pulsation during the night when photosynthesis ceases.<\/jats:p>","DOI":"10.1242\/jeb.250262","type":"journal-article","created":{"date-parts":[[2025,7,18]],"date-time":"2025-07-18T09:15:00Z","timestamp":1752830100000},"source":"Crossref","is-referenced-by-count":2,"title":["Spatio-temporal patterns of the incoming water flow in pulsating corals"],"prefix":"10.1242","volume":"228","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6753-714X","authenticated-orcid":false,"given":"Dror","family":"Malul","sequence":"first","affiliation":[{"name":"Faculty of Civil and Environmental Engineering, Technion IIT 1 , Haifa 32000 ,","place":["Israel"]},{"name":"The Interuniversity Institute for Marine Sciences 2 , PO Box\u00a0469, Eilat 88103 ,","place":["Israel"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2334-2551","authenticated-orcid":false,"given":"Roi","family":"Holzman","sequence":"additional","affiliation":[{"name":"School of Zoology 3 , Faculty of Life Sciences , , Tel Aviv 69978 ,","place":["Israel"]},{"name":"Tel Aviv University 3 , Faculty of Life Sciences , , Tel Aviv 69978 ,","place":["Israel"]},{"name":"The Interuniversity Institute for Marine Sciences 2 , PO Box\u00a0469, Eilat 88103 ,","place":["Israel"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7757-1237","authenticated-orcid":false,"given":"Uri","family":"Shavit","sequence":"additional","affiliation":[{"name":"Faculty of Civil and Environmental Engineering, Technion IIT 1 , Haifa 32000 ,","place":["Israel"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"237","published-online":{"date-parts":[[2025,7,30]]},"reference":[{"key":"2025073003424581600_JEB250262C1","doi-asserted-by":"publisher","first-page":"1116","DOI":"10.4319\/lo.1993.38.6.1116","article-title":"Flow patterns induced by substrata and body morphologies of benthic organisms, and their roles in determining availability of food particles","volume":"38","author":"Abelson","year":"1993","journal-title":"Limnol. Oceanogr."},{"key":"2025073003424581600_JEB250262C2","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1016\/S0022-5193(87)80201-1","article-title":"Paddles and rakes: fluid flow through bristled appendages of small organisms","volume":"129","author":"Cheer","year":"1987","journal-title":"J. Theor. Biol."},{"key":"2025073003424581600_JEB250262C3","first-page":"5559","article-title":"Modular interactive video object segmentation: Interaction-to-mask, propagation and difference-aware fusion","author":"Cheng","year":"2021"},{"key":"2025073003424581600_JEB250262C4","doi-asserted-by":"publisher","first-page":"28821","DOI":"10.1038\/srep28821","article-title":"Nitrogen and phosphorus uptake rates of different species from a coral reef community after a nutrient pulse","volume":"6","author":"Den Haan","year":"2016","journal-title":"Sci. Rep."},{"key":"2025073003424581600_JEB250262C5","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1086\/BBLv217n3p283","article-title":"Functional morphology and fluid interactions during early development of the scyphomedusa Aurelia aurita","volume":"217","author":"Feitl","year":"2009","journal-title":"Biol. Bull."},{"key":"2025073003424581600_JEB250262C6","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1007\/s00338-005-0055-8","article-title":"Water flow influences oxygen transport and photosynthetic efficiency in corals","volume":"25","author":"Finelli","year":"2006","journal-title":"Coral Reefs"},{"key":"2025073003424581600_JEB250262C8","first-page":"1381","article-title":"FFTW: an adaptive software architecture for the FFT","author":"Frigo","year":"1998"},{"key":"2025073003424581600_JEB250262C9","doi-asserted-by":"publisher","first-page":"68","DOI":"10.1016\/j.jembe.2009.08.003","article-title":"Feeding behavior, cirral fan anatomy, Reynolds numbers, and leakiness of Balanus glandula, from post-metamorphic juvenile to the adult","volume":"379","author":"Geierman","year":"2009","journal-title":"J. Exp. Mar. Biol. Ecol."},{"key":"2025073003424581600_JEB250262C10","doi-asserted-by":"publisher","first-page":"jeb222083","DOI":"10.1242\/jeb.222083","article-title":"Cool your jets: biological jet propulsion in marine invertebrates","volume":"224","author":"Gemmell","year":"2021","journal-title":"J. Exp. Biol."},{"volume-title":"Digital Image Processing","year":"2009","author":"Gonzalez","key":"2025073003424581600_JEB250262C11"},{"key":"2025073003424581600_JEB250262C12","doi-asserted-by":"publisher","first-page":"247","DOI":"10.2307\/1540115","article-title":"Reef corals: autotrophs or heterotrophs?","volume":"141","author":"Goreau","year":"1971","journal-title":"Biol. Bull."},{"key":"2025073003424581600_JEB250262C13","doi-asserted-by":"publisher","first-page":"jeb244520","DOI":"10.1242\/jeb.244520","article-title":"Multiscale flow between the branches and polyps of gorgonians","volume":"226","author":"Hamlet","year":"2023","journal-title":"J. Exp. Biol."},{"key":"2025073003424581600_JEB250262C14","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1016\/S0022-0981(96)02687-1","article-title":"Morphological variation in coral aggregations: branch spacing and mass flux to coral tissues","volume":"209","author":"Helmuth","year":"1997","journal-title":"J. Exp. Mar. Biol. Ecol."},{"key":"2025073003424581600_JEB250262C15","doi-asserted-by":"publisher","first-page":"e10483","DOI":"10.1002\/ece3.10483","article-title":"Physiology of the widespread pulsating soft coral Xenia umbellata is affected by food sources, but not by water flow","volume":"13","author":"Hill","year":"2023","journal-title":"Ecol. Evol."},{"key":"2025073003424581600_JEB250262C16","doi-asserted-by":"crossref","first-page":"e0225676","DOI":"10.1371\/journal.pone.0225676","article-title":"Effects of coral colony morphology on turbulent flow dynamics","volume":"15","author":"Hossain","year":"2020","journal-title":"PLoS ONE"},{"key":"2025073003424581600_JEB250262C17","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1111\/j.1469-185X.2008.00058.x","article-title":"Heterotrophy in tropical scleractinian corals","volume":"84","author":"Houlbr\u00e8que","year":"2009","journal-title":"Biol. Rev."},{"key":"2025073003424581600_JEB250262C18","doi-asserted-by":"publisher","first-page":"69","DOI":"10.3354\/meps034069","article-title":"The bivalve pump","volume":"34","author":"J\u00f8rgensen","year":"1986","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"2025073003424581600_JEB250262C19","doi-asserted-by":"publisher","first-page":"2328","DOI":"10.1103\/PhysRevLett.77.2328","article-title":"Effect of nutrient diffusion and flow on coral morphology","volume":"77","author":"Kaandorp","year":"1996","journal-title":"Phys. Rev. Lett."},{"key":"2025073003424581600_JEB250262C20","first-page":"952","article-title":"Effect of fluid viscosity on the liquid-feeding flow phenomena of a female mosquito","volume":"216","author":"Kim","year":"2013","journal-title":"J. Exp. Biol."},{"key":"2025073003424581600_JEB250262C21","doi-asserted-by":"publisher","first-page":"8978","DOI":"10.1073\/pnas.1301826110","article-title":"Benefit of pulsation in soft corals","volume":"110","author":"Kremien","year":"2013","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2025073003424581600_JEB250262C22","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1006\/jtbi.1994.1087","article-title":"The sponge pump","volume":"168","author":"Larsen","year":"1994","journal-title":"J. Theor. Biol."},{"key":"2025073003424581600_JEB250262C23","doi-asserted-by":"publisher","first-page":"jeb246541","DOI":"10.1242\/jeb.246541","article-title":"The fluid dynamics of barnacle feeding","volume":"227","author":"Maar","year":"2024","journal-title":"J. Exp. Biol."},{"key":"2025073003424581600_JEB250262C24","doi-asserted-by":"publisher","first-page":"20200180","DOI":"10.1098\/rspb.2020.0180","article-title":"Coral tentacle elasticity promotes an out-of-phase motion that improves mass transfer","volume":"287","author":"Malul","year":"2020","journal-title":"Proc. R. Soc. B"},{"key":"2025073003424581600_JEB250262C25","doi-asserted-by":"publisher","first-page":"2527","DOI":"10.1073\/pnas.0912348107","article-title":"Flow enhances photosynthesis in marine benthic autotrophs by increasing the efflux of oxygen from the organism to the water","volume":"107","author":"Mass","year":"2010","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2025073003424581600_JEB250262C26","doi-asserted-by":"publisher","first-page":"4125","DOI":"10.1242\/jeb.00649","article-title":"The ontogenetic scaling of hydrodynamics and swimming performance in jellyfish (Aurelia aurita)","volume":"206","author":"McHenry","year":"2003","journal-title":"J. Exp. Biol."},{"key":"2025073003424581600_JEB250262C27","doi-asserted-by":"publisher","first-page":"22135","DOI":"10.1038\/s41598-022-26325-5","article-title":"The widely distributed soft coral Xenia umbellata exhibits high resistance against phosphate enrichment and temperature increase","volume":"12","author":"Mezger","year":"2022","journal-title":"Sci. Rep."},{"key":"2025073003424581600_JEB250262C28","doi-asserted-by":"publisher","first-page":"583188","DOI":"10.3389\/fmars.2021.583188","article-title":"In situ pumping rate of 20 marine demosponges is a function of osculum area","volume":"8","author":"Morganti","year":"2021","journal-title":"Front. Mar. Sci."},{"key":"2025073003424581600_JEB250262C29","doi-asserted-by":"publisher","first-page":"1021679","DOI":"10.3389\/fmars.2023.1021679","article-title":"Xenia umbellata (Octocorallia): a novel model organism for studying octocoral regeneration ability","volume":"10","author":"Nadir","year":"2023","journal-title":"Front. Mar. Sci."},{"key":"2025073003424581600_JEB250262C30","doi-asserted-by":"publisher","first-page":"1027","DOI":"10.1007\/s00348-008-0488-2","article-title":"Primary and secondary vortical structures contribution in the entrainment of low Reynolds number jet flows","volume":"44","author":"Nastase","year":"2008","journal-title":"Exp. Fluids"},{"key":"2025073003424581600_JEB250262C31","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1146\/annurev-fluid-120710-101048","article-title":"Flow and transport in regions with aquatic vegetation","volume":"44","author":"Nepf","year":"2012","journal-title":"Annu. Rev. Fluid Mech."},{"key":"2025073003424581600_JEB250262C32","doi-asserted-by":"publisher","first-page":"657759","DOI":"10.3389\/fmicb.2021.657759","article-title":"Dissolved nitrogen acquisition in the symbioses of soft and hard corals with Symbiodiniaceae: a key to understanding their different nutritional strategies?","volume":"12","author":"Pupier","year":"2021","journal-title":"Front. Microbiol."},{"key":"2025073003424581600_JEB250262C33","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1098\/rsta.1895.0004","article-title":"IV. On the dynamical theory of incompressible viscous fluids and the determination of the criterion","volume":"186","author":"Reynolds","year":"1895","journal-title":"Philos. Trans. R. Soc. Lond. A"},{"key":"2025073003424581600_JEB250262C34","doi-asserted-by":"publisher","first-page":"726","DOI":"10.1038\/35099547","article-title":"Endoscopic exploration of red sea coral reefs reveals dense populations of cavity-dwelling sponges","volume":"413","author":"Richter","year":"2001","journal-title":"Nature"},{"key":"2025073003424581600_JEB250262C35","doi-asserted-by":"publisher","first-page":"129","DOI":"10.3354\/meps047129","article-title":"The ascidian pump: properties and energy cost","volume":"47","author":"Riisgard","year":"1988","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"2025073003424581600_JEB250262C36","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1007\/s11538-020-00741-y","article-title":"Collective pulsing in xeniid corals: part ii-using computational fluid dynamics to determine if there are benefits to coordinated pulsing","volume":"82","author":"Samson","year":"2020","journal-title":"Bull. Math. Biol."},{"key":"2025073003424581600_JEB250262C37","doi-asserted-by":"publisher","first-page":"jeb192518","DOI":"10.1242\/jeb.192518","article-title":"A novel mechanism of mixing by pulsing corals","volume":"222","author":"Samson","year":"2019","journal-title":"J. Exp. Biol."},{"key":"2025073003424581600_JEB250262C38","doi-asserted-by":"publisher","first-page":"2369","DOI":"10.1242\/jeb.053744","article-title":"Flow structure and transport characteristics of feeding and exchange currents generated by upside-down Cassiopea jellyfish","volume":"215","author":"Santhanakrishnan","year":"2012","journal-title":"J. Exp. Biol."},{"key":"2025073003424581600_JEB250262C39","doi-asserted-by":"publisher","first-page":"659","DOI":"10.1093\/icb\/22.3.659","article-title":"Nutritional strategies of cnidarians: the absorption, translocation and utilization of dissolved nutrients by Heteroxenia fuscescens","volume":"22","author":"Schlichter","year":"1982","journal-title":"Am. Zool."},{"key":"2025073003424581600_JEB250262C40","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1007\/s002270050328","article-title":"Water flow and prey capture by three scleractinian corals, Madracis mirabilis, Montastr","volume":"131","author":"Sebens","year":"1998","journal-title":"Mar. Biol."},{"article-title":"The Israel National Monitoring Program in the Northern Gulf of Aqaba. Interuniversity Institute for Marine Sciences in Eilat","year":"2023","author":"Shaked","key":"2025073003424581600_JEB250262C41"},{"key":"2025073003424581600_JEB250262C42","doi-asserted-by":"publisher","first-page":"20180448","DOI":"10.1098\/rsif.2018.0448","article-title":"The role of turbulent hydrodynamics and surface morphology on heat and mass transfer in corals","volume":"15","author":"Stocking","year":"2018","journal-title":"J. R. Soc. Interface"},{"key":"2025073003424581600_JEB250262C43","doi-asserted-by":"publisher","first-page":"755","DOI":"10.1007\/s00227-009-1359-y","article-title":"A comparison of filtration rates among pelagic tunicates using kinematic measurements","volume":"157","author":"Sutherland","year":"2010","journal-title":"Mar. Biol."},{"year":"2004","author":"Sveen","key":"2025073003424581600_JEB250262C44"},{"key":"2025073003424581600_JEB250262C45","doi-asserted-by":"publisher","first-page":"e0294470","DOI":"10.1371\/journal.pone.0294470","article-title":"Short-term ocean acidification decreases pulsation and growth of the widespread soft coral Xenia umbellata","volume":"18","author":"Tilstra","year":"2023","journal-title":"PLoS ONE"},{"key":"2025073003424581600_JEB250262C46","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1016\/j.jbiomech.2018.02.011","article-title":"The fluid dynamics of Balanus glandula barnacles: adaptations to sheltered and exposed habitats","volume":"71","author":"Vo","year":"2018","journal-title":"J. Biomech."},{"key":"2025073003424581600_JEB250262C47","doi-asserted-by":"publisher","first-page":"1018","DOI":"10.1038\/s41586-023-06442-5","article-title":"Reef-building corals farm and feed on their photosynthetic symbionts","volume":"620","author":"Wiedenmann","year":"2023","journal-title":"Nature"}],"container-title":["Journal of Experimental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.250262\/3657514\/jeb250262.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.250262\/3658850\/jeb250262.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.250262\/3658850\/jeb250262.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,7,30]],"date-time":"2025-07-30T07:42:54Z","timestamp":1753861374000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jeb\/article\/228\/15\/jeb250262\/368730\/Spatio-temporal-patterns-of-the-incoming-water"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,30]]},"references-count":46,"journal-issue":{"issue":"15","published-print":{"date-parts":[[2025,8,1]]}},"URL":"https:\/\/doi.org\/10.1242\/jeb.250262","relation":{},"ISSN":["0022-0949","1477-9145"],"issn-type":[{"type":"print","value":"0022-0949"},{"type":"electronic","value":"1477-9145"}],"subject":[],"published":{"date-parts":[[2025,7,30]]},"article-number":"jeb250262"}}