Marine microbiome - Knowledge

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settlement cue. For example, the settlement of zoospores from the green alga Ulva intestinalis onto the biofilms of specific bacteria is mediated by their attraction to the quorum-sensing molecule, acyl-homoserine lactone, secreted by the bacteria. Classic examples of marine host–microbe developmental dependence include the observation that algal cultures grown in isolation exhibited abnormal morphologies and the subsequent discovery of morphogenesis-inducing compounds, such as thallusin, secreted by epiphytic bacterial symbionts. Bacteria are also known to influence the growth of marine plants, macroalgae, and phytoplankton by secreting phytohormones such as indole acetic acid and cytokinin-type hormones. In the marine choanoflagellate Salpingoeca rosetta, both multicellularity and reproduction are triggered by specific bacterial cues, offering a view into the origins of bacterial control over animal development (reviewed by Woznica and King. The benefit to the bacteria, in return, is that they receive physical space to colonize at particular points in the water column typically accessible only to planktonic microbes. Perhaps the best-studied example of intimate host–microbe interactions controlling animal development is the Hawaiian bobtail squid Euprymna scolopes. It lives in a mutualistic symbiosis with the bioluminescent bacteria Aliivibrio fischeri. The bacteria are fed a solution of sugars and amino acids by the host and, in return, provide bioluminescence for countershading and predator avoidance. This mutualism with microbes provides a selective advantage for the squid in predator–prey interactions. Another invertebrate example can be found in tubeworms, in which Hydroides elegans metamorphosis is mediated by a bacterial inducer and mitogen-activated protein kinase (MAPK) signaling in biofilms.

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by sequestering nutrients in the form of polyphosphate granules in the tissue of their host and nitrogen cycling, e.g., through nitrification, denitrification, and ammonia oxidation.]. Many macroalgal-associated bacteria are specifically adapted to degrade complex algal polysaccharides (e.g., fucoidan, porphyran, and laminarin ) and modify both the quality and quantity of organic carbon supplied to the ecosystem. The sulfur-oxidizing gill endosymbionts of lucinid clams contribute to primary productivity through chemosynthesis and facilitate the growth of seagrasses (important foundation species) by lowering sulfide concentrations in tropical sediments. Gammaproteobacterial symbionts of lucinid clams and stilbonematid nematodes were also recently shown to be capable of nitrogen fixation (bacterial symbiont genomes encode and express nitrogenase genes, highlighting the role of symbiotic microbes in nutrient cycling in shallow marine systems.

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and have a relatively simple body plan that commonly associates with bacteria, archaea, algal protists, fungi, and viruses. Sponge microbiomes are composed of specialists and generalists, and complexity of their microbiome appears to be shaped by host phylogeny. Studies have shown that the sponge microbiome contributes to nitrogen cycling in the oceans, especially through the oxidation of ammonia by archaea and bacteria. Most recently, microbial symbionts of tropical sponges were shown to produce and store polyphosphate granules, perhaps enabling the host to survive periods of phosphate depletion in oligotrophic marine environments. The microbiomes of some sponge species do appear to change in community structure in response to changing environmental conditions, including temperature and ocean acidification, as well as synergistic impacts.

1609: 1360: 1521:, and cellular debris derived from the linings of the airways which, when released into the relatively cooler outdoor air, condense to form a visible mass of vapor, which can be collected. There are various methods for collecting exhaled breath samples, one of the most recent is through the use of aerial drones. This method provides a safer, quieter, and less invasive alternative and often a cost-effective option for monitoring fauna and flora. Once obtained, the blow samples are taken to the laboratory and we proceed with the amplification and sequencing of the respiratory tract microbiota. The use of aerial drones has been more successful with large cetaceans due to slow swim speeds and larger blow sizes. 1351:

bacterial endosymbionts that reside upon coordinated use of sulfur present in the environment. This system has benefited from some of the most sophisticated 'omics and visualization tools. For example, multi-labeled probing has improved visualization of the microbiome and transcriptomics and proteomics have been applied to examine host–microbiome interactions, including energy transfer between the host and microbes and recognition of the consortia by the worm's innate immune system. The major strength of this system is that it does offer the ability to study host–microbiome interactions with a low diversity microbial consortium, and it also offers a number of host and microbial genomic resources

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bacteriostatic antibiotics but also compounds like halogenated furanones, cyclic dipeptides, and acyl-homoserine lactone mimics that disrupt bacterial quorum sensing and inhibit biofilm formation. The bacteria likely are able to utilize the carbon-rich exudates from their hosts. For example, in the case of giant kelp, the alga emits approximately 20% of primary production as dissolved organic carbon. Whereas these prior examples illustrate how the microbiomes can protect hosts from surface colonization, a similar phenomenon has also been observed internally in the shipworm Bankia setacea, in which symbionts produce a boronated

1581: 717: 926: 786:, which themselves can vary throughout the ontogeny of the host and as a result of environmental perturbations. Rather than host-associated microbes functioning independently, complex multi-assemblage microbiomes have major impact on the fitness and function of their hosts. Studying these complex interactions and biological outcomes is difficult, but to understand the origin and evolution of organisms and populations and the structure and function of communities and ecosystems, the understanding of symbioses in host–microbiome systems needs advancing. 894:, while the coral provides the algae with a protected environment and limiting compounds (e.g., nitrogen species) needed for photosynthesis. However, this is a classic example of a mutualistic symbiosis that is sensitive to environmental disturbances, which can disrupt the fragile interactions between host and microbe. When reefs become warm and eutrophic, mutualistic Symbiodiniaceae may induce cellular damage to the host and/or sequester more resources for their own growth, thereby injuring and parasitizing their hosts. 1459: 1595: 1482: 732: 656: 105: 1138: 790:

biodiversity (i.e., the richness of species and their interactions) pervasively influences the functioning of Earth's ecosystems, including ecosystem productivity. However, this research has focused almost exclusively on macroorganisms. Because microbial symbionts are integral parts of most living organisms, the understanding of how microbial symbionts contribute to host performance and adaptability needs broadening.

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framework from the fields of microbiology, evolutionary biology, community ecology, and oceanography. Individual taxa within the microbiome may help hosts withstand a wide range of environmental conditions, including those predicted under scenarios of climate change. Next, we explore two different avenues of how interdisciplinary collaborations could advance this line of research.

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surrounding environment. Knowing the microbiome of the skin of marine mammals under ''normal'' conditions has allowed us to understand how these communities are different from the free microbial communities found in the sea and how they can change according to abiotic and biotic variations, and also ''

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The access of microbial samples from the gut out of marine mammals is limited because most species are rare, endangered, and deep divers. There are different techniques for sampling the cetacean's gut microbiome. The most common is collecting fecal samples from the environment and taking a probe from

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Studies have also suggested that resident bacteria, archaea, and fungi additionally contribute to nutrient and organic matter cycling within the coral, with viruses also possibly playing a role in structuring the composition of these members, thus providing one of the first glimpses at a multi-domain

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are common members of the ocean's diverse benthic habitats and their abundance and ability to filter large volumes of seawater have led to the awareness that these organisms play critical roles in influencing benthic and pelagic processes in the ocean. They are one of the oldest lineages of animals,

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are one of the more common examples of an animal host whose symbiosis with microalgae can turn to dysbiosis, and is visibly detected as bleaching. Coral microbiomes have been examined in a variety of studies, which demonstrate how variations in the ocean environment, most notably temperature, light,

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organisms. Along with the coral–Symbiodiniaceae mutualism, this sponge-bacterial symbiosis helps explain Darwin's paradox, i.e., how highly productive coral reef ecosystems exist within otherwise oligotrophic tropical seas. Some sponge symbionts play a significant role in the marine phosphorus cycle

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Extending beyond nutritional symbioses, microbial symbionts can alter the reproduction, development, and growth of their hosts. Specific bacterial strains in marine biofilms often directly control the recruitment of planktonic larvae and propagules, either by inhibiting settlement or by serving as a

49:. The potential for microbiomes to influence the health, physiology, behavior, and ecology of marine animals could alter current understandings of how marine animals adapt to change, and especially the growing climate-related and anthropogenic-induced changes already impacting the ocean environment. 1504:

are in danger because they are affected by multiple stress factors which make them more vulnerable to various diseases. These animals have been noted to show high susceptibility to airway infections, but very little is known about their respiratory microbiome. Therefore, the sampling of the exhaled

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is one of the best studied symbiotic relationships in the sea and is a choice system for general symbiosis research. This relationship has provided insight into fundamental processes in animal-microbial symbioses, and especially biochemical interactions and signaling between the host and bacterium.

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Relationships are generally thought to exist in a symbiotic state, and are normally exposed to environmental and animal-specific factors that may cause natural variations. Some events may change the relationship into a functioning but altered symbiotic state, whereas extreme stress events may cause

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There are many outstanding questions in ecology and evolution that could be addressed by expanding the phylogenetic and ecological breadth of host-associated microbiome studies, including all possible interactions throughout the microbiome. There is strong empirical evidence and new consensus that

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is another relatively well-studied marine host to microbes. These three centimetre long worms reside within shallow marine sediments of the Mediterranean Sea. The worms do not contain a mouth or a digestive or excretory system, but are instead nourished with the help of a suite of extracellular

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Host-associated microbiomes also influence biogeochemical cycling within ecosystems with cascading effects on biodiversity and ecosystem processes. For example, microbial symbionts comprise up to 40% of the biomass of their sponge hosts. Through a process termed the "sponge-loop," they convert

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of marine macroalgae excrete a diverse chemical arsenal capable of selectively shaping further bacterial colonization and deterring the settlement of biofouling marine invertebrates such as bryozoans. As in corals, these diverse, microbially secreted compounds include not only bactericidal and

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group) These illustrate the incorporation of various new biochemical functions, such as photosynthesis, nitrogen fixation and recycling, and methanogenesis, into protist hosts by endosymbionts. Endosymbiosis in protists is widespread and represents an important source of innovation. Previously

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These examples demonstrate the importance of microbial symbioses for the functioning of ocean ecosystems. Understanding symbioses with this same level of detail in the context of complex communities (i.e., whole microbiomes) remains ripe for exploration and, indeed, requires a more integrated

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and regulate microbiome assembly and maintenance in many marine organisms, including sponges, macroalgae, and corals. For example, tropical corals harbor diverse bacteria in their surface mucus layer that produce quorum-sensing inhibitors and other antibacterial compounds as a defense against

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The outermost epidermal layer, i.e. the skin, is the first barrier that protects the individual from the outside world and the epidermal microbiome on it is considered an indicator not only of the health of the animal but is also considered an ecological indicator that shows the state of the

782: have revealed some of the profoundly important symbiotic roles microbes play in the lives of their hosts. These studies, however, have tended to focus on a small number of specific microbial taxa. In contrast, most hosts retain groups of many hundreds of different microbes (i.e., a 833:

nutritional symbioses with microbes. There are many examples of marine nutritional mutualisms in which microbes enable hosts to utilize resources or substrates otherwise unavailable to the host alone. Such symbioses have been described in detail in reduced and anoxic sediments (e.g.,

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Kleiner, M., Wentrup, C., Lott, C., Teeling, H., Wetzel, S., Young, J., Chang, Y.J., Shah, M., VerBerkmoes, N.C., Zarzycki, J. and Fuchs, G. (2012) "Metaproteomics of a gutless marine worm and its symbiotic microbial community reveal unusual pathways for carbon and energy use".

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All animals on Earth form associations with microorganisms, including protists, bacteria, archaea, fungi, and viruses. In the ocean, animal–microbial relationships were historically explored in single host–symbiont systems. However, new explorations into the diversity of

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Thomas, T., Moitinho-Silva, L., Lurgi, M., Björk, J.R., Easson, C., Astudillo-García, C., Olson, J.B., Erwin, P.M., López-Legentil, S., Luter, H. and Chaves-Fonnegra, A. (2016) "Diversity, structure and convergent evolution of the global sponge microbiome".

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Wippler, J., Kleiner, M., Lott, C., Gruhl, A., Abraham, P.E., Giannone, R.J., Young, J.C., Hettich, R.L. and Dubilier, N. (2016) "Transcriptomic and proteomic insights into innate immunity and adaptations to a symbiotic lifestyle in the gutless marine worm

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Morrow, K.M., Bourne, D.G., Humphrey, C., Botté, E.S., Laffy, P., Zaneveld, J., Uthicke, S., Fabricius, K.E. and Webster, N.S. (2015) "Natural volcanic CO 2 seeps reveal future trajectories for host–microbial associations in corals and sponges".

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is emerging as a central member of the coral's microbiome, with flexibility in its lifestyle. Given the recent mass bleaching occurring on reefs, corals will likely continue to be a useful and popular system for symbiosis and dysbiosis research.

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Dubilier, N., Mülders, C., Ferdelman, T., de Beer, D., Pernthaler, A., Klein, M., Wagner, M., Erséus, C., Thiermann, F., Krieger, J. and Giere, O. (2001) "Endosymbiotic sulphate-reducing and sulphide-oxidizing bacteria in an oligochaete worm".

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Woyke, T., Teeling, H., Ivanova, N.N., Huntemann, M., Richter, M., Gloeckner, F.O., Boffelli, D., Anderson, I.J., Barry, K.W., Shapiro, H.J. and Szeto, E. (2006) "Symbiosis insights through metagenomic analysis of a microbial consortium".

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Hughes, T.P., Kerry, J.T., Álvarez-Noriega, M., Álvarez-Romero, J.G., Anderson, K.D., Baird, A.H., Babcock, R.C., Beger, M., Bellwood, D.R., Berkelmans, R. and Bridge, T.C. (2017) "Global warming and recurrent mass bleaching of corals".

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Webster, N.S., Negri, A.P., Botté, E.S., Laffy, P.W., Flores, F., Noonan, S., Schmidt, C. and Uthicke, S. (2016) "Host-associated coral reef microbes respond to the cumulative pressures of ocean warming and ocean acidification".

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antibiotic thought to keep the wood-digesting cecum clear of bacterial foulants. By producing antimicrobial compounds, these microbes are able to defend their niche space to prevent other organisms from crowding them out.

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Schimak, M.P., Kleiner, M., Wetzel, S., Liebeke, M., Dubilier, N. and Fuchs, B.M. (2016) "MiL-FISH: Multilabeled oligonucleotides for fluorescence in situ hybridization improve visualization of bacterial cells".

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Ruehland, C., Blazejak, A., Lott, C., Loy, A., Erséus, C. and Dubilier, N. (2008) "Multiple bacterial symbionts in two species of co‐occurring gutless oligochaete worms from Mediterranean sea grass sediments".

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Neave, M.J., Michell, C.T., Apprill, A. and Voolstra, C.R. (2017) "Endozoicomonas genomes reveal functional adaptation and plasticity in bacterial strains symbiotically associated with diverse marine hosts".

4926: 1875:"Phylogenetic characterization and in situ localization of the bacterial symbiont of shipworms (Teredinidae: Bivalvia) by using 16S rRNA sequence analysis and oligodeoxynucleotide probe hybridization" 1930:

Ruehland C, Blazejak A, Lott C, Loy A, Erséus C, Dubilier N (2008). "Multiple bacterial symbionts in two species of co-occurring gutless oligochaete worms from Mediterranean sea grass sediments".

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sequences, since sequences produced by the Illumina platform are of insufficient length (approximately 250 base pairs) for the design of primers and probes. In 2019, Goldsmith et al demonstrated

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and spirilloxanthin. The endosymbionts are photosynthetically active; hence, this symbiosis represents an evolutionary transition of an aerobic organism to an anaerobic one while incorporating

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Zhang, F., Blasiak, L.C., Karolin, J.O., Powell, R.J., Geddes, C.D. and Hill, R.T. (2015) "Phosphorus sequestration in the form of polyphosphate by microbial symbionts in marine sponges".

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The microbiomes of diverse marine animals are currently under study, from simplistic organisms including sponges and ctenophores to more complex organisms such as sea squirts and sharks.

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Johnson WR, Torralba M, Fair PA, Bossart GD, Nelson KE, Morris PJ (December 2009). "Novel diversity of bacterial communities associated with bottlenose dolphin upper respiratory tracts".

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Acevedo-Whitehouse K, Rocha-Gosselin A, Gendron D (April 2010). "A novel non-invasive tool for disease surveillance of free-ranging whales and its relevance to conservation programs".

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Simister, R., Taylor, M.W., Tsai, P., Fan, L., Bruxner, T.J., Crowe, M.L. and Webster, N. (2012) "Thermal stress responses in the bacterial biosphere of the Great Barrier Reef sponge,

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De Goeij JM, Van Oevelen D, Vermeij MJ, Osinga R, Middelburg JJ, De Goeij AF, et al. (2013). "Surviving in a Marine Desert: The Sponge Loop Retains Resources within Coral Reefs".

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Lima N, Rogers T, Acevedo-Whitehouse K, Brown MV (February 2012). "Temporal stability and species specificity in bacteria associated with the bottlenose dolphins respiratory system".

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Peixoto, R.S., Rosado, P.M., Leite, D.C.D.A., Rosado, A.S. and Bourne, D.G. (2017) "Beneficial microorganisms for corals (BMC): proposed mechanisms for coral health and resilience".

902:, are important in fostering coral recovery in the wake of disturbance. Epulopiscium bacteria in the guts of surgeonfishes produce enzymes that allow their hosts to digest complex 5502:"Microbial diversity and structure in the gastrointestinal tracts of two stranded short-finned pilot whales (Globicephala macrorhynchus) and a pygmy sperm whale (Kogia breviceps)" 3359:

Amin SA, Hmelo LR, Van Tol HM, Durham BP, Carlson LT, Heal KR, et al. (2015). "Interaction and signalling between a cosmopolitan phytoplankton and associated bacteria".

766:, i.e., persistent interactions between host and microbe in which none of the partners gets harmed and at least one of them benefits, are ubiquitous from shallow reefs to 680: 4795:

Anthony, K.R., Kline, D.I., Diaz-Pulido, G., Dove, S. and Hoegh-Guldberg, O.(2008) "Ocean acidification causes bleaching and productivity loss in coral reef builders".

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the center that is non-contaminated. Besides there are studies from rectal swabs and rare studies from stranded dead or living animals direct from the intestine.

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Ribes, M., Calvo, E., Movilla, J., Logares, R., Coma, R. and Pelejero, C. (2016) "Restructuring of the sponge microbiome favors tolerance to ocean acidification

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Black circle: macronucleus, white big circle: food vacuoles, green circles: phototrophs, brown circles: chemoautotrophs, yellow ovals: heterotrophic prokaryotes

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Radax, R., Hoffmann, F., Rapp, H.T., Leininger, S. and Schleper, C. (2012) "Ammonia‐oxidizing archaea as main drivers of nitrification in cold‐water sponges".

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image of the surface and scales of the fish, with arrows pointing to bacterial-sized cells and larger cells (which are not noted) are presumably phytoplankton.

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are critical to the functioning of Indo-Pacific coral reefs, as they are among the only fishes capable of consuming large macroalgae that bloom in the wake of

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effects on biodiversity and ecosystem processes. The microbiomes of diverse marine animals are currently under study, from simplistic organisms including

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Hoey AS, Bellwood DR (2009). "Limited Functional Redundancy in a High Diversity System: Single Species Dominates Key Ecological Process on Coral Reefs".

5794:"Respiratory Microbiome of Endangered Southern Resident Killer Whales and Microbiota of Surrounding Sea Surface Microlayer in the Eastern North Pacific" 935:

depends on symbiotic bacteria living under its cuticle as its source of food. The bacteria are responsible for the bright white appearance of the worms.

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Bayer, K., Schmitt, S. and Hentschel, U. (2008) "Physiology, phylogeny and in situ evidence for bacterial and archaeal nitrifiers in the marine sponge

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Quigley KM, Bay LK, Willis BL (2018). "Leveraging new knowledge of Symbiodinium community regulation in corals for conservation and reef restoration".

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Bourne, D.G., Morrow, K.M. and Webster, N.S. (2016) "Insights into the coral microbiome: underpinning the health and resilience of reef ecosystems".

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Apprill A, Mooney TA, Lyman E, Stimpert AK, Rappé MS (April 2011). "Humpback whales harbour a combination of specific and variable skin bacteria".

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Lesser, M.P., Fiore, C., Slattery, M. and Zaneveld, J. (2016) "Climate change stressors destabilize the microbiome of the Caribbean barrel sponge,

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Neave, M.J., Apprill, A., Ferrier-Pagès, C. and Voolstra, C.R. (2016) "Diversity and function of prevalent symbiotic marine bacteria in the genus

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Pfister CA, Altabet MA, Weigel BL (2019). "Kelp beds and their local effects on seawater chemistry, productivity, and microbial communities".

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and create the structural habitats and nutrient resources that are the foundation of their respective ecosystems. All of these taxa engage in

3139:"Revisiting the STEC Testing Approach: Using espK and espV to Make Enterohemorrhagic Escherichia coli (EHEC) Detection More Reliable in Beef" 2617:

Croft MT, Lawrence AD, Raux-Deery E, Warren MJ, Smith AG (2005). "Algae acquire vitamin B12 through a symbiotic relationship with bacteria".

1168: 1004: 3190:"Acyl-homoserine lactones modulate the settlement rate of zoospores of the marine alga Ulva intestinalis via a novel chemokinetic mechanism" 1371:. OTUs from next-generation sequencing are displayed if the OTU contained more than two sequences in the unrarefied OTU table (3626 OTUs). 4485:

Givens, C.E., Ransom, B., Bano, N. and Hollibaugh, J.T. (2015) "Comparison of the gut microbiomes of 12 bony fish and 3 shark species".

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unrecognized metabolic innovations of marine microbial symbioses that are ecologically important are discovered regularly. For example,

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Gast RJ, Sanders RW, Caron DA (2009). "Ecological strategies of protists and their symbiotic relationships with prokaryotic microbes".

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breaching and (H) a SEM image of a humpback's skin surface associated bacteria, with arrows indicating two different cell morphologies.

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Blasiak, L.C., Zinder, S.H., Buckley, D.H. and Hill, R.T. (2014) "Bacterial diversity associated with the tunic of the model chordate

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Duffy JE, Godwin CM, Cardinale BJ (2017). "Biodiversity effects in the wild are common and as strong as key drivers of productivity".

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USGS scientists publish long-read microbiome sequences from temperate coral, providing community resource for probe and primer design

6159:"Rhodoliths holobionts in a changing ocean: host-microbes interactions mediate coralline algae resilience under ocean acidification" 4778:

Dubinsky, Z. and Jokiel, P.L. (1994) "Ratio of energy and nutrient fluxes regulates symbiosis between zooxanthellae and corals".

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Provasoli L, Pintner IJ (1980). "Bacteria Induced Polymorphism in an Axenic Laboratory Strain of Ulva Lactuca (Chlorophyceae)1".

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Suzuki A, Ueda K, Segawa T, Suzuki M (June 2019). "Fecal microbiota of captive Antillean manatee Trichechus manatus manatus".

6247: 1437:, while also producing longer sequences useful to the research community for probe and primer design (see diagram on right). 687: 1560:), and associated bacteria and viruses. Co-evolutionary patterns exist for coral microbial communities and coral phylogeny. 5651:"Interannual comparison of core taxa and community composition of the blow microbiota from East Australian humpback whales" 3272:

Matsuo Y, Imagawa H, Nishizawa M, Shizuri Y (2005). "Isolation of an Algal Morphogenesis Inducer from a Marine Bacterium".

287: 5394:"Microbiome Composition and Function in Aquatic Vertebrates: Small Organisms Making Big Impacts on Aquatic Animal Health" 393: 1651:

Apprill, A. (2017) "Marine animal microbiomes: toward understanding host–microbiome interactions in a changing ocean".

1580: 906:, enabling the host fish to feed on tough, leathery red and brown macroalgae. This trophic innovation has facilitated 6232: 1505:

breath or "blow" of the cetaceans can provide an assessment of the state of health. Blow is composed of a mixture of

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McFall-Ngai, M.J. (2000) "Negotiations between animals and bacteria: the 'diplomacy'of the squid-vibrio symbiosis".

2519:"The importance of sponges and mangroves in supporting fish communities on degraded coral reefs in Caribbean Panama" 1204: 292: 3896:"Comparing and Evaluating Metagenome Assembly Tools from a Microbiologist's Perspective - Not Only Size Matters!" 45:

hosts is moving the field into studies that address interactions between the animal host and a more multi-member

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Knowlton N, Rohwer F (October 2003). "Multispecies microbial mutualisms on coral reefs: the host as a habitat".

3844:"Patterns and controls of reef-scale production of dissolved organic carbon by giant kelp M acrocystis pyrifera" 1615: 3622:"Characterization of thegacA-dependent surface and coral mucus colonization by an opportunistic coral pathogen 473: 1310: 2921:"Caribbean Spiny Lobster Fishery is Underpinned by Trophic Subsidies from Chemosynthetic Primary Production" 56:

that do not live in close relationship with a microbial partner. Host-associated microbiomes also influence

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to study microbial community interactions associated with symbiotic state. However, the ability to develop

1246: 1154: 1114: 572: 57: 2570:"Host-Microbe Coevolution: Applying Evidence from Model Systems to Complex Marine Invertebrate Holobionts" 4116:"Evolution of a Vegetarian Vibrio: Metabolic Specialization of Vibrio breoganii to Macroalgal Substrates" 1283: 1119: 226: 6047:"Microbes in the coral holobiont: partners through evolution, development, and ecological interactions" 4730:"Season, but not symbiont state, drives microbiome structure in the temperate coral Astrangia poculata" 1434: 1418: 1368: 1290: 767: 701: 199: 4293: 3072:"Recruitment in the sea: Bacterial genes required for inducing larval settlement in a polychaete worm" 1409:, widely documented along the eastern coast of the United States. The coral can live with and without 1379:

and inorganic nutrients, affect the abundance and performance of the microalgal symbionts, as well as

798: 3955:"Boronated tartrolon antibiotic produced by symbiotic cellulose-degrading bacteria in shipworm gills" 1099: 1009: 4324:

Petersen JM, Kemper A, Gruber-Vodicka H, Cardini U, Van Der Geest M, Kleiner M, et al. (2017).

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Van Der Heide T, Govers LL, De Fouw J, Olff H, Van Der Geest M, Van Katwijk MM, et al. (2012).

5602:"Extensive Core Microbiome in Drone-Captured Whale Blow Supports a Framework for Health Monitoring" 1770:"Coral-associated micro-organisms and their roles in promoting coral health and thwarting diseases" 739: 639: 634: 342: 4539:

McFall-Ngai, M. (2014) "Divining the essence of symbiosis: insights from the squid-vibrio model".

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Seah BK, Antony CP, Huettel B, Zarzycki J, Schada von Borzyskowski L, Erb TJ, et al. (2019).

2107:"Marine Animal Microbiomes: Toward Understanding Host–Microbiome Interactions in a Changing Ocean" 1190: 1594: 5443:

Bik EM, Costello EK, Switzer AD, Callahan BJ, Holmes SP, Wells RS, et al. (February 2016).

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McFall-Ngai M, Hadfield MG, Bosch TC, Carey HV, Domazet-Lošo T, Douglas AE, et al. (2013).

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Cavalcanti GS, Shukla P, Morris M, Ribeiro B, Foley M, Doane MP, et al. (September 2018).

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Geoghegan JL, Pirotta V, Harvey E, Smith A, Buchmann JP, Ostrowski M, et al. (June 2018).

3412:"Effects of epibiotic bacteria on leaf growth and epiphytes of the seagrass Posidonia oceanica" 1327: 1200: 567: 2401: 1425:

to more specifically target key microbial groups has been hindered by the lack of full length

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Pollock FJ, McMinds R, Smith S, Bourne DG, Willis BL, Medina M, et al. (November 2018).

5735:"The use of Unmanned Aerial Vehicles (UAVs) to sample the blow microbiome of small cetaceans" 5392:

Sehnal L, Brammer-Robbins E, Wormington AM, Blaha L, Bisesi J, Larkin I, et al. (2021).

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Pirotta V, Smith A, Ostrowski M, Russell D, Jonsen ID, Grech A, et al. (December 2017).

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Corzett CH, Elsherbini J, Chien DM, Hehemann JH, Henschel A, Preheim SP, et al. (2018).

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Elshahawi SI, Trindade-Silva AE, Hanora A, Han AW, Flores MS, Vizzoni V, et al. (2013).

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Daniels, C. and Breitbart, M. (2012) "Bacterial communities associated with the ctenophores

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