Aliivibrio fischeri

Last updated

Aliivibrio fischeri
Aliivibrio fischeri.jpg
Aliivibrio fischeri glowing on a petri dish
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Bacteria
Phylum: Pseudomonadota
Class: Gammaproteobacteria
Order: Vibrionales
Family: Vibrionaceae
Genus: Aliivibrio
Species:
A. fischeri
Binomial name
Aliivibrio fischeri
(Beijerinck 1889) Urbanczyk et al. 2007
Synonyms [1]

Aliivibrio fischeri (also called Vibrio fischeri) is a Gram-negative, rod-shaped bacterium found globally in marine environments. [2] This species has bioluminescent properties, and is found predominantly in symbiosis with various marine animals, such as the Hawaiian bobtail squid. It is heterotrophic, oxidase-positive, and motile by means of a single polar flagella. [3] Free-living A. fischeri cells survive on decaying organic matter. The bacterium is a key research organism for examination of microbial bioluminescence, quorum sensing, and bacterial-animal symbiosis. [4] It is named after Bernhard Fischer, a German microbiologist. [5]

Contents

Ribosomal RNA comparison led to the reclassification of this species from genus Vibrio to the newly created Aliivibrio in 2007. [6] However, the name change is not generally accepted by most researchers, who still publish Vibrio fischeri (see Google Scholar for 2018–2019).

Genome

The genome for A. fischeri was completely sequenced in 2004 [7] and consists of two chromosomes, one smaller and one larger. Chromosome 1 has 2.9 million base pairs (Mbp) and chromosome 2 has 1.3 Mbp, bringing the total genome to 4.2 Mbp. [7]

A. fischeri has the lowest G+C content of 27 Vibrio species, but is still most closely related to the higher-pathogenicity species such as V. cholerae. [7] The genome for A. fischeri also carries mobile genetic elements. [7]

Ecology

The Hawaiian bobtail squid, its photophores populated with Allivibrio fischeri Euprymna scolopes, South shore of Oahu, Hawaii.tiff
The Hawaiian bobtail squid, its photophores populated with Allivibrio fischeri

A. fischeri are globally distributed in temperate and subtropical marine environments. [8] They can be found free-floating in oceans, as well as associated with marine animals, sediment, and decaying matter. [8] A. fischeri have been most studied as symbionts of marine animals, including squids in the genus Euprymna and Sepiola , where A. fischeri can be found in the squids' light organs. [8] This relationship has been best characterized in the Hawaiian bobtail squid (Euprymna scolopes), where A. fischeri is the only species of bacteria inhabiting the squid's light organ. [9]

Symbiosis with the Hawaiian bobtail squid

A. fischeri colonization of the light organ of the Hawaiian bobtail squid is currently studied as a simple model for mutualistic symbiosis, as it contains only two species and A. fischeri can be cultured in a lab and genetically modified. This mutualistic symbiosis functions primarily due to A. fischeri bioluminescence. A. fischeri colonizes the light organ of the Hawaiian bobtail squid and luminesces at night, providing the squid with counter-illumination camouflage, which prevents the squid from casting a shadow on the ocean floor.

A. fischeri colonization occurs in juvenile squids and induces morphological changes in the squids light organ. Interestingly, certain morphological changes made by A. fischeri do not occur when the microbe cannot luminesce, indicating that bioluminescence (described below) is truly essential for symbiosis. In the process of colonization, ciliated cells within the animals' photophores (light-producing organs) selectively draw in the symbiotic bacteria. These cells promote the growth of the symbionts and actively reject any competitors. The bacteria cause these cells to die off once the light organ is sufficiently colonized.

The light organs of certain squid contain reflective plates that intensify and direct the light produced, due to proteins known as reflectins. They regulate the light for counter-illumination camouflage, requiring the intensity to match that of the sea surface above. [10] Sepiolid squid expel 90% of the symbiotic bacteria in their light organ each morning in a process known as "venting". Venting is thought to provide the source from which newly hatched squid are colonized by A. fischeri.

Bioluminescence

The bioluminescence of A. fischeri is caused by transcription of the lux operon, which is induced through population-dependent quorum sensing. [2] The population of A. fischeri needs to reach an optimal level to activate the lux operon and stimulate light production. The circadian rhythm controls light expression, where luminescence is much brighter during the day and dimmer at night, as required for camouflage.

The bacterial luciferin-luciferase system is encoded by a set of genes labelled the lux operon. In A. fischeri, five such genes (luxCDABEG) have been identified as active in the emission of visible light, and two genes (luxR and luxI) are involved in regulating the operon. Several external and intrinsic factors appear to either induce or inhibit the transcription of this gene set and produce or suppress light emission.

A. fischeri is one of many species of bacteria that commonly form symbiotic relationships with marine organisms. [11] Marine organisms contain bacteria that use bioluminescence so they can find mates, ward off predators, attract prey, or communicate with other organisms. [12] In return, the organism the bacteria are living within provides the bacteria with a nutrient-rich environment. [13] The lux operon is a 9-kilobase fragment of the A. fischeri genome that controls bioluminescence through the catalytic activity of the enzyme luciferase. [14] This operon has a known gene sequence of luxCDAB(F)E, where luxA and luxB code for the protein subunits of the luciferase enzyme, and the luxCDE codes for a fatty acid reductase complex that makes the fatty acids necessary for the luciferase mechanism. [14] luxC codes for the enzyme acyl-reductase, luxD codes for acyl-transferase, and luxE makes the proteins needed for the enzyme acyl-protein synthetase. Luciferase produces blue/green light through the oxidation of reduced flavin mononucleotide and a long-chain aldehyde by diatomic oxygen. The reaction is summarized as: [15]

FMNH2 + O2 + R-CHO → FMN + R-COOH + H2O + light.

The reduced flavin mononucleotide (FMNH) is provided by the fre gene, also referred to as luxG. In A. fischeri, it is directly next to luxE (giving luxCDABE-fre) from 1042306 to 1048745

To generate the aldehyde needed in the reaction above, three additional enzymes are needed. The fatty acids needed for the reaction are pulled from the fatty acid biosynthesis pathway by acyl-transferase. Acyl-transferase reacts with acyl-ACP to release R-COOH, a free fatty acid. R-COOH is reduced by a two-enzyme system to an aldehyde. The reaction is: [13]

R-COOH + ATP + NADPH → R-CHO + AMP + PP + NADP+.

Quorum sensing

Quorum sensing in Aliivibrio fischeri Green pentagons denote AHL autoinducer that LuxI produces (3OC6-homoserine lactone). Transcriptional regulator, LuxR, modulates expression of AHL synthase, LuxI, and the lux operon, leading to luciferase-mediated light emission Quorum sensing in Vibrio fischeri.png
Quorum sensing in Aliivibrio fischeri 
Green pentagons denote AHL autoinducer that LuxI produces (3OC6-homoserine lactone). Transcriptional regulator, LuxR, modulates expression of AHL synthase, LuxI, and the lux operon, leading to luciferase-mediated light emission

One primary system that controls bioluminescence through regulation of the lux operon is quorum sensing, a conserved system across many microbial species that regulates gene expression in response to bacterial concentration. Quorum sensing functions through the production of an autoinducer, usually a small organic molecule, by individual cells. As cell populations increase, levels of autoinducers increase, and specific proteins that regulate transcription of genes bind to these autoinducers and alter gene expression. This system allows microbial cells to "communicate" amongst each other and coordinate behaviors like luminescence, which require large amounts of cells to produce an effect. [16]

In A. fischeri, there are two primary quorum sensing systems, each of which respond to slightly different environments. The first system is commonly referred to as the lux system, as it is encoded within the lux operon, and uses the autoinducer 3OC6-HSL. [17] The protein LuxI synthesizes this signal, which is subsequently released from the cell. This signal, 3OC6-HSL, then binds to the protein LuxR, which regulates the expression of many different genes, but is most known for upregulation of genes involved in luminescence. [18] The second system, commonly referred to as the ain system, uses the autoinducer C8-HSL, which is produced by the protein AinS. Similar to the lux system, the autoinducer C8-HSL increases activation of LuxR. In addition, C8-HSL binds to another transcriptional regulator, LitR, giving the ain and lux systems of quorum sensing slightly different genetic targets within the cell. [19]

The different genetic targets of the ain and lux systems are essential, because these two systems respond to different cellular environments. The ain system regulates transcription in response to intermediate cell density cell environments, producing lower levels of luminescence and even regulating metabolic processes like the acetate switch. [20] On the other hand, the lux quorum sensing system occurs in response to high cell density, producing high levels of luminescence and regulating the transcription of other genes, including QsrP, RibB, and AcfA. [21] Both of the ain and lux quorum sensing systems are essential for colonization of the squid and regulate multiple colonization factors in the bacteria. [18]

Activation of the lux operon by LuxR and LuxI in Aliivibrio fischeri (A) At low cell density, the autoinducers (3OC6-HSL - red dots), produced by LuxI, diffuse through the cell membrane into the growth medium (B) As the cell growth continues, the autoinducers in the medium start to accumulate in a confined environment. A very low intensity of light can be detected. (C) When enough autoinducers have accumulated in the medium, they can re-enter the cell where they directly bind the LuxR protein to activate luxICDABEG expression. (D) High levels of autoinducers activate the luminescent system of A. fischeri. A high intensity of light can be detected. Activation of the lux operon in Aliivibrio fischeri.jpg
Activation of the lux operon by LuxR and LuxI in Aliivibrio fischeri 
(A) At low cell density, the autoinducers (3OC6-HSL – red dots), produced by LuxI, diffuse through the cell membrane into the growth medium
(B) As the cell growth continues, the autoinducers in the medium start to accumulate in a confined environment. A very low intensity of light can be detected.
(C) When enough autoinducers have accumulated in the medium, they can re-enter the cell where they directly bind the LuxR protein to activate luxICDABEG expression.
(D) High levels of autoinducers activate the luminescent system of A. fischeri. A high intensity of light can be detected.

Natural transformation

Natural bacterial transformation is an adaptation for transferring DNA from one individual cell to another. Natural transformation, including the uptake and incorporation of exogenous DNA into the recipient genome, has been demonstrated in A. fischeri. [24] This process requires induction by chitohexaose and is likely regulated by genes tfoX and tfoY. Natural transformation of A. fischeri facilitates rapid transfer of mutant genes across strains and provides a useful tool for experimental genetic manipulation in this species.

State microbe status

In 2014, Hawaiʻian State Senator Glenn Wakai submitted SB3124 proposing Aliivibrio fischeri as the state microbe of Hawaiʻi. [25] The bill was in competition with a bill to make Flavobacterium akiainvivens the state microbe, but neither passed. In 2017, legislation similar to the original 2013 F. akiainvivens bill was submitted in the Hawaiʻi House of Representatives by Isaac Choy [26] and in the Hawaiʻi Senate by Brian Taniguchi. [27]

List of synonyms

See also

Related Research Articles

In biology, quorum sensing or quorum signaling (QS) is the ability to detect and respond to cell population density by gene regulation. Quorum sensing is a type of cellular signaling, and more specifically can be considered a type of paracrine signaling. However, it also contains traits of both autocrine signaling: a cell produces both the autoinducer molecule and the receptor for the autoinducer. As one example, QS enables bacteria to restrict the expression of specific genes to the high cell densities at which the resulting phenotypes will be most beneficial, especially for phenotypes that would be ineffective at low cell densities and therefore too energetically costly to express. Many species of bacteria use quorum sensing to coordinate gene expression according to the density of their local population. In a similar fashion, some social insects use quorum sensing to determine where to nest. Quorum sensing in pathogenic bacteria activates host immune signaling and prolongs host survival, by limiting the bacterial intake of nutrients, such as tryptophan, which further is converted to serotonin. As such, quorum sensing allows a commensal interaction between host and pathogenic bacteria. Quorum sensing may also be useful for cancer cell communications.

<span class="mw-page-title-main">Bioluminescence</span> Emission of light by a living organism

Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some bioluminescent bacteria, and terrestrial arthropods such as fireflies. In some animals, the light is bacteriogenic, produced by symbiotic bacteria such as those from the genus Vibrio; in others, it is autogenic, produced by the animals themselves.

<span class="mw-page-title-main">Luciferase</span> Enzyme family

Luciferase is a generic term for the class of oxidative enzymes that produce bioluminescence, and is usually distinguished from a photoprotein. The name was first used by Raphaël Dubois who invented the words luciferin and luciferase, for the substrate and enzyme, respectively. Both words are derived from the Latin word lucifer, meaning "lightbearer", which in turn is derived from the Latin words for "light" (lux) and "to bring or carry" (ferre).

<i>Vibrio harveyi</i> Species of bacterium

Vibrio harveyi is a Gram-negative, bioluminescent, marine bacterium in the genus Vibrio. V. harveyi is rod-shaped, motile, facultatively anaerobic, halophilic, and competent for both fermentative and respiratory metabolism. It does not grow below 4 °C. V. harveyi can be found free-swimming in tropical marine waters, commensally in the gut microflora of marine animals, and as both a primary and opportunistic pathogen of marine animals, including Gorgonian corals, oysters, prawns, lobsters, the common snook, barramundi, turbot, milkfish, and seahorses. It is responsible for luminous vibriosis, a disease that affects commercially farmed penaeid prawns. Additionally, based on samples taken by ocean-going ships, V. harveyi is thought to be the cause of the milky seas effect, in which, during the night, a uniform blue glow is emitted from the seawater. Some glows can cover nearly 6,000 sq mi (16,000 km2).

<i>N</i>-Acyl homoserine lactone Class of chemical compounds

N-Acyl homoserine lactones are a class of signaling molecules involved in bacterial quorum sensing, a means of communication between bacteria enabling behaviors based on population density.

Luminescent bacteria emit light as the result of a chemical reaction during which chemical energy is converted to light energy. Luminescent bacteria exist as symbiotic organisms carried within a larger organism, such as many deep sea organisms, including the Lantern Fish, the Angler fish, certain jellyfish, certain clams and the Gulper eel. The light is generated by an enzyme-catalyzed chemoluminescence reaction, wherein the pigment luciferin is oxidised by the enzyme luciferase. The expression of genes related to bioluminescence is controlled by an operon called the lux operon.

<i>Euprymna scolopes</i> Species of cephalopods known as the Hawaiian bobtail squid

Euprymna scolopes, also known as the Hawaiian bobtail squid, is a species of bobtail squid in the family Sepiolidae native to the central Pacific Ocean, where it occurs in shallow coastal waters off the Hawaiian Islands and Midway Island. The type specimen was collected off the Hawaiian Islands and is deposited at the National Museum of Natural History in Washington, D.C.

Autoinducers are signaling molecules that are produced in response to changes in cell-population density. As the density of quorum sensing bacterial cells increases so does the concentration of the autoinducer. Detection of signal molecules by bacteria acts as stimulation which leads to altered gene expression once the minimal threshold is reached. Quorum sensing is a phenomenon that allows both Gram-negative and Gram-positive bacteria to sense one another and to regulate a wide variety of physiological activities. Such activities include symbiosis, virulence, motility, antibiotic production, and biofilm formation. Autoinducers come in a number of different forms depending on the species, but the effect that they have is similar in many cases. Autoinducers allow bacteria to communicate both within and between different species. This communication alters gene expression and allows bacteria to mount coordinated responses to their environments, in a manner that is comparable to behavior and signaling in higher organisms. Not surprisingly, it has been suggested that quorum sensing may have been an important evolutionary milestone that ultimately gave rise to multicellular life forms.

<span class="mw-page-title-main">Autoinducer-2</span> Chemical compound

Autoinducer-2 (AI-2), a furanosyl borate diester or tetrahydroxy furan, is a member of a family of signaling molecules used in quorum sensing. AI-2 is one of only a few known biomolecules incorporating boron. First identified in the marine bacterium Vibrio harveyi, AI-2 is produced and recognized by many Gram-negative and Gram-positive bacteria. AI-2 arises by the reaction of 4,5-dihydroxy-2,3-pentanedione, which is produced enzymatically, with boric acid and is recognized by the two-component sensor kinase LuxPQ in Vibrionaceae.

<span class="mw-page-title-main">John Woodland Hastings</span>

John Woodland "Woody" Hastings, was a leader in the field of photobiology, especially bioluminescence, and was one of the founders of the field of circadian biology. He was the Paul C. Mangelsdorf Professor of Natural Sciences and Professor of Molecular and Cellular Biology at Harvard University. He published over 400 papers and co-edited three books.

<span class="mw-page-title-main">LuxR-type DNA-binding HTH domain</span>

In molecular biology, the LuxR-type DNA-binding HTH domain is a DNA-binding, helix-turn-helix (HTH) domain of about 65 amino acids. It is present in transcription regulators of the LuxR/FixJ family of response regulators. The domain is named after Vibrio fischeri luxR, a transcriptional activator for quorum-sensing control of luminescence. LuxR-type HTH domain proteins occur in a variety of organisms. The DNA-binding HTH domain is usually located in the C-terminal region of the protein; the N-terminal region often containing an autoinducer-binding domain or a response regulatory domain. Most luxR-type regulators act as transcription activators, but some can be repressors or have a dual role for different sites. LuxR-type HTH regulators control a wide variety of activities in various biological processes.

Interspecies quorum sensing is a type of quorum sensing in which bacteria send and receive signals to other species besides their own. This is accomplished by the secretion of signaling molecules which trigger a response in nearby bacteria at high enough concentrations. Once the molecule hits a certain concentration it triggers the transcription of certain genes such as virulence factors. It has been discovered that bacteria can not only interact via quorum sensing with members of their own species but that there is a kind of universal molecule that allows them to gather information about other species as well. This universal molecule is called autoinducer 2 or AI-2.

Vibrio campbellii is a Gram-negative, curved rod-shaped, marine bacterium closely related to its sister species, Vibrio harveyi. It is an emerging pathogen in aquatic organisms.

Acyl-homoserine-lactone synthase is an enzyme with systematic name acyl-(acyl-carrier protein):S-adenosyl-L-methionine acyltranserase . This enzyme catalyses the following chemical reaction

<span class="mw-page-title-main">Bioluminescent bacteria</span>

Bioluminescent bacteria are light-producing bacteria that are predominantly present in sea water, marine sediments, the surface of decomposing fish and in the gut of marine animals. While not as common, bacterial bioluminescence is also found in terrestrial and freshwater bacteria. These bacteria may be free living or in symbiosis with animals such as the Hawaiian Bobtail squid or terrestrial nematodes. The host organisms provide these bacteria a safe home and sufficient nutrition. In exchange, the hosts use the light produced by the bacteria for camouflage, prey and/or mate attraction. Bioluminescent bacteria have evolved symbiotic relationships with other organisms in which both participants benefit close to equally. Another possible reason bacteria use luminescence reaction is for quorum sensing, an ability to regulate gene expression in response to bacterial cell density.

<span class="mw-page-title-main">Marine microbial symbiosis</span>

Microbial symbiosis in marine animals was not discovered until 1981. In the time following, symbiotic relationships between marine invertebrates and chemoautotrophic bacteria have been found in a variety of ecosystems, ranging from shallow coastal waters to deep-sea hydrothermal vents. Symbiosis is a way for marine organisms to find creative ways to survive in a very dynamic environment. They are different in relation to how dependent the organisms are on each other or how they are associated. It is also considered a selective force behind evolution in some scientific aspects. The symbiotic relationships of organisms has the ability to change behavior, morphology and metabolic pathways. With increased recognition and research, new terminology also arises, such as holobiont, which the relationship between a host and its symbionts as one grouping. Many scientists will look at the hologenome, which is the combined genetic information of the host and its symbionts. These terms are more commonly used to describe microbial symbionts.

Everett Peter Greenberg is an American microbiologist. He is the inaugural Eugene and Martha Nester Professor of Microbiology at the Department of Microbiology of the University of Washington School of Medicine. He is best known for his research on quorum sensing, and has received multiple awards for his work.

Margaret McFall-Ngai is an American animal physiologist and biochemist best-known for her work related to the symbiotic relationship between Hawaiian bobtail squid, Euprymna scolopes and bioluminescent bacteria, Vibrio fischeri. Her research helped expand the microbiology field, primarily focused on pathogenicity and decomposition at the time, to include positive microbial associations. She currently is a professor at PBRC’s Kewalo Marine Laboratory and director of the Pacific Biosciences Research Program at the University of Hawaiʻi at Mānoa.

Karen Visick is an American microbiologist and expert in bacterial genetics known for her work on the role of bacteria to form biofilm communities during animal colonization. She conducted doctoral research with geneticist Kelly Hughes at the University of Washington, where she identified a key regulatory checkpoint during construction of the bacterial flagellum. She conducted postdoctoral research on development of the Vibrio fischeri-Euprymna scolopes symbiosis with Ned Ruby at University of Southern California and University of Hawaiʻi. The bacteria are bioluminescent and provide light to the host. Visick and Ruby revealed that bacteria that do not produce light exhibit a defect during host colonization.

References

  1. 1 2 "Aliivibrio fischeri". NCBI taxonomy. Bethesda, MD: National Center for Biotechnology Information. Retrieved 6 December 2017. Other names: genbank synonym: Vibrio fischeri (Beijerinck 1889) Lehmann and Neumann 1896 (Approved Lists 1980) synonym: Vibrio noctiluca Weisglass and Skreb 1963 synonym: Photobacterium fischeri Beijerinck 1889 synonym: Microspira marina (Russell 1892) Migula 1900 synonym: Microspira fischeri (Beijerinck 1889) Chester 1901 synonym: Einheimischer Leuchtbacillus Fischer 1888 synonym: Bacillus phosphorescens indigenus Eisenberg 1891 synonym: Bacillus fischeri (Beijerinck 1889) Trevisan 1889 synonym: Achromobacter fischeri (Beijerinck 1889) Bergey et al. 1930
  2. 1 2 Madigan M, Martinko J, eds. (2005). Brock Biology of Microorganisms (11th ed.). Prentice Hall. ISBN   978-0-13-144329-7.
  3. ↑ Bergey, DH (1994). Holt, JG (ed.). Bergey's Manual of Determinative Bacteriology (9th ed.). Baltimore: Williams & Wilkins.
  4. ↑ Holt JG, ed. (1994). Bergey's Manual of Determinative Bacteriology (9th ed.). Williams & Wilkins. ISBN   978-0-683-00603-2.
  5. ↑ George M. Garrity: Bergey's Manual of Systematic Bacteriology . 2. Auflage. Springer, New York, 2005, Volume 2: The Proteobacteria, Part B: The Gammaproteobacteria ISBN   0-387-24144-2
  6. ↑ Urbanczyk, H.; Ast, J. C.; Higgins, M. J.; Carson, J.; Dunlap, P. V. (2007). "Reclassification of Vibrio fischeri, Vibrio logei, Vibrio salmonicida and Vibrio wodanis as Aliivibrio fischeri gen. nov., comb. nov., Aliivibrio logei comb. nov., Aliivibrio salmonicida comb. nov. and Aliivibrio wodanis comb. nov". International Journal of Systematic and Evolutionary Microbiology. 57 (12): 2823–2829. doi: 10.1099/ijs.0.65081-0 . PMID   18048732.
  7. 1 2 3 4 Ruby, E. G.; Urbanowski, M.; Campbell, J.; Dunn, A.; Faini, M.; Gunsalus, R.; Lostroh, P.; Lupp, C.; McCann, J. (2005-02-22). "Complete genome sequence of Vibrio fischeri: A symbiotic bacterium with pathogenic congeners". Proceedings of the National Academy of Sciences of the United States of America. 102 (8): 3004–3009. Bibcode:2005PNAS..102.3004R. doi: 10.1073/pnas.0409900102 . ISSN   0027-8424. PMC   549501 . PMID   15703294.
  8. 1 2 3 McFall-Ngai M (2014). "The Importance of Microbes in Animal Development: Lessons from the Squid-Vibrio Symbiosis - Supplemental Material". Annual Review of Microbiology. 68: 177–194. doi:10.1146/annurev-micro-091313-103654. PMC   6281398 . PMID   24995875.
  9. ↑ Norsworthy AN, Visick KL (November 2013). "Gimme shelter: how Vibrio fischeri successfully navigates an animal's multiple environments". Frontiers in Microbiology. 4: 356. doi: 10.3389/fmicb.2013.00356 . PMC   3843225 . PMID   24348467.
  10. ↑ Jones, B. W.; Nishiguchi, M. K. (2004). "Counterillumination in the hawaiian bobtail squid, Euprymna scolopes Berry (Mollusca : Cephalopoda)" (PDF). Marine Biology . 144 (6): 1151–1155. doi:10.1007/s00227-003-1285-3. S2CID   86576334.
  11. ↑ (Distal, 1993).
  12. ↑ Widder, E. A. (2010-05-07). "Bioluminescence in the Ocean: Origins of Biological, Chemical, and Ecological Diversity". Science. 328 (5979): 704–708. Bibcode:2010Sci...328..704W. doi:10.1126/science.1174269. ISSN   0036-8075. PMID   20448176. S2CID   2375135.
  13. 1 2 Winfrey, Michael R. (1997-01-01). Unraveling DNA: Molecular Biology for the Laboratory. Prentice-Hall. ISBN   9780132700344.
  14. 1 2 (Meighen, 1991).
  15. ↑ Silverman et al., 1984
  16. 1 2 Waters, Christopher M.; Bassler, Bonnie L. (2005). "Quorum sensing: Cell-to-cell communication in bacteria". Annual Review of Cell and Developmental Biology. 21: 319–346. doi:10.1146/annurev.cellbio.21.012704.131001. PMID   16212498.
  17. ↑ Eberhad, A (1981). "Structural identification of autoinducer of Photobacterium fischeri luciferase". PubMed. Retrieved 2020-04-26.
  18. 1 2 Lupp, Claudia (2005). "Vibrio fischeri uses two quorum-sensing systems for the regulation of early and late colonization factors". J Bacteriol. 187 (11): 3620–3629. doi: 10.1128/JB.187.11.3620-3629.2005 . PMC   1112064 . PMID   15901683.
  19. ↑ Lupp, C (2003). "The Vibrio fischeri quorum-sensing systems ain and lux sequentially induce luminescence gene expression and are important for persistence in the squid host". Mol. Microbiol. 50 (1): 319–31. doi: 10.1046/j.1365-2958.2003.t01-1-03585.x . PMID   14507383.
  20. ↑ Studer, Sarah (2008). "AinS Quorum Sensing Regulates the Vibrio fischeri Acetate Switch". J Bacteriol. 190 (17): 5915–5923. doi:10.1128/JB.00148-08. PMC   2519518 . PMID   18487321.
  21. ↑ Qin, N (2007). "Analysis of LuxR regulon gene expression during quorum sensing in Vibrio fischeri". J Bacteriol. 189 (11): 4127–34. doi:10.1128/JB.01779-06. PMC   1913387 . PMID   17400743.
  22. ↑ Li, Zhi; Nair, Satish K. (2012). "Quorum sensing: How bacteria can coordinate activity and synchronize their response to external signals?". Protein Science. 21 (10): 1403–1417. doi:10.1002/pro.2132. PMC   3526984 . PMID   22825856.
  23. ↑ Tanet, Lisa; Tamburini, Christian; Baumas, Chloé; Garel, Marc; Simon, Gwénola; Casalot, Laurie (2019). "Bacterial Bioluminescence: Light Emission in Photobacterium phosphoreum is Not Under Quorum-Sensing Control". Frontiers in Microbiology. 10: 365. doi: 10.3389/fmicb.2019.00365 . PMC   6409340 . PMID   30886606. CC-BY icon.svg Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License
  24. ↑ Pollack-Berti A, Wollenberg MS, Ruby EG (2010). "Natural transformation of Vibrio fischeri requires tfoX and tfoY". Environ. Microbiol. 12 (8): 2302–11. doi:10.1111/j.1462-2920.2010.02250.x. PMC   3034104 . PMID   21966921.
  25. ↑ Cave, James (3 April 2014). "Hawaii, Other States Calling Dibs On Official State Bacteria". Huffington Post. Retrieved 24 October 2017.
  26. ↑ Choy, Isaac (25 January 2017). "HB1217". Hawaii State Legislature. Honolulu, HI: Hawaii State Legislature . Retrieved 22 October 2017.
  27. ↑ Taniguchi, Brian (25 January 2017). "SB1212". Hawaii State Legislature. Honolulu, HI: Hawaii State Legislature . Retrieved 22 October 2017.
This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.