Wikipedia

Aliivibrio fischeri

Aliivibrio fischeri(formerlyVibrio fischeri) is aGram-negative,rod-shapedbacteriumfound globally inmarineenvironments.[2]This bacterium grows most effectively in water with a salt concentration at around 20g/L, and at temperatures between 24 and 28°C.[3]This species is non-pathogenic[3]and hasbioluminescentproperties. It is found predominantly insymbiosiswith various marine animals, such as theHawaiian bobtail squid.It isheterotrophic,oxidase-positive,andmotileby means of a tuft of polarflagella.[4]Free-livingA. fischericellssurvive on decayingorganic matter.The bacterium is a key research organism for examination of microbialbioluminescence,quorum sensing,and bacterial-animal symbiosis.[5]It is named afterBernhard Fischer,a German microbiologist.[6]

Aliivibrio fischeri
Aliivibrio fischeriglowing on a petri dish
Scientific classificationEdit this classification
Domain: Bacteria
Phylum: Pseudomonadota
Class: Gammaproteobacteria
Order: Vibrionales
Family: Vibrionaceae
Genus: Aliivibrio
Species:
A. fischeri
Binomial name
Aliivibrio fischeri
(Beijerinck1889)Urbanczyket al. 2007
Synonyms[1]

Aliivibrio fischeriis the familyVibrionaceae.This family of bacteria tend to have adaptable metabolisms that can adjust to diverse circumstances. This flexibility may contribute toA. fischeri'sability to survive both alone and in symbiotic relationships.[7]

Ribosomal RNAcomparison led to the reclassification of this species from genusVibrioto the newly createdAliivibrioin 2007.[8]The change is recognized as avalid publication,and according to the List of Prokaryotic names with Standing in Nomenclature (LPSN), thecorrect name.[9]However, the name change has not been universally adopted by most researchers, who still publish using the nameVibrio fischeri.[citation needed]

Genome

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ThegenomeofA. fischeriwas completelysequencedin 2004 and consists of twochromosomes,one smaller and one larger. Chromosome 1 has 2.9 millionbase pairs(Mbp) and chromosome 2 has 1.5 Mbp, bringing the total genome to 4.4 Mbp.[10]

A. fischerihas the lowestG+C contentof 27Vibriospecies but is still related to higher-pathogenicity species such asV. cholerae.The genome forA. fischerialso carriesmobile genetic elements.[11]The precise functions of these elements inA. fischeriare not fully understood. However, they are known to acquire new genes that are associated with virulence and resistance to environmental stresses in other bacterial genomes.[12]

Some strains ofA. fischeri,such as strain ES114, contain a plasmid. Theplasmidin strain ES114 is called pES100 and is most likely used for conjugation purposes. This purpose was determined based on the 45.8 kbp gene sequence, most of which codes for a type IV section system. The ability to preform conjugation can be helpful for both beneficial and pathogenic strains, as it allows for DNA exchange.[13]

There is evidence that the genome ofA. fischeriincludes pilus gene clusters. These clusters encode for many different kinds ofpili,which serve a variety of functions. In this species, there are pili used for pathogenesis, twitching motility, tight adhesion, and toxin-coregulation, and more.[13]

Ecology

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TheHawaiian bobtail squid,itsphotophorespopulated withAliivibrio fischeri

A. fischeriare globally distributed intemperateandsubtropicalmarine environments.[14]They can be foundfree-floatingin oceans, as well as associated with marine animals, sediment, and decaying matter.[14]A. fischerihave been most studied assymbiontsof marine animals, includingsquidsin the genusEuprymnaandSepiola,whereA. fischerican be found in the squids'light organs.[14]This relationship has been best characterized in theHawaiian bobtail squid(Euprymna scolopes).A. fischeriis the only species of bacteria inhabiting the squid's light organ,[15]despite an environment full of other bacteria.[7]

Symbiosis with the Hawaiian bobtail squid

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A. fischericolonization of the light organ of theHawaiian bobtail squid(Euprymna scolopes[16])is currently studied as a simple model for mutualistic symbiosis, as it contains only two species andA. fischerican be cultured in a lab and genetically modified.Aliivibrio fischeriutilizeschitinas a primary carbon and nitrogen source in its symbiosis with the Hawaiian bobtail squid. In the squid’s light organ,A. fischeribreaks down chitin intoN-acetylglucosamine(GlcNAc), which acts as both a nutrient and achemoattractant,guiding colonization.Chitinasesfacilitate this breakdown, while the regulatory protein NagC controls gene expression for chitin and GlcNAc use. The bacteria metabolize GlcNAc throughfermentationor respiration, supporting energy needs and bioluminescence, which are crucial for the mutualistic relationship with the squid.[7]This mutualistic symbiosis providesA. fischeriwith nutrients and a protected environment and helps the squid avoid predation using bioluminescence.

A. fischeriprovides luminescence by colonizing the light organ of the Hawaiian bobtail squid,[17]which is on its ventral side.[7]The organ luminesces at night, providing the squid withcounter-illumination camouflage.The light organs of some squid contain reflective plates that intensify and direct the light produced, due toproteinsknown asreflectins.They regulate the light intensity to match that of the sea surface below.[17]This strategy prevents the squid from casting a shadow on the ocean floor, helping it avoid predation during feeding.[7][17]TheA. fischeripopulation is maintained by daily cycles. About 90% ofA. fischeriare ejected by the squid every morning in a process known as "venting". The 10% of bacteria remaining in the squid replenish the bacterial population before the following night.[7]

A. fischeriarehorizontally acquiredby young squids from their environment. Venting is thought to provide the source from which newly hatched squid are colonized. This colonization induces developmental and morphological changes in the squid's light organ, which is translucent.[7][17]Morphological changes made byA. fischerido not occur when the microbe cannot luminesce, such as a decrease in the number of pores in the light organ. Additionally, if colonization byA. fischeriis abruptly removed by antibiotics, the ciliated epithelium of the light organ will regress.[16]These changes show that bioluminescence is truly essential for symbiosis.

In the process of colonization,ciliatedcells within the animals'photophores(light-producing organs) selectively draw in the symbiotic bacteria. These cells create microcurrents that, when combined with mucus,[16]promote the growth of the symbionts and actively reject any competitors. The bacteria cause the ciliated cells to die once the light organ is sufficiently colonized.[17]

Bioluminescence

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ThebioluminescenceofA. fischeriis caused bytranscriptionof theluxoperon,and the following translation of the lux proteins, which produce the light. This process is induced through population-dependentquorum sensing.[2]The population ofA. fischerineeds to reach an optimal level to activate theluxoperon and stimulate light production. Thecircadian rhythmcontrols light expression, where luminescence is much brighter during the day and dimmer at night, as required for camouflage.[18]

The bacterialluciferin-luciferasesystem is encoded by a set of genes labelled theluxoperon. InA. fischeri,five such genes (luxCDABEG) have been identified as active in the emission of visible light, and two genes (luxRandluxI) are involved in regulating theoperon.Several external and intrinsic factors appear to eitherinduceorinhibitthe transcription of this gene set and produce or suppresslight emission.[citation needed]

A. fischeriis one of many species of bacteria that commonly formsymbiotic relationshipswith marine organisms.[19]Marine organisms contain bacteria that use bioluminescence so they can find mates, ward off predators, attract prey, or communicate with other organisms.[20]In return, the organism the bacteria are living within provides the bacteria with a nutrient-rich environment.[21]Theluxoperon is a 9-kilobase fragment of theA. fischerigenome that controls bioluminescence through the catalytic activity of the enzyme luciferase.[22]This operon has a known gene sequence ofluxCDAB(F)E,whereluxAandluxBcode for the protein subunits of the luciferase enzyme, and theluxCDEcodes for a fatty acidreductasecomplex that makes thefatty acidsnecessary for the luciferase mechanism.[22]luxCcodes for the enzyme acyl-reductase,luxDcodes foracyl-transferase,andluxEmakes the proteins needed for the enzyme acyl-protein synthetase. Luciferase produces blue/green light through the oxidation of reducedflavin mononucleotideand a long-chainaldehydebydiatomic oxygen.The reaction is summarized as:[23]

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

The reduced flavin mononucleotide (FMNH) is provided by thefregene, also referred to asluxG.InA. fischeri,it is directly next toluxE(givingluxCDABE-fre) from 1042306 to 1048745.[24]

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-ACPto release R-COOH, a free fatty acid. R-COOH is reduced by a two-enzyme system to an aldehyde. The reaction is:[21]

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

Quorum sensing

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Quorum sensing inAliivibrio fischeri [25]
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 theluxoperonisquorum sensing,a conserved mechanism across many microbial species that regulates gene expression in response to bacterial concentration. Quorum sensing functions through the production of anautoinducer,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, altering gene expression. This system allows microbial cells to "communicate" amongst each other and coordinate behaviors, such as luminescence, which require large amounts of cells to produce a noticeable effect.[25]

InA. fischeri,there are two primary quorum sensing systems, each of which responds to slightly different environments. The first system is commonly referred to as theluxsystem, as it is encoded within theluxoperon, and uses the autoinducer 3OC6-HSL.[26]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 most notably upregulation of genes involved in luminescence.[27]The second system, commonly referred to as theainsystem, uses the autoinducer C8-HSL, which is produced by the protein AinS. Similar to theluxsystem, the autoinducer C8-HSL increases activation of LuxR. In addition, C8-HSL binds to another transcriptional regulator, LitR, giving theainandluxsystems of quorum sensing slightly different genetic targets within the cell.[28]

The different genetic targets of theainandluxsystems are essential, because these two systems respond to different cellular environments. Theainsystem regulates transcription in response to intermediate cell density cell environments, producing lower levels of luminescence and even regulating metabolic processes such as theacetate switch.[29]In contrast, theluxquorum sensing system occurs in response to high cell densities, producing high levels of luminescence and regulating the transcription of additional genes, including QsrP, RibB, and AcfA.[30]Both of theainandluxquorum sensing systems are essential for colonization of the squid and regulate multiple colonization factors in the bacteria.[27]

Activation of theluxoperon by LuxR and LuxI inAliivibrio fischeri [31][32]
(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 ofA. fischeri.A high intensity of light can be detected.

Research applications

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A. fischerihas broad applications inecotoxicologyand environmental research. Its bioluminescence is observed in oxygen-rich environments and thus is sensitive to toxicants.[33]Reductions in light emissions are used inbioassayssuch as theMicrotox testto assess water quality.[34]It plays a key role in studying the effects of chemical mixtures, helping identify synergistic or antagonistic toxic interactions.[35]In biotechnology, its light-producing mechanism is harnessed for developingbiosensorsthat detect environmental pollutants in real time, making it a valuable tool in pollution monitoring and water treatment studies.[36]Bioluminescence inhibition assays ofA. fischerican be used to measure for organicsolvents,heavy metals,[37]polycyclic aromatic hydrocarbons(PAH's),pesticides,[38]andtotal petroleum hydrocarbons(TPH's).[39]The bacteria’s adaptation to competitive marine environments, where they may produce unique bioactive compounds, may also position them as useful organisms for discovering novel antibiotics from marine sources.[36]

Natural transformation

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Natural bacterialtransformationis an adaptation for transferring DNA from one individual cell to another. Natural transformation, including the uptake and incorporation of exogenousDNAinto the recipientgenome,has been demonstrated inA. fischeri.[40]This process is induced bychitohexaoseand is likely regulated by genestfoXandtfoY.Natural transformation ofA. fischerifacilitates rapid transfer of mutant genes across strains and provides a valuable tool for experimental genetic manipulation in this species.[citation needed]

State microbe status

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In 2014,Hawaiʻian State SenatorGlenn Wakaisubmitted SB3124, proposingAliivibrio fischerias thestate microbeofHawaiʻi.[41]The bill competed with a bill advocating forFlavobacterium akiainvivensto receive the same designation; ultimately, neither bill passed. In 2017, similar legislation similar to the original 2013F. akiainvivensbill was submitted in theHawaiʻi House of RepresentativesbyIsaac Choy[42]and in theHawaiʻi SenatebyBrian Taniguchi,butA. fischeridid not appear in this or any later proposals.[43]

List of synonyms

  • Achromobacter fischeri(Beijerinck 1889) Bergey et al. 1930
  • Bacillus fischeri(Beijerinck 1889) Trevisan 1889
  • Bacterium phosphorescens indigenus(Eisenberg 1891) Chester 1897
  • Einheimischer leuchtbacillusFischer 1888
  • Microspira fischeri(Beijerinck 1889) Chester 1901
  • Microspira marina(Russell 1892) Migula 1900
  • Photobacterium fischeriBeijerinck 1889
  • Vibrio noctilucaWeisglass and Skreb 1963[1]

See also

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References

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