In order to provide environmentally friendly light sources, bioluminescent organisms such as the species of bacteria called Vibrio fischeri, could be utilized; however, the natural light from bioluminescent organisms is not bright enough to be effective. Download as PDF. Two‐Component Signaling Systems, Part B. Although the ecological rationale for bioluminescence in V. The natural bioluminescent bacteria can. BACTERIAL BIOLUMINESCENCE 497 lished data). All exhibit high specificity for FMNH2andalong-chain aliphatic aldehyde (8 to 16 carbons) (41, 97). For luciferases from different strains ofa given species, the kinetics of decay with a given aldehyde are similar (J. M.Fitzgerald, Ph.D.thesis, MonashUniversity.

Abstract

We report two Australian patients with soft tissue infections due to Photorhabdus species. Recognized as important insect pathogens, Photorhabdus spp. are bioluminescent gram-negative bacilli. Bacteria belonging to the genus are emerging as a cause of both localized soft tissue and disseminated infections in humans in the United States and Australia. The source of infection in humans remains unknown.

Bioluminescence is the production of visible light by a chemical reaction in a living organism. The property is rarely reported in the clinical bacteriology laboratory because bacterial bioluminescence is seen primarily in marine species. Photorhabdus spp (family: Enterobacteriaceae) are the only terrestrial bacteria known to exhibit this property (1). The classification within the genus is complex with three currently recognized species: P. luminescens, P. temperate, and P. asymbiotica (). Several subspecies are recognized.

Photorhabdus spp. have been the subject of intensive study by agricultural scientists because of the role these bacteria play in controlling insects. Insects, like humans, are subject to infestation by nematodes (3). Photorhabdus spp. inhabit the gut of some insect-pathogenic nematodes (Heterorhabditis spp.), where they form a symbiotic relationship. Nematode species of this type are able to invade the larvae of susceptible insects and release Photorhabdus spp. The bacteria proliferate and promote nematode reproduction by killing the insect larvae.

Insect-pathogenic nematodes harboring Photorhabdus spp are used as biopesticides in a number of countries, including the United States and Australia. Agricultural scientists are also attempting to develop insect-resistant transgenic crops by using insecticidal toxin genes derived from Photorhabdus spp ().

Genes encoding homologues of insecticidal toxins from Photorhabdus spp occur naturally within the genome of Yersinia pestis, the cause of plague. Lateral transfer of genetic material between Photorhabdus and Yersinia species is thought to have resulted from their common association with insects as bacterial pathogens ().

Human infection with Photorhabdus spp. has been described in two previous publications—six cases from the United States () and four cases from South Eastern Australia (Victoria and New South Wales) (). We report two additional recent human cases of Photorhabdus infection from the Australian state of Queensland.

The Study

Patient 1

A 39-year-old male pest controller from Gladstone on a routine visit to his general practitioner in April 2001 inquired about the recent appearance of a red macule, 8 mm in diameter, on the medial aspect of his right ankle. No specific treatment was given. When he was seen again 18 days later, a painful, necrotic ulcer, about 12 mm in diameter, had developed at the original site of the red spot. An exudates was produced, although polymorphs were observed rarely, and no microorganisms were identified on Gram stain. A gram-negative organism later identified as Photorhabdus sp. was isolated in pure growth from the exudate. The patient began a 10-day course of oral cephalexin. When he was observed again 11 days later, he exhibited a persistent discharge with surrounding cellulitis. He was therefore prescribed a 10-day course of oral amoxycillin-clavulanate. Three weeks later, the ulcer appeared to be healing; after another 6 weeks, signs of infection had again developed. A gram-negative organism was isolated from the exudate but was not formally identified.

The patient was prescribed an additional 7-day course of oral cephalexin. When he was observed 3 months later, the infection had resolved. In his recent work as a pest controller, he had been spraying chemical insecticides under houses and in foreign cargo ships. He had never used insect pathogenic nematodes as a biopesticide.

Patient 2

A 78-year-old man from the Queensland Gold Coast sought treatment in January 1999 with a 3-day history of a painful, swollen right foot. The patient had a history of polymyalgia rheumatica for which he was taking prednisone, 8 mg daily. In January 1999, after working barefoot in the garden, the man noted intense pain in his right forefoot and a very small amount of bloody discharge from the web space between his fourth and fifth toes.

The next day he was seen by his general practitioner who treated him with oral dicloxacillin. Two days later he was admitted to the hospital with increasingly severe pain with extensive redness and swelling extending to his right knee. He was noted to be afebrile with a mild neutrophil leukocytosis. He was started on a regimen of intravenous dicloxacillin and gentamicin.

Surgical debridement of the right foot was required on three occasions during the first 8 days of his admission. Pus was collected for culture on two of these occasions, and tissue was obtained during the third. An organism identified as Photorhabdus sp. was isolated in pure culture from each of these operative specimens. The same organism was also isolated, together with Staphylococcus aureus, from a superficial swab collected in the emergency department on presentation. No bacterial growth was obtained from blood cultures collected on admission.

The patient was treated with intravenous gentamicin for 2 weeks and ceftazidime for 1 week. He was discharged on a 6-week course of oral ciprofloxacin. The foot remained healed on follow-up 3 months later.

Photorhabdus spp. can be isolated and identified to genus level by using techniques available in most clinical bacteriology laboratories. A total of five isolates from the two patients described in the current report were examined in our laboratories with standard techniques (one from patient 1 and four from patient 2). The phenotypic characteristics that the isolates displayed were typical of the genus.

Colonies were formed after 24–48 hours on tryptic soy agar containing either 5% sheep or horse blood (bioMérieux, Baulkham Hills, Australia) at both 35°C and at room temperature, with a tendency to “swarm” (Figure 1). The isolates also grew on MacConkey agar. On sheep and horse blood agar, a thin line of annular hemolysis was observed 4–12 mm from the colony edge. The hemolysis was more apparent when the isolates were incubated at room temperature (Figure 2). The organisms were motile, gram-negative, rod-shaped bacteria. They were facultatively anaerobic, oxidase negative, and strongly catalase positive. Other biochemical reactions were as described previously ().

Photorhabdus isolate from patient 2, growing on tryptic soy agar containing 5% sheep blood, after 48 hours’ incubation at 35°C. Arrows indicate “swarming.” The colonies could be seen to glow faintly with the naked eye under conditions of total darkness after 10 minutes of adjustment.

Photorhabdus isolate from patient 2 after 5 days growth at room temperature on sheep blood agar. Arrows indicate the characteristic thin line of “annular” hemolysis surrounding the colonies.

The defining characteristic was the presence of faint luminescence, which could be clearly seen with the naked eye when the colonies were examined under conditions of total darkness. It was critical to this examination that the observer’s eyes be allowed to adjust to the darkness for 10 minutes.

Two commercially available automated bacterial identification systems were used in our laboratories: MicroScan Walkaway (Dade Behring Inc., MicroScan Division, West Sacramento, CA) and bioMerieux Vitek (bioMérieux; Hazelwood, MO). Photorhabdus spp. do not currently appear on the databases of either of these systems, which leads to misidentification (Table 1).

Table 1

Photorhabdus spp. have been shown to form a heterogeneous group based on DNA-DNA hybridization studies, 16S rDNA sequencing and polymerase chain reaction ribotyping (). A polyphasic approach is now applied to classifying isolates within the genus, dividing it into three species and several subspecies. The American clinical isolates described by Farmer et al. () belong to a new species, Photorhabdus asymbiotica (). A specific epithet has not yet been assigned to the Australian clinical isolates but they also may form a new species within the genus (7).

Antimicrobial sensitivity was assessed by using broth microdilution. The isolates were sensitive to a broad range of antimicrobial agents with activity against gram-negative bacteria including ciprofloxacin, gentamicin, tetracycline, ceftriaxone, and amoxycillin-clavulanate. Isolates from both patients were resistant to cephalothin and ampicillin.

Conclusions

Publication of information about these two cases brings to a total of 12 the number of human infections with Photorhabdus spp. documented in the medical literature (Table 2 and Figure 3). The clinical picture described in the 12 cases has generally been one of localized or more commonly multifocal skin/soft tissue infection. Such infection has had a tendency to relapse. The disseminated distribution of skin/soft tissue infection in several cases suggests hematogenous spread. Bacteremia was documented in 4/12 case-patients. Cough was documented in two of the bacteremic case-patients. In one of these, isolates of a Photorhabdus sp. were obtained from sputum as well as from blood and skin/soft tissue.

Table 2

Australian and American clinical isolates of Photorhabdus.

Given the very limited clinical experience, making definitive recommendations about treatment is not possible. Antimicrobial therapy should be guided by in vitro sensitivities. The tendency for Photorhabdus infection to relapse suggests that prolonged therapy for a period of weeks would be prudent, perhaps with an oral fluoroquinolone.

Photorhabdus spp are not human commensals. The patients apparently acquired the pathogen from an unidentified source in the terrestrial environment. This hypothesis is supported by the observations that at least 4/6 of the Australian patients were engaged in outdoor activities around the time of acquisition and that the initial site of infection was on the lower limbs in more than half of Australian and American case-patients.

Photorhabdus spp. have never been shown to live freely in soil, although they will survive in soil under laboratory conditions (). Photorhabdus spp. have only been isolated naturally from two nonclinical sources: insect-pathogenic nematodes (Heterorhabditis spp) and the insects they parasitize (beetles, moths, and the like). It seems likely therefore that Photorhabdus spp are transmitted to humans by a terrestrial invertebrate (nematode or arthropod), but that vector has not yet been identified.

Biography

Dr. Gerrard is Director of Infectious Diseases at the Gold Coast Hospital and a clinical senior lecturer at the University of Queensland, Australia. His research interests include clinical and laboratory aspects of emerging bacterial pathogens.

References

Bioluminescent bacteria are light-producingbacteria 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.^[1] These bacteria may be free living (such as Vibrio harveyi) or in symbiosis with animals such as the Hawaiian Bobtail squid (Aliivibrio fischeri) or terrestrial nematodes (Photorhabdus luminescens). 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.^[2] Another possible reason bacteria use luminescence reaction is for quorum sensing, an ability to regulate gene expression in response to bacterial cell density.^[3]

• 6Evolution of bio-luminescence
• 8Bacterial groups that exhibit bioluminescence
• 10Mechanism

History[edit]

Records of bioluminescent bacteria existed for thousands of years.^[4] They appear in the folklore of many regions, including Scandinavia and the Indian subcontinent. Both Aristotle and Charles Darwin have described the phenomenon of the oceans glowing.^[4] Since its discovery less than 30 years ago, the enzyme luciferase and its regulatory gene, lux, have led to major advances in molecular biology, through use as a reporter gene.^[5]Luciferase was first purified by McElroy and Green in 1955.^[6] It was later discovered that there were two subunits to luciferase, called subunits α and β. The genes encoding these enzymes, luxA and luxB, respectively, were first isolated in the lux operon of Aliivibrio fisheri^[4].

Purpose of bio-luminescence[edit]

The wide-ranged biological purposes of bio-luminescence include but are not limited to attraction of mates,^[7] defense against predators, and warning signals. In the case of bioluminescent bacteria, bio-luminescence mainly serves as a form of dispersal. It has been hypothesized that enteric bacteria (bacteria that survive in the guts of other organisms) - especially those prevalent in the depths of the ocean - employ bio-luminescence as an effective form of distribution.^[8] After making their way into the digestive tracts of fish and other marine organisms and being excreted in fecal pellets, bioluminescent bacteria are able to utilize their bio-luminescent capabilities to lure in other organisms and prompt ingestion of these bacterial-containing fecal pellets.^[9] The bio-luminescence of bacteria thereby ensures their survival, persistence, and dispersal as they are able to enter and inhabit other organisms.

Regulation of bio-luminescence[edit]

The regulation of bio-luminescence in bacteria is achieved through the regulation of the oxidative enzyme called luciferase. It is important that bio-luminescent bacteria decrease production rates of luciferase when the population is sparse in number in order to conserve energy. Thus, bacterial bioluminescence is regulated by means of chemical communication referred to as quorum sensing.^[10] Essentially, certain signaling molecules named autoinducers^[11] with specific bacterial receptors become activated when the population density of bacteria is high enough. The activation of these receptors leads to a coordinated induction of luciferase production that ultimately yields visible luminescence.^[12]

Biochemistry of bio-luminescence[edit]

The chemical reaction that is responsible for bio-luminescence is catalyzed by the enzyme luciferase. In the presence of oxygen, luciferase catalyzes the oxidation of an organic molecule called luciferin.^[13] Though bio-luminescence across a diverse range of organisms such as bacteria, insects, and dinoflagellates function in this general manner (utilizing luciferase and luciferin), there are different types of luciferin-luciferase systems. For bacterial bio-luminescence specifically, the biochemical reaction involves the oxidation of an aliphaticaldehyde by a reduced flavin mononucleotide.^[14] The products of this oxidation reaction include an oxidized flavin mononucleotide, a fatty acid chain, and energy in the form of a blue-green visible light.^[15]

Reaction: FMNH2 + O2 + RCHO → FMN + RCOOH + H2O + light

Bioelectromagnetics of bio-luminescence[edit]

How Bioluminescence Works

It has been proposed that bioelectromagnetics may be involved in biological processes behind bio-luminescence, which may function as a pump. This pump may involve mm and sub-mm wave coupling of bio-luminescence radiation for quorum sensing regulation. This proposal arises from the observation that mm-wave radiation exposure has been reported to induce changes in DNA conformation and possibly gene expression. Agents that change DNA conformation also increase light emission in bio-luminescence of bacteria.^[16]

Evolution of bio-luminescence[edit]

Of all light emitters in the ocean, bio-luminescent bacteria is the most abundant and diverse. However, the distribution of bio-luminescent bacteria is uneven, which suggests evolutionary adaptations. The bacterial species in terrestrial genera such as Photorhabdus are bio-luminescent. On the other hand, marine genera with bio-luminescent species such as Vibrio and Shewanella oneidensis have different closely related species that are not light emitters.^[17] Nevertheless, all bio-luminescent bacteria share a common gene sequence: the enzymatic oxidation of Aldehyde and reduced Flavin mononucleotide by luciferase which are contained in the lux operon.^[18] Bacteria from distinct ecological niches contain this gene sequence; therefore, the identical gene sequence evidently suggests that bio-luminescence bacteria result from evolutionary adaptations.

Use as laboratory tool[edit]

After the discovery of the lux operon, the use of bioluminescent bacteria as a laboratory tool is claimed to have revolutionized the area of environmental microbiology.^[4] The applications of bioluminescent bacteria include biosensors for detection of contaminants, measurement of pollutant toxicity ^[4][19] and monitoring of genetically engineered bacteria released into the environment.^[20][21][22] Biosensors, created by placing a lux gene construct under the control of an inducible promoter, can be used to determine the concentration of specific pollutants.^[4] Biosensors are also able to distinguish between pollutants that are bioavailable and those that are inert and unavailable.^[4] For example, Pseudomonas fluorescens has been genetically engineered to be capable of degrading salicylate and naphthalene, and is used as a biosensor to assess the bioavailability of salicylate and naphthalene.^[4] Biosensors can also be used as an indicator of cellular metabolic activity and to detect the presence of pathogens.^[4]

Evolution[edit]

The light-producing chemistry behind bioluminescence varies across the lineages of bioluminescent organisms.^[17] Based on this observation, bioluminescence is believed to have evolved independently at least 40 times.^[17] In bioluminescent bacteria, the reclassification of the members ofVibrio fischeri species group as a new genus, Aliivibrio, has led to increased interest in the evolutionary origins of bioluminescence^[17]. Among bacteria, the distribution of bioluminescent species is polyphyletic. For instance, while all species in the terrestrial genus Photorhabdus are luminescent, the genera Aliivibrio, Photobacterium, Shewanella and Vibrio contain both luminous and non-luminous species.^[17] Despite bioluminescence in bacteria not sharing a common origin, they all share a gene sequence in common. The appearance of the highly conserved lux operon in bacteria from very different ecological niches suggests a strong selective advantage despite the high energetic costs of producing light. DNA repair is thought to be the initial selective advantage for light production in bacteria.^[17] Consequently, the lux operon may have been lost in bacteria that evolved more efficient DNA repair systems but retained in those where visible light became a selective advantage.^[17][23] The evolution of quorum sensing is believed to have afforded further selective advantage for light production. Quorum sensing allows bacteria to conserve energy by ensuring that they do not synthesize light-producing chemicals unless a sufficient concentration are present to be visible.^[17]

Bacterial groups that exhibit bioluminescence[edit]

All bacterial species that have been reported to possess bioluminescence belong within the families Vibrionaceae, Shewanellaceae, or Enterobacteriaceae, all of which are assigned to the class Gammaproteobacteria.^[24]

Bioluminescence For Kids

(List from Dunlap and Henryk (2013), 'Luminous Bacteria', The Prokaryotes^[24])

Distribution[edit]

Bioluminescent bacteria are most abundant in marine environments during spring blooms when there are high nutrient concentrations. These light-emitting organisms are found mainly in coastal waters near the outflow of rivers, such as the northern Adriatic Sea, Gulf of Trieste, northwestern part of the Caspian Sea, coast of Africa and many more.^[25] These are known as milky seas. Bioluminescent bacteria are also found in freshwater and terrestrial environments but are less wide spread than in seawater environments. They are found globally, as free-living, symbiotic or parasitic forms ^[1] and possibly as opportunistic pathogens.^[24] Factors that affect the distribution of bioluminescent bacteria include temperature, salinity, nutrient concentration, pH level and solar radiation.^[26] For example, Aliivibriofischeri grows favourably in environments that have temperatures between 5 to 30 °C and a pH that is less than 6.8; whereas, Photobacterium phosphoreum thrives in conditions that have temperatures between 5 and 25 °C and a pH that is less than 7.0.^[27]

Genetic diversity[edit]

All bioluminescent bacteria share a common gene sequence: the lux operon characterized by the luxCDABE gene organization.^[24]LuxAB codes for luciferase while luxCDE codes for a fatty-acid reductase complex that is responsible for synthesizing aldehydes for the bioluminescent reaction. Despite this common gene organization, variations, such as the presence of other lux genes, can be observed among species. Based on similarities in gene content and organization, the lux operon can be organized into the following four distinct types: the Aliivibrio/Shewanella type, the Photobacterium type, theVibrio/Candidatus Photodesmus type, and the Photorhabdus type. While this organization follows the genera classification level for members of Vibrionaceae (Aliivibrio, Photobacterium, and Vibrio), its evolutionary history is not known.^[24]

With the exception of the Photorhabdus operon type, all variants of the lux operon contain the flavin reductase-encoding luxG gene.^[24] Most of the Aliivibrio/Shewanella type operons contain additional luxI/luxR regulatory genes that are used for autoinduction during quorum sensing.^[28] The Photobacterum operon type is characterized by the presence of rib genes that code for riboflavin, and forms the lux-rib operon. TheVibrio/Candidatus Photodesmus operon type differs from both the Aliivibrio/Shewanella and the Photobacterium operon types in that the operon has no regulatory genes directly associated with it.^[24]

Mechanism[edit]

All bacterial luciferases are approximately 80 KDa heterodimers containing two subunits: α and β. The α subunit is responsible for light emission.^[4] The luxA and luxB genes encode for the α and β subunits, respectively. In most bioluminescent bacteria, the luxA and luxB genes are flanked upstream by luxC and luxD and downstream by luxE.^[4]

The bioluminescent reaction is as follows:

FMNH₂ + O₂ + R-CHO -> FMN + H₂O + R-COOH + Light (~ 495 nm)

Molecular oxygen reacts with FMNH2 (reduced flavin mononucleotide) and a long-chain aldehyde to produce FMN (flavin mononucleotide), water and a corresponding fatty acid. The blue-green light emission of bioluminescence, such as that produced by Photobacterium phosphoreum and Vibro harveyi, results from this reaction.^[4] Because light emission involves expending six ATP molecules for each photon, it is an energetically expensive process. For this reason, light emission is not constitutively expressed in bioluminescent bacteria; it is expressed only when physiologically necessary.

Quorum sensing[edit]

Bioluminescence in bacteria can be regulated through a phenomenon known as autoinduction or quorum sensing.^[4]Quorom sensing is a form of cell-to-cell communication that alters gene expression in response to cell density. Autoinducer is a diffusible pheromone produced constitutively by bioluminescent bacteria and serves as an extracellular signalling molecule.^[4] When the concentration of autoinducer secreted by bioluminescent cells in the environment reaches a threshold (above 10⁷ cells per mL), it induces the expression of luciferase and other enzymes involved in bioluminescence.^[4] Bacteria are able to estimate their density by sensing the level of autoinducer in the environment and regulate their bioluminescence such that it is expressed only when there is a sufficiently high cell population. A sufficiently high cell population ensures that the bioluminescence produced by the cells will be visible in the environment.

A well known example of quorum sensing is that which occurs between Aliivibrio fischeri and its host. This process is regulated by LuxI and LuxR, encoded by luxI and luxR respectively. LuxI is autoinducer synthase that produces autoinducer (AI) while LuxR functions as both a receptor and transcription factor for the lux operon.^[4] When LuxR binds AI, LuxR-AI complex activates transcription of the lux operon and induces the expression of luciferase.^[28] Using this system, A. fischeri has shown that bioluminescence is expressed only when the bacteria are host-associated and have reached sufficient cell densities.^[29]

Another example of quorum sensing by bioluminescent bacteria is by Vibrio harveyi, which are known to be free-living. Unlike Aliivibrio fischeri,V. harveyi do not possess the luxI/luxR regulatory genes and therefore have a different mechanism of quorum sensing regulation. Instead, they use the system known as three-channel quorum sensing system.^[30]

Role[edit]

The uses of bioluminescence and its biological and ecological significance for animals, including host organisms for bacteria symbiosis, have been widely studied. Its benefits for bacteria, however, still remain unclear.^[4]

One explanation for the role of bacterial bioluminescence is from the biochemical aspect. Several studies have shown the biochemical roles of the luminescence pathway. It can function as an alternate pathway for electron flow under low oxygen concentration, which can be advantageous when no fermentable substrate is available.^[1] In this process, light emission is a side product of the metabolism.

Evidence also suggests that bacterial luciferase contributes to the resistance of oxidative stress. In laboratory culture, luxA and luxB mutants of Vibrio harveyi, which lacked luciferase activity, showed impairment of growth under high oxidative stress compared to wild type. The luxD mutants, which had an unaffected luciferase but were unable to produce luminescence, showed little or no difference. This suggests that luciferase mediates the detoxification of reactive oxygen.^[31]

Bacterial bioluminescence has also been proposed to be a source of internal light in photoreactivation, a DNA repair process carried out by photolyase.^[32] Experiments have shown that non-luminescent V. harveyi mutants are more sensitive to UV irradiation, suggesting the existence of a bioluminescent-mediated DNA repair system.^[23]

Another hypothesis, called the “bait hypothesis”, is that bacterial bioluminescence attracts predators who will assist in their dispersal.^[32] They are either directly ingested by fish or indirectly ingested by zooplankton that will eventually be consumed by higher trophic levels. Ultimately, this may allow passage into the fish gut, a nutrient-rich environment where the bacteria can divide, be excreted, and continue their cycle. Experiments using luminescent Photobacterium leiognathi and non-luminescent mutants have shown that luminescence attracts zooplankton and fish, thus supporting this hypothesis.^[32]

Symbiosis with other organisms[edit]

The symbiotic relationship between the Hawaiian bobtail squid Euprymna scolopes and the marine gram-negative bacterium Aliivibrio fischeri has been well studied. The two organisms exhibit a mutualistic relationship in which bioluminescence produced by A. fischeri helps to attract pray to the squid host, which provides nutrient-rich tissues and a protected environment forA. fischeri.^[33] Bioluminescence provided by A. fischeri also aids in the defense of the squid E. scolopes by providing camouflage during its nighttime foraging activity.^[34] Following bacterial colonization, the specialized organs of the squid undergo developmental changes and a relationship becomes established. The squid expels 90% of the bacterial population each morning, because it no longer needs to produce bioluminescence in the daylight.^[4] This expulsion benefits the bacteria by aiding in their dissemination. A single expulsion by one bobtail squid produces enough bacterial symbionts to fill 10,000m³ of seawater at a concentration that is comparable to what is found in coastal waters.^[34] Thus, in at least some habitats, the symbiotic relationship between A. fischeri and E. scolopes plays a key role in determining the abundance and distribution of E. scolopes. There is a higher abundance of A. fischeri in the vicinity of a population of E. scolopes and this abundance markedly decreases with increasing distance from the host’s habitat.^[34]

Bioluminescent Photobacterium species also engage in mutually beneficial associations with fish and squid.^[35] Dense populations of P. kishitanii, P. leiogathi, and P. mandapamensis can live in the light organs of marine fish and squid, and are provided with nutrients and oxygen for reproduction^[35] in return for providing bioluminescence to their hosts, which can aid in sex-specific signaling, predator avoidance, locating or attracting prey, and schooling.^[35]

Bioluminescence In Bacteria Pdf

See also[edit]

References[edit]

Pathogenic Bacteria Pdf

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Bioluminescence Bacteria Definition

Further reading[edit]

• Hastings, J W; Nealson, K H (1977). 'Bacterial Bioluminescence'. Annual Review of Microbiology. 31 (1): 549–595. doi:10.1146/annurev.mi.31.100177.003001. ISSN0066-4227. PMID199107.

Bacteria