Autoinducer-2 (AI-2), a furanosyl borate diester or tetrahydroxy furan (species dependent), is a member of a family of

signaling molecule In biology, cell signaling (cell signalling in British English) or cell communication is the ability of a Cell (biology), cell to receive, process, and transmit signals with its environment and with itself. Cell signaling is a fundamental property ...

s used in

quorum sensing In biology, quorum sensing or quorum signalling (QS) is the ability to detect and respond to cell population density by gene regulation. As one example, QS enables bacteria to restrict the expression of specific genes to the high cell densities at ...

. AI-2 is one of only a few known

biomolecules A biomolecule or biological molecule is a loosely used term for molecules present in organisms that are essential to one or more typically biological processes, such as cell division, morphogenesis, or development. Biomolecules include lar ...

incorporating boron. First identified in the marine bacterium '' Vibrio harveyi'', AI-2 is produced and recognized by many

Gram-negative Gram-negative bacteria are bacteria that do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. They are characterized by their cell envelopes, which are composed of a thin peptidoglycan cell wa ...

and Gram-positive bacteria. AI-2 arises by the reaction of

4,5-Dihydroxy-2,3-pentanedione 4,5-Dihydroxy-2,3-pentanedione (DPD) is an organic compound that occurs naturally but exists as several related structures. The idealized formula for this species is CH3C(O)C(O)CH(OH)CH2OH, but it is known to exist as several other forms resultin ...

, which is produced enzymatically with

boric acid Boric acid, more specifically orthoboric acid, is a compound of boron, oxygen, and hydrogen with formula . It may also be called hydrogen borate or boracic acid. It is usually encountered as colorless crystals or a white powder, that dissolve ...

, and is recognized by the two-component sensor kinase LuxPQ in

Vibrionaceae The Vibrionaceae are a family of Pseudomonadota given their own order, Vibrionales. Inhabitants of fresh or salt water, several species are pathogenic, including the type species ''Vibrio cholerae'', which is the agent responsible for cholera. ...

. AI-2 is actively transported by the Lsr ABC-type transporter into the cell in Enterobacteriaceae and few other bacterial taxa (''

Pasteurella __NOTOC__ ''Pasteurella'' is a genus of Gram-negative, facultatively anaerobic bacteria. ''Pasteurella'' species are nonmotile and pleomorphic, and often exhibit bipolar staining ("safety pin" appearance). Most species are catalase- and oxidase ...

'', '' Photorhabdus'', ''

Haemophilus ''Haemophilus'' is a genus of Gram-negative, pleomorphic, coccobacilli bacteria belonging to the family Pasteurellaceae. While ''Haemophilus'' bacteria are typically small coccobacilli, they are categorized as pleomorphic bacteria because of t ...

'', ''

Bacillus ''Bacillus'' (Latin "stick") is a genus of Gram-positive, rod-shaped bacteria, a member of the phylum '' Bacillota'', with 266 named species. The term is also used to describe the shape (rod) of other so-shaped bacteria; and the plural ''Bacill ...

''), where it is phosphorylated by LsrK. Then, Phospho-AI-2 binds the transcriptional repressor protein, LsrR, which subsequently is released from the promoter/operator region of the lsr operon – and transcription of the ''lsr'' genes is initiated. AI-2 signalling is also regulated by glucose and cAMP/CRP via the ''lsr'' operon. In the presence of glucose, low levels of cAMP/CRP result in almost no ''lsr'' operon (''lsrABCDFG'') expression. Without glucose, cAMP-CRP is needed to stimulate the ''lsr'' expression, while LsrR represses its expression in the absence of the inducer, phospho-AI-2. As AI-2 accumulates, more AI-2 is taken in via LsrABCD, phosphorylated via LsrK, and the ''lsr'' transcription is de-repressed, enabling even more AI-2 uptake. Doubts have been expressed regarding AI-2's status as a universal signal. Although the ''luxS'' gene, which encodes the protein responsible for AI-2 production is widespread, the latter has mainly a primary metabolic role in the recycling of

S-adenosyl-L-methionine ''S''-Adenosyl methionine (SAM), also known under the commercial names of SAMe, SAM-e, or AdoMet, is a common cosubstrate involved in methyl group transfers, transsulfuration, and aminopropylation. Although these anabolic reactions occur through ...

, with AI-2 being a by-product of that process. An unequivocally AI-2 related behavior was found to be restricted primarily to organisms bearing known AI-2 receptor genes. Thus, while it is certainly true that some bacteria respond to AI-2, it is doubtful that it is always being produced for purposes of signaling.

References

{{Reflist Signal transduction Cell biology Cell communication Cell signaling Tetrahydroxyborate esters