Lambda phage - Knowledge

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The Q gene is a regulator gene found on this operon, which will control the expression of later genes for rightward transcription. Once the gene's regulatory proteins allow for expression, the Q protein will then act as an anti-terminator. This will then allow for the rest of the operon to be read through until it reaches the transcription terminator. Thus allowing expression of later genes in the operon, and leading to the expression of the lytic cycle.

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competitive inhibitor to the relevant proteases. This means that a cell "in trouble", i.e. lacking in nutrients and in a more dormant state, is more likely to lysogenise. This would be selected for because the phage can now lie dormant in the bacterium until it falls on better times, and so the phage can create more copies of itself with the additional resources available and with the more likely proximity of further infectable cells.

132: 36: 1088: 1066: 870: 824: 815: 669: 2971:. The 48 kb DNA fragment of lambda phage is not essential for productive infection and can be replaced by foreign DNA, which can then be replicated by the phage. Lambda phage will enter bacteria more easily than plasmids, making it a useful vector that can either destroy or become part of the host's DNA. Lambda phage can also be manipulated and used as an anti-cancer vaccine that targets human 77: 2871:, DNA binding protein and RNApol cofactor, binds DNA (at Qut sites) and transfers onto the initiating RNApol. This RNApol modification alters its recognition of termination sequences, so normal ones are ignored; special Q termination sequences some 20,000 bp away are effective. Q-extended transcripts include phage structural proteins (A-F, Z-J) and lysis genes ( 2847:, RNA-binding protein and RNA polymerase cofactor, binds RNA (at Nut sites) and transfers onto the nascent RNApol that just transcribed the nut site. This RNApol modification prevents its recognition of termination sites, so normal RNA polymerase termination signals are ignored and RNA synthesis continues into distal phage genes ( 968:, an enzyme that escapes through the S holes and cleaves the cell wall. Rz and Rz1 are membrane proteins that form a complex that somehow destroys the outer membrane, after the endolysin has degraded the cell wall. For wild-type lambda, lysis occurs at about 50 minutes after the start of infection and releases around 100 virions. 2159: 562: 2934:

A full biophysical model for lambda's lysis-lysogeny decision remains to be developed. Computer modeling and simulation suggest that random processes during infection drive the selection of lysis or lysogeny within individual cells. However, recent experiments suggest that physical differences among

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In cells with sufficient nutrients, protease activity is high, which breaks down cII. This leads to the lytic lifestyle. In cells with limited nutrients, protease activity is low, making cII stable. This leads to the lysogenic lifestyle. cIII appears to stabilize cII, both directly and by acting as a

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The prophage is duplicated with every subsequent cell division of the host. The phage genes expressed in this dormant state code for proteins that repress expression of other phage genes (such as the structural and lysis genes) in order to prevent entry into the lytic cycle. These repressive proteins

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Pr promoter has been found to activate the origin in the use of rightward transcription, but the whole picture of this is still somewhat misunderstood. Given there are some caveats to this, for instance this process is different for other phages such as N15 phage, which may encode for DNA polymerase.

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Pr is the promoter for rightward transcription, and the cro gene is a regulator gene. The cro gene will encode for the Cro protein that will then repress Prm promoter. Once Pr transcription is underway the Q gene will then be transcribed at the far end of the operon for rightward transcription.

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Lambda phage is a non-contractile tailed phage, meaning during an infection event it cannot 'force' its DNA through a bacterial cell membrane. It must instead use an existing pathway to invade the host cell, having evolved the tip of its tail to interact with a specific pore to allow entry of its DNA

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The virus particle consists of a head and a tail that can have tail fibers. The whole particle consists of 12–14 different proteins with more than 1000 protein molecules total and one DNA molecule located in the phage head. However, it is still not entirely clear whether the L and M proteins are part

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An important distinction here is that between the two decisions; lysogeny and lysis on infection, and continuing lysogeny or lysis from a prophage. The latter is determined solely by the activation of RecA in the SOS response of the cell, as detailed in the section on induction. The former will also

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The tail of lambda phages is made of at least 6 proteins (H, J, U, V, Stf, Tfa) and requires 7 more for assembly (I, K, L, M, Z, G/T). This assembly process begins with protein J, which then recruits proteins I, L, K, and G/T to add protein H. Once G and G/T leave the complex, protein V can assemble

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Multiplicity reactivation (MR) is the process by which multiple viral genomes, each containing inactivating genome damage, interact within an infected cell to form a viable viral genome. MR was originally discovered with phage T4, but was subsequently found in phage λ (as well as in numerous other

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N protein binds to boxB in each transcript, and contacts the transcribing RNA polymerase via RNA looping. The N-RNAP complex is stabilized by subsequent binding of several host Nus (N utilisation substance) proteins (which include transcription termination/antitermination factors and, bizarrely, a

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The Q protein modifies the RNA polymerase at the promoter region and is recruited to RNA polymerase. The Q protein turns into a RNA polymerase subunit after it is recruitment to RNAP and modifies the enzyme into a processive state. Note that NusA can stimulate the activity of the Q

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Survival of UV-irradiated phage λ is increased when the E. coli host is lysogenic for an homologous prophage, a phenomenon termed prophage reactivation. Prophage reactivation in phage λ appears to occur by a recombinational repair process similar to that of MR.

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promoter's reading frame, so no lytic or structural proteins are made. Elevated levels of int (much higher than that of xis) result in the insertion of the lambda genome into the hosts genome (see diagram). Production of cI leads to the binding of cI to the

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genes. Gam proteins are involved in recombination. Gam is also important in that it inhibits the host RecBCD nuclease from degrading the 3’ ends in rolling circle replication. Int and xis are integration and excision proteins vital to lysogeny.

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result in no insertion or excision of phage genomes, the evolutionarily favoured action - leaving any pre-inserted phage genomes inserted (so reducing competition) and preventing the insertion of the phage genome into the genome of a doomed

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phase, during which it kills and lyses the cell to produce offspring. Lambda strains, mutated at specific sites, are unable to lysogenize cells; instead, they grow and enter the lytic cycle after superinfecting an already lysogenized cell.

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be affected by this; a cell undergoing an SOS response will always be lysed, as no cI protein will be allowed to build up. However, the initial lytic/lysogenic decision on infection is also dependent on the cII and cIII proteins.

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Iwagami, Yoshifumi; Casulli, Sarah; Nagaoka, Katsuya; Kim, Miran; Carlson, Rolf I.; Ogawa, Kosuke; Lebowitz, Michael S.; Fuller, Steve; Biswas, Biswajit; Stewart, Solomon; Dong, Xiaoqun; Ghanbari, Hossein; Wands, Jack R. (2017).

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Lysogen repression all 3 sites bound is a low occurrence due to OR3 weak binding affinity. OR1 repression increases binding affinity to OR2 due to repressor-repressor interaction. Increased concentrations of repressor increase

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The life cycle of lambda phages is controlled by cI and Cro proteins. The lambda phage will remain in the lysogenic state if cI proteins predominate, but will be transformed into the lytic cycle if cro proteins predominate.

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found in the phage lambda is a notable example of the level of control possible over gene expression by a very simple system. It forms a 'binary switch' with two genes under mutually exclusive expression, as discovered by

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bacterial and mammalian viruses). MR of phage λ inactivated by UV light depends on the recombination function of either the host or of the infecting phage. Absence of both recombination systems leads to a loss of MR.

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similar to that used by phage M13. The DNA is nicked and the 3’ end serves as a primer. Note that this does not release single copies of the phage genome but rather one long molecule with many copies of the genome: a

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transcribes these regions, it recruits N and forms a complex with several host Nus proteins. This complex skips through most termination sequences. The extended transcripts (the 'late early' transcripts) include the

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promoters are no longer repressed and switch on, and the cell returns to the lytic sequence of expression events (note that cII is not stable in cells undergoing the SOS response). There is however one notable

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N Antitermination requires the assembly of a large ribonucleoprotein complex to effectively prolong the anti-termination process, without the full complex the RNA polymerase is able to bypass only a single

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Diagram showing the retro-regulation process that yields a higher concentration of xis compared to int. The mRNA transcript is digested by bacterial RNase starting from the cleaved hairpin loop at sib.

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onto the J/H scaffold. Then, protein U is added to the head-proximal end of the tail. Protein Z is able to connect the tail to the head. Protein H is cleaved due to the actions of proteins U and Z.

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This is the lifecycle that the phage follows following most infections, where the cII protein does not reach a high enough concentration due to degradation, so does not activate its promoters.

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This occurs without the N protein interacting with the DNA; the protein instead binds to the freshly transcribed mRNA. Nut sites contain 3 conserved "boxes", of which only BoxB is essential.

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contains 48,502 base pairs of double-stranded, linear DNA, with 12-base single-strand segments at both 5' ends. These two single-stranded segments are the "sticky ends" of what is called the

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Forms a membrane protein complex that destroys the outer cell membrane following the cell wall degradation by endolysin. Spanin, Rz1(outer membrane subunit) and Rz(inner membrane subunit).

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Bacteriophage lambda virion (schematic). Protein names and their copy numbers in the virion particle are shown. The presence of the L and M proteins in the virion is still unclear.

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gene resulting in a lack of growth of lambda phage. This is due to RNA polymerase attaching to pL promoter site instead of the pR promotor site. Leftward transcription results in

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The boxB RNA sequences are located close to the 5' end of the pL and pR transcripts. When transcribed, each sequence forms a hairpin loop structure that the N protein can bind to.

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sites and the resulting complex can ignore terminators, however the mechanism is very different; the Q protein first associates with a DNA sequence rather than an mRNA sequence.

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site circularizes the DNA in the host cytoplasm. In its circular form, the phage genome, therefore, is 48,502 base pairs in length. The lambda genome can be inserted into the

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Burz DS, Beckett D, Benson N, Ackers GK (July 1994). "Self-assembly of bacteriophage lambda cI repressor: effects of single-site mutations on the monomer-dimer equilibrium".

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promoters are left active, the former producing cI protein and the latter a short inactive transcript. The genome remains inserted into the host genome in a dormant state.

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Another example is the P22 phage may replace the p gene, which encodes for an essential replication protein for something that is capable of encoding for a DnaB helices.

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proteins. In lysogenic cells, this response is hijacked, and RecA* stimulates cI autocleavage. This is because cI mimics the structure of LexA at the autocleavage site.

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operator is very similar, except that it has no direct effect on cI transcription. As an additional repression of its own expression, however, cI dimers bound to

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of the phage and host DNA. The original B-O-B' sequence is changed by the integration to B-O-P'-phage DNA-P-O-B'. The phage DNA is now part of the host's genome.

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of the virion. All characterized lambdoid phages possess an N protein-mediated transcription antitermination mechanism, with the exception of phage HK022.

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genes. O and P are responsible for initiating replication, and Q is another antiterminator that allows the expression of head, tail, and lysis genes from

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regulation ensures a stable minimum concentration of the repressor molecule and, should SOS signals arise, allows for more efficient prophage induction.

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site, and P protein binds the DnaB subunit of the host replication machinery as well as binding O. This effectively commandeers the host DNA polymerase.

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gene, resulting in high concentrations of Int protein. This int protein integrates the phage DNA into the host chromosome (see "Prophage Integration").

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The lysogenic lifecycle begins once the cI protein reaches a high enough concentration to activate its promoters, after a small number of infections.

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Lambda phage DNA injection into the cell membrane using Mannose PTS permease (a sugar transporting system) as a mechanism of entry into the cytoplasm

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The classic induction of a lysogen involved irradiating the infected cells with UV light. Any situation where a lysogen undergoes DNA damage or the

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Equal concentrations of xis and int result in the excision of the inserted genome from the host genome for replication and later phage production.

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The DNA passes through the mannose permease complex in the inner membrane (encoded by the manXYZ genes) and immediately circularises using the

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gene is absent. The lack of growth of lambda phage is believed to occur due to a temperature sensitivity resulting in inhibition of growth.

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Blanco M, Devoret R (March 1973). "Repair mechanisms involved in prophage reactivation and UV reactivation of UV-irradiated phage lambda".

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The host cell, containing a dormant phage genome, experiences DNA damage due to a high stress environment, and starts to undergo the

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are broken down when the host cell is under stress, resulting in the expression of the repressed phage genes. Stress can be from

4962: 4034: 3141: 2012:. It binds DNA in the helix-turn-helix binding motif. It regulates the transcription of the cI protein and the Cro protein. 480:" ensues, where the lambda DNA is replicated and new phage particles are produced within the cell. This is followed by cell 476:, causing DNA in the head of the phage to be ejected through the tail into the cytoplasm of the bacterial cell. Usually, a " 4158:

Pitre, Emmanuelle; te Velthuis, Aartjan J. W. (2021-01-01), Cameron, Craig E.; Arnold, Jamie J.; Kaguni, Laurie S. (eds.),

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Visual representation of repressor tetramer/octamer binding to phage lambda L and R operator sites (stable lysogenic state)

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promoter mRNA results in no hairpin loop on the 3' end, and the transcript is no longer targeted for RNAaseIII degradation.

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have high (and approximately equal) concentrations this leads to the excision of phage genomes from the host's genome.

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2 a loop is induced in the DNA, allowing these dimers to bind together to form an octamer. This is a phenomenon called

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operons, and consists of the parts B-O-B', whereas the complementary sequence in the circular phage genome is called

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Gene N acts a regulatory gene that results in RNA polymerase to be unable to recognize translation-termination sites.

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RecA (a cellular protein) detects DNA damage and becomes activated. It is now RecA*, a highly specific co-protease.

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and hence cI stimulates its own transcription. When it is present at a much higher concentration, it also binds to

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when a prophage is present. This prophage may enter the lytic cycle when the lysogen enters a stressed condition.

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2 are almost always simultaneously occupied by cI. However, this does not increase the affinity between cI and O

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The function of LexA in the SOS response. LexA expression leads to inhibition of various genes including LexA.

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The integration of phage λ takes place at a special attachment site in the bacterial and phage genomes, called

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Transcription of the phage chromosome proceeds leftward when the host RNA polymerase attaches to promotor site

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promoter, close enough that the σ factor has not been released from the RNA polymerase holoenzyme. Part of the

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proteins are produced. This results (initially) in the excision of any inserted genomes from the host genome.

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The phage genome is still inserted in the host genome and needs excision for DNA replication to occur. The

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repressor), activating LexA auto-protease activity, which destroys LexA repressor, allowing production of

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Lysogen repressors and polymerase bound to OR1 and recruits OR2, which will activate PRM and shutdown PR.

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is degraded the phage will go into the lytic pathway. Low temperature, starvation of the cells and high

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cell by means of its J protein in the tail tip. The J protein interacts with the maltose outer membrane

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promoter (preventing cro production). At high concentrations downregulates its own production through O

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sites as they are packaged. Packaging cannot occur from circular phage DNA, only from concatomeric DNA.

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Friedman DI, Court DL (April 2001). "Bacteriophage lambda: alive and well and still doing its thing".

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The presence of cI causes immunity to superinfection by other lambda phages, as it will inhibit their

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The cIII protein acts to protect the cII protein from proteolysis by FtsH (a membrane-bound essential

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Some base pairs with serve a dual function with promoter and operator for either cl and cro proteins.

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sites, 12-base G-C-rich cohesive "sticky ends". The single-strand viral DNA ends are ligated by host

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Protein cl turned ON, with repressor bound to OR2 polymerase binding is increased and turn OFF OR1.

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Linear layout of lambda phage genome with major operons, promoter regions and capsid coding genes.

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Dodd IB, Shearwin KE, Egan JB (April 2005). "Revisited gene regulation in bacteriophage lambda".

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cells, that exist prior to infection, predetermine whether a cell will lyse or become a lysogen.

1271:) end of the mRNA for RNAaseIII degradation, which results in a lower effective concentration of 793:

determines the lifestyle of the phage; stable cII will lead to the lysogenic pathway, whereas if

142: 4706:"Evolutionary relationships among diverse bacteriophages and prophages: all the world's a phage" 5598: 5507: 5490: 5183: 5068: 5019: 4878:(illustrates infection and lytic/lysogenic pathways with some protein and transcription detail) 3848:

Michod RE, Bernstein H, Nedelcu AM (May 2008). "Adaptive value of sex in microbial pathogens".

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Brammar WJ, Hadfield C (November 1984). "A programme for the construction of a lambda phage".

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A simplified representation of the integration/excision paradigm and the major genes involved.

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Thomason LC, Schiltz CJ, Court C, Hosford CJ, Adams MC, Chappie JS, Court DL (October 2021).

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The O and P proteins initiate replication of the phage chromosome (see "Lytic Replication").

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Huskey RJ (April 1969). "Multiplicity reactivation as a test for recombination function".

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protein regulator, manages excision and insertion of phage genome into the host's genome.

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promoter transcript is, however, no longer included in this reading frame (see diagram).

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and consists of the parts P-O-P'. The integration itself is a sequential exchange (see

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Q protein then binds and displaces part of the σ factor and transcription re-initiates.

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in the circular chromosome, causing A-T-rich regions to unwind and drive transcription.

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2 (which are over 2 kb downstream from the R operators). When cI dimers are bound to O

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site on the mRNA) continues transcription, and can skip through termination sequences.

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The lambda repressor gene system consists of (from left to right on the chromosome):

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The phage chromosome is inserted to the host bacterial chromosome through DNA ligase.

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The head and tail genes are transcribed and the corresponding proteins self-assemble.

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promoter can now extend to produce mRNA for the lysis and the head and tail proteins.

55: 4909: 3920: 1602: 5201: 5049: 4818: 4794: 4774: 4735: 4725: 4682: 4678: 4674: 4641: 4612: 4581: 4546: 4536: 4493: 4483: 4389: 4379: 4358:"Lambda phage-based vaccine induces antitumor immunity in hepatocellular carcinoma" 4313: 4239: 4223: 4171: 4130: 4120: 4071: 4063: 3999: 3943: 3900: 3857: 3822: 3785: 3777: 3709: 3668: 3660: 3619: 3611: 3542: 3532: 3483: 3475: 3434: 3393: 3385: 3297: 3289: 3243: 3233: 3177: 3102: 3094: 3041: 2988: 1956: 1946:

Protein interactions that lead to either Lytic or Lysogenic cycles for Lambda phage

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DNA repair protein, functions as a co-protease during SOS response, auto-cleaving

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Groth AC, Calos MP (January 2004). "Phage integrases: biology and applications".

3781: 3537: 3464:"Structure of the mannose transporter of the bacterial phosphotransferase system" 2956: 2779:, manages insertion of phage genome into the host's genome. In Conditions of low 1325: 1221:

II transcription on the left operon. Bar positive phenotype is present when the

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genome. During infections, the phage particle recognizes and binds to its host,

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Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

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Diagram of temperate phage life cycle, showing both lytic and lysogenic cycles.

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Henkin, Tina M.; Peters, Joseph E. (2020). "Bacteriophages and Transduction".

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production). At high concentrations downregulates its own production through O

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promoter transcript, turning down cro production, and has a transcript of the

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are found on the same piece of mRNA, so approximately equal concentrations of

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Lambda phage inserts chromosome into the cytoplasm of the host bacterial cell.

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Replication protein O. Initiates Phage Lambda DNA replication by binding at

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Structural proteins and phage genomes self-assemble into new phage particles.

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state, which favours lysogeny. cIII also directly stabilises the cII protein.

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protease) by acting as a competitive inhibitor. This inhibition can induce a

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Leftward transcription is believed to result in a deletion mutation of the

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Cleaved cI can no longer dimerise, and loses its affinity for DNA binding.

612: 5539: 5347: 5193: 5060: 4976: 4269:"Bacteriophage Lambda Terminase and the Mechanism of Viral DNA Packaging" 3342:

Campbell AM (1996). "Bacteriophages". In Neidhardt FC, Curtiss R (eds.).

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producing the 'immediate early' transcripts. At first, these express the

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chromosome and is then called a prophage. See section below for details.

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Ptashne, M. "A Genetic Switch: Phage Lambda Revisited", 3rd edition 2003

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will push the phage toward integration and lysogeny while low levels of

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At high concentration of cI, transcriptions of both genes are repressed.

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promoter regions during a lysogenic state vs induced, early lytic state.

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and requires both the phage protein Int and the bacterial protein IHF (

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subunit. These bindings provide control over the host DNA polymerase.

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At high concentrations of cI, the dimers will also bind to operators O

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Lysogeny is maintained solely by cI. cI represses transcription from

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provides insufficient context for those unfamiliar with the subject

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Hendrix RW, Smith MC, Burns RN, Ford ME, Hatfull GF (March 1999).

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3, which will be occupied only when the cI concentration is high.

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The lambda repressor is a self assembling dimer also known as the

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binding. This activates transcription in the other direction from

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For the first few replication cycles, the lambda genome undergoes

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gene. This is expressed, turning on cI repressor production. The

591: 497: 2480:, a membrane protein that perforates the membrane during lysis. 1875: 1632:. It is therefore the only protein expressed by lysogenic phage. 1491:

As cII and cIII concentrations decrease, transcription from the

869: 5445: 5440: 5390: 4890:(basic visual demonstration of Lambda bacteriophage life cycle) 3130:

Griffiths A, Miller J, Suzuki D, Lewontin R, Gelbart W (2000).

2964: 2452: 1539: 1334:

The 'late early' transcripts continue being written, including

887:

The 'late early' transcripts continue being written, including

597:

The linear phage genome is injected through the outer membrane.

529: 493: 465: 449: 3689: 2791:

high then this leads to the insertion of the phage genome. If

2387:

1 binding. No cooperative binding (c.f. below for cI binding)

2375:

3). At low concentrations blocks the pRM promoter (preventing

5248: 4999: 3129: 2477: 1391:

promoter transcript, thereby switching off Q production. The

961: 481: 457: 290: 4757:

Kitano H (March 2002). "Systems biology: a brief overview".

4210:

Reece-Hoyes, John S.; Walhout, Albertha J. M. (2018-01-01).

3166:"Structural basis of bacteriophage lambda capsid maturation" 3038:"Bacteriophage Lambda Structural Model at Atomic Resolution" 668: 257: 4906:(3D display of protein structures for bacteriophage Lambda) 3974: 3646: 2972: 2886: 2325: 2319:

3 binding. Repressor also inhibits transcription from the P

1659:

operators. Both operators have three binding sites for cI:

1625:

while upregulating and controlling its own expression from

1398:

promoter produces antisense mRNA to the cro section of the

4517:"Lambda repressor turns off transcription of its own gene" 1349:

The stabilized cII acts to promote transcription from the

801:(MOI) are known to favor lysogeny (see later discussion). 729:

at specific sites of the nascently transcribed mRNA. When

403:

Bacteriophage Lambda Structural Model at Atomic Resolution

5310: 5131: 4166:, Viral Replication Enzymes and their Inhibitors Part A, 3372:

Werts C, Michel V, Hofnung M, Charbit A (February 1994).

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and form an intasome, a DNA-protein-complex designed for

1488:

promoters leads to no further production of cII and cIII.

496:

without apparent harm to the host. The host is termed a

469: 4884:

from MIT showing both lysis and lysogeny by phage lambda

4807:"Regulation of transcription: from lambda to eukaryotes" 4464:"The operators controlled by the lambda phage repressor" 4354: 3282:"Bacteriophage lambda: Early pioneer and still relevant" 1820:

of the host is otherwise stimulated leads to induction.

1319: 4446:

Watson J, Baker T, Bell S, Gann A, Levine M, Losick R.

3989: 1715:

greatly increases the affinity of the binding of cI to

914:

promoter (which is a promoter of the lysogenic cycle).

464:

The phage particle consists of a head (also known as a

3847: 3597: 3510: 1450:

and other early gene expression. cI also binds to the

565:

Lambda phage J protein interaction with the LamB porin

448:

life cycle that allows it to either reside within the

4939: 4863:(illustrates genes active at all stages in lifecycle) 4703: 4448:

Molecular Biology of the Gene (International Edition)

3222:"The protein interaction map of bacteriophage lambda" 3219: 3083:"Genetic Studies of Lysogenicity in Escherichia Coli" 2859:

Antitermination for Transcription of Late Lytic Genes

2835:

Antitermination for Transcription of Late Early Genes

2425:

Initiates Phage Lambda DNA replication by binding to

664:

genes, producing N, Cro and a short inactive protein.

1512:

stop being promoted since they are no longer needed.

3572:

Snyder and Champness molecular genetics of bacteria

3220:Rajagopala SV, Casjens S, Uetz P (September 2011). 1925:

Multiplicity reactivation and prophage reactivation

1558:. The sequence of the bacterial att site is called 1259:promoter forms a stable secondary structure with a 1204: 265:Electron micrograph of a virus particle of species 156:. Unsourced material may be challenged and removed. 4209: 3420: 3080: 2250:. Low stability due to susceptibility to cellular 2019:The cI dimer may bind to any of three operators, O 4660: 4514: 4418:"7.14E: Bacteriophage Lambda as a Cloning Vector" 4267:Feiss, Michael; Catalano, Carlos Enrique (2013), 3739:Journal of Embryology and Experimental Morphology 3695: 2196:Regulatory protein CIII. Lysogeny, cII Stability 1176: 5613: 4602: 4157: 4049: 3511:Kobiler O, Rokney A, Oppenheim AB (April 2007). 3461: 1909:The new intact transcript has one copy of both 1229:xis and int regulation of insertion and excision 899:and genes for replication of the lambda genome ( 4839: 4515:Meyer BJ, Kleid DG, Ptashne M (December 1975). 4445: 3763: 3736: 3593: 3591: 3337: 3335: 3035: 1711:. As cI easily dimerises, the binding of cI to 1346:and genes for replication of the lambda genome. 4567: 4454: 3462:Liu X, Zeng J, Huang K, Wang J (August 2019). 3279: 1457:promoter, turning off transcription there too. 488:pathway. In this state, the λ DNA is called a 5076: 4925: 4631: 4605:Current Opinion in Genetics & Development 4266: 4098: 3933: 3215: 3213: 3211: 3209: 3065:Lederberg E (January 1950). "Lysogenicity in 1876:Control of phage genome excision in induction 1471:promoter may occur, maintaining levels of cI. 725:, and functions by engaging the transcribing 4899:Lambda Phage Reference Proteome from UniProt 3588: 3569: 3421:Erni B, Zanolari B, Kocher HP (April 1987). 3332: 3163: 2230:(Clear 2) Activates transcription from the P 2171: 2047:1 enhances binding of a second cI dimer to O 1719:, and this happens almost immediately after 1103:Q is similar to N in its effect: Q binds to 4661:Gottesman ME, Weisberg RA (December 2004). 4050:Arkin A, Ross J, McAdams HH (August 1998). 3983: 3730: 3326:"Escherichia phage Lambda, complete genome" 1704:; binding here inhibits transcription from 672:Early activation events involving N protein 632:Transcription starts from the constitutive 444:in 1950. The wild type of this virus has a 64:Learn how and when to remove these messages 27:Bacteriophage that infects Escherichia coli 5090: 5083: 5069: 4932: 4918: 4667:Microbiology and Molecular Biology Reviews 3812: 3598:Santangelo TJ, Artsimovitch I (May 2011). 3206: 3081:Lederberg EM, Lederberg J (January 1953). 2283:(Clear 1) Transcription inhibitor, binds O 1734:tightens the binding of RNA polymerase to 1597: 1267:section of the mRNA. This targets the 3' ( 972: 256: 4822: 4739: 4729: 4686: 4585: 4550: 4540: 4497: 4487: 4393: 4383: 4243: 4134: 4124: 4075: 3789: 3713: 3696:Deighan P, Hochschild A (February 2007). 3672: 3623: 3546: 3536: 3487: 3438: 3397: 3301: 3247: 3237: 3181: 3136:(7th ed.). New York: W. H. Freeman. 3106: 3064: 2121:In the presence of cI proteins, only the 1774:bend the DNA between them to tetramerise. 1295:sequence), so a higher concentrations of 1144: 903:). Cro dominates the repressor site (see 590:, a porin molecule, which is part of the 234:Learn how and when to remove this message 216:Learn how and when to remove this message 114:Learn how and when to remove this message 4904:Lambda Phage Protein Structures in NCBI 4027:A genetic switch: phage lambda revisited 3341: 2947:and has been an excellent tool first in 2943:Lambda phage has been used heavily as a 2917: 2106:3, repressing transcription of cI. This 1961: 1941: 1866: 1799: 1635: 1601: 1464:site unbound, so transcription from the 1380:promoter produces antisense mRNA to the 1148: 868: 850:The entire complex (including the bound 667: 611: 560: 519: 507: 398: 4804: 4024: 2187: 2181: 2178: 1549: 1192:L resulting in the translation of gene 1047:These concatemers are cleaved at their 789:On initial infection, the stability of 688:promoter, preventing expression of the 14: 5614: 4756: 4568:Brüssow H, Hendrix RW (January 2002). 3882: 3275: 3273: 3271: 3269: 3267: 2913: 2456:nuclease from degrading 3' ends—allow 2338:Lysis, Control of Repressor's Operator 2330:in cells undergoing the SOS response. 2311:1). At low concentrations blocks the P 2184: 1752:, thus regulating its own levels in a 977:Rightward transcription expresses the 5506: 5505: 5064: 4913: 4291: 4099:St-Pierre F, Endy D (December 2008). 3164:Wang C, Zeng J, Wang J (April 2022). 2783:concentration there is no effect. If 2323:promoter. Susceptible to cleavage by 1320:Lysogenic (or lysenogenic) life cycle 1157:Leftward transcription expresses the 492:and stays resident within the host's 430:, that infects the bacterial species 96:providing more context for the reader 4876:Basic Animation of Lambda Lifecyecle 4570:"Phage genomics: small is beautiful" 3764:Guzmán P, Guarneros G (March 1989). 2973:aspartyl (asparaginyl) β-hydroxylase 2938: 1730:, as the N terminal domain of cI on 1423:No Q results in no extension of the 1055: 1007: 804: 154:adding citations to reliable sources 125: 70: 29: 3427:The Journal of Biological Chemistry 3280:Casjens SR, Hendrix RW (May 2015). 3264: 3133:An Introduction to Genetic Analysis 2114:In the absence of cI proteins, the 858: 24: 4438: 2270:Repressor, Maintenance of Lysogeny 1134:, causing the holoenzyme to pause. 696:(N utilisation) sites, one in the 25: 5643: 5207:Micro 4868: 4273:Madame Curie Bioscience Database 4212:"Gateway Recombinational Cloning" 3850:Infection, Genetics and Evolution 2963:; and the application of its Red 2221:Lysogeny, Transcription activator 2209:(FtsH) binding protein, protects 907:), repressing synthesis from the 873:Lysis plaques of lambda phage on 574:Bacteriophage Lambda binds to an 45:This article has multiple issues. 5486: 5485: 4851:"Bacteriophage Lambda Infection" 3715:10.1111/j.1365-2958.2006.05563.x 3351:: Cellular and Molecular Biology 3036:Padilla-Sanchez V (2021-07-16). 2881:antisense mRNA during lysogeny. 2347:Transcription inhibitor, binds O 2158: 2146: 2134: 1745:, inhibiting transcription from 1205:Leftward Transcription mutations 1086: 1075: 1064: 822: 813: 277: 130: 75: 34: 5162:precursor, heterogenous nuclear 4663:"Little lambda, who made thee?" 4634:Current Opinion in Microbiology 4410: 4348: 4285: 4260: 4203: 4151: 4092: 4043: 4018: 3962: 3927: 3876: 3841: 3806: 3757: 3640: 3563: 3504: 3455: 3414: 2213:from degradation by proteases. 1474:Lack of transcription from the 141:needs additional citations for 53:or discuss these issues on the 5292:Trans-acting small interfering 5256:Enhancer RNAs 5174:Transfer 4894:Lambda Phage genome in GenBank 4842:Molecular Genetics of Bacteria 4840:Snyder L, Champness W (2007). 4811:Trends in Biochemical Sciences 4679:10.1128/MMBR.68.4.796-813.2004 3365: 3318: 3157: 3123: 3058: 3029: 2371:1, i.e. preferentially binds O 2307:3, i.e. preferentially binds O 1804:Transcriptional state of the P 1177:Leftward transcription process 1035:Soon, the phage switches to a 904: 707:reading frame, and one in the 13: 1: 5179:Ribosomal 5157:Messenger 4646:10.1016/S1369-5274(00)00189-2 4587:10.1016/S0092-8674(01)00637-7 4385:10.1016/j.heliyon.2017.e00407 3440:10.1016/S0021-9258(18)61180-9 3390:10.1128/jb.176.4.941-947.1994 3353:. Washington, DC: ASM Press. 3023: 2755:Tail fiber assembly protein. 2687:Tail tape measure protein H. 2043:3. Binding of a cI dimer to O 1834:Normally RecA* binds LexA (a 551: 4216:Cold Spring Harbor Protocols 3948:10.1016/0027-5107(73)90001-8 3905:10.1126/science.164.3877.319 3862:10.1016/j.meegid.2008.01.002 3815:Journal of Molecular Biology 3604:Nature Reviews. Microbiology 3538:10.1371/journal.pone.0000363 2999:Molecular weight size marker 2849:cII, cIII, xis, int, O, P, Q 2787:is low in concentration and 2763:Genome Integration, Excision 2636:Host specificity protein J. 2534:Phage capsid head proteins. 1937: 1795: 1586:). Both Int and IHF bind to 556: 7: 5516:Enterobacteria phage lambda 4882:Time-lapse microscopy video 4068:10.1093/genetics/149.4.1633 3294:10.1016/j.virol.2015.02.010 3071:Microbial Genetics Bulletin 2982: 1700:cI binds most favorably to 1645:This is coordinated by the 1592:site-specific recombination 681:, preventing access to the 426:) is a bacterial virus, or 10: 5648: 5358:Multicopy single-stranded 5202:Interferential 4824:10.1016/j.tibs.2005.04.003 4176:10.1016/bs.enz.2021.07.005 4170:, Academic Press: 83–113, 3782:10.1093/genetics/121.3.401 2458:rolling circle replication 1885:section beyond the normal 1323: 1119:site is very close to the 1037:rolling circle replication 1028:gene. O protein binds the 862: 503: 5514: 5481: 5416: 5366: 5309: 5272:Guide 5264: 5192: 5147: 5130: 5099: 4952: 4617:10.1016/j.gde.2005.02.001 4292:Smith, George P. (1985). 3827:10.1016/j.jmb.2003.09.082 3480:10.1038/s41422-019-0194-z 3183:10.1016/j.str.2021.12.009 2738:Tail attachment protein. 2704:Tail assembly protein I. 2619:Large terminase protein. 2551:Head decoration protein. 2172:Protein function overview 1306:Higher concentrations of 1283:cistron is nearer to the 1020:This is initiated at the 960:cause cell lysis. S is a 799:multiplicity of infection 692:gene. N binds to the two 440:). It was discovered by 272: 264: 255: 248: 5546:Escherichia virus Lambda 5234:Small nuclear 3239:10.1186/1471-2180-11-213 3099:10.1093/genetics/38.1.51 2977:specialized transduction 2907:and facilitating lysis. 2721:DNA-packing protein FI. 2670:Probable endopeptidase. 2242:promoters, transcribing 2125:gene may be transcribed. 2118:gene may be transcribed. 2096:long-range cooperativity 423:Escherichia virus Lambda 390:Escherichia virus Lambda 267:Escherichia virus Lambda 250:Escherichia virus Lambda 5348:Genomic 4888:Lambda Phage Life cycle 4805:Ptashne M (June 2005). 4779:10.1126/science.1069492 4542:10.1073/pnas.72.12.4785 4318:10.1126/science.4001944 4126:10.1073/pnas.0808831105 3378:Journal of Bacteriology 2182:Function in life cycle 1598:Maintenance of lysogeny 1584:integration host factor 1460:Lack of cro leaves the 1446:promoter, turning off 1416:promoter expresses the 1130:site resembles the -10 973:Rightward transcription 931:Transcription from the 5451:Artificial chromosomes 5239:Small nucleolar 4731:10.1073/pnas.96.5.2192 4489:10.1073/pnas.60.4.1282 3702:Molecular Microbiology 3653:Molecular Microbiology 3349:Salmonella typhimurium 3046:10.5281/zenodo.5134493 2923: 2653:Minor tail protein M. 2585:Minor capsid protein. 2441:Lysis, DNA replication 2416:Lysis, DNA Replication 2395:Lysis, DNA replication 2262:will result in lysis. 1967: 1947: 1872: 1813: 1641: 1607: 1154: 1145:Leftward transcription 944:Products of the genes 880: 770:genes discussed later. 673: 617: 566: 525: 513: 409:Enterobacteria phage λ 404: 5244:Small Cajal Body RNAs 4228:10.1101/pdb.top094912 2921: 2875:). Downregulated by P 1965: 1945: 1870: 1803: 1639: 1605: 1576:genetic recombination 1152: 872: 671: 615: 564: 523: 511: 402: 5297:Subgenomic messenger 5212:Small interfering 5184:Transfer-messenger 4222:(1): pdb.top094912. 3288:. 479–480: 310–330. 3014:Corynebacteriophages 2951:, and then later in 2568:Major head protein. 2051:2, an effect called 1550:Prophage integration 1384:gene message of the 1024:site located in the 721:The N protein is an 582:(the product of the 452:of its host through 274:Virus classification 150:improve this article 18:Bacteriophage lambda 5622:Genetics techniques 5013:Medicago truncatula 4771:2002Sci...295.1662K 4765:(5560): 1662–1664. 4722:1999PNAS...96.2192H 4533:1975PNAS...72.4785M 4480:1968PNAS...60.1282P 4376:2017Heliy...300407I 4310:1985Sci...228.1315S 4304:(4705): 1315–1317. 4275:, Landes Bioscience 4117:2008PNAS..10520705S 4111:(52): 20705–20710. 4004:10.1021/bi00194a003 3897:1969Sci...164..319H 3616:10.1038/nrmicro2560 3529:2007PLoSO...2..363K 2994:Lambda holin family 2914:Lytic vs. lysogenic 1017:(circle-to-circle). 905:"Repressor" section 92:improve the article 5326:Chloroplast 5169:modified Messenger 5132:Ribonucleic acids 4422:Biology LibreTexts 4025:Ptashne M (2004). 3970:"Barbara J. Meyer" 2953:molecular genetics 2949:microbial genetics 2924: 2602:Portal protein B. 1968: 1948: 1873: 1814: 1759:cI binding to the 1642: 1608: 1291:cistron is to the 1287:sequence than the 1252:The mRNA from the 1155: 881: 847:ribosome subunit). 674: 618: 567: 526: 514: 405: 5609: 5608: 5508:Taxon identifiers 5499: 5498: 5376:Xeno 5338:Complementary 5311:Deoxyribonucleic 5305: 5304: 5282:Small hairpin 5058: 5057: 4861:on 19 March 2014. 4036:978-0-87969-716-7 3998:(28): 8399–8405. 3936:Mutation Research 3891:(3877): 319–320. 3665:10.1111/mmi.14792 3433:(11): 5238–5247. 3328:. 6 January 2020. 3176:(4): 637–645.e3. 3143:978-0-7167-3520-5 3009:Zygotic induction 2939:As a genetic tool 2911: 2910: 2031:3, in the order O 1754:negative feedback 1580:Holliday junction 1056:Q antitermination 1008:Lytic replication 805:N antitermination 742:genes along with 397: 396: 244: 243: 236: 226: 225: 218: 200: 124: 123: 116: 68: 16:(Redirected from 5639: 5602: 5601: 5589: 5588: 5576: 5575: 5563: 5562: 5550: 5549: 5548: 5535: 5534: 5533: 5503: 5502: 5489: 5488: 5466:Yeast 5287:Small temporal 5217:Piwi-interacting 5145: 5144: 5141: 5122:Deoxynucleotides 5085: 5078: 5071: 5062: 5061: 4934: 4927: 4920: 4911: 4910: 4862: 4857:. Archived from 4845: 4836: 4826: 4798: 4753: 4743: 4733: 4716:(5): 2192–2197. 4700: 4690: 4657: 4628: 4599: 4589: 4564: 4554: 4544: 4511: 4501: 4491: 4451: 4433: 4432: 4430: 4429: 4414: 4408: 4407: 4397: 4387: 4352: 4346: 4345: 4289: 4283: 4282: 4281: 4280: 4264: 4258: 4257: 4247: 4207: 4201: 4200: 4199: 4198: 4155: 4149: 4148: 4138: 4128: 4096: 4090: 4089: 4079: 4062:(4): 1633–1648. 4047: 4041: 4040: 4022: 4016: 4015: 3987: 3981: 3980: 3966: 3960: 3959: 3931: 3925: 3924: 3880: 3874: 3873: 3845: 3839: 3838: 3810: 3804: 3803: 3793: 3761: 3755: 3754: 3745:(Suppl): 75–88. 3734: 3728: 3727: 3717: 3693: 3687: 3686: 3676: 3659:(4): 1044–1063. 3644: 3638: 3637: 3627: 3595: 3586: 3585: 3567: 3561: 3560: 3550: 3540: 3508: 3502: 3501: 3491: 3459: 3453: 3452: 3442: 3418: 3412: 3411: 3401: 3369: 3363: 3362: 3345:Escherichia coli 3339: 3330: 3329: 3322: 3316: 3315: 3305: 3277: 3262: 3261: 3251: 3241: 3226:BMC Microbiology 3217: 3204: 3203: 3185: 3161: 3155: 3154: 3152: 3150: 3127: 3121: 3120: 3110: 3078: 3067:Escherichia coli 3062: 3056: 3055: 3053: 3052: 3033: 2989:Esther Lederberg 2873:S, R, Rz and Rz1 2185:Promoter region 2176: 2175: 2162: 2150: 2138: 1957:Barbara J. Meyer 1090: 1079: 1068: 859:Lytic life cycle 826: 817: 456:or enter into a 442:Esther Lederberg 433:Escherichia coli 282: 281: 260: 246: 245: 239: 232: 221: 214: 210: 207: 201: 199: 158: 134: 126: 119: 112: 108: 105: 99: 79: 78: 71: 60: 38: 37: 30: 21: 5647: 5646: 5642: 5641: 5640: 5638: 5637: 5636: 5627:Model organisms 5612: 5611: 5610: 5605: 5597: 5592: 5584: 5579: 5571: 5566: 5558: 5553: 5544: 5543: 5538: 5529: 5528: 5523: 5510: 5500: 5495: 5477: 5418:Cloning vectors 5412: 5398:Locked 5362: 5312: 5301: 5260: 5188: 5135: 5134: 5126: 5095: 5089: 5059: 5054: 4948: 4942:model organisms 4938: 4871: 4866: 4849: 4844:(3rd ed.). 4527:(12): 4785–89. 4462:(August 1968). 4450:(6th ed.). 4441: 4439:Further reading 4436: 4427: 4425: 4416: 4415: 4411: 4353: 4349: 4290: 4286: 4278: 4276: 4265: 4261: 4208: 4204: 4196: 4194: 4156: 4152: 4097: 4093: 4048: 4044: 4037: 4023: 4019: 3988: 3984: 3968: 3967: 3963: 3932: 3928: 3881: 3877: 3846: 3842: 3811: 3807: 3762: 3758: 3735: 3731: 3694: 3690: 3645: 3641: 3596: 3589: 3582: 3568: 3564: 3509: 3505: 3460: 3456: 3419: 3415: 3370: 3366: 3340: 3333: 3324: 3323: 3319: 3278: 3265: 3218: 3207: 3162: 3158: 3148: 3146: 3144: 3128: 3124: 3063: 3059: 3050: 3048: 3034: 3030: 3026: 2985: 2957:recombinant DNA 2941: 2916: 2879: 2865: 2841: 2817: 2813: 2807:Genome Excision 2773: 2769: 2752: 2735: 2718: 2701: 2684: 2667: 2650: 2633: 2616: 2599: 2582: 2565: 2548: 2531: 2514: 2494: 2474: 2447: 2422: 2401: 2386: 2382: 2374: 2370: 2366: 2362: 2358: 2354: 2350: 2344: 2322: 2318: 2314: 2310: 2306: 2302: 2298: 2294: 2290: 2286: 2280: 2276: 2241: 2237: 2233: 2227: 2202: 2174: 2167: 2163: 2154: 2151: 2142: 2139: 2105: 2101: 2093: 2089: 2085: 2081: 2077: 2073: 2066: 2062: 2058: 2050: 2046: 2042: 2038: 2034: 2030: 2026: 2022: 1997: 1990: 1983: 1940: 1927: 1904: 1890: 1878: 1860: 1853: 1811: 1807: 1798: 1789: 1782: 1764: 1750: 1739: 1728: 1709: 1695: 1676: 1657: 1650: 1630: 1623: 1616: 1600: 1552: 1527: 1520: 1510: 1503: 1496: 1486: 1479: 1469: 1455: 1444: 1428: 1414: 1403: 1396: 1389: 1378: 1368: 1361: 1354: 1328: 1326:Lysogenic cycle 1322: 1257: 1231: 1207: 1179: 1147: 1124: 1101: 1100: 1099: 1098: 1093: 1092: 1091: 1082: 1081: 1080: 1071: 1070: 1069: 1058: 1010: 994: 975: 936: 912: 867: 861: 837: 836: 835: 834: 829: 828: 827: 819: 818: 807: 716: 705: 686: 651: 644: 637: 559: 554: 506: 393: 276: 240: 229: 228: 227: 222: 211: 205: 202: 159: 157: 147: 135: 120: 109: 103: 100: 89: 80: 76: 39: 35: 28: 23: 22: 15: 12: 11: 5: 5645: 5635: 5634: 5629: 5624: 5607: 5606: 5604: 5603: 5590: 5577: 5564: 5551: 5536: 5520: 5518: 5512: 5511: 5497: 5496: 5494: 5493: 5482: 5479: 5478: 5476: 5475: 5474: 5473: 5468: 5463: 5458: 5448: 5443: 5438: 5433: 5428: 5422: 5420: 5414: 5413: 5411: 5410: 5405: 5403:Peptide 5400: 5395: 5394: 5393: 5388: 5383: 5381:Glycol 5372: 5370: 5364: 5363: 5361: 5360: 5355: 5350: 5345: 5340: 5335: 5334: 5333: 5328: 5317: 5315: 5307: 5306: 5303: 5302: 5300: 5299: 5294: 5289: 5284: 5279: 5274: 5268: 5266: 5262: 5261: 5259: 5258: 5253: 5252: 5251: 5246: 5241: 5236: 5226: 5221: 5220: 5219: 5214: 5209: 5198: 5196: 5190: 5189: 5187: 5186: 5181: 5176: 5171: 5166: 5165: 5164: 5153: 5151: 5142: 5128: 5127: 5125: 5124: 5119: 5114: 5109: 5103: 5101: 5097: 5096: 5093:nucleic acids 5088: 5087: 5080: 5073: 5065: 5056: 5055: 5053: 5052: 5047: 5042: 5037: 5030: 5023: 5016: 5009: 5002: 4997: 4990: 4985: 4980: 4973: 4966: 4959: 4953: 4950: 4949: 4937: 4936: 4929: 4922: 4914: 4908: 4907: 4901: 4896: 4891: 4885: 4879: 4870: 4869:External links 4867: 4865: 4864: 4847: 4837: 4817:(6): 275–279. 4802: 4799: 4754: 4701: 4673:(4): 796–813. 4658: 4640:(2): 201–207. 4629: 4611:(2): 145–152. 4600: 4565: 4512: 4452: 4442: 4440: 4437: 4435: 4434: 4409: 4347: 4284: 4259: 4202: 4150: 4091: 4042: 4035: 4017: 3982: 3961: 3942:(3): 293–305. 3926: 3875: 3856:(3): 267–285. 3840: 3821:(3): 667–678. 3805: 3776:(3): 401–409. 3756: 3729: 3708:(3): 911–920. 3688: 3639: 3610:(5): 319–329. 3587: 3580: 3562: 3503: 3474:(8): 680–682. 3454: 3413: 3384:(4): 941–947. 3364: 3331: 3317: 3263: 3205: 3156: 3142: 3122: 3079:; followed by 3069:strain K-12". 3057: 3027: 3025: 3022: 3021: 3020: 3011: 3006: 3001: 2996: 2991: 2984: 2981: 2969:recombineering 2961:gateway method 2945:model organism 2940: 2937: 2915: 2912: 2909: 2908: 2897: 2894: 2889: 2883: 2882: 2877: 2869:Antiterminator 2866: 2863: 2860: 2857: 2853: 2852: 2845:Antiterminator 2842: 2839: 2836: 2833: 2829: 2828: 2818: 2815: 2811: 2808: 2805: 2801: 2800: 2774: 2771: 2767: 2764: 2761: 2757: 2756: 2753: 2750: 2747: 2744: 2740: 2739: 2736: 2733: 2730: 2727: 2723: 2722: 2719: 2716: 2713: 2710: 2706: 2705: 2702: 2699: 2696: 2693: 2689: 2688: 2685: 2682: 2679: 2676: 2672: 2671: 2668: 2665: 2662: 2659: 2655: 2654: 2651: 2648: 2645: 2642: 2638: 2637: 2634: 2631: 2628: 2625: 2621: 2620: 2617: 2614: 2611: 2608: 2604: 2603: 2600: 2597: 2594: 2591: 2587: 2586: 2583: 2580: 2577: 2574: 2570: 2569: 2566: 2563: 2560: 2557: 2553: 2552: 2549: 2546: 2543: 2540: 2536: 2535: 2532: 2529: 2526: 2523: 2519: 2518: 2515: 2512: 2509: 2506: 2502: 2501: 2495: 2492: 2489: 2486: 2482: 2481: 2475: 2472: 2469: 2466: 2462: 2461: 2450:Inhibits host 2448: 2445: 2442: 2439: 2435: 2434: 2423: 2420: 2417: 2414: 2410: 2409: 2402: 2399: 2396: 2393: 2389: 2388: 2384: 2380: 2372: 2368: 2364: 2360: 2356: 2352: 2348: 2345: 2342: 2339: 2336: 2332: 2331: 2320: 2316: 2312: 2308: 2304: 2300: 2296: 2292: 2288: 2284: 2281: 2278: 2274: 2271: 2268: 2264: 2263: 2239: 2235: 2231: 2228: 2225: 2222: 2219: 2215: 2214: 2203: 2200: 2197: 2194: 2190: 2189: 2186: 2183: 2180: 2173: 2170: 2169: 2168: 2164: 2157: 2155: 2152: 2145: 2143: 2140: 2133: 2130: 2129: 2126: 2119: 2103: 2099: 2091: 2087: 2083: 2079: 2075: 2071: 2064: 2060: 2056: 2048: 2044: 2040: 2036: 2032: 2028: 2024: 2020: 2006: 2005: 1999: 1995: 1992: 1988: 1985: 1981: 1978: 1939: 1936: 1926: 1923: 1922: 1921: 1918: 1907: 1902: 1899:domain on the 1893: 1888: 1877: 1874: 1865: 1864: 1858: 1851: 1846: 1843: 1832: 1829: 1809: 1805: 1797: 1794: 1793: 1792: 1787: 1780: 1775: 1762: 1757: 1748: 1737: 1726: 1707: 1698: 1693: 1674: 1655: 1648: 1634: 1633: 1628: 1621: 1614: 1599: 1596: 1562:, between the 1551: 1548: 1531: 1530: 1525: 1518: 1513: 1508: 1501: 1494: 1489: 1484: 1477: 1472: 1467: 1458: 1453: 1442: 1426: 1421: 1412: 1401: 1394: 1387: 1376: 1371: 1366: 1359: 1352: 1347: 1324:Main article: 1321: 1318: 1317: 1316: 1304: 1255: 1250: 1230: 1227: 1206: 1203: 1202: 1201: 1200: 1199: 1186: 1183: 1178: 1175: 1146: 1143: 1142: 1141: 1138: 1135: 1122: 1105:RNA polymerase 1095: 1094: 1085: 1084: 1083: 1074: 1073: 1072: 1063: 1062: 1061: 1060: 1059: 1057: 1054: 1053: 1052: 1045: 1033: 1018: 1009: 1006: 992: 974: 971: 970: 969: 942: 939: 934: 929: 922:antiterminator 918: 915: 910: 863:Main article: 860: 857: 856: 855: 848: 844: 831: 830: 821: 820: 812: 811: 810: 809: 808: 806: 803: 787: 786: 783:bacteriostatic 771: 731:RNA polymerase 727:RNA polymerase 723:antiterminator 719: 718:reading frame. 714: 703: 684: 666: 665: 649: 642: 635: 630: 625:puts negative 610: 609: 598: 595: 570:to the hosts. 558: 555: 553: 550: 505: 502: 395: 394: 387: 385: 381: 380: 373: 369: 368: 361: 357: 356: 349: 345: 344: 341:Caudoviricetes 337: 333: 332: 325: 321: 320: 317:Heunggongvirae 313: 309: 308: 301: 294: 293: 288: 284: 283: 270: 269: 262: 261: 253: 252: 242: 241: 224: 223: 165:"Lambda phage" 138: 136: 129: 122: 121: 104:September 2013 83: 81: 74: 69: 43: 42: 40: 33: 26: 9: 6: 4: 3: 2: 5644: 5633: 5630: 5628: 5625: 5623: 5620: 5619: 5617: 5600: 5595: 5591: 5587: 5582: 5578: 5574: 5569: 5565: 5561: 5556: 5552: 5547: 5541: 5537: 5532: 5526: 5522: 5521: 5519: 5517: 5513: 5509: 5504: 5492: 5484: 5483: 5480: 5472: 5469: 5467: 5464: 5462: 5459: 5457: 5454: 5453: 5452: 5449: 5447: 5444: 5442: 5439: 5437: 5434: 5432: 5429: 5427: 5424: 5423: 5421: 5419: 5415: 5409: 5406: 5404: 5401: 5399: 5396: 5392: 5389: 5387: 5386:Threose 5384: 5382: 5379: 5378: 5377: 5374: 5373: 5371: 5369: 5365: 5359: 5356: 5354: 5351: 5349: 5346: 5344: 5343:Deoxyribozyme 5341: 5339: 5336: 5332: 5331:Mitochondrial 5329: 5327: 5324: 5323: 5322: 5319: 5318: 5316: 5314: 5308: 5298: 5295: 5293: 5290: 5288: 5285: 5283: 5280: 5278: 5275: 5273: 5270: 5269: 5267: 5263: 5257: 5254: 5250: 5247: 5245: 5242: 5240: 5237: 5235: 5232: 5231: 5230: 5227: 5225: 5222: 5218: 5215: 5213: 5210: 5208: 5205: 5204: 5203: 5200: 5199: 5197: 5195: 5191: 5185: 5182: 5180: 5177: 5175: 5172: 5170: 5167: 5163: 5160: 5159: 5158: 5155: 5154: 5152: 5150: 5149:Translational 5146: 5143: 5139: 5133: 5129: 5123: 5120: 5118: 5115: 5113: 5110: 5108: 5105: 5104: 5102: 5098: 5094: 5086: 5081: 5079: 5074: 5072: 5067: 5066: 5063: 5051: 5048: 5046: 5043: 5041: 5038: 5036: 5035: 5031: 5029: 5028: 5024: 5022: 5021: 5017: 5015: 5014: 5010: 5008: 5007: 5003: 5001: 4998: 4996: 4995: 4991: 4989: 4988:Fission yeast 4986: 4984: 4983:Budding yeast 4981: 4979: 4978: 4974: 4972: 4971: 4970:Chlamydomonas 4967: 4965: 4964: 4960: 4958: 4955: 4954: 4951: 4947: 4943: 4935: 4930: 4928: 4923: 4921: 4916: 4915: 4912: 4905: 4902: 4900: 4897: 4895: 4892: 4889: 4886: 4883: 4880: 4877: 4873: 4872: 4860: 4856: 4852: 4848: 4843: 4838: 4834: 4830: 4825: 4820: 4816: 4812: 4808: 4803: 4800: 4796: 4792: 4788: 4784: 4780: 4776: 4772: 4768: 4764: 4760: 4755: 4751: 4747: 4742: 4737: 4732: 4727: 4723: 4719: 4715: 4711: 4707: 4702: 4698: 4694: 4689: 4684: 4680: 4676: 4672: 4668: 4664: 4659: 4655: 4651: 4647: 4643: 4639: 4635: 4630: 4626: 4622: 4618: 4614: 4610: 4606: 4601: 4597: 4593: 4588: 4583: 4579: 4575: 4571: 4566: 4562: 4558: 4553: 4548: 4543: 4538: 4534: 4530: 4526: 4522: 4518: 4513: 4509: 4505: 4500: 4495: 4490: 4485: 4481: 4477: 4474:(4): 1282–7. 4473: 4469: 4465: 4461: 4457: 4453: 4449: 4444: 4443: 4423: 4419: 4413: 4405: 4401: 4396: 4391: 4386: 4381: 4377: 4373: 4370:(9): e00407. 4369: 4365: 4364: 4359: 4351: 4343: 4339: 4335: 4331: 4327: 4323: 4319: 4315: 4311: 4307: 4303: 4299: 4295: 4288: 4274: 4270: 4263: 4255: 4251: 4246: 4241: 4237: 4233: 4229: 4225: 4221: 4217: 4213: 4206: 4193: 4189: 4185: 4181: 4177: 4173: 4169: 4165: 4161: 4154: 4146: 4142: 4137: 4132: 4127: 4122: 4118: 4114: 4110: 4106: 4102: 4095: 4087: 4083: 4078: 4073: 4069: 4065: 4061: 4057: 4053: 4046: 4038: 4032: 4028: 4021: 4013: 4009: 4005: 4001: 3997: 3993: 3986: 3978: 3976: 3971: 3965: 3957: 3953: 3949: 3945: 3941: 3937: 3930: 3922: 3918: 3914: 3910: 3906: 3902: 3898: 3894: 3890: 3886: 3879: 3871: 3867: 3863: 3859: 3855: 3851: 3844: 3836: 3832: 3828: 3824: 3820: 3816: 3809: 3801: 3797: 3792: 3787: 3783: 3779: 3775: 3771: 3767: 3760: 3752: 3748: 3744: 3740: 3733: 3725: 3721: 3716: 3711: 3707: 3703: 3699: 3692: 3684: 3680: 3675: 3670: 3666: 3662: 3658: 3654: 3650: 3643: 3635: 3631: 3626: 3621: 3617: 3613: 3609: 3605: 3601: 3594: 3592: 3583: 3581:9781555819750 3577: 3573: 3566: 3558: 3554: 3549: 3544: 3539: 3534: 3530: 3526: 3522: 3518: 3514: 3507: 3499: 3495: 3490: 3485: 3481: 3477: 3473: 3469: 3468:Cell Research 3465: 3458: 3450: 3446: 3441: 3436: 3432: 3428: 3424: 3417: 3409: 3405: 3400: 3395: 3391: 3387: 3383: 3379: 3375: 3368: 3360: 3356: 3352: 3348: 3344: 3338: 3336: 3327: 3321: 3313: 3309: 3304: 3299: 3295: 3291: 3287: 3283: 3276: 3274: 3272: 3270: 3268: 3259: 3255: 3250: 3245: 3240: 3235: 3231: 3227: 3223: 3216: 3214: 3212: 3210: 3201: 3197: 3193: 3189: 3184: 3179: 3175: 3171: 3167: 3160: 3145: 3139: 3135: 3134: 3126: 3118: 3114: 3109: 3104: 3100: 3096: 3092: 3088: 3084: 3076: 3072: 3068: 3061: 3047: 3043: 3039: 3032: 3028: 3019: 3015: 3012: 3010: 3007: 3005: 3002: 3000: 2997: 2995: 2992: 2990: 2987: 2986: 2980: 2978: 2974: 2970: 2966: 2962: 2958: 2954: 2950: 2946: 2936: 2932: 2928: 2920: 2906: 2902: 2898: 2895: 2893: 2890: 2888: 2885: 2884: 2880: 2874: 2870: 2867: 2861: 2858: 2855: 2854: 2850: 2846: 2843: 2837: 2834: 2831: 2830: 2826: 2822: 2819: 2809: 2806: 2803: 2802: 2798: 2794: 2790: 2786: 2782: 2778: 2775: 2765: 2762: 2759: 2758: 2754: 2748: 2745: 2742: 2741: 2737: 2731: 2728: 2725: 2724: 2720: 2714: 2711: 2708: 2707: 2703: 2697: 2694: 2691: 2690: 2686: 2680: 2677: 2674: 2673: 2669: 2663: 2660: 2657: 2656: 2652: 2646: 2643: 2641:M V U G L T Z 2640: 2639: 2635: 2629: 2626: 2623: 2622: 2618: 2612: 2609: 2606: 2605: 2601: 2595: 2592: 2589: 2588: 2584: 2578: 2575: 2572: 2571: 2567: 2561: 2558: 2555: 2554: 2550: 2544: 2541: 2538: 2537: 2533: 2527: 2524: 2521: 2520: 2516: 2510: 2507: 2504: 2503: 2499: 2496: 2490: 2487: 2484: 2483: 2479: 2476: 2470: 2467: 2464: 2463: 2459: 2455: 2454: 2449: 2443: 2440: 2437: 2436: 2432: 2428: 2424: 2418: 2415: 2412: 2411: 2407: 2403: 2397: 2394: 2391: 2390: 2378: 2359:1 (affinity O 2346: 2340: 2337: 2334: 2333: 2329: 2327: 2295:3 (affinity O 2282: 2272: 2269: 2266: 2265: 2261: 2257: 2253: 2249: 2245: 2229: 2223: 2220: 2217: 2216: 2212: 2208: 2204: 2198: 2195: 2192: 2191: 2177: 2161: 2156: 2149: 2144: 2137: 2132: 2131: 2127: 2124: 2120: 2117: 2113: 2112: 2111: 2109: 2097: 2068: 2054: 2053:cooperativity 2017: 2013: 2011: 2003: 2000: 1993: 1986: 1979: 1976: 1973: 1972: 1971: 1964: 1960: 1958: 1953: 1944: 1935: 1931: 1919: 1916: 1912: 1908: 1905: 1898: 1894: 1891: 1884: 1880: 1879: 1869: 1861: 1854: 1847: 1844: 1841: 1837: 1836:transcription 1833: 1830: 1827: 1823: 1822: 1821: 1819: 1802: 1790: 1783: 1776: 1773: 1769: 1765: 1758: 1755: 1751: 1744: 1740: 1733: 1729: 1722: 1718: 1714: 1710: 1703: 1699: 1696: 1689: 1685: 1681: 1677: 1670: 1666: 1662: 1658: 1651: 1644: 1643: 1638: 1631: 1624: 1617: 1610: 1609: 1604: 1595: 1593: 1589: 1585: 1581: 1577: 1573: 1569: 1565: 1561: 1557: 1547: 1545: 1541: 1537: 1528: 1521: 1514: 1511: 1504: 1497: 1490: 1487: 1480: 1473: 1470: 1463: 1459: 1456: 1449: 1445: 1439:sites in the 1438: 1434: 1429: 1422: 1419: 1415: 1408: 1404: 1397: 1390: 1383: 1379: 1372: 1369: 1362: 1355: 1348: 1345: 1341: 1337: 1333: 1332: 1331: 1327: 1313: 1309: 1305: 1302: 1298: 1294: 1290: 1286: 1282: 1279:mRNA (as the 1278: 1274: 1270: 1266: 1262: 1258: 1251: 1248: 1244: 1240: 1236: 1233: 1232: 1226: 1224: 1220: 1216: 1212: 1197: 1196: 1195: 1191: 1187: 1184: 1181: 1180: 1174: 1171: 1167: 1163: 1160: 1151: 1139: 1136: 1133: 1129: 1125: 1118: 1114: 1113: 1112: 1110: 1106: 1089: 1078: 1067: 1050: 1046: 1043: 1038: 1034: 1031: 1027: 1023: 1019: 1016: 1015:θ replication 1012: 1011: 1005: 1001: 997: 995: 988: 984: 980: 967: 963: 959: 955: 951: 947: 943: 940: 937: 930: 927: 923: 919: 916: 913: 906: 902: 898: 894: 890: 886: 885: 884: 878: 877: 871: 866: 853: 849: 845: 842: 841: 840: 825: 816: 802: 800: 796: 792: 784: 780: 776: 772: 769: 765: 761: 757: 753: 749: 745: 741: 737: 732: 728: 724: 720: 717: 710: 706: 699: 695: 691: 687: 680: 677:Cro binds to 676: 675: 670: 663: 659: 655: 652: 645: 638: 631: 628: 624: 620: 619: 614: 607: 603: 599: 596: 593: 589: 585: 581: 577: 573: 572: 571: 563: 549: 545: 543: 539: 535: 531: 522: 518: 510: 501: 499: 495: 491: 487: 483: 479: 475: 471: 467: 462: 459: 455: 451: 447: 443: 439: 435: 434: 429: 428:bacteriophage 425: 424: 420:, officially 419: 415: 411: 410: 401: 392: 391: 386: 383: 382: 379: 378: 374: 371: 370: 367: 366: 362: 359: 358: 355: 354: 350: 347: 346: 343: 342: 338: 335: 334: 331: 330: 326: 323: 322: 319: 318: 314: 311: 310: 307: 306: 305:Duplodnaviria 302: 299: 296: 295: 292: 289: 286: 285: 280: 275: 271: 268: 263: 259: 254: 251: 247: 238: 235: 220: 217: 209: 198: 195: 191: 188: 184: 181: 177: 174: 170: 167: – 166: 162: 161:Find sources: 155: 151: 145: 144: 139:This article 137: 133: 128: 127: 118: 115: 107: 97: 93: 87: 84:This article 82: 73: 72: 67: 65: 58: 57: 52: 51: 46: 41: 32: 31: 19: 5632:Siphoviridae 5515: 5461:Bacterial 5436:Lambda phage 5435: 5100:Constituents 5032: 5025: 5018: 5011: 5004: 4992: 4975: 4968: 4961: 4957:Lambda phage 4956: 4874:Life Cycle, 4859:the original 4854: 4841: 4814: 4810: 4762: 4758: 4713: 4709: 4670: 4666: 4637: 4633: 4608: 4604: 4580:(1): 13–16. 4577: 4573: 4524: 4520: 4471: 4467: 4447: 4426:. Retrieved 4424:. 2017-05-17 4421: 4412: 4367: 4361: 4350: 4301: 4297: 4287: 4277:, retrieved 4272: 4262: 4219: 4215: 4205: 4195:, retrieved 4167: 4163: 4153: 4108: 4104: 4094: 4059: 4055: 4045: 4026: 4020: 3995: 3992:Biochemistry 3991: 3985: 3973: 3964: 3939: 3935: 3929: 3888: 3884: 3878: 3853: 3849: 3843: 3818: 3814: 3808: 3773: 3769: 3759: 3742: 3738: 3732: 3705: 3701: 3691: 3656: 3652: 3642: 3607: 3603: 3571: 3565: 3520: 3516: 3506: 3471: 3467: 3457: 3430: 3426: 3416: 3381: 3377: 3367: 3350: 3346: 3343: 3320: 3285: 3229: 3225: 3173: 3169: 3159: 3147:. Retrieved 3132: 3125: 3093:(1): 51–64. 3090: 3086: 3074: 3070: 3066: 3060: 3049:. Retrieved 3031: 3017: 3004:Sankar Adhya 2942: 2933: 2929: 2925: 2904: 2900: 2896:Host protein 2892:SOS Response 2876: 2872: 2848: 2824: 2796: 2792: 2788: 2784: 2780: 2460:to proceed. 2451: 2430: 2426: 2405: 2376: 2324: 2259: 2255: 2251: 2247: 2243: 2210: 2206: 2188:Description 2122: 2115: 2108:autonegative 2107: 2095: 2069: 2018: 2014: 2007: 2001: 1974: 1969: 1949: 1932: 1928: 1914: 1910: 1900: 1896: 1886: 1882: 1856: 1849: 1826:SOS response 1818:SOS response 1815: 1785: 1778: 1771: 1767: 1760: 1746: 1742: 1735: 1731: 1724: 1720: 1716: 1712: 1705: 1701: 1691: 1687: 1683: 1679: 1672: 1668: 1664: 1660: 1653: 1646: 1626: 1619: 1612: 1587: 1583: 1571: 1567: 1563: 1559: 1555: 1553: 1532: 1523: 1516: 1506: 1499: 1492: 1482: 1475: 1465: 1461: 1451: 1447: 1440: 1436: 1432: 1424: 1417: 1410: 1406: 1399: 1392: 1385: 1381: 1374: 1364: 1357: 1350: 1343: 1339: 1335: 1329: 1311: 1307: 1303:is observed. 1300: 1296: 1292: 1288: 1284: 1280: 1276: 1272: 1268: 1264: 1253: 1246: 1242: 1238: 1234: 1222: 1218: 1214: 1210: 1208: 1193: 1189: 1169: 1165: 1161: 1158: 1156: 1127: 1120: 1116: 1108: 1102: 1048: 1029: 1025: 1021: 1002: 998: 990: 986: 982: 978: 976: 957: 953: 949: 945: 932: 925: 908: 900: 896: 892: 888: 882: 874: 851: 838: 788: 778: 774: 767: 763: 759: 755: 751: 747: 743: 739: 735: 712: 711:gene in the 708: 701: 700:gene in the 697: 693: 689: 682: 678: 661: 657: 647: 640: 633: 601: 587: 583: 575: 568: 546: 541: 537: 533: 527: 515: 473: 463: 437: 431: 422: 421: 417: 414:lambda phage 413: 408: 407: 406: 389: 388: 376: 365:Siphoviridae 364: 353:Caudovirales 352: 340: 328: 316: 304: 297: 287:(unranked): 266: 249: 230: 212: 206:January 2012 203: 193: 186: 179: 172: 160: 148:Please help 143:verification 140: 110: 101: 90:Please help 85: 61: 54: 48: 47:Please help 44: 5540:Wikispecies 5456:P1-derived 5224:Antisense 5117:Nucleotides 5112:Nucleosides 5107:Nucleobases 5006:Arabidopsis 4977:Tetrahymena 4164:The Enzymes 3977:Interactive 3523:(4): e363. 3018:Lambdavirus 2821:Excisionase 1863:difference. 1544:antibiotics 1132:Pribnow box 924:, binds to 920:Q, another 865:Lytic cycle 750:genes, and 478:lytic cycle 418:coliphage λ 377:Lambdavirus 329:Uroviricota 5616:Categories 5408:Morpholino 5321:Organellar 5229:Processual 5194:Regulatory 5138:non-coding 5027:Drosophila 5020:C. elegans 4994:Neurospora 4428:2023-11-28 4279:2023-11-28 4197:2023-11-28 3359:1156862867 3051:2021-07-24 3024:References 2505:Rz and Rz1 2205:(Clear 3) 2055:. Thus, O 2010:cI protein 1840:DNA repair 1791:promoters. 1536:starvation 1370:promoters. 1275:mRNA than 1042:concatemer 833:terminator 627:supercoils 623:DNA gyrase 606:DNA ligase 552:Life cycle 536:site. 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