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==As a genetic tool== Lambda phage has been used heavily as a [[model organism]] and has been an excellent tool first in [[microbial genetics]], and then later in [[molecular genetics]].<ref>{{Citation |last1=Pitre |first1=Emmanuelle |title=Chapter Four - Understanding viral replication and transcription using single-molecule techniques |date=2021-01-01 |url=https://www.sciencedirect.com/science/article/pii/S1874604721000159 |journal=The Enzymes |volume=49 |pages=83β113 |editor-last=Cameron |editor-first=Craig E. |access-date=2023-11-28 |series=Viral Replication Enzymes and their Inhibitors Part A |publisher=Academic Press |last2=te Velthuis |first2=Aartjan J. W. |doi=10.1016/bs.enz.2021.07.005 |pmid=34696840 |s2cid=239573473 |editor2-last=Arnold |editor2-first=Jamie J. |editor3-last=Kaguni |editor3-first=Laurie S.}}</ref> Some of its uses include its application as a vector for the cloning of [[recombinant DNA]]; the use of its site-specific recombinase (int) for the shuffling of cloned DNAs by the [[Gateway Technology|gateway method]];<ref>{{Cite journal |last1=Reece-Hoyes |first1=John S. |last2=Walhout |first2=Albertha J. M. |date=2018-01-01 |title=Gateway Recombinational Cloning |url=http://cshprotocols.cshlp.org/content/2018/1/pdb.top094912 |journal=Cold Spring Harbor Protocols |language=en |volume=2018 |issue=1 |pages=pdb.top094912 |doi=10.1101/pdb.top094912 |issn=1940-3402 |pmid=29295908|pmc=5935001 }}</ref> and the application of its Red [[operon]], including the proteins Red alpha (also called 'exo'), beta and gamma in the DNA engineering method called [[recombineering]]. The 48 kb DNA fragment of lambda phage is not essential for productive infection and can be replaced by foreign DNA,<ref>{{Citation |last1=Feiss |first1=Michael |title=Bacteriophage Lambda Terminase and the Mechanism of Viral DNA Packaging |date=2013 |url=https://www.ncbi.nlm.nih.gov/books/NBK6485/ |work=Madame Curie Bioscience Database [Internet] |access-date=2023-11-28 |publisher=Landes Bioscience |language=en |last2=Catalano |first2=Carlos Enrique}}</ref> 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.<ref>{{Cite journal |last=Smith |first=George P. |date=1985 |title=Filamentous Fusion Phage: Novel Expression Vectors that Display Cloned Antigens on the Virion Surface |url=https://www.jstor.org/stable/1694587 |journal=Science |volume=228 |issue=4705 |pages=1315β1317 |doi=10.1126/science.4001944 |jstor=1694587 |pmid=4001944 |bibcode=1985Sci...228.1315S |issn=0036-8075}}</ref> Lambda phage can also be manipulated and used as an anti-cancer vaccine that targets human [[ASPH|aspartyl (asparaginyl) Ξ²-hydroxylase]] (ASPH, HAAH), which has been shown to be beneficial in cases of hepatocellular carcinoma in mice.<ref>{{Cite journal |last1=Iwagami |first1=Yoshifumi |last2=Casulli |first2=Sarah |last3=Nagaoka |first3=Katsuya |last4=Kim |first4=Miran |last5=Carlson |first5=Rolf I. |last6=Ogawa |first6=Kosuke |last7=Lebowitz |first7=Michael S. |last8=Fuller |first8=Steve |last9=Biswas |first9=Biswajit |last10=Stewart |first10=Solomon |last11=Dong |first11=Xiaoqun |last12=Ghanbari |first12=Hossein |last13=Wands |first13=Jack R. |year=2017 |title=Lambda phage-based vaccine induces antitumor immunity in hepatocellular carcinoma |journal=[[Heliyon]] |language=en |volume=3 |issue=9 |pages=e00407 |doi=10.1016/j.heliyon.2017.e00407 |doi-access=free |pmc=5619992 |pmid=28971150|bibcode=2017Heliy...300407I }}</ref> Lambda phage has also been of major importance in the study of [[Transduction (genetics)|specialized transduction]].<ref>{{Cite web |date=2017-05-17 |title=7.14E: Bacteriophage Lambda as a Cloning Vector |url=https://bio.libretexts.org/Bookshelves/Microbiology/Microbiology_(Boundless)/07%3A_Microbial_Genetics/7.14%3A_Cloning_Techniques/7.14E%3A_Bacteriophage_Lambda_as_a_Cloning_Vector |access-date=2023-11-28 |website=Biology LibreTexts |language=en}}</ref>
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