TECHNOLOGY REPORT
Generation of a Conditional Knockout Allele for the Janus
Kinase 2 (Jak2) Gene in Mice
Andrea Krempler,
1
Yongyue Qi,
1
Aleata A. Triplett,
1
Jianqiong Zhu,
2
Hallgeir Rui,
2
and
Kay-Uwe Wagner
1
*
1
Eppley Institute for Research in Cancer and Allied Diseases and the Department of Pathology and Microbiology,
University of Nebraska Medical Center, Omaha, Nebraska
2
Department of Oncology, Lombardi Comprehensive Cancer Center, Georgetown University Medical Center,
Washington, DC
Received 13 April 2004; Accepted 4 June 2004
Summary: To study biologically relevant functions of the
Janus kinase 2 (Jak2) in multiple cytokine and hormone
receptor signal transduction pathways, we generated a
conditional knockout (floxed) allele of this gene by plac-
ing loxP sites around the first coding exon of Jak2. Ho-
mozygous floxed animals developed normally and ex-
hibited no phenotypic abnormalities. The conversion of
the floxed allele into a null mutation was achieved by
transmitting the targeted allele through the female
germline of MMTV-Cre (line A) mice. Embryos that carry
two Jak2 null alleles died around midgestation and ex-
hibited impaired definitive erythropoiesis, which is a
hallmark of Jak2 deficiency reported previously in con-
ventional knockouts. This observation suggested that
the Cre-mediated deletion of the first coding exon re-
sults in a true null mutation that is incapable of mediat-
ing signals through the erythropoietin receptor. Using
mouse embryonic fibroblasts derived from Jak2 null em-
bryos and their wildtype littermate controls, we dem-
onstrated that Jak2-deficiency decouples growth
hormone-receptor signaling from its downstream me-
diators, the signal transducer and activator of tran-
scription (Stat) 5a and 5b. genesis 40:52–57, 2004.
© 2004 Wiley-Liss, Inc.
Key words: Janus kinase 2; Stat5; embryonic development;
Cre recombinase; gene targeting
Jak-Stat pathways mediate extracellular polypeptide sig-
nals, such as hormones and cytokines, from transmem-
brane receptors to target gene promoters in the nucleus.
Consequently, cells respond to these signals by altering
their gene expression and, in turn, change their growth
properties and their physiological activities (Aaronson et
al., 2002). Four members of the Janus kinase (Jak) family
have been identified in mammals: Jak1, Jak2, Tyk2, and
Jak3. Genetic studies in mice revealed that the biological
functions of individual Janus kinases are pleiotropic,
which is consistent with their suggested coupling to
multiple cytokine receptors in specific target cells cre-
viewed by Kisseleva et al., 2002). In particular, Jak2 is
suggested to mediate signals through single-chain recep-
tors for ligands such as prolactin (PRL), growth hormone
(GH), erythropoietin (EPO), and thrombopoietin (TPO)
as well as to the multichain IL-3 receptor family (e.g.,
IL-3R and GM-CSF-R), and members of the gp130 recep-
tor family. Although more than one Jak family member
was suggested to interact with certain cytokine recep-
tors in vitro, knockout studies in mice revealed a remark-
able physiological specificity of individual Janus kinases
and their associated Stat proteins (Ihle, 2001; Kisseleva
et al., 2002; Levy et al., 2002).
Among gene deletion models of Janus kinase family
members, Jak2-deficient mice exhibit the most severe
phenotype. Embryos with a complete null mutation of
Jak2 die around day 12.5 of gestation due to impaired
definitive erythropoiesis (Neubauer et al., 1998; Parga-
nas et al., 1998). Conventional knockout models for Jak2
exhibit similarities to EPO and EPO-R-deficient mice (Wu
et al., 1995), suggesting that Jak2 is essential for trans-
ducing signals through the EPO receptor. In vitro differ-
A. Krempler and Y. Qi contributed equally to this work.
Present address for A. Krempler: Department of Medical Biochemistry
and Molecular Biology, University of the Saarland, Building 44, Homburg,
66424, Germany.
* Correspondence to: Dr. Kay-Uwe Wagner, Eppley Institute for Research
in Cancer and Allied Diseases, University of Nebraska Medical Center,
986805 Nebraska Medical Center, Rm. 8009, Omaha, NE 68198-6805.
E-mail: kuwagner@unmc.edu
Contract grant sponsor: Public Health Service, Contract grant number:
CA93797 (to K.U.W.), Contract grant sponsor: National Cancer Institute,
Contract grant number: CA101841 (to H.R., K.U.W.), Contract grant spon-
sor: Public Health Service, National Institutes of Health, Contract grant
number: DK052013 (to H.R.), Contract grant sponsor: Deutsche For-
schungsgemeinschaft, Contract grant number: DFG, KR 2107/1-1 (to A.K.),
Contract grant sponsor: Nebraska Cancer and Smoking Disease Research
Program, Contract grant number: NE DHHS LB595 (to K.U.W.), Contract
grant sponsor: Cattlemen’s Ball of Nebraska (for generation of the Jak2-
deficient animal model).
Published online in
Wiley InterScience (www.interscience.wiley.com)
DOI: 10.1002/gene.20063
© 2004 Wiley-Liss, Inc. genesis 40:52–57 (2004)