Clock Gene in Animal Update: 2009.5.8 Site Map
List of clock genes Outline of clock genes Animal model List of animals

Mutant of Clock Genes

Core Loop
PERIOD CLOCK ARNTL (BMAL) CRY (mammal) timeless (fly)
Interlocking Loop
DEC1,2 ID1-4 E4BP4 (vri) Pdp1 (fly) REV-ERB ROR SIRT1
Regulator
cry (fly) GSK3B CSNK1E CSNK2A BTRC (slmb) FBXL3 JET PP
See also JAX Mice Strain for Circadian Rhythms

PERIOD →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Per1 Per1tm1Brd(Per1Brdm1) KO (null) Short (-1.1h) Zheng2001
Per1tm1Drw (Per1ldc) KO (null) AR (10~14 day) Bae2001
Per1tm1Saco KO (null) Short (-0.69~0.77h) Cermakian2001
Mouse Per2  Per2tm1Brd(Per2Brdm1, Per2m) KO (del: PASB) Short (-1.6h)→AR (2~18 day)
Cancer prone (Fu2002)
Zheng1999
Per2tm1Drw (Per2ldc) KO (null) AR (~20 day) Bae2001
Per2tm1Ccl KO (null) Same to Per2m (data not shown) Fu2002
Per2tm1Jt(Per2Luciferase) Knock-in Per2::Luc. Luc gene was fused to C-terminal of Per2 gene. Normal circadian activity. Yoo2004
Mouse Per3 Per3tm1Drw KO (null) Short (-0.5h) Shearman2000
Drosophila per per01 (per0) EMS (Q464@) AR Konopka1971
perL1 (perL) EMS (V243D) Long (+4~6h) Konopka1971
pers (perS) EMS (S589N) Short (-5h) Konopka1971
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CLOCK, NPAS2 →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Clock Clockm1Jt (Clockdelta19) ENU (SDN) Long ( h)→AR (-day) Vitaterna1994
Clocktm1.1Drw (Clockdelta5-6) KO (null) Normal. AR with Npas2-KO Debruyne2006
Mouse Npas2 Npas2tm1Slm (NPAS-deltabHLH, NPAS2-lacZ) KO Impaired memory and sleep.
AR with Clock-KO
Garcia2000
Zebrafish clock1 clockdg3 ENU (DN)(I254N) Short (-h) Tan2008
Drosophila Clk ClkJrk EMS (Q776@) (SDN) ClkJrk/Jrk: AR
ClkJrk/+: Long (~24.67h), AR (47%)
Allada1998
Clkar EMS (Missense, exon 2/intron 2 splice site) Clkar/+: Normal
Clkar/ar, Clkar/Jrk: AR
Allada2003
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ARNTL (BMAL) →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse ARNTL Arntltm1Bra KO Homo: AR. Hetero: Normal Bunger2000
Arntltm1.1Weit KO conditional Homo+ubiquitous Cre: AR Storch2007
Mouse ARNTL2 NA - - -
Drosophila cyc cyc01 EMS (A586T) Homo: AR. Hetero: Normal Rutial1998
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BHLHE40,41 (DEC1, 2), cwo →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Bhlhe40 (Dec1, Sharp2)
Bhlhe40tm1Tan KO Defects in  CD4+ T cell activation. Altered clock-controlled gene expression Sun2001
Gréchez-Cassiau2003
Bhlhe40tm1Rhli KO Increased hippocampal exon 4-specific Bdnf mRNA levels  Jiang2008
Bhlhe40tm1.1Rhli KO
Mouse Bhlhe41 (Dec2, Sharp1) Bhlhe41tm1Mjro KO Bglhe40, 41 W-KO mouse: slightly Long, altered response to light. Rossner2008
Drosophila cwo PBac{WH}stich1f05073 PBac Reduced-amplitude molecular and behavioral rhythms Lim2007, Matsumoto2007, Kadener2007
PBac{RB}stich1e04207 PBac
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ID1-4 →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
MouseId1 Id1tm1Zhu KO Tumorigenesis. Lethal, postnatal. Yan1997
Id1tm1Xhsu KO (Reporter) Perry2007
Mouse Id2 Id2tm1Yyk KO Lack of lymph nodes and Peyer's patches. Defect in NK-cell. Yokota1999
Id2tm1Ago KO Defect in CD8(+) T cell maturation. Cannarile2006
Id2tm1Mcha KO Defect in olfactory neurogenesis. Havrda2008
Id2tm1*** KO Abnormally rapid entrainment. 1/4 was arrhythmic. Duffield2009
Mouse Id3 Id3tm1Zhu KO Impaired immune responses and B-cell proliferation. Pan1999
Id3tm1Cmu KO Defect in H-Y TCR-mediated negative selection. Rivera2000
Mouse Id4 Id4tm1Mais KO Defect in neural cell proliferation. Yun2004
Normal entrainment responses. Duffield2009
Id4tm1Fsky KO Defect in neural differentiation. Bedford2005
Drosophila emc 84 alleles and 6 Transgenics - - -
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NFIL3(E4BP4), vrille →Homolog list; → OutlineengJPN

Gene Allele Mutagen Phenotype Reference
Mouse Nfil3NANANANA
Drosophila vri vri1 EMS (DN) Homo, lethal. Hetero, -0.4~0.8 h Blau1999
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DBP, TEF, HLF, Pdp1 →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Dbp Dbptm1Schb (Dbpnull) KO (null) Triple mutant for Dbp, Tef, Hlf cause lethal seizures, but showed normal circadian rhythm. Lopez-Molina1997
Mouse Tef Teftm1Schb KO (null) Gachon2004
Mouse Hlf Hlftm1Schb KO (null)
Drosophila Pdp1 Pdp1P205 delta2-3 Homo, lethal. Hetero, Long (+0.5h) Cyran2003
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NR1D1, NR1D2 (REV-ERB-ALPHA, BETA) →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Nr1D1 (Rev-erba Nr1d1tm1Ven KO Increased cell death and delayed development in the cerebellum. Chomez2000
Nr1d1tm1Schb KOShort (-0.38h). Altered phase shifting properties. Preitner2002
Mouse Nr1D2 (Rev-erbb NA NA NA NA
Drosophila Eip75B Eip75B1, etc. -Lethal Bialecki2002
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RORA, B, C→Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Rora Rorasg(staggerer) Spontaneous Short (0.35h)(Sato2004).  Ataxia (defect in Purkinje cells). Sidman1962
Hamilton1996
Roratm1Vgi KO (null)Die between 24 to 28 days after birth. staggerer like phenotypes. Dussault1998
Roratm1Mba KO (null) Half of homozygote died shortly after weaning. staggerer like phenotypes. Steinmayr1998
Mouse Rorb Rorbtm1Mba KOLong (0.59h). Blind (severely disorganized retina) but entrained by LD. Andre1998
Mouse Rorc Rorctm1Litt KO Defect in thymopoiesis. Sun2000
Rorctm1Amj KO Kurebayashi2000
Drosophila Hr46 - - Lethal -
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SIRT1 →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Sirt1 -
KO Developmental defect. Lethal (early postnatal). Sterile (Survivor). Chen2003McBurney2003
Drosophila Sir2 Sir24.5, Sir25.26 etc. delta2-3Viable, Fertile Newman2002
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CRY→Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Cry1 Cry1tm1Asn KO (Reporter) Short (~1h). AR with Cry2 KO.  Vitaterna1999
Cry1tm1Jhjh KO Short (-1.26h). AR with Cry2 KO.  van der Horst1999
Mouse Cry2 Cry2tm1Asn KO Long (~+1h). Thresher1998
Cry2tm1Jhjh KO Long (+0.86h). AR with Cry2 KO.  van der Horst1999
Drosophila cry cryb (crybaby) EMS (D410N) Rhythmic under LL. Phase response to short light pulse was eliminated. Do not entrain to shifted LD (16 lx, blue) with norpA39(=P41). Stanewsky1998
crym EMS (truncates the last 19 aa) Rhythmic under LL. Phase response to short light pulse was reduced. Busza2004
cry01, cry 02, cry 03 KOSame to cryb. Dolezelova2007
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TIMELESS →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Timeless Timelesstm1Rep KO Prenatal lethal Gotter2000
Drosophila timeless tim01 (tim0) delta2-3 Arrhythmic Sehgal1994, Myers1995
timS1 EMS Homo, 21.1 +/- 0.2 h; hetero, 22.2 +/- 0.1 Rothenfluh2000Genetics
timL1  EMS (K586M) Homo, 27.8; hetero, 25.5 Rothenfluh2000Genetics
timUL EMS (E260K) Homo, 32.8; hetero, 25.6 Rothenfluh 2000Neuron
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GSK3, shaggy →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Gsk3a Gsk3atm1.1Jrw KO Enhanced glucose and insulin sensitivity. Reduced fat mass MacAulay2007
Mouse Gsk3b Gsk3btm1Jrw KO Lethal E13.5 - E14.5 Hoeflich2000
Gsk3btm1Grc KO LethalStankunas2003
Gsk3btm1.1Atak KO Hetero, impaired memory. Kimura2008
Drosophila sgg sgg32/sggM11; hs-sgg (Conditional rescue) 26 h Martinek2001
tim-Gal4 + sggEP1576 (Over expression) 21 h
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CSNK1E, dco (dbt) →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Hamster Csnk1e tau Spontaneous +/+, 24h; +/tau, 22 h; tau/tau, 20 h Ralph1988Lowrey2000
Mouse Csnk1e Csnk1etm1Asil KO (tau) tau/+, 21.8 h;
tau/tau, 20.0 h
Meng2008
Csnk1etm1.1Asil KO (null) Normal
Mouse Csnk1d Under construction KO Liver specific, +2h (Per2::luc) Etchegaray2009
Drosophila
double-time (dco)
dcoS (dbtS) EMS dcoS/+, 21.8 h; dcoS/dcoS, 18h Price1998Kloss1998
dcoL (dbtL) EMS dcoL/+, 24.9 h; dcoL/dcoL, 26.8 h
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CSNK2A1, CkIIalpha →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Csnk2a1 Csnk2a1tm1Dcs KO Lethal, mid-gestation.Lou2008
Drosophila CkIIalpha CkIIalphaTik EMS (DN. M161K, E165D) Hetero, Long (+2.8h).Lin2002
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BTRC, slmb →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Btrc Btrctm1Paga KO Reduced fertility. Abnormality in mitosis. Guardavaccaro2003
Btrctm1Kin KO Defect in degradation of I kappa B. Normal period under DD. Nakayama2003,
Ohsaki2008
Mouse Fbxw11NANANANA
Drosophila slmb slmb8 slmbL.hs (Conditional rescue) ArrhythmicGrima2002
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FBXL3 (Overtime, After-hours)→Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Fbxl3 Fbxl3Afh ENU Long: homo, +3h; hetero, +0.5~1h  
Fbxl3Ovtm ENU Long: homo, +3h; hetero, +0.7hSiepka2007
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FBXL15, jet →Homolog list; → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Mouse Fbxl15NANANANA
Drosophila jet jetr Spontaneous Rhythmic under LL.Koh2006
jetc Spontaneous Rhythmic under LL with jetc/jetr.
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Protein Phosphatase → OutlineengJPN

MGI/FlyBase Allele Mutagen Phenotype Reference
Drosophila wdb
(PP2A regulator)
tim-GAL4 + P{EP}wdbEP3559 -
Long (~0.7h). AR (5%)Sathyanarayanan2004
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Foot note
Note. Name of allele should be italicized (Geneallele), but was shown in roman font (Geneallele) because superscripted italic font is not clear.
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