Table 3.

Molecular genetic modifiers of Apc knockout mouse models

Modifier affects	Modifier gene(s)	Modifier allele(s)	Allele property	Apc Model	Effect of mutant allele on intestinal tumor multiplicity	Factor of effect	Reference
Karyotypic stability	BubR1	Bub1bGt(neo-btk)1Dai	Knockout (het)	Min	Decrease/Increasea	2/10a	Rao et al., 2005138
	Cdx2	Cdx2tm1Mmt	Knockout (het)	Δ716	Decrease/Increasea	9/6a	Aoki et al., 2003141
	Terc	Terctm1Rdp	Knockout	Min	Decrease (at G4)	10	Rudolph et al., 2001153
DNA mutation rate	Pms2	Pms2tm1Lisk	Knockout	Min	Increase	3	Baker et al., 1998154
	Mlh1	Mlh1tm1Lisk	Knockout	Min	Increase	3	Shoemaker et al., 2000155
	Msh2	Msh2tm1Mak	Knockout	Min	Increase	7	Reitmair et al., 1996156
	Msh3/ Msh6	Msh3tm1Rak Msh6tm1Rak	Knockout	1638N	Increase	12	Kuraguchi et al., 2001157
	Fen1	Fen1tm1Rak	Knockout (het)	1638N	Increase	1.5	Kucherlapati et al., 2002158
	Myh	Mutyhtm1Jhmi	Knockout	Min	Increase	1.5	Sieber et al., 2004159
Recombination rates	Rb9	Rb(7.18)9Lub	Translocation	Min	Decrease	19	Haigis and Dove, 2003100
	Recql4	Recql4tm1Glu	Knockout	Min	Increase	2	Mann et al., 2005144
	Blm	Blmtm3Brd	Hypomorph	Min	Increase	3	Luo et al., 2000143
Differentiation	EphB2	ΔcyEphB2	Dom neg Tg	Min	Decrease	3	Batlle et al., 2005147
	EphB3	EphB3tm1Kln	Knockout	Min	Increase	2	Batlle et al., 2005147
DNA methylation	Mbd2	Mbd2tm1Bh	Knockout	Min	Decrease	10	Sansom et al., 2003145
	Mbd4	Mbd4tm1Bird	Knockout	Min	Increase	2	Millar et al., 2002146
	Dnmt1	Dnmt1tm1Jae	Knockout (het)	Min	Decrease	2	Cormier and Dove, 200085
Stromal regulation	Foxl1	Foxl1tm1Khk	Knockout	Min	Increase	8	Perrault et al., 2005160
	TSP1	Thbs1tm1Hyn	Knockout	Min	Increase	2	Gutierrez et al., 2003161
Cell growth and proliferation	c-Jun	Juntm2.1Wag	Hypomorph	Min	Decrease	2	Nateri et al., 2005162
	Cyclin D1	Ccnd1tm1Wbg	Knockout	Min	Decrease	6	Hulit et al., 2004163
	Egfr	Egfrwa2	Hypomorph	Min	Decrease	10	Roberts et al., 2002164
	p21	Cdkn1atm1Led	Knockout	1638N	Increase	2	Yang et al., 2001165
	p27	Cdkn1btm1Mlf	Knockout	Min	Increase	5	Philippp-Staheli et al., 2002166
	p53	Trp53tm1Ldo	Knockout	Min	Increase	2	Halberg et al., 2000167
	Igf2	H19tm1Tilg	Activates Ifg2	Min	Increase	2	Sakatani et al., 2005139
Pleiotropic	Matrilysin	Mmp7tm1Lmm	Knockout	Min	Decrease	2	Wilson et al., 1997168
	Pla2g2a	Pla2g2aAKR	Tg	Min	Decrease	2	Cormier et al., 1997125
	BAH	Asphtm1Jed	Knockout	Min	Increase	2	Dinchuk et al., 2002169
	E-cadherin	Cdh1tm1Cbm	Knockout (het)	1638N	Increase	9	Smits et al., 2000170
	PPAR-δ	Ppardtm1Jps	Knockout	Min	Increase	1.5	Harman et al., 2004171
	Netrin-1	Tg-netrin-1	Tg	1638N	Enhances progression	N/A	Mazelin et al., 2004172
	Smad4	Smad4tm1Mmt	Knockout	Δ716	Enhances progression	N/A	Takaku et al., 1999173

^aEffects on the small intestine and colon, respectively ;

^bThe Robertsonian translocation is centromeric fusion of chromosomes 7 and 18.

Note: The Mom (Modifier of Min) and Scc (Susceptibility to colon cancer)¹⁷⁴ loci are in general not yet fully defined in molecular detail (see text) and are therefore not included in Table 3.