Table 1 Open in new tab Mouse models...

Genotype

Y chromosome contribution* (Zfy contribution)

Meiotic pairing partner for X

Phenotype

Offspring obtained

References^#

Intact (Zfy1 and Zfy2)

Normal spermatogenesis

Normal mating

N/A

XOSry

Sry tg

None

Spermatogonial proliferation arrest

None

Mazeyrat et al. (2001)

XSxr^bO

Sxr^b (Zfy2/1 fusion gene)

None

Spermatogonial proliferation arrest

None

Vernet et al. (2011, 2012)

XSxr^aO

Sxr^a (Zfy1 and Zfy2)

None

Apoptotic elimination of MI spermatocytes due to X chromosome univalency
Production of haploid and diploid spermatids in small numbers (due to efficient MI apoptosis)
Very few abnormal spermatozoa

N/A

Vernet et al. (2011, 2012)

XSxr^aY *^X

Sxr^a (Zfy1 and Zfy2)

None

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

ROSI

Vernet et al. (2011, 2012)

XY *^XSxr^a

Sxr^a (Zfy1 and Zfy2)

None

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

ROSI and ICSI

Yamauchi et al. (2009)

X^ESxr^bO

Sxr^b (Zfy2/1 fusion gene)

None

Partial elimination of MI spermatocytes due to X chromosome univalency
Production of haploid and diploid spermatids
Decreased number of postmeiotic cells
Incomplete abnormal spermiogenesis

ROSI

Vernet et al. (2011, 2016b) and Yamauchi et al. (2014, 2015)

X^ESxr^bY*^X

Sxr^b (Zfy2/1 fusion gene)

Y*^X

Production of haploid and diploid spermatids
Decreased number of postmeiotic cells
Incomplete abnormal spermiogenesis

ROSI

Vernet et al. (2014, 2016b) and Yamauchi et al. (2014, 2015)

X^EOSry

Sry tg, Eif2s3y tg

None

No elimination of MI spermatocytes due to X chromosome univalency
Production of haploid and diploid spermatids
Decreased number of postmeiotic cells and no spermiogenesis

ROSI

Vernet et al. (2011, 2012, 2016b), Yamauchi et al. (2014, 2015), and Ruthig et al. (2017)

X^EY*^XSry

Sry tg, Eif2s3y tg

Y*^X

Production of haploid and diploid spermatids
Decreased number of postmeiotic cells and no spermiogenesis

ROSI

Vernet et al. (2014, 2016b) and Yamauchi et al. (2014, 2015)

X^E,Z2OSry

Sry tg, Eif2s3y tg, Zfy2 tg

None

Production of haploid and diploid spermatids
Decreased number of postmeiotic cells and no spermiogenesis

ROSI

Vernet et al. (2011) and this study

X^E,Z1/UOSry

Sry tg, Eif2s3y tg, Zfy1/Uba1y tg

None

Production of haploid and diploid spermatids
Decreased number of postmeiotic cells and no spermiogenesis

None

Vernet et al. (2011) and this study

X^E,Z1/U^,^Z2OSry

Sry tg, Eif2s3y tg, Zfy1/Uba1y tg, Zfy2 tg

None

Production of haploid and diploid spermatids
Decreased number of postmeiotic cells and no spermiogenesis

ROSI (poor efficiency)

This study

X^E,Z2Y*^XSry

Sry tg, Eif2s3y tg, Zfy2 tg

Y*^X

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

ROSI and ICSI

Vernet et al. (2014, 2016b) and Yamauchi et al. (2015)

X^E,Z1/UY*^XSry

Sry tg, Eif2s3y tg, Zfy1/Uba1y tg

Y*^X

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

Not tested

Vernet et al. (2014, 2016b)

X^E,Z1/U^,^Z2 Y*^XSry

Sry tg, Eif2s3y tg, Zfy1/Uba1y tg, Zfy2 tg

Y*^X

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

Not tested

Vernet et al. (2014)

X^E,Z2Sxr^bY*^X

Sxr^b, Zfy2 tg

Y*^X

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

ROSI

Yamauchi et al. (2015)

XY Zfy1 KO

Intact (Zfy2) but lacking Zfy1

Normal spermatogenesis

Normal mating

Yamauchi et al. (2022) and Nakasuji et al. (2017)

XY Zfy2 KO

Intact (Zfy1) but lacking Zfy2

Grossly normal spermatogenesis except mild sperm headshape and minor fertility defects

Normal mating

Yamauchi et al. (2022) and Nakasuji et al. (2017)

XY Zfy DKO

Intact but lacking Zfy1 and Zfy2

Decreased number of postmeiotic cells
Production of very few and abnormal spermatozoa

ROSI and ICSI

Yamauchi et al. (2022) and Nakasuji et al. (2017)

Genotype

Y chromosome contribution* (Zfy contribution)

Meiotic pairing partner for X

Phenotype

Offspring obtained

References^#

Intact (Zfy1 and Zfy2)

Normal spermatogenesis

Normal mating

N/A

XOSry

Sry tg

None

Spermatogonial proliferation arrest

None

Mazeyrat et al. (2001)

XSxr^bO

Sxr^b (Zfy2/1 fusion gene)

None

Spermatogonial proliferation arrest

None

Vernet et al. (2011, 2012)

XSxr^aO

Sxr^a (Zfy1 and Zfy2)

None

Apoptotic elimination of MI spermatocytes due to X chromosome univalency
Production of haploid and diploid spermatids in small numbers (due to efficient MI apoptosis)
Very few abnormal spermatozoa

N/A

Vernet et al. (2011, 2012)

XSxr^aY *^X

Sxr^a (Zfy1 and Zfy2)

None

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

ROSI

Vernet et al. (2011, 2012)

XY *^XSxr^a

Sxr^a (Zfy1 and Zfy2)

None

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

ROSI and ICSI

Yamauchi et al. (2009)

X^ESxr^bO

Sxr^b (Zfy2/1 fusion gene)

None

Partial elimination of MI spermatocytes due to X chromosome univalency
Production of haploid and diploid spermatids
Decreased number of postmeiotic cells
Incomplete abnormal spermiogenesis

ROSI

Vernet et al. (2011, 2016b) and Yamauchi et al. (2014, 2015)

X^ESxr^bY*^X

Sxr^b (Zfy2/1 fusion gene)

Y*^X

Production of haploid and diploid spermatids
Decreased number of postmeiotic cells
Incomplete abnormal spermiogenesis

ROSI

Vernet et al. (2014, 2016b) and Yamauchi et al. (2014, 2015)

X^EOSry

Sry tg, Eif2s3y tg

None

No elimination of MI spermatocytes due to X chromosome univalency
Production of haploid and diploid spermatids
Decreased number of postmeiotic cells and no spermiogenesis

ROSI

Vernet et al. (2011, 2012, 2016b), Yamauchi et al. (2014, 2015), and Ruthig et al. (2017)

X^EY*^XSry

Sry tg, Eif2s3y tg

Y*^X

Production of haploid and diploid spermatids
Decreased number of postmeiotic cells and no spermiogenesis

ROSI

Vernet et al. (2014, 2016b) and Yamauchi et al. (2014, 2015)

X^E,Z2OSry

Sry tg, Eif2s3y tg, Zfy2 tg

None

Production of haploid and diploid spermatids
Decreased number of postmeiotic cells and no spermiogenesis

ROSI

Vernet et al. (2011) and this study

X^E,Z1/UOSry

Sry tg, Eif2s3y tg, Zfy1/Uba1y tg

None

Production of haploid and diploid spermatids
Decreased number of postmeiotic cells and no spermiogenesis

None

Vernet et al. (2011) and this study

X^E,Z1/U^,^Z2OSry

Sry tg, Eif2s3y tg, Zfy1/Uba1y tg, Zfy2 tg

None

Production of haploid and diploid spermatids
Decreased number of postmeiotic cells and no spermiogenesis

ROSI (poor efficiency)

This study

X^E,Z2Y*^XSry

Sry tg, Eif2s3y tg, Zfy2 tg

Y*^X

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

ROSI and ICSI

Vernet et al. (2014, 2016b) and Yamauchi et al. (2015)

X^E,Z1/UY*^XSry

Sry tg, Eif2s3y tg, Zfy1/Uba1y tg

Y*^X

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

Not tested

Vernet et al. (2014, 2016b)

X^E,Z1/U^,^Z2 Y*^XSry

Sry tg, Eif2s3y tg, Zfy1/Uba1y tg, Zfy2 tg

Y*^X

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

Not tested

Vernet et al. (2014)

X^E,Z2Sxr^bY*^X

Sxr^b, Zfy2 tg

Y*^X

Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids)
Decreased number of postmeiotic cells
Abnormal spermatozoa

ROSI

Yamauchi et al. (2015)

XY Zfy1 KO

Intact (Zfy2) but lacking Zfy1

Normal spermatogenesis

Normal mating

Yamauchi et al. (2022) and Nakasuji et al. (2017)

XY Zfy2 KO

Intact (Zfy1) but lacking Zfy2

Grossly normal spermatogenesis except mild sperm headshape and minor fertility defects

Normal mating

Yamauchi et al. (2022) and Nakasuji et al. (2017)

XY Zfy DKO

Intact but lacking Zfy1 and Zfy2

Decreased number of postmeiotic cells
Production of very few and abnormal spermatozoa

ROSI and ICSI

Yamauchi et al. (2022) and Nakasuji et al. (2017)

Table 1

Open in new tab

Mouse models used to investigate the Zfy genes.

Genotype	Y chromosome contribution* (Zfy contribution)	Meiotic pairing partner for X	Phenotype	Offspring obtained	References^#
XY	Intact (Zfy1 and Zfy2)	Y	Normal spermatogenesis	Normal mating	N/A

XOSry	Sry tg	None	Spermatogonial proliferation arrest	None	Mazeyrat et al. (2001)

XSxr^bO	Sxr^b (Zfy2/1 fusion gene)	None	Spermatogonial proliferation arrest	None	Vernet et al. (2011, 2012)

XSxr^aO	Sxr^a (Zfy1 and Zfy2)	None	Apoptotic elimination of MI spermatocytes due to X chromosome univalency Production of haploid and diploid spermatids in small numbers (due to efficient MI apoptosis) Very few abnormal spermatozoa	N/A	Vernet et al. (2011, 2012)

XSxr^aY *^X	Sxr^a (Zfy1 and Zfy2)	None	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	ROSI	Vernet et al. (2011, 2012)

XY *^XSxr^a	Sxr^a (Zfy1 and Zfy2)	None	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	ROSI and ICSI	Yamauchi et al. (2009)

X^ESxr^bO	Sxr^b (Zfy2/1 fusion gene)	None	Partial elimination of MI spermatocytes due to X chromosome univalency Production of haploid and diploid spermatids Decreased number of postmeiotic cells Incomplete abnormal spermiogenesis	ROSI	Vernet et al. (2011, 2016b) and Yamauchi et al. (2014, 2015)

X^ESxr^bY*^X	Sxr^b (Zfy2/1 fusion gene)	Y*^X	Production of haploid and diploid spermatids Decreased number of postmeiotic cells Incomplete abnormal spermiogenesis	ROSI	Vernet et al. (2014, 2016b) and Yamauchi et al. (2014, 2015)

X^EOSry	Sry tg, Eif2s3y tg	None	No elimination of MI spermatocytes due to X chromosome univalency Production of haploid and diploid spermatids Decreased number of postmeiotic cells and no spermiogenesis	ROSI	Vernet et al. (2011, 2012, 2016b), Yamauchi et al. (2014, 2015), and Ruthig et al. (2017)

X^EY*^XSry	Sry tg, Eif2s3y tg	Y*^X	Production of haploid and diploid spermatids Decreased number of postmeiotic cells and no spermiogenesis	ROSI	Vernet et al. (2014, 2016b) and Yamauchi et al. (2014, 2015)

X^E,Z2OSry	Sry tg, Eif2s3y tg, Zfy2 tg	None	Production of haploid and diploid spermatids Decreased number of postmeiotic cells and no spermiogenesis	ROSI	Vernet et al. (2011) and this study

X^E,Z1/UOSry	Sry tg, Eif2s3y tg, Zfy1/Uba1y tg	None	Production of haploid and diploid spermatids Decreased number of postmeiotic cells and no spermiogenesis	None	Vernet et al. (2011) and this study
X^E,Z1/U^,^Z2OSry	Sry tg, Eif2s3y tg, Zfy1/Uba1y tg, Zfy2 tg	None	Production of haploid and diploid spermatids Decreased number of postmeiotic cells and no spermiogenesis	ROSI (poor efficiency)	This study

X^E,Z2Y*^XSry	Sry tg, Eif2s3y tg, Zfy2 tg	Y*^X	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	ROSI and ICSI	Vernet et al. (2014, 2016b) and Yamauchi et al. (2015)

X^E,Z1/UY*^XSry	Sry tg, Eif2s3y tg, Zfy1/Uba1y tg	Y*^X	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	Not tested	Vernet et al. (2014, 2016b)

X^E,Z1/U^,^Z2 Y*^XSry	Sry tg, Eif2s3y tg, Zfy1/Uba1y tg, Zfy2 tg	Y*^X	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	Not tested	Vernet et al. (2014)

X^E,Z2Sxr^bY*^X	Sxr^b, Zfy2 tg	Y*^X	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	ROSI	Yamauchi et al. (2015)

XY Zfy1 KO	Intact (Zfy2) but lacking Zfy1	X	Normal spermatogenesis	Normal mating	Yamauchi et al. (2022) and Nakasuji et al. (2017)

XY Zfy2 KO	Intact (Zfy1) but lacking Zfy2	X	Grossly normal spermatogenesis except mild sperm headshape and minor fertility defects	Normal mating	Yamauchi et al. (2022) and Nakasuji et al. (2017)

XY Zfy DKO	Intact but lacking Zfy1 and Zfy2	X	Decreased number of postmeiotic cells Production of very few and abnormal spermatozoa	ROSI and ICSI	Yamauchi et al. (2022) and Nakasuji et al. (2017)

Genotype	Y chromosome contribution* (Zfy contribution)	Meiotic pairing partner for X	Phenotype	Offspring obtained	References^#
XY	Intact (Zfy1 and Zfy2)	Y	Normal spermatogenesis	Normal mating	N/A

XOSry	Sry tg	None	Spermatogonial proliferation arrest	None	Mazeyrat et al. (2001)

XSxr^bO	Sxr^b (Zfy2/1 fusion gene)	None	Spermatogonial proliferation arrest	None	Vernet et al. (2011, 2012)

XSxr^aO	Sxr^a (Zfy1 and Zfy2)	None	Apoptotic elimination of MI spermatocytes due to X chromosome univalency Production of haploid and diploid spermatids in small numbers (due to efficient MI apoptosis) Very few abnormal spermatozoa	N/A	Vernet et al. (2011, 2012)

XSxr^aY *^X	Sxr^a (Zfy1 and Zfy2)	None	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	ROSI	Vernet et al. (2011, 2012)

XY *^XSxr^a	Sxr^a (Zfy1 and Zfy2)	None	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	ROSI and ICSI	Yamauchi et al. (2009)

X^ESxr^bO	Sxr^b (Zfy2/1 fusion gene)	None	Partial elimination of MI spermatocytes due to X chromosome univalency Production of haploid and diploid spermatids Decreased number of postmeiotic cells Incomplete abnormal spermiogenesis	ROSI	Vernet et al. (2011, 2016b) and Yamauchi et al. (2014, 2015)

X^ESxr^bY*^X	Sxr^b (Zfy2/1 fusion gene)	Y*^X	Production of haploid and diploid spermatids Decreased number of postmeiotic cells Incomplete abnormal spermiogenesis	ROSI	Vernet et al. (2014, 2016b) and Yamauchi et al. (2014, 2015)

X^EOSry	Sry tg, Eif2s3y tg	None	No elimination of MI spermatocytes due to X chromosome univalency Production of haploid and diploid spermatids Decreased number of postmeiotic cells and no spermiogenesis	ROSI	Vernet et al. (2011, 2012, 2016b), Yamauchi et al. (2014, 2015), and Ruthig et al. (2017)

X^EY*^XSry	Sry tg, Eif2s3y tg	Y*^X	Production of haploid and diploid spermatids Decreased number of postmeiotic cells and no spermiogenesis	ROSI	Vernet et al. (2014, 2016b) and Yamauchi et al. (2014, 2015)

X^E,Z2OSry	Sry tg, Eif2s3y tg, Zfy2 tg	None	Production of haploid and diploid spermatids Decreased number of postmeiotic cells and no spermiogenesis	ROSI	Vernet et al. (2011) and this study

X^E,Z1/UOSry	Sry tg, Eif2s3y tg, Zfy1/Uba1y tg	None	Production of haploid and diploid spermatids Decreased number of postmeiotic cells and no spermiogenesis	None	Vernet et al. (2011) and this study
X^E,Z1/U^,^Z2OSry	Sry tg, Eif2s3y tg, Zfy1/Uba1y tg, Zfy2 tg	None	Production of haploid and diploid spermatids Decreased number of postmeiotic cells and no spermiogenesis	ROSI (poor efficiency)	This study

X^E,Z2Y*^XSry	Sry tg, Eif2s3y tg, Zfy2 tg	Y*^X	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	ROSI and ICSI	Vernet et al. (2014, 2016b) and Yamauchi et al. (2015)

X^E,Z1/UY*^XSry	Sry tg, Eif2s3y tg, Zfy1/Uba1y tg	Y*^X	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	Not tested	Vernet et al. (2014, 2016b)

X^E,Z1/U^,^Z2 Y*^XSry	Sry tg, Eif2s3y tg, Zfy1/Uba1y tg, Zfy2 tg	Y*^X	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	Not tested	Vernet et al. (2014)

X^E,Z2Sxr^bY*^X	Sxr^b, Zfy2 tg	Y*^X	Potentiated meiosis II: production of haploid spermatids (no/few diploid spermatids) Decreased number of postmeiotic cells Abnormal spermatozoa	ROSI	Yamauchi et al. (2015)

XY Zfy1 KO	Intact (Zfy2) but lacking Zfy1	X	Normal spermatogenesis	Normal mating	Yamauchi et al. (2022) and Nakasuji et al. (2017)

XY Zfy2 KO	Intact (Zfy1) but lacking Zfy2	X	Grossly normal spermatogenesis except mild sperm headshape and minor fertility defects	Normal mating	Yamauchi et al. (2022) and Nakasuji et al. (2017)

XY Zfy DKO	Intact but lacking Zfy1 and Zfy2	X	Decreased number of postmeiotic cells Production of very few and abnormal spermatozoa	ROSI and ICSI	Yamauchi et al. (2022) and Nakasuji et al. (2017)

See Fig. 1 for gene content of Y, Sxr^a, and Sxr^b. See text for more details regarding specific genotypes. Abbreviations used: tg, transgene; E, Eif2s3y transgene; Z2, Zfy2 transgene; Z1/U, Zfy1/Uba1y transgene; KO, knock-out; DKO, double knock-out; ROSI: round spermatid injection; ICSI: intracytoplasmic sperm injection. Only X chromosome-encoded Zfy transgenes are included (autosomally encoded Zfy transgenes lead to pachytene arrest preventing spermatogenesis progression).

Only selected, most relevant references describing phenotypic characterization of these mice are shown.

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