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. 2019 Oct 8;10:962. doi: 10.3389/fgene.2019.00962

Table 2.

Mouse models related with the histone-to-protamine transition.

Gene Phenotype Function Reference
H1t Fertility and no spermatogenesis abnormalities Dispensable for histone-to-protamine transition Drabent et al., 2000; Fantz et al., 2001
H1t2 Reduced fertility with delayed nuclear condensation and aberrant elongation of spermatids Indispensable for the incorporation of PRMs and proper chromatin condensation Martianov et al., 2005; Tanaka et al., 2005
Th2b Fertility with normal spermatogenesis in Th2b-null mice Destabilize chromatin and regulate TPs and PRMs incorporation Montellier et al., 2013
Infertility with abnormal spermatozoa in TH2B C-terminus modified mice
Th2a/Th2b Infertility with accumulated spermatocytes at interkinesis and abnormal spermatozoa Indispensable for cohesin release and TP2 incorporation Shinagawa et al., 2015
H2al2 Infertility and the sperm chromatin show a compaction defects Assemble open nucleosomes and allow TPs incorporation Barral et al., 2017
H2a.b Reduced fertility with abnormal spermatozoa Destabilize chromatin and modulate the dynamics of H2AL2 removal and TP1 incorporation Anuar et al., 2019
H3f3a Reduced fertility with dysmorphic spermatozoa Require for normal development of some spermatids Tang et al., 2015
H3f3b Infertility with abnormal spermatozoa and reduced sperm count Indispensable for spermatogenesis related genes expression, TP1 removal and PRM1 incorporation Yuen et al., 2014
H3t Infertility with azoospermia Require for spermatogonial differentiation and ensures entry into meiosis Ueda et al., 2017
Epc1 Infertility with abnormal round spermatids to elongating spermatids transition Require for round spermatids maturation by regulating histone acetylation and TP2 incorporation Dong et al., 2017
Tip60 The germ cell is arrested at the RS stage Contribute to round spermatids maturation by regulating histone acetylation and TP2 incorporation Dong et al., 2017
Sirt1 Reduced fertility with abnormal spermatozoa and decreased sperm count Require for acetylation of H4K5, H4K8 and H4K12 histone, and TP2 incorporation Bell et al., 2014
Brdt Infertility with complete absence of post-meiotic cells in Brdt-null mice Control the chromatin organization and meiotic sex chromosome inactivation; the first bromodomain of BRDT is essential to link histone removal and TPs, PRMs incorporation Shang et al., 2007; Dhar et al., 2012; Gaucher et al., 2012; Manterola et al., 2018
Infertility with abnormal spermatids in Brdt ΔBD1/ΔBD1 mice
Pa200 Reduced fertility with abnormal spermatozoa and decreased sperm count Recognize acetylated histones and mediate the core histones for acetylation dependent degradation through proteasomes Khor et al., 2006; Qian et al., 2013
Rnf8 Infertility with abnormal spermatozoa and reduced sperm count Require for histone ubiquitination and modulate H4K16ac to facilitate histone removal and TPs, PRMs incorporation Lu et al., 2010
Miwi Infertility with abnormal spermatozoa and reduced sperm count in Miwi D-box mutations mice Essential for nuclear translocation of RNF8 and facilitates the histone ubiquitination and further histone removal Gou et al., 2017
L3mbtl2 Reduced fertility with abnormal spermatozoa and decreased sperm count Require for the RNF8-UbH2A pathway and further PRM1 incorporation Meng et al., 2019
Phf7 Infertility with abnormal spermatozoa and decreased sperm count Recognize the H3K4me3/me2 and catalyze H2A ubiquitination to facilitate the histone removal Wang et al., 2019
Pygo2 Infertile with abnormal spermatozoa and decreased sperm count in Pygo2 reduced mice Recognize H3K4me3 and recruit HAT to facilitate H3 acetylation and expression of Prms, Tnp2, and H1fnt. Nair et al., 2008
Jhdm2a Infertile with abnormal spermatozoa and decreased sperm count Control H3K9 methylation at the promoter of Tnp1 and Prm1 genes and regulate their expression Okada et al., 2007
Setd2 Infertility with round spermatid arrest Catalyze H3K36me3 and facilitate the activation of Tnps and Prms genes Zuo et al., 2018
Tssk6 Infertility with abnormal spermatozoa Mediate γH2AX to possess normal quantities of histone H3, H4 and PRM2 precursor and intermediate Spiridonov et al., 2005; Jha et al., 2017
Cdyl Reduced fertility with decreased sperm count and motility Regulate histone crotonylation to facilitate TP1 and PRM2 incorporation Liu et al., 2017b
Parp11 Infertility with teratozoospermia Modulate PARsylation to facilitate chromatin condensation Meyer-Ficca et al., 2015
Parg110 Reduced fertility with poor sperm chromatin quality Dispensable for histone removal and TP1 incorporation Meyer-Ficca et al., 2009; Meyer-Ficca et al., 2011
Tnp1 Reduced fertility with subtle abnormal spermatozoa and decreased spermatozoa Dispensable for histone displacement as the compensation by TP2 and PRM2 precursor Yu et al., 2000
Tnp2 Reduced fertility with abnormal spermatozoa Dispensable for histone displacement but necessary for maintaining the normal processing of PRM2 and the completion of chromatin condensation Zhao et al., 2001
Tnp1/Tnp2 Infertile with abnormal spermatozoa and decreased sperm count Indispensable for PRM2 incorporation and chromatin condensation Shirley et al., 2004
Prm1 Infertile with abnormal spermatozoa and decreased sperm count Indispensable for spermiogenesis and chromatin condensation Cho et al., 2001
Prm2 Infertile with abnormal spermatozoa and decreased sperm count
Camk4 Infertile with abnormal spermatozoa and decreased sperm count Mediate the phosphorylation of PRM2 and facilitate basic nuclear proteins removal Wu et al., 2000