Table 1.

Genes related to spermatogenesis unearthed by gene editing technology.

Gene	Species	Techniques Used for Function Analysis	Fertility	Phenotype/Clinical Symptoms	References
Akap4	Mus musculus	KO 1	Male infertility	Abnormal sperm morphology and reduced motility	[32]
Amh	Danio rerio	KO	-	Dysregulation of germ cell development and the over-proliferation of spermatogonia	[33]
Armc2	M. musculus	KO	Male infertility	Multiple morphological abnormalities of the flagella	[34]
Asb17	M. musculus	KO	Fertile	Oligospermia and a disorganized ES junction	[35]
Bcorl1	M. musculus	KO	Male infertility	Impaired sperm viability and abnormal mitochondrial structure of sperm cells	[36]
Cabs1	M. musculus	KO	Significantly impaired fertility	Defective sperm flagellum differentiation and abnormal sperm tail structure	[37]
Ccdc63	M. musculus	KO	Male infertility	Shortened flagella	[38]
Cct6b	M. musculus	KO	-	No differences in development, fertility, appearance, testis weight, or sperm counts. Nuclear base bending abnormality	[39]
Cdc14a	M. musculus	KO	Significantly impaired fertility	Low sperm count, impaired sperm motility and high percentage of morphologically abnormal sperm	[40]
Cib4	M. musculus	KO	Male infertility	Impaired haploid differentiation and absence of elongated spermatozoa in the epididymal tail	[41]
Cmtm4	M. musculus	KO	Significantly impaired fertility	Decreased sperm count, decreased epididymal sperm motility, increased percentage of abnormal backward bending of sperm head and bending of sperm mid-section	[42]
CSR-1a	Caenorhabditis elegans	KI 2/KO	-	A transgenerational loss of sperm-based fertility in hermaphrodites	[43]
Cyp11c1	Danio rerio	KO	-	Exhibits female secondary sexual characteristics, severe deficiency of androgens and cortisol, impaired spermatogenesis and characteristic reproductive behavior, disturbed arrangement of spermatogenic tubules, and abnormal differentiation of spermatogonia.	[44]
Ddx4	M. musculus	cKO 3(Cre-loxP)	-	Spermatogonia developed and became arrested at the round spermatid stage	[45]
Defb23/26/42	R. norvegicus	KO	No clear phenotype for single knockout, but 23/26 or 23/26/42 combined knockout is infertile.	Impaired sperm motility, the sperm showed precocious capacitation and increased spontaneous acrosome reaction.	[46]
Dmrt1	Danio rerio	KO	-	Severe testicular developmental defects and gradual loss of all Vasa-positive germ cells	[33]
Dmrt6	Oreochromis mossambicus	KO	-	Fewer spermatocytes	[47]
Dnah17	M. musculus	KO	-	Asthenozoospermia, abnormal sperm flagellar morphology and low sperm activity.	[48,49]
Dpy19l2	M. musculus	KO (NA) 9	Male infertility	The NDL facing the acrosome, the acro-plaxome, caudal descent and acrosome spreading are defective.	[50]
Ephb2	M. musculus	KO (SSCs) 7	-	Proliferation and stem cell activity are impaired.	[51]
Fam170a	M. musculus	KO	Significantly impaired fertility	Abnormal spermiation, abnormal head morphology, and reduced progressive sperm motility.	[52]
Fto	M. musculus	KO (spermatogonia)	-	Chromosome instability and G2/M arrest	[53]
Gh1	Danio rerio	Point mutation	-	Delayed spermatogenesis	[54]
HIF-1α	R. norvegicus	KD 4	-	The distribution of germ cells was disordered and apoptosis of spermatogenic cells increased significantly.	[55,56]
Hsf5	Danio rerio	KO	Male infertility	Reduced sperm count, increased sperm head size, and abnormal tail architecture	[57]
Hydin	M. musculus	Biallelic mutations (ESCs)	-	Hydin-disrupted sperm obtained from the chimeric mice possessed short tails and were immotile, but it can produce viable pups.	[58]
		KO (NA)	-	Die within 3 weeks before sexual maturation due to hydrocephaly.	[58]
Igf3	Oreochromis niloticus	KO	Male infertility	The proliferation and differentiation of spermatogonia are severely inhibited at the beginning of meiosis, and semen volume and sperm count are drastically reduced.	[59]
Lipocalin8	M. musculus	KO	Normal fertility	There was no significant effect on the morphological appearance of the testes but epididymal sperm maturation defects.	[60]
		cKI 5	Normal fertility	-	[61]
Mct8	R. norvegicus	KO	Fertile, lower fertilization rate	Serum THs (T3 and T4) level were significantly increased, growth delay along with thyroid dysfunction, testis maldevelopment and impaired spermiogenesis.	[62]
Meig1	M. musculus	Y68 point mutation	Male infertility	The sperm count is significantly reduced, and a few developed sperm fail to move and exhibit a variety of abnormalities.	[63]
Pick1	M. musculus	KO (NA)	Male infertility	Fragmentation of acrosomes in the early stages of spermiogenesis, round-headed sperm, reduced sperm count, and severely impaired sperm motility.	[64]
Pmfbp1	Bombyx mori	Point mutation	Male infertility	Defects in the development of eupyrene sperm bundles	[65]
Prss55	M. musculus	KO/DKO 6	Male infertility	Impaired migration from the uterus to the oviduct and impaired ability to bind the zona pellucida (ZP) of oocytes	[66]
Rln3a	Oreochromis niloticus	KO	Significantly impaired fertility	Hypogonadism, sperm deformation and a significant decrease in sperm motility.	[67]
Rnf216	M. musculus	KO	Male mice are sterile and females are capable of reproduction.	Smaller testes, defective meiosis, and reduced number of germ cells.	[68,69]
Sox30	Oreochromis niloticus	KO	Significantly impaired fertility	Abnormal spermiogenesis, reduction of sperm motility	[70]
	M. musculus	cKO (Cre-loxP)	Male infertility	Stagnant germ cell development, abnormal acrosome and axon development and complete cessation of spermatogenesis.	[71]
Spata16	M. musculus	851G→A/R284Q point mutation	Fertile	-	[72]
		781-bp deletion	Male infertility	Spermio-genic arrest, with impaired differentiation of round spermatids into the mature sperm.	[72]
Spata3	M. musculus	KO	Normal fertility with reduced in vitro fertility	Acrosome defects and excessive lipid droplet residues in the cytoplasm.	[73]
Spatc1l	M. musculus	KO	Male infertility	Separation of sperm head from tail	[74]
Ssmem1	M. musculus	KO	Male infertility	Globozoospermia, loss of sperm motility and abnormal localization of Golgi at steps eight and nine of spermatid development.	[75]
Sun3	M. musculus	KO	Male infertility	Reduced sperm counts and a globozoospermia-like phenotype.	[76]
Tcfl5	M. musculus	KO	Male infertility	Sperm cells and spermatozoa of Tcfl5+/- mice (infertility) have been abnormal.	[77]
Tle6	M. musculus	KO (spermatogonia, CRISPR/Cas9, Tet-on) 8	-	Spermatogonia proliferation and cell cycle are inhibited.	[78]
Tmprss12	M. musculus	KO	Male infertility	Normal spermatogenesis and sperm morphology, but ejaculated spermatozoa failed to migrate from the uterus to the oviduct.	[79]
Tsga10	M. musculus	KO	Male infertility	Disordered mitochondrial sheath formation and reduced sperm motility.	[80]
Tssk3	M. musculus	KO	Male infertility	Reduced sperm count and abnormal morphology.	[81]
Ttc21a	M. musculus	Frameshift mutation	Male infertility (78%)	The motility and progressive motility of spermatozoa were significantly reduced. Morphological abnormalities of sperm. The structural abnormalities of the connecting piece during spermiogenesis and multiple structural defects of the flagella.	[82]
Ythdf2	M. musculus	KO (spermatogonia)	-	Cell proliferation, cell adhesion and cell spread were inhibited.	[83]
Zfp628	M. musculus	KO	Male infertility	Post-meiotic germ cell arrest at the round spermatid stage in the seminiferous tubules of the testis.	[84]
Zfy1/Zfy2	M. musculus	KO	Normal fertility	-	[85,86]
		DKO	Infertility	Abnormal sperm morphology, fertilization failure and early embryo development failure.
Zmym3	M. musculus	KO	Male infertility	Abnormal spindle assembly at mid-meiotic division.	[87]
1700102P08Rik	M. musculus	KO	Male infertility	Smaller testes and epididymis, stagnation of spermatogenesis at the spermatocyte stage, absence of spermatozoa in the epididymis, and apoptosis of testicular cells.	[88]

¹ KO: CRISPR/Cas9-mediated knockout; ² KI: CRISPR/Cas9-mediated knock-in; ³ cKO: CRISPR/Cas9-mediated conditional knockout; ⁴ KD: CRISPR/Cas9-mediated knockdown; ⁵ cKI: CRISPR/Cas9-mediated conditional knock-in; ⁶ DKO: CRISPR/Cas9-mediated double knockout; ⁷ The corresponding cells on which gene editing was performed are indicated in parentheses; ⁸ The corresponding cells on which gene editing was performed are indicated in parentheses; ⁹ NA: The technique of mediated gene knockout is unknown or not mentioned in the original article.