TABLE 1.

Summary of RNA-binding proteins involved in miRNA biogenesis.

RNA-binding protein	Target miRNA(s)	Binding motif in RNA	Stage of biogenesis	Interactions with biogenesis proteins	Effect on miRNA biogenesis	Cell line	References
DDX5 (p68)	Most miRNA		Pri-miRNA cleavage	Microprocessor	Promote	HeLa, 293FT	Shiohama et al. (2007), Beezhold et al. (2010)
DDX17 (p72)	Most miRNA		Pri-miRNA cleavage	Microprocessor	Promote	HeLa, 293FT	Shiohama et al. (2007), Beezhold et al. (2010)
SRSF3		CNNC	Pri-miRNA cleavage		Promote	HEK293	Auyeung et al. (2013)
KSRP	Let-7 family, miR-196a, miR-155	G-rich regions of the terminal loop	Pri-miRNA cleavage and pre-miRNA cleavage	Drosha	Promote	HeLa, U2OS, P19, NIH-3T3, bone marrow derived macrophages, RAW 264.7 macrophages	Ruggiero et al. (2009), Trabucchi et al. (2009), Castilla-Llorente et al. (2013)
hnRNPA1	miR-18a, Let-7a	G-rich regions in the pri-miRNA terminal loop	Pri-miRNA cleavage		Promote or inhibit	HeLa	Guil and Cáceres (2007), Michlewski et al. (2008), Michlewski and Cáceres (2010), Castilla-Llorente et al. (2013)
TDP-43	miR-143, miR-574	GU-rich regions in the terminal loop	Pri-miRNA cleavage and pre-miRNA cleavage	Drosha	Promote	HEK293T, SK-N-BE(2)-C, SH-SY5Y	Kawahara and Mieda-Sato (2012), Di Carlo et al. (2013)
LIN28B	Let-7	Pri-miRNA terminal loop and GGAG in the stem	Pri-miRNA cleavage		Inhibit	HEK293, HeLa, H1299, Igrov1, HepG2, T47D, MDA-MB-231, SK-Mel-28	Heo et al. (2009), Nam et al. (2011), Piskounova et al. (2011), Mayr et al. (2012), Mayr and Heinemann (2013)
HuR/MIS2	miR-7	AUAA in Pri-miRNA terminal loop	Pri-miRNA cleavage		Inhibit	HeLa, human brain astrocytoma 1321N1, HEK293T, SH-SY5Y	Choudhury et al. (2013)
YB-1	miR-29b-2	UYAUC in miRNA terminal loop	Pri-miRNA cleavage and pre-miRNA cleavage		Inhibit	U251-MG	Wu et al. (2015)
Matr3	miR-138-2	Pri-miRNA terminal loop	Pri-miRNA cleavage or nuclear export		Inhibit	E18 Sprague-Dawley rat cells	Weiss et al. (2019)
EWS	miR-34a, miR-122, mir-222, etc.	Pri-miRNA terminal loop flanking regions	Pri-miRNA cleavage		Promote	HeLa	Ouyang et al. (2017)
QKI5	miR-124-1	QKI response element	Pri-miRNA cleavage	DGCR8	Promote	CD34+ HSC	Wang et al. (2017)
Exportin 5	Most miRNA	Double stranded pre-miRNA stem and 3’ 2-nt overhang	Nuclear Export		Promote	HeLa	Bohnsack et al. (2004), Lund et al. (2004), Okada et al. (2009)
Exportin 1	miR-320, miR-484	m7G cap	Nuclear Export		Promote	HEK293	Xie et al. (2013)
TRBP	Most miRNA	Regions of pre-miRNA stem-loop with tight base pairing	Pre-miRNA cleavage	DExD/H-box helicase domain of Dicer	Promote	HEK293	Haase et al. (2005), Takahashi et al. (2018), Yoshida et al. (2021)
ADAR1	Most miRNA		Pre-miRNA cleavage	DExD/H-box helicase domain of Dicer	Promote	HEK293	Ota et al. (2013)
DAZL	Several miRNAs	GUU	Pre-miRNA cleavage		Promote	hESC line H9, HSF6	Yan et al. (2022)
LIN28A	Let-7	Pre-miRNA terminal loop	Pre-miRNA cleavage		Inhibit	HEK293, HeLa, H1299, Igrov1, HepG2, T47D, MDA-MB-231, SK_Mel_28, HepG2, Huh7, Hep3B	Heo et al. (2008), Heo et al. (2009), Lightfoot et al. (2011), Nam et al. (2011), Piskounova et al. (2011)
FMRP			Processing in the cytoplasm and promotion of translational repression	AGO1, Dicer	Promote	HeLa, human lymphoblastoid cell lines	Jin et al. (2004a), Jin et al. (2004b), Basu and Bahadur (2016)
CSDE1	miR-451	UGAU	Processing in the cytoplasm	AGO2, PARN	Promote	HEK293T, MEL cells	Kakumani et al. (2023)
SYNCRIP	miR-3470a, miR-194-2	hEXO motif (GGCU)	miRNA sorting into EVs	DGCR8	Promote	Murine hepatocyte 3A cells	Santangelo et al. (2016), Chen et al. (2020)
	Let-7a	UAGAAU in pri-miRNA terminal loop	Pri-miRNA cleavage			HEK293
hnRNPA2B1	miR-198	GAGG	miRNA sorting into EVs		Promote	Human peripheral blood mononuclear cells, HEK293	Villarroya-Beltri et al. (2013)
Cx43	miR-133b		miRNA sorting into EVs		Promote	HEK293, C33a	Martins‐Marques et al. (2022)
SAFB2			Cluster assistance phenomenon		Promote	HEK293T	Hutter et al. (2020)
EHR			Cluster assistance phenomenon	DGCR8	Promote	HEK293T, Expi293F, HEK293E, K562, HCT116	Fang and Bartel (2020), Hutter et al. (2020), Kwon et al. (2020)