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. 2023 Feb 10;42(7):e112699. doi: 10.15252/embj.2022112699

Table 9.

Factors involved in cytoplasmic pre‐40S maturation.

Yeast 40S 60S Human 40S 60S Function in cytoplasmic pre‐40S maturation Citation
Enp1 x ENP1/BYSL x Released by Hrr25/CKI phosphorylation, Release allows stable incorporation of Rps3/uS3, Rps10/eS10 and Rps20/uS10, Formation of the beak structure Schäfer et al (2006), Zemp et al (2014) and Ameismeier et al (2018)
Ltv1 x LTV1 x
Tsr1 x TSR1 x Released, occludes binding sites for mRNA and translation initiation factors Larburu et al (2016) and Scaiola et al (2018)
Dim1 x DIMT1 x Methyltransferase, yeast: modifies subsequent adenines near 3′ end of 18S rRNA; human: function occurs already in the nucleus Strunk et al (2011) and Zorbas et al (2015)
Rrp12 x RRP12 x Released, which allows incorporation of Asc1 (RACK1 in humans) Wyler et al (2011) and Larburu et al (2016)
Dim2/Pno1 x PNO1/DIM2 x Inhibits binding of eIF3; release contributes to Major structural rearrangements, allowing Nob1 activity Ameismeier et al (2018) and Scaiola et al (2018)
Nob1 x NOB1 x Endonucleolytic cleavage of 20S pre‐rRNA (18S‐E pre‐rRNA in humans) to yield mature 18S rRNA Fatica et al (2003) and Lamanna and Karbstein (2011)
Hrr25 x CKI‐δ/CSNK1D x Phosphorylation of Enp1‐Ltv1‐Rps3/uS3 complex, which triggers release of Ltv1 Schäfer et al (2006) and Zemp et al (2014)
CKI‐ε/CSNK1E x
Rio1 x RIOK1/RIO1 x Orchestrates structural changes, pre‐rRNA maturation and trans‐acting factor release Vanrobays et al (2003) and Widmann et al (2012)
Rio2 x RIOK2/RIO2 x Kinase contributes to pre‐rRNA maturation and trans‐acting factor release Geerlings et al (2003) and Zemp et al (2009)
RIOK3 x Promotes 18S‐E processing Geerlings et al (2003)
Prp43 x DHX15 Conformational pre‐rRNA switch, which allows Nob1 activity Pertschy et al (2009)
Pfa1/Sqs1 x SON x Cofactor for Prp43 Sloan et al (2013b)
Pxr1/Gno1 x PINX1 x Cofactor for Prp43 Guglielmi and Werner (2002) and Chen et al (2014)
Hbs1 x HBS1 In yeast: facilitates formation of 80S like particle Lebaron et al (2012), Strunk et al (2012) and Hector et al (2014)
Dom34 x PELO
Rli1 x ABCE1
Fun12 x EIF5B
USP16 x Deubiquitylates lysine 113 in RPS27A/eS31 Montellese et al (2020)
LRRC47 x Bound to late cytoplasmic pre‐40S particle Ameismeier et al (2020)
EIF1AD x Supports trans‐acting factor release and pre‐rRNA maturation Ameismeier et al (2020)