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Abiel Rindisbacher 28.05.2013 How to escape gene loss: new func?on or dosage benefit? • Ancient gene duplica?on event in Saccharomyces cerevisiae • Most duplicated genes got lost • Some escaped (10%) → new func?on OR dosage benefit • 59 of 79 yeast ribosome genes encoded by 2 genes • Previously thought to be func?onally redundant (dosage benefit) Wikipedia ASH1 • Ash1 localises at the bud-‐?p • Needed for ASH1 mRNA localisa?on: Transla?on AND Transla?onal regula?on (repression) • Some factors needed for localisa?on: → involved in ribosomal assembly (Loc1, Puf6) Effect of Loc1 on ASH1 mRNA localiza?on… …a consequence of its role in ribosomal assembly? → Loc1 regulates the transla?on of ASH1 mRNA not only localisa?on to the bud ?p Cells expressing the E3 reporter = ASH1 – Bud-‐?p localisa?on → bud-‐site posi?oning? Localisa?on of ASH1 mRNA in different mutants with random bud-‐ site selec<on phenotype: Factors unrelated to transla?on → unique defects Factors related to transla?on → like in loc1Δ “Genes required for bud-‐site selec<on are needed for the localiza?on of ASH1 mRNA” Func?onal specificity between duplicated ribosomal proteins? • 14/15 ribosomal proteins (RPs) implicated in bud-‐site selec<on have duplicate • Only one each needed for bud-‐ site selec?on and ASH1 mRNA localisa?on • “Transla?on of ASH1 mRNA requires a specific subset of duplicated RP paralogs” Data mining • Transcrip<onal profiles of cells lacking the RPs involved in ASH1 mRNA localiza?on → RP paralogs have specialized cellular roles beyond their effects on ASH1 • High-‐throughput data (compiling phenotypic effects of absence of all non-‐essen<al RP) → Duplicated RPs show different phenotypes from their paralogs ! No difference in expression level Paralog-‐specific localiza?on? • loc1Δ: Rpl7b and Rps18 → ER • puf6Δ: Rpl7b → ER • loc1Δpuf6Δ: o Rps18b →ER o Rpl7b → discrete cytoplasmic foci o Rps18a → nucleolus → synthe?c defect “Paralogous RPs require different factors for their assembly and exhibit paralog-‐ specific aberrant localiza<ons in the absence of certain factors” Are duplicated RPs playing dis?nct func?onal roles? • Localized transla?on of ASH1 mRNA requires a specific subset of duplicated ribosomal protein genes • Addi?onal levels of func<onal divergence between paralogs • Func?onal specificity occurs in all duplicated ribosomal protein genes • Paralog-‐specific localiza?ons and assembly defects • Not dosage effect → Yes Discussion Model: • ‘‘Loc1 is required for the assembly of ribosomes containing a specific subset of duplicated ribosomal proteins and this specialized ribosome is required for the regulated transla?on of ASH1 mRNA’’ • The model does not provide a mechanism • Specific ribosomes involved in other processes as well • There are indica?ons for specific ribosomes in other organisms as well Ribosome code Specific ribosomes: • Func?onally dis?nct RPs • Different forms of rRNA • Posiransla?onal modifica?ons → “Ribosome code”; similar to the “histone code” Transla?ng the Histone Code Thomas Jenuwein and C. David Allis Ø New level of complexity in the regula<on of gene expression Thank you for your aien?on!